© Steve Trudell
Danny’s DNA Discoveries – Psathyrella s.l. of the PNW
by Danny Miller
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Danny’s DNA Discoveries – Psathyrella s.l. of the PNW
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The Psathyrellaceae family is known for black spored, very fragile mushrooms with dry, usually hygrophanous caps and gills that are barely attached and easily come free. There used to be two kinds:
The problem is, the ability to turn to ink didn't just evolve once. Perhaps it's a gene that may or may not get expressed in different lineages (they seem to be able to evolve back and forth between being able to "ink" and not fairly readily). Since the inky ones are not all in one related group, but sprinkled throughout the family tree, we can't just use one name for the inky species and one name for the non-inky species, so both groups have many different genera in them now. This page addresses those that don't turn to ink with non-pleated caps (some inky caps hardly ink at all, but like the strongly inky species, they usually have a pleated cap). The spores are not always black, they might be dark brown or have a hint of purple like some of the Strophariaceae. Not all of them are very fragile, but they all do have a cellular cap cuticle (explained on my Agrocybe page). The entire mushroom is usually so thin, that not only the top layer of the cap can break in any direction, that usually extends to the entire cap breaking easily in any direction. If no matter how carefully you try to handle your mushroom, odd shaped chunks are missing from the cap by the time to carry it somewhere, it just might be a Psathyrella. Another clue to a cellular cap cuticle is it has a tendancy to wrinkle more easily as the spherical cells collapse in on each other, and they sparkle in the sun as the light reflects off the spheres. Panaeolus is the genus most easily confused with the Psathyrellas. They are unrelated mushrooms that evolved to look almost identical - but are more likely to be found on dung or in grass and have a stiffer stem that is less likely to be white. There is very often a characteristic dark band around the outside of the cap as the centre of the cap dries out first (usually the edges of a cap are thinner and dry out first). The black spores mature at random times instead of altogether, so if you look closely, a Panaeolus will usually have splotchy, mottled gill faces, something that is rarer is Psathyrella. According to Smith, Panaeolus can be told apart microscopically by "brown basidiole-like bodies embedded in the hymenium".
The standard reference for Psathyrella here in America is the 1972 The North American Species of Psathyrella by Alexander Smith, probably the most prolific American mycologist of all time. This phone book sized monograph discusses 414 species found in North America that all pretty much look alike. Saying that a microscope is usually required to identify a Psathyrella is an understatement. It's interesting to see how his concepts have held up to DNA study.
That being said, this is still an unbelievably valuable resource and must be the starting point from which further Psathyrella studies are done. Since some collections are mixed, each type collection must be sequenced and compared to more recent collections. There has been great progress made - I would guess that more than 100 of his type sequences have been sequenced by now. Matt Gordon has been doing some of the important work of sequencing many of our local type sequences. Approximately 160 'Psathyrellas' have been reported from the PNW, and up until now very little was known about most of them. With this report I'm happy to say that we now have a handle on what about 91 of them might be genetically, with only 67 or so left in the "Little known" category. Identifying your 'Psathyrella' through DNA is now somewhat possible. To identify a Psathyrella through traditional methods is still quite difficult. This page is not yet going to help you reliably get to new genus, never mind to species. While I do show photographs for many species, even the photos are not going to be that helpful. I recommend Smith's key. Wachter has named 18 sections of Psathyrella, with descriptions, plus descriptions of the segregate genera, but there is net yet a good key to figuring out what section a species is in. If you can get a species to section or genus, however, you can see below what the possible species might be. How common they are is colour coded, relative to how many time Smith found other species, and whether or not we've been successful in finding it recently. Nobody really understands most species' true rarity yet. 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. This report is also heavily supported by excellent papers by Orstadius, Wachter and Voto. References:
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 Psathyrella s.s. - click to expand
This section contains the species we know something about genetically.
Explanation of terms: The type area is noted after the name. The number of times Smith found it in his study in each state is noted. Newer possible synonyms are in brackets, along with their type area and where Smith found them. This does not mean that all the collections were actually the same species, we now know that often they were not, but this will hopefully give some idea of a species' scarcity. We also don't for sure that species whose holotypes are from out of the area really represent what is to be found in the PNW. The number of sequences of recent collections that have been found of this species is also given, to give further information about its scarcity. Example: Psathyrella amazica WA (WAx2 ORx1) - recent: WAx1 - means that this species was described from WA, so exists there by definition. It was only found 3 times (twice in WA and once in OR) during Smith's study, so it seems to be fairly uncommon (I would have said rare, but we did find it recently). Recently, we found a collection in WA with a sequence that matches the type sequence or another reliable sequence. Psathyrella subamazica MI (WAx12 ORx8) - means that this species was described from MI, and Smith thought he found it a lot in the PNW, more than a dozen times, so I have marked it abundant. There is a common species in the PNW here, but it may not be P. subamazica unless we have proof that DNA from both coasts is the same. We must have DNA from at least one coast, providing at least some information about it, or I would have put it under the "Little known Psathyrellas" category.
Species that aren't actually in Psathyrella Psathyrella conopilea EU - now in Parasola, which is surprising since it is not pleated and it does not turn to ink at all, but it can be recognized as a Parasola by the lack of any universal veil material, and from very small brown cap hairs. Psathyrella canoceps OR - it took DNA to discover that this belongs in Coprinopsis because there were really no clues that I know of. Psathyrella uliginicola WY/Psathyrella subagraria NY/Psathyrella longipes CA/Psathyrella fragilissima OR/Psathyrella elwhaensis WA/Psathyrella marcescibilis EU - these are relatively large, stocky mushrooms with spores >10u long, so even though they don't turn to ink, that is a clue that they really belong in Coprinopsis. Small, delicate species with spores >10u usually belong in Psathyrella.
The section names are based on Wachter.
/Microrhizae - delicate species with long thin stems (<3 mm thick) and large spores (>10u). Pink in the cap and on the bottom of the gill faces is possible. This section is separated from other similar sections by the presence of a partial and universal veil leaving material on the caps or hanging from the cap margin when young. Psathyrella microrhiza EU - recent: PNWx16 - (Psathyrella semivestita UK WAx2) - Smith did not believe in using the name of this very abundant species in NA, so it begs the question: what did he call the dozens of collections he must have made of it? I have yet to figure that out. Wachter has given us a reliable sequence, correcting an error by Orstadius, and 16 modern local collections match the sequence! All 16 are very close to each other and differ from the group of four species names mentioned below by only 4 bp or so. Psathyrella semivestita is said to be a synonym I can find no reason to question that. No one has reported a collection of it that seemed to differ from P. microrhiza in recent memory. I don't know what Psathyrella bifrons UK (WAX3) is, but the one sequence with that name, from France, sequenced the same as P. microrhiza, It may not have been labeled correctly, but it may indicate that it is at least in this section.
Psathyrella rufogrisea OR (ORx1) (P. alluviana WAx1 ORx8, P. barlowiana ORx2, P. uskensis WAx1) - recent: WAx2 - all four of the type sequence are almost identical. Nobody has studied the micro and ecology to determine for sure that they are consepecific. P. rufogrisea is the oldest name. It should be studied if P. rufogrisea var. riparia ID (IDx1) (differs microscopically) and P. rufogrisea var. bonnerensis ID (IDx1) (found on wood) are the same. All four species have unusually large spores, so I could believe they're all the same, although P. uskensis has even larger spores than the other three. Recently, 2 collections have been sequenced from WA.
Psathyrella 'alluviana PNW10' - recent: ORx2 - One of Smith's WA P. alluviana collections turned out to be a sister species, 3% different (and another mentioned below was completely unrelated). I do not yet know how to tell it apart from the others. It occurs in the EU as well. Two recent OR collections have been found. No great photos yet.
/Psathyrella - separated from the /Microrhizae by no veil material on the cap edges. Psathyrella bipellis is distinct and may deserve its own section. Psathyrella aquatica OR - the famous underwater mushroom from the Rogue River. One Smith collection of the older Psathyrella fontinalis (but not the type) from MI has a very similar sequence, and is described from wet springs.
Psathyrella pseudogracilis EU - recent: ORx2 - never reported from the PNW, but we've got two recent OR sequences that match Wachter's EU sequences, so it's here.
Psathyrella corrugis EU (=P. amarescens EU) (WAx39 ORx16 IDx1) - recent: PNWx5 - this is now the type species of the genus, since the name Psathyrella gracilis has fallen out of favour for some reason. Smith found it everywhere, it seems to be our most common species. Oddly, some collections have the same 6-8 differences near the beginning of ITS1 and some don't. It is odd for differences that are usually random in ITS to all be clustered together. When you see that happen, it often means your sequence was bad. In 2003, those with the different sequence (including most PNW ones) were described as Psathyrella amarescens. But as no clearly defined features exist between them, Wachter stated that P. amarescens is "clearly a form of P. corrugis".
Psathyrella longicauda EU - recent: WAx1 ORx3 - sequenced recently from WA and OR matching a reliable Wachter sequence, correcting Orstadius' sense of the species. Smith may have called it Psathyrella caudata EU (ORx1), which he reported once from OR. If this isn't what he found, then what was?
Psathyrella bipellis EU (WAx5) - recent: ORx1 - also known as Psathyrella barlae. Unlike other section members, it is not delicate, has a burgundy cap, and has a sweet odor. The other species with a sweet odor is Psathyrella suavissima in section /Pennatae, but instead of a burgundy cap it has a yellow-brown cap with white veil material all over it and hanging off the cap margin. We have one modern collection sequenced from OR. This is all by itself on the edge of this section, and it may deserve a section of its own. There are 2 clades of sequences in the EU. The second one doesn't seem to be a distinct species, but differs by some noise in one particular spot.
probable Psathyrella bipellis © Noah Siegel
/Atomatae - difficult to tell apart from /Microrhizae and /Psathyrella, with sparse veil material. Differs microscopically and more likely to be growing in rich soil or dung. Psathyrella prona EU (WAx7) - recent: WAx2 ORx2 - our sequence exactly matches the EU type sequence. Note that you cannot recognize this species by a pink cap alone when fresh (as sometimes claimed), as at least P. microrhiza can have a pink cap as well. But the somewhat olive toned upper stem may help distinguish this species. Psathyrella orbitarum EU (WAx3) - we need local sequences to compare with our reliable EU sequence to see if it really is here Psathyrella tenera NY (WAx1 IDx1) - we have local and EU DNA of this east coast species, which differ by 4 bp or so, the same amount that distinct species like P. prona vary by. We need east coast DNA to see if ours is the real species or not. Psathyrella potteri MI - recent: WAx1 - not known from the PNW until one recent WA collection from dung (and seemingly more likely to be in dung than the other species). P. hirta, in a different section, grows on dung but has scales. Wachter provides sequences of this species, and it has been determined that Smith collection AH33721 originally identified as Psathyrella kauffmanii var. exannulata represents this species.
Psathyrella prona (2 images) and P. potteri © Yi-Min Wang
/Hydrophilae - at least most species have especially small spores (<6 x 3.5u) and they often grow clustered Psathyrella piluliformis complex (WAx8 ORx12 BCx1) Psathyrella piluliformis? EU (clade 1) - recent: PNWx4 Psathyrella carinthiaca EU (clade 2) - recent: PNWx6 This complex is easy to ID - clustered on hardwood with a veil and tiny spores <6 x 3.5u. Smith knew these as Psathyrella hydrophila. Large clusters of mushrooms with larger spores may be Britzelmayria. There are 2 clades of sequences in the EU and all over the world, differing in 8 positions from each other. However, at least one sequence has 4 amibiguous nucleotides where the alleles of both clades are present in the same individual. So there are only 4 differences unaccounted for between clades. This could mean they're all the same species, although the fact that most sequences fall into one of the 2 clades and not in between is curious. We have local sequences in both clades. Some light was shed on the topic when Dr. Mary Berbee at UBC discovered that sequences in clade 2 match the Austrian type sequence of Psathyrella carinthiaca. We need a type sequence of Psathyrella piluliformis to see if that is clade 1 and if we can resolve the matter. The description of P. carinthiaca gave differences between it and P. piluliformis, but those differences don't seem to always bear out as a way to separate the two clades so it is probably not that simple.
Psathyrella fragrans ID (IDx1) - similar small spores, but clustered on conifer debris and at least the type collection was fragrant. We have the type sequence. Psathyrella maculata WA (WAx6 ORx5) - similarly small spores, but distinctively stocky (for a Psathyrella) developing a patchy/streaky cap growing in clusters on alder. We only have EU sequences that Wachter thinks might be this, so we need a type sequence to make sure.
Psathyrella mucrocystis ID (IDx1) - we only have an EU sequence that Wachter thinks could be this, so we need the type sequence to make sure. The spores are not as small as the other section members, so it is difficult to recognize.
/Lutenses Psathyrella lutensis EU - recent: BCx1 WAx1 ORx1 - never reported by Smith from the PNW, but recently found once in BC, twice in WA and once in OR, matching reliable UNITE and Wachter EU sequences. Our latest collection, below, had yellow tones so perhaps this can be recognized by having colours usually only found in Candolleomyces. But a photo of young fruitbodies in OR did not have any yellow cap colouring, although one stem was yellowish, so this colouration cannot be relied on.
/Saponaceae Psathyrella abieticola ID (WAx1 IDx1) - (P. subnuda WAx20 ORx15, P. vesiculocystis IDx1) - recent: ORx12 - a Smith collection of P. subnuda var subnuda (but not the type), the type collection of P. subnuda var velosa, and the type collection of P. vesiculocystis all match the type sequence of the older P. abieticola, and in the absence of other differences, Voto has synonymized them all. It seems to be quite common, but is very nondescript and requires a microscope for identification. We should consider that P. cuspidata WA (WAx3) is an older name for all of them. The current synonymies show that growth on the ground or on wood, and whether or not veil remains are hanging on the margin may not be taxonomically significant. They all share utriform cystidia with P. cuspidata. We are left with the primary difference of P. cuspidata being larger spores, 9-12x6-8 vs 8-10x4.5-7 for the above three. But one collection shown below had the larger spores.
/Pseudostropharia Psathyrella caput-medusae EU (WAx1) - (Psathyrella gruberi ORx1) - This is a pretty distinct species with a cap and stem covered in white scales that turn brown. The cap scales can rub off. The type of P. gruberi matches the type of P. caput-medusae, so they may be the same species. There is a second sister species in Europe, that may or may not be distinct.
Psathyrella pseudocotonea WA (WAx1) - it should be investigated if this is really a distinct species. We have no DNA, but I'm guessing it's in this section, since it's a section with a distinct appearance, so I did not place it in the "Little known species" section, even though we don't know much.
/Pennatae - contains some burned ground species and some ringed species Psathyrella pennata EU - recent: WAx3 - (=Psathyrella carbonicola OR) - this not uncommon burned ground species may be easy to recognize by habitat. The two type sequences differ only by 1 indel, and many people think they are the same species. Psathyrella CA02 - one OR sequence is similar, but distinct. It is also a burn related species, but was highly decorated with universal veil fluff and veil material around the cap margin.
Psathyrella rainierensis WA (WAx1) - the very short type sequence only has about 70 bits of useful ITS1 data, but those bits line up well with some EU sequences of Psathyrella flexispora, which is a complex with 2% genetic variation in ITS. We need better sequences. Psathyrella flexispora complex EU - recent: WAx3 - a species complex varying by 2% in ITS in the EU, one local collection is within this complex. It should be investigated if P. rainierensis above is a newer synonym. As the name implies, the spores are variable.
Psathyrella cf incondita WA (WAx1) - the WA holotype was not sequenced but two MI paratypes are 2 different species. Voto has a guess as to which represents the real thing, but we don't know that it will match the WA holotype. If it does, it is in this section. It is only 2 bp and 1 indel from the type sequence of the newer Chinese species Psathyrella jilinensis. We need the type sequence. Psathyrella jilinensis China - one WA sequence is 3 bp different than the type and may be this. Psathyrella 'jilinensis PNW07' - recent: WAx2 - more often, we find a sister species to the type sequence of Psathyrella jilinensis (and even more distinct from P. cf incondita), differing in almost a dozen places, most of them in ITS1. It is a stocky species on wood that has copious veil material that seems to hang around the cap margin more than form a ring on the stem.
Psathyrella paradoxa OR (WAx15 ORx5 IDx5) - the type sequence only got about 45 ITS2 characters, but the LSU sequence shows it is in the /pennatae. For something so abundant, nobody really knows anything about it and it is never reported or recognized. I guess it has an apt name. Psathyrella flexuosipes WA (WAx3 ORx1) - no modern collections, but we have the type sequence. Psathyrella fibrillosa EU (WAx2 ORx1) - recent: ORx1 - we have reliable EU sequences from Wachter, Orstadius and Nagy and now, one recent OR collection with a good photograph!
Psathyrella fibrillosa © Ann Goddard Psathyrella subpurpurea OR (ORx1) - recent: WAx1 ORx1 - It has a sometimes fleeting ring and a purple tinged cap with a hoary overcast when fresh, which fades as it dries out (it is hygrophanous) but reportedly becomes dark again when dehydrated. The best character is probably the hoary cap and ring/ring zone combination. We have the type sequence and a modern OR collection that matches it well. The WA collection had dried out and lost the fresh colour and only showed a ring zone.
Psathyrella sphagnicola EU (IDx1) - It supposedly also has a ring. We have reliable EU sequences so we can check if it is really here or not.
Psathyrella longistriata group (more ringed species)
Psathyrella 'longistriata PNW01'
Psathyrella
'longistriata
PNW02'Psathyrella
ellenae var. yubaensis
Psathyrella longistriata WA (WAx22 ORx20 IDx13) is the ringed species Smith reported by far the most. PNW01 was examined microscopically, and while not a perfect match to anything, it might best match P. longistriata. Psathyrella annulata ID (IDx1), Psathyrella ellenae ID (IDx1) and Psathyrella solheimii WY (IDx1) are probably in this complex. The aforementioned Psathyrella subpurpurea and Psathyrella sphagnicola also have a ring, but are not in the same tight genetic clade, and are not considered part of this group.
Psathyrella 'longistriata PNW12' © Yi-Min Wang
Psathyrella lanatipes OR (WAx8 ORx4) is right in the middle of the P. longistriata group clade, yet it doesn't have a ring, so that's odd. No modern collections have been found of it. Psathyrella wapinitaensis OR (ORx1 IDx3) - we have the type sequence. Psathyrella suavissima EU - recent: WAx1 - never reported from the PNW, but recently found in WA with sequence perfectly matching reliable EU sequences. It has also been called Psathyrella sacchariolens, but that name was never validly published. It smells very sweet and has a yellow-brown cap with white veil material all over the cap and on the cap margin when young. The similarly sweet smelling Psathyrella bipellis in a different section has a burgundy cap without white veil material.
Psathyrella hirta EU (WAx6 ORx1) - recognizable by growing on dung (like section /Atomatae) and having white cap and stem scales when young, that can rub off. we have a reliable EU sequence, but no recent collections to verify that's what Smith found here.
Psathyrella subcaespitosa OR (ORx1) - we have the Oregon type sequence, and one modern WA collection. Psathyrella cortinarioides UK - recent: WAx1 - this UK species was never reported in the PNW before until we found it at Bridle Trails. Another Bridle Trails first! One theory is that this is the abundant species that Smith called Psathyrella frustulenta EU (WAx11 ORx12 IDx19) (as it is unclear what species that name refers to).
Psathyrella PNW09 ('cf frustulenta' EU) - recent: WAx1 ORx1 - this is a name that's not being used much anymore, but has been used for a species with lots of shaggy white cap and stem material (like P. hirta but not found on dung). We have a recently found unnamed species that somewhat resembled it when fresh, found in WA, plus another record that differs by 3 bp from OR. We need to both figure out what P. frustulenta is, and what this present species is. Some think that it's just another name for P. cortinarioides, above, but we do know what that is, and the present species is distinct.
Psathyrella squamosa EU (ORx1) - recent: WAx1 - this species has a bit of genetic variation in the EU, so our recent collection, 7 bp different from some EU sequences, may still be the same species. Psathyrella PNW13 - one WA collection and a bunch of CA collections have sequences between 2 and 6 bp different from each other. One CA collection keyed out microscopically to Psathyrella seymourensis MI, but we have no type area sequences of that to compare to.
/Nolitangerae Psathyrella undulatipes OR (ORx1) (Psathyrella directa OR - ORx1) - recent: WAx1 - these two have the same type sequence, P. undulatipes being described earlier in the book. But the spore lengths and therefore shapes are somewhat different. The possibly synonymy should be investigated.
Psathyrella noli-tangere EU (WAx3 IDx4) - we have the EU type sequence, but no recent local collections to make sure this was what Smith found. Psathyrella fulvescens EU (WAx2) - we have a reliable EU sequence from Wachter, but no local collections to make sure this was what Smith found. Psathyrella PNW05 (ORx1) - recent: WAx2 ORx4 -sequencing of Smith's original P. alluviana collections found he had actually collected three different species. One of them, this one, is not even in the same section as P. alluviana. It is probably undescribed. Spores are ~7-9x4.5-5u.
Psathyrella warrenensis ID (IDx1) - we have the type sequence and a recent WA collection. Psathyrella senex EU (ORx1) - recent: WAx1 ORx1 - our sequence is only 2bp different than a reliable EU sequence, plus what appears to be noise at the beginning of one the longer sequence which I am ignoring.
/Confusae Psathyrella epimyces EU - this distinct species grows on shaggy mane mushrooms. Smith never found it here, but others have. We need local sequences to see if they match our reliable EU sequences.
/Cystopsathra Psathyrella sphaerocystis UK (IDx1) - an especially granulose section. We need local collections to compare with our reliable EU sequences.
/Pygmaeae Psathyrella olympiana WA (WAx8 ORx1) - stem and cap are white-fibrillose when fresh, but notable for having thick walled encrusted metuloid cystidia. There are two genetic species in the EU. Our first local collection matched concept #1.
/Obtusatae Psathyrella subcaerulea OR (ORx6) - we have the type sequence Psathyrella quercicola OR (ORx3) - this type sequence is only 1 bp different (+ 2 ambiguous locations that probably don't count) from the type sequence of Psathyrella subcaerulea var velata OR (ORx3). Since that variety's sequence is far away from its type variety sequence, it needs its own species name, so P. quercicola is the best name, assuming these are indeed the same species. The spore sizes as measured by Smith are different, so this should be double checked, perhaps they're not synonymous. Psathyrella nitens WA (WAx1) - recent: ORx3 - we have the type sequence and three recent sequences from OR.
Psathyrella 'nitens PNW06' (WAx2) - recent: ORx1 - two subsequent WA collections by Smith differ by 7 bp from the type sequence, and so does a recent OR collection and a recent WA collection.
Psathyrella 'nitens PNW06' © Sharon Squazzo Psathyrella obtusata EU (ORx3 IDx1) - (Psathyrella subargillacea MI (WAx1)) - The ITS type sequence of P. subargillacea is only 2 bp different from reliable EU sequences of P. obtusata, so they may be the same species, but the spore sizes are reported to be somewhat different, so maybe not. We should check collections for overlap.
/Spadiceogriseae Psathyrella spadiceogrisea EU (WAx1 IDx1) - we have an epitype sequence (not as good as a holotype, but pretty reliable) of this EU species but need local collections to prove it's here. In the EU and elsewhere, it's a complex with DNA varying up to 2%, so I expect our DNA might not exactly match. At least one of Smith's collections was mislabeled. Psathyrella kauffmanii MI group - (P. kauffmanii var exannulata ORx1, P. alnicola IDx5, P. oregonensis ORx1) - recent: P. oregonensis ORx1 CAx1 - Voto says the single collection of P. kauffmanii var. exannulata from Oregon is a micro match to the east coast type variety, except lacking a veil, so it is not known if it should be considered a distinct variety or not. It needs to be sequenced. The type sequences of all three species are almost identical, but as P. kauffmanii is the only one said to have a ring and P. oregonensis is the only conifer species, each one has unique properties, so Voto did not synonymize them all yet. P. kauffmanii was described first in the monograph, so that makes it the oldest name. Note that this genetic species has never been recorded in the PNW with a ring, but it has back east.
Psathyrella solheimii WY (IDx1) - this is one of the only species without DNA evidence that is not in the "Little known" section, because Voto analyzed the type microscopically and determined that it seems to be a distinct species in this section. We need a type sequence to see in detail what it is. Psathyrella albescens TN (IDx1) - we have the type sequence from TN, but we need local sequences to see if it really does occur here too. Psathyrella owyheensis ID (=P. carminei EU) (IDx1) - recent: WAx1 ORx1 - we have the type sequence. Voto determined that ITS DNA of this species can vary by 1% or so with no ecological or morphological differences, and found that the type sequence of Psathyrella carminei EU was inside this clade and therefore was a junior synonym (the two Tef1 sequences were also similar enough for him to think them conspecific). Wachter noticed that sequences of P. carminei could vary by up to 1%, although he didn't realize that P. owyheensis fell into this clade and didn't address that. Recent WA and OR sequences are within 1% of each other in ITS and inside this clade and probably represent P. owyheensis.
Psathyrella owyheensis © Yi-Min Wang Psathyrella umbrosa NM - (P. fulvoumbrina IDx1) - The type sequences are the same and the micro and habitats are a good match according to Voto, so we should use the older name. Psathyrella elliptispora OR (ORx1) - (P. griseopallida CA) - recent: WAx1 ORx2 - the types differ by 1 bp. Smith's spelling P. elliptospora has been corrected. The spore shapes are somewhat different for the two names, so I don't know for sure that they're synonymous. The photographed collection has spores matching P. elliptispora.
Psathyrella elliptispora © Yi-Min Wang Psathyrella 'elliptispora CA01' - one recent WA collection and some CA collections are 4 bp different from the type sequence of P. elliptispora, and 5 bp different than P. praetenuis. (Another odd recent OR sequence is 5% different in ITS1 but only 3 bp different in ITS2).
Psathyrella praetenuis ID (IDx2) - (P. atrifolia WAx12 ORx12 IDx12) - recent: WAx1 ORx5 - Smith's spelling has been corrected and the name was changed from P. atrofolia to P. atrifolia. We don't have the type sequence of P. atrifolia, but Voto sequenced one of Smith's original collections and it matched P. praetenuis, and the morphologic and ecologic differences between species was also determined to be very slight. A second of Smith's collections didn't successfully sequence, but was morphologically and ecologically analyzed and determined to match as well. A third original collection was an oddball that seemed to be different. We'll need the type sequence to prove that P. atrifolia is a synonym of P. praetenuis, but that's the current reasonable assumption and declaration by Voto.
Psathyrella PNW11 - this one OR sequence is in between P. praetenuis, P. umbrosa and P. elliptispora, about 6-8 differences between all of them. Psathyrella pseudolimicola ID (IDx1) - we don't have an ITS sequence, but a Tef-1a sequence shows it is sister to P. griseopallida. Psathyrella septentrionalis MI (WAx1) - recent: WAx1 - grows on wood with reddish-brown colours and ample veil remnants hanging off the cap margin. It's a nice looking species complex. We have one recent collection from WA that is a pretty good ITS DNA match to this and the next species. We have the MI type sequence and a similar sequence of P. septentrionalis var. vesiculosa MI. Psathyrella velibrunnescens WA (WAx20 ORx5) - supposedly differs from P. septentrionalis by a veil that discolors brown. No type sequence, but we have a sequence from the EU of a collection that seemed a good micro match by Voto that is a guess as to what it might be. It is a close sister species of P. septentrionalis (ITS is not that differentiable). Of course, it's an EU sequence and a WNA type area, so the real thing could be slightly different genetically, even if Voto's sequence is identified correctly. It was very common in Smith's day. The recent collection of P. septentrionalis could be this.
Psathyrella septentrionalis/velibrunnescens complex © Yi-Min Wang Psathyrella aff ammophila EU - Smith never found it in the PNW, but others have since reported a species in sand dunes. One CA collection differs from EU sequences by 6 bp, so it may be a sister species if any other differences can be found, but Voto analyzed ENA material and thought that both ENA sand dune species, Psathyrella velatipes and Psathyrella arenulina, were morphologically and ecologically the same species as Psathyrella ammophila, so perhaps our species is also the same too. Not every sand dune species is going to be this one. Another CA sand dune collection had a unique sequence quite distant from every other species. And we also have a sand Panaeolus.
/Jacobssoniorum Psathyrella PNW08 ('under water') ID - a recent sequence is in this section, found under water in Idaho, but not at all closely related to P. aquatica.
/??? Psathyrella ramicola OR (ORx1) - the type sequence falls outside of all other sections, this is somewhat suspicious and should be investigated. |
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Little known Psathyrellas - click to expand
Species nobody knows anything about, but have been reported by Smith and/or others from the PNW (in alphabetical order). The ones that aren't rare have been coloured in, and should be studied first.
Psathyrella acuticystis ID (IDx2) Psathyrella alboalutacea WA (WAx7) Psathyrella argentata ID (IDx1) Psathyrella avellaneifolia MI (ORx3) - possibly a Homophron, but not necessarily Psathyrella borealis MI (IDx1) Psathyrella bifrons UK (WAx3) - as stated above, one sequence with this name was actually P. microrhiza, but it was likely misidentified. It could mean that this species is in section /Microrhizae, but I'm not sure. Psathyrella boulderensis ID (IDx3) Psathyrella candidissima WA (WAx1) Psathyrella fusca EU (WAx3 ORx3) - should perhaps be abandoned as a name (or at the very least we need a type), as nobody is sure what sequences represent it. One Smith collection turned out to be P. kauffmanii, we should sequence the rest to see what they really are. Many sequences of what people are calling this in the EU turn out to be 3 bp different than the type sequence of Psathyrella abieticola, a local species described above. So if this name isn't abandoned, one of the possibilities is that it may be a sister species, or even a valid older name for P. abieticola. Psathyrella frustulenta EU (WAx11 ORx12 IDx19) - nobody seems to be sure what this species is. We need to figure out what Smith found that he called this, because it's very common. One theory is that it is P. cortinariodes. Another possibility is P. sp. 'cf frustulenta', above. It is likely in section /Pennatae. Psathyrella cascadensis WA (WAx15) Psathyrella circellatipes EU (IDx1) Psathyrella columbiana BC (BCx1) - Glacier National Park near the Alberta border Psathyrella crassulistipes ID (IDx4) Psathyrella cuspidata WA (WAx3) - see P. abieticola in the section above for a discussion on how this could be an older synonym for P. abieticola. Psathyrella deserticola ID (IDx1) Psathyrella distans WA (WAx1 ORx1) Psathyrella duplicata WA (WAx1 ORx1) Psathyrella equina ID (IDx1) Psathyrella fulva ID (IDx1) Psathyrella fuscifolia NY (WAx2 ORx2) - Smith's spelling of Psathyrella fuscofolia has been corrected. Psathyrella fuscospora ID (IDx1) - for some reason the 'o' was OK this time Psathyrella grasmerensis ID (IDx1) Psathyrella idahoensis ID (IDx1) Psathyrella indecorosa WA (WAx1 IDx1) Psathyrella iterata WA (WAx18 IDx2) Psathyrella luteovelata WA (WAx6 ORx1 IDx1) Psathyrella mazamensis WA (WAx1) Psathyrella mesocystis ID (IDx1) Psathyrella minima NY (IDx3) Psathyrella monticola ID (IDx1) Psathyrella nezpercii ID (IDx3) Psathyrella ochracea EU - never found by Smith, I do not believe Phillips' report from the PNW is true. Psathyrella pallida ID (IDx1) Psathyrella payettensis ID (IDx1) Psathyrella polycephala EU (WAx2 ORx1) - some consider it a Psilocybe (Deconica) but I believe Smith. Psathyrella populorum ID (IDx1) Psathyrella pseudofrustulenta WA (WAx1) Psathyrella pseudofulvescens WA (WAx1) Psathyrella pseudolongipes WA (WAx1) Psathyrella pseudotrepida ID (IDx4) Psathyrella pseudovernalis WA (WAx1) Psathyrella psilocyboides ID (WAx2 IDx2) Psathyrella roothaanensis ID (IDx1) Psathyrella rubescens WA (WAx6) Psathyrella rubicola WA (WAx3) Psathyrella salictaria ID (IDx1) Psathyrella similis MO (IDx1) Psathyrella stuntzii WA (WAx1) Psathyrella subagraria NY (ORx1) - this could possibly be a sister species of Coprinopsis uliginicola, or at least belong in Coprinopsis. Psathyrella subalpina WA (WAx3) Psathyrella subhepatica MI (ORx2) Psathyrella sublongipes ID (IDx1) Psathyrella subradicata ID (IDx1) Psathyrella subrubella MI (ORx1) Psathyrella subtenacipes WA (WAx1) Psathyrella tahomensis WA (WAx4) Psathyrella trepida EU (WAx7) - if one EU sequence is to be believed (but it probably shouldn't), it is in section /Jacobssoniorum. This sequence, however, matches the type of P. sublatispora. I have no idea if Smith's 7 WA collections are either of these species or neither. |
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Lacrymaria - click to expand
These species are often more sturdy and less fragile, with a well developed veil, rough spores and cap hairs/scales that don't rub off (like the smooth spored and not quite as stocky Cystoagaricus). Often, water drops form on the gills edges (Lacrymaria means "tears"). It is necessary to separate this from Psathyrella, or the entire family would be one genus, inky caps and non-inky caps together. Species mentioned: Lactrymaria lacrymabunda, velutina, echiniceps, subcinnamomea, rigidipes
Lacrymaria lacrymabunda EU (WAx1) - Wachter provided sequences of it, and it took finding over a dozen sequences of L. velutina, discussed next, before finding a sequence that matched L. lacrymabund. in BC. Adding to the confusion is that Smith though it was a species with small spores (<7.5x4u) but countless other sources say otherwise. We need to measure our collection. What did Smith find with small spores? Lacrymaria velutina EU (ORx1 IDx3) recent: PNWx6 - The handful of GenBank sequences with this name all match our sequences fairly well, and the spores are the right shape and size (large and warty spores up to 11x6.5u). The sequences are also found in Asia and Europe, where the species was described, but unlike the NA and Asian collections, the matching sequences from Europe were not labeled P. velutina, but other various species instead. We'll need a type sequence to prove that's what this species is. All half dozen sequences of Lacrymaria from the PNW so far have been in this clade. There is some variation in the sequences, up to 6 bp, so although Wachter considers them all one species, we should keep an open mind about it. Other species in this genus seem to share similar ITS so maybe there's more than one species hiding out in here, but all collections seem to microscopically match L. velutina. The photos show the light cap fibrels can turn dark like in L. echiniceps, so I think the colour of the cap fibrels is probably not taxonomically important. Lacrymaria 'rigidipes PNW02' (WAx1 ORx2 IDx10 BCx1) - L. rigidipes is from NY but we only have one local sequence from BC and an EU sequence with that name. It's possible our local sequence is not this, so I am giving it the moniker PNW02 for now. We need reliable NY type area sequences of what that species really is. It is supposedly differentiated by a depression in the shape of the spores, which are otherwise large like L. velutina. The spores are said to be sometimes almost smooth and sometimes distinctly warty, so that doesn't help. We don't have a local photo yet.
Lacrymaria subcinnamomea MI (WAx1 ORx4 IDx2) - This smaller? species' type sequence has almost the same ITS DNA as L. lacrymabunda (no bp differences but 5 indels). Other sequences from the EU and ENA share the same 5 indels, so it is being treated as a distinct species, with the possibility that other genes may need to be sequenced to properly show them as distinct species. Smith reported it from the PNW so we need local specimens to confirm if this species is really here. It has medium sized spores (<10x5u) that are barely warty. Lacrymaria echiniceps NY (WAx1 ORx1 IDx1) - is also rarely reported here, but we need reliable sequences from back east before we can start comparing what we find here. It has the same medium sized, almost smooth spores of L. subcinnamomea, but the hairs on the cap are said to be darker. This may not be taxonomically significant. If they are the same, this is the older name.
Summary Judging by Smith's reports, large spored species (L. velutina and L. rigidipes) were reported in WA or OR 4x and ID 13x (perhaps more common inland than near the coast?) and medium spored species (L. subcinnamomea and L. echiniceps) were reported in WA or OR 7x and ID 3x (perhaps more common near the coast?). There are 2 major clades of Lacrymaria. Most of the known medium spored species are in the bottom clade, and the our large spored local genetic species found so far is in the top clade. My guess is that the genetic species we have found is L. velutina but L. echiniceps is in fact here waiting to be verified, and that dark cap fibrils or not are not indicative of species. |
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Homophron - click to expand
These species never have any veils (neither a partial veil nor a universal veil leaving material on the cap or stem), the spore colour is pale for the family (never black) and the cystidia are metuloids (thick walled with crystals, like many Inocybe). Our common local species is a large, stocky, clustered species growing on wood. Species of Psathyrella grow in clusters too, but are either found on the ground, or if on wood, have darker spore prints and a veil. It is necessary to separate Homophron from Psathyrella, or the entire family would be one genus, inky caps and non-inky caps together. Species mentioned: Homophron spadiceum (Psathyrella spadicea), Psathyrella variata, Psathyrella sublateritia, Homophron camptopodum (Psathyrella camptopoda, Psathyrella camptopus), Homophron naucoria (Psathyrella naucoria), Homophron naucorioides (Psathyrella naucoriodes)
Homophron spadiceum EU (WAx9 ORx2 IDx3) - a large, stocky, clustered species on or near hardwoods like cottonwood or alder. This European species has been reported commonly throughout the PNW. The Idaho type collection of Psathyrella variata ID (IDx1) has been sequenced (with a sequence matching well to H. spadiceum's type sequence) and analyzed by Orstadius and officially been declared a synonym. The recent PNW collections hae sequences 3-5 bp or so different from the types and in one case from each other (plus a few ambiguous locations), so it's possible there's more than one species in this group, but so far the ambiguous locations account for much of the differences and the other differences aren't consistent enough to indicate a high likelihood of another species. 'Psathyrella' sublateritia MI (WAx9 ORx5 IDx1) is almost identical, but with a brick red spore print. One presumably red spored collection from presumably MN was sequenced and is almost identical (3 indels) in LSU from the type sequence of H. spadiceum, but LSU mutates slower than ITS and that does not mean it is the same species. P. sublateritia was not moved to Homophron when the rest of these species were. We need local collections.
Homophron cernuum EU (=Psathyrella crenulata AK?) - not as distinctive as H. spadiceum (not as stocky and not as clustered). It has somewhat smaller spores than H. spadiceum and more rounded metuloid cystidia. Smith found them in Alaska, but they weren't known from the PNW until one recent WA collection matched closely with the type sequence of P. crenulata (2 bp different). Orstadius synonymized P. crenulata with the older Homophron cernuum, and said the ITS sequences of the two were identical, but that's not strictly true. There are more than a half dozen differences between the type sequences, but other sequences of each don't show much or any difference, and no significant ecological or morphological differences were found between the species (Smith spoke of more voluminous cheilocystidia in P. crenulata as the only difference). So for now, the synonymy stands, even with type sequences 1.5% different from each other.
Homophron cernuum © Yi-Min Wang
Homophron camptopodum NY (WAx2) - a much smaller species, not always as clustered, also on hardwood. Also known as Psathyrella camptopoda and Psathyrella camptopus. When this east coast species was moved to Homophron, Smith collections of this presumably rare species found in WA and later ID were examined and seemed to be the same as ones from the east coast, as was a collection from Russia. However, the two sequences we have (from the EU and ??) are 5 bp and 3 indels different from each other in ITS (about 1% different) so it's not certain that there's only one species around the world. Homophron naucoria WA (WAx3 IDx1) and Homophron naucorioides ID (WAx2 IDx3) differ by either a slightly different spore shape or spore print colour, and might be distinct species in this group, or they might just be newer synonyms. When they were moved to Homophron I'm not sure the work was done to figure that out. We have one LSU sequence of a Smith collection of each of them (but not the types). The latter species is very similar to the LSU of H. camptopodum (no differences out of 200 locations from one sequence, and 2 differences out of 400 locations from another), and the former species is somewhat different but not a clean sequence, so this needs further study. |
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Candolleomyces - click to expand
No pleurocystidia, although pleurocystidia in Psathyrella may be very hard to find. Learn the species, as the genus is hard to describe. The common one is often honey coloured, a bit brighter than the usual greys, and can be identified by colour alone. This genus and the following ones were separated to avoid paraphyly if Coprinellus s.l. is removed. You could be forgiven for still thinking of them as a kind of Psathyrella. Species mentioned: Candolleomyces candolleanus (Psathyrella candolleana), typhae (Psathyrella typhae). Psathyrella hymenocephala, roguiana, singeri, Psathyrella incerta Candolleomyces candolleanus EU (WAx41 ORx8 IDx5) - recent: almost 20 -this abundant European species is found worldwide. Usually somewhat honey coloured, urban or wild, no pleurocystidia, veil material often hanging off the cap margin when young. ITS1 varies unusually. Our 15 or so local sequences have between 7-10 differences in ITS1 from the EU type sequence, but 0-3 differences in ITS2. It has the same kind of genetic variation worldwide, but no ecological, macro or micro differences can be spotted, so the species is not being split up. Psathyrella roguiana OR (ORx1)'s type has been sequenced and it is a very close match, and likely represents the same species. Psathyrella hymenocephala NY (WAx1 ORx1) is also considered to be the same species, as Smith admitted collections intergraded between the two and collections from both the east and west that better match P. hymenocephala sequence comfortably inside the C. candolleanus sequences. On the other hand, at least 5 additional Candolleomyces species have been described whose ITS cannot be distinguished from C. candolleanus, but which supposedly each have at least 1 microscopic difference. (They are 'Psathyrella' cladii-marisci, C. badhyzensis, C. badiophyllus, C. trinitatensis, and C. incanus). Both Bau's and Zhou's multi-gene tree showed each species distinct, but most species only had one sequence, so he did not show that even 5 genes can distinguish the species, and thus I don't think he proved that the species are distinct. For now, I am going to refer to everything in this clade as Candolleomyces candollanus.
Candolleomyces candolleanus © Daniel Winkler
Candolleomyces typhae EU (WAx1) - This European species does not resemble C. candolleanus, but it does not have pleurocystidia. We need local collections to verify that it really does occur here, as Smith reported it from WA. Formerly Psathyrella typhae. Candolleomyces singeri FL - there is a sequence from China, which Wachter believes may represent this species. A variant was found in Oregon by Smith that he thought might be the same species, but it remains somewhat speculative that this species is here. We need local collections. Psathyrella incerta NY (WAx3 ORx1 IDx1) - is said to have no pleurocystidia. If that is true, it may be in this genus. This NY species was rarely reported by Smith in the PNW. |
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Britzelmayria - click to expand
Growing in large clusters on the ground, not wood, with rooting stems that often fuse. It has larger spores and very little veil material (probably never visible) unlike the similar clustered but wood inhabiting Psathyrella piluliformis. Species mentioned: Britzelmayria multipedata (Psathyrella multipedata)
Britzelmayria multipedata MS - recent: WAx1 - this Missouri species was never found in the PNW by Smith, but recently was found in WA, with DNA a pretty good match to reliable sequences from Europe. However, we need type area sequences from the east coast before we can be certain that ours is the real thing. Britzelmayria PNW01 - one tightly clustered collection of stocky fruitbodies were quite strigose on the stem underneath what appeared to be a ring zone. It seems to be new to science.
Britzelmayria multipedata © Buck McAdoo (2 images), Britzelmayria PNW01 © Andrew Parker |
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Typhrasa and Kauffmania - click to expand
The definitions of these genera are pretty obscure - in Typhrasa the cystidia have large refractive globules, and in Kauffmania the spores are rather pale without a discernable germ pore. Our one species in each genus are hard to differentiate from our many Psathyrellas. Species mentioned: Typhrasa gossypina (Psathyrella gossypina), Psathyrella delineata, Psathyrella canadensis, Kauffmania larga (Psathyrella larga)
Typhrasa gossypina EU - small but stocky. Orstadius examined the NY type of Psathyrella delineata NY (ORx2) and it was a perfect micro match for the older European species Typhrasa gossypina. Four subsequent east coast sequences of newer collections turned out to be 6 bp and 2 indels different, so Wachter reserves the right that they may turn out to be a different species. But for now, with no other differences found, I am assuming they are the same. Psathyrella canadensis ON (WAx1) is assumed to be another synonym. Orstadius examined the one Smith collection from WA and found that it too was a micro match. Typhrasa PNW01 - I think this is a distinct species. The ITS sequence was dirty, but the LSU sequence was clean and seemed distinct from other Typhrasa LSU sequences. More collections are needed of local Typhrasa.
probable Typhrasa gossypina © George Barron, T. PNW01 © Michael Beug
Kauffmania larga MI - recent: WAx1 ORx1 -This dark brown capped rather stately species is said to be the only one in the genus, although worldwide sequences of it can vary by a dozen bp or so. Smith never found it here, but we did find recent collections in WA and OR that are within the typical genetic diversity of the species 4 bp from some EU sequences. We need east coast type area sequences, which are missing.
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Cystoagaricus - click to expand
Large, wood inhabiting species with a hairy/scaly cap and odd-shaped spores. Like Lacrymaria, the scales are not removable. Lacrymaria usually has warty spores, while these have smooth spores. Also, Lacrymaria is not wood inhabiting and is usually stockier, from what I can tell. Species mentioned: Cystoagaricus hirtosquamulosus (Psathyrella hirtosquamulosus), Psathyrella lepidotoides, Cystoagaricus weberi (Psathyrella weberi)
Cystoagaricus cf hirtosquamulosus NY (WAx4) - One CA sequence is 3 bp and 1 indel different than an official ENA sequence from TN, but some have said two west coast collections look distinct from east coast collections, so maybe that's enough ITS difference for ours to be a distinct species. EU sequences are 2.5% different, mostly in ITS1, which is odd. Wachter knew that yet he assumes that it is all one species worldwide. Cystoagaricus lepidotoides ID (IDx1) -We have the ID type sequence and matching sequences from OR and UT. Cystoagaricus weberi FL (IDx1) - Voto moved this here, so it probably is a Cystoagaricus, but no sequences are available.
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