Danny’s DNA Discoveries – Inky Caps of the PNW
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
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 turn to ink (or at least tatter in age). Those with a pleated cap are likely inkies and not "Psathyrella". The spores are usually black, but not always -they might be very dark brown. Not all of them are very fragile, but they all do have a cellular cap cuticle (explained on my Psathyrella 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.
A few species of Coprinopsis and Parasola do not deliquesce in the least, and so are also described on the Psathyrella page.
Parasola - click to expand
These species are often extremely pleated, occasionally with tiny cap hairs sticking out, which may have more of a tendency to open up and flatten in age than other genera. There's often a bright brown eye on the disc. There is no universal veil material on the cap (nor any other pileal cystidial elements), and they don't really turn to ink, just tatter. The gills are often distinctly free. They are often found in grass and gardens. They can be very difficult to separate from Coprinellus and Tulosesus. Coprinellus is more likely to have granular veil particles on the cap. Tulosesus is even harder to differentiate, but probably doesn't have as distinct an eye on the disc. One 'Psathyrella' species that is not pleated turned out to be a new section of Parasola. It is very difficult to tell apart from Psathyrella.
Species mentioned: Parasola plicatilis, auricoma, lactea, neoplicatilis and conopilea
The following have been reported in the PNW but not confirmed by DNA yet, so we need collections.
Parasola plicatilis EU - (no cap hairs)
Parasola auricoma EU - (more brightly coloured, with very tiny cap hairs). This seems to be a complex so there may be more than one species going by that name.
Most collections are sequencing to be species not expected from the PNW.
Parasola 'neoplicatilis PNW01' - Interestingly, the first typical pleated Parasola that I had sequenced from Bridle Trails in WA turned out to be none of the species reported from here, but a sister (3% different) of a new species that is currently being described (I think from Japan), Parasola neoplicatilis n.p. It is possible it will need its own name.
Parasola lactea MI - some say it is the same as Parasola leiocephala, but the DNA appears to be quite distinct.
Parasola 'lilatincta PNW02' - Also interestingly, the second typical Parasola sequenced, from Oregon, also turned out to be none of the above that it was supposed to be. This one is a sister species of the P. lilatincta complex, which in the EU has ITS DNA that varies by 2%. This unnamed sister species is also a complex with ITS DNA that varies by >2%.
Parasola 'neoplicitilis PNW01' © Daniel Winkler, P. lactea © Yi-Min Wang (2 images), P. 'lilatincta PNW02' © Bruce Newhouse
Parasola aporos EU (=P. plicatilis-simils EU?) - continuing the tradition, the third Parasola sequenced was a third species unknown to the PNW, and we've since found it again, both times in WA. The sequences are 3-4 bp in ITS1 from the type sequence of P. aporos and from each other (but identical in ITS2). P. aporos differs from the type of P. plicatilis-similis by 1 bp in ITS2. If they are synonyms, P. aporos appears to be older.
Parasola aporos © Yi-Min Wang
Parasola PNW03 - one WA sequence is a distinct species from anything known. Nothing much is known about it yet besides apparently rather distant gills.
Parasola PNW04 - one OR sequence is also a distinct species from anything known.
Parasola PNW03 © Yi-Min Wang (2 images) and P. PNW04 © Connor Dooley (2 images)
Parasola conopilea - (sometimes misspelled P. conopila or P. conopilus). This former Psathyrella 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. There are 2 clades of this European species, and ours is closest to the clade with the type sequence, so this complex would have to be split up into more than 2 species for ours to need a new name, and I don't think that's going to happen. This picture shows the cap cuticle clearly and easily breaking across (not radially) due to the cellular cap cuticle.
Parasola conopilea © Alan Rockefeller
Coprinellus - click to expand
Usually also golden brown in youth, and also strongly pleated, also often free gills, very much like Parasola, although there is not usually as sharp an eye on the disc. These are species with a universal veil material made up of roundish cells often visible as particles on the young, fresh cap. The caps may turn to ink, or they just might tatter. The segregated Tulosesus is more likely to have no universal veil, but microscopic cap hairs. (as well as angular spores). However, it should be noted that of all the segregate genera in the Psathyrellaceae family, the segregation of Tulosesus from Coprinellus is the least backed up by data.
Species mentioned: Coprinellus micaceus, saccharinus, domesticus, radians, xanthothrix and flocculosus. Coprinus arachnoideus, bubalinus and alutaceivelatus. Coprinopsis bulalina and alutaceivelata.
Coprinellus micaceus - This worldwide species has glistening particles like mica all over the cap when young. It is said to have miniscule pale hairs on the stem, unlike the following lookalike. You should suspect any similar species you find without the mica particles is probably this species, because they wash off easily and the species without the particles are much rarer.
Wachter (see the introduction bibliography on the Psathyrella page) points out that many American sequences are quite distinct genetically from many European sequences, although both have now travelled the world. Interestingly, the 4 PNW sequences I have match the real European sequences, and only one California sequence I've seen represents the American species. No difference can be found between them yet, so there is not yet a call to treat them as two separate species.
probable Coprinellus micaceus © Christian Schwarz
Coprinellus saccharinus - This very, very similar species is said to have a smooth stem. It has genetics that are slightly different in America versus the EU, and the one OR sequence I have matches the Americas. Again, without any ecological or morphological differences, there is no call to separate the species yet, especially since the genetic difference is so slight - only about 3 bp plus 5 ambiguous positions. This species was not known from the PNW until the Oregon sequence, so I don't know how to recognize it.
Coprinellus saccharinus © Jonathan Frank
Coprinellus disseminatus EU complex - This usually forms a large cluster of small fruitbodies, with smooth to ever so slightly scurfy caps. It does not deliquesce, but it's quite pleated. Different collections have different sequences that seem to differ by >3% from each in ITS, but that usually involves big chunks of indels, so I would say the differences could be accounted for by a half dozen bp and a half dozen chunks of indels. One study found no morphological or ecological differences among different collections, so perhaps they will continue to be the same species. The two local OR collections we have sequence differently from each other and from EU sequences by the above amount. Instead of labeling each sequence with its own code, I'm going to consider this all one complex for now, but it could be useful to get additional local sequences to see if they match one of the existing two or if they are unique.
Coprinellus disseminatus complex © Danny Miller
Coprinellus domesticus group (3 species?) - The particles are larger and resist wearing off a little more, but the main feature of these European species is an orange felty mat over the substrate. This should be confirmed in our local collections, as so far it has not been noted in them.
We have a good understanding of what C. domesticus is, but two other European species, Coprinellus radians and Coprinellus xanthothrix are poorly understood. There are perhaps a half dozen undescribed species in this complex in Europe, and so far I have found DNA of three undescribed species in the group here in the PNW. So far, there are no consistent ecological or morphological differences found between any of the these species, except for Coprinellus domesticus, which seems to be able to be identified somewhat reliably with a microscope, but we have not confirmed any of the named species from the PNW. #1 was found in WA, OR and CA. #2 and #3 are only known from a single sequence in OR.
Coprinellus section /Domestici © Christian Schwarz
Coprinellus flocculosus EU (=C. arachnoideus WA, =C. bubalinus CA, =C. alutaceivelatus WA) - This European wood chip species has the largest felty patches on the cap, even larger than the above. Fred Van de Bogart (see my Coprinus page) described three species that have almost identical ITS DNA and are probably synonyms: Coprinus arachnoideus, Coprinus bubalinus and Coprinus alutaceivelatus. The latter two were later mistakenly placed in Coprinopsis, which is interesting, perhaps partially because large felty patches is usually a trait of Coprinopsis instead of the more particle-like veils of Coprinellus.
probable Coprinellus flocculosus © Christian Schwarz
Tulosesus - click to expand
This new genus is hard to characterize. Again, the usually pleated caps may deliquesce or they may just tatter. They don't usually grow on wood, are often golden brown, fading to grey in age except in the centre, but the eye on the disc won't usually be as distinct as in Parasola. The caps may be slightly granular, but do not usually have the mica particles or veil material of the Coprinellus species above (more usually just very tiny hairs (setae) instead). They are not usually as finely pleated as Parasola. They were segregated from Coprinellus, but it should be noted that of all the segregate genera in the Psathyrellaceae family, the segregation of Tulosesus from Coprinellus may be the least backed up by data, so feel free to still think of these as Coprinellus section Setulosi for now.
Species mentioned: Tulosesus impatiens, heterosetulosus, hiascens, congregatus, ephemerus, sclerocystidiosus, eurysporus, velatopruinatus
The following European species are reported from the PNW, without genetic proof yet: T. heterosetulosus, T. hiascens, T. congregatus and T. ephemerus. We need collections of all of these. I do have sequences of all of them from the EU to compare to, except for T. ephemerus, which has no reliable sequences yet. Once again, ironically, as in Parasola, most local collections that were sequenced turned out to be none of the above.
Tulosesus 'impatiens PNW01' - young caps have a somewhat unique broad umbo and are yellow-brown. Mature caps change to grey-brown (although this colour change can be common in this family). CA and WA sequences are at least 7bp different than official Wachter and UNITE EU sequences, so our local west coast species may be undescribed. This find is the closest to finding one of the reported species here instead of something surprising.
Tulosesus sclerocystidiosus EU - this EU species varies by 1% or so in ITS, according to Wachter. Three OR sequences and 1 WA sequence match some EU sequences fairly well. Some of them are about 1% different, and just outside the clade of EU sequences, but since that kind of variation exists within EU sequences, I'm comfortable calling our species that for now.
Tulosesus velatopruinatus EU - We had one environmental sample from OR of this species, and now a fruiting body was finally found there too, but it was in a flower pot, not from the wild. Wachter provides us EU sequences and ours match exactly.
Tulosesus eurysporus OR - described from Oregon, so it is definitely here, but we don't have any sequences to know how to recognize it.
Tulosesus 'impatiens PNW01' © Yi-Min Wang and Alan Rockefeller, Tulosesus sclerocystidiosus © Richard Morrison, T. velatopruinatus © Bruce Newhouse
Coprinopsis - click to expand
Microscopically, the pileipellis is a cutis, which means the top layer of the cap has strings of cells laying flat. To the naked eye, they often have shaggy veil material on the cap, but not always, and so do some other genera occasionally. The caps are often greyish, and they usually completely deliquesce. The sizes can range from miniscule to very large (caps 1 mm to 10 cm). Not all Coprinopsis turn to ink. Some former Psathyrellas that do not deliquesce may be recognized by their relatively stocky stature and large spores >10u across.
Species mentioned: Coprinopsis atramentaria, atramentaria var. crassivelata, depressiceps, striata, pinguispora, acuminata, romagnesiana, strossmayeri, picacea, lagopus, lagopides, marcida, brunneistragulata, jonesii, pachyderma, cinerea, fimetarius, friesii, nivea, stercorea, tetraspora, subdomestica, undulata, sylvicola, kubickae, phaeospora, psychromorbida, uliginicola, canoceps. Psathyrella longipes, fragilissima and elwhaensis. Coprinus alnivorus.
Coprinopsis atramentaria EU - Alcohol inky (=Coprinopsis depressiceps WA?) - The alcohol inkies contain coprine, a substance with similar effects as disulfram (the active ingredient in antabuse) that make you quite sick if ingested when you have also ingested alcohol. This most famous alcohol inky cap, with a large, grey-brown cap with a broad or depressed cap apex (not narrow and/or umbonate like Coprinopsis striata, next). The cap and lower stem may be bald or have brown scales, but they are different than the orange-brown concentric scales of C. romagnesiana, below, which has a scalier stem too. Fred Van de Bogart described local collections with a depressed cap centre and a slightly different shape of spore as Coprinopsis depressiceps, and reported both species locally. The ITS type sequence of C. depressiceps is about 6 bp different from many EU sequences of Coprinopsis atramentaria, but Wachter showed that throughout Europe and the world you can find that ITS can vary by that amount within C. atramentaria, so perhaps they are synonyms. We should make local collections matching C. atramentaria and see what their sequence is.
Coprinopsis striata - Alcohol inky (=Coprinopsis pinguispora WA) - Very similar alcohol inky, but with a narrower and/or pointier cap and not quite as large. It is practically smooth with even fewer visible scales. Our local species was described by Fred from WA, and most sequences from OR, WA and BC of the "atramentaria group" are this species, so it appears to be our most common species by far. Fred also described Coprinopsis pinguispora a few pages later, but its type sequence is the same, so it is probably not a distinct species. It only differed by not having striations on the cap, which is probably a variable feature.
Coprinopsis acuminata EU - a sister species, with more "normal" shaped spores than C. striata, that Fred also reported from the PNW, so it's possible it is here too and should be looked for.
Coprinopsis atramentaria © Keifer Chalk, Coprinopsis striata © NAMA and the Field Museum of Natural History
Coprinopsis romagnesiana WA - Alcohol inky - This alcohol inky has coloured, orange-brown scales, concentrically arranged as the cap expands (unlike in C. atramentaria), with the scales built in to the cap so that they can't be washed off. The stem is quite scaly. It's an EU species that has been found and sequenced in WA three times now.
Coprinopsis strossmayeri EU/variegata NY - Another largish, grey, clustered, narrow capped species (like C. striata) but with a cap covered in whitish universal scale patches that can wash off the cap and with thick orange-brown rhizomorphs that smell mouldy. The European species has variable ITS DNA that varies by about 2.5% in Europe, but so far it's all considered one species. A couple of east coast sequences that might be C. variegata fall in this complex. This species complex was not known in the PNW until one poisoning report in BC where the mushroom they ate was sequenced and turned out to be somewhere in this complex (but not matching the east coast C. variegata exactly). Interestingly, it is not considered poisonous, as it is not in the same clade as the alcohol inkies.
Coprinopsis atramentaria var. crassivelata WA - this is one of the two above species (we do not have the type sequence). Fred did not note either of those species for the PNW, but did describe this variety, which is probably one of those two (the other probably not having been noticed by him). At first I thought it was C. romagnesiana as that is closely related to C. atramentaria and found in WA. But Fred's description is of universal veil material that can wash off, which is more like C. variegata. He does not note the colour of the veil patches. The spore sizes more closely matches C. variegata as well. I suppose it does not matter which one it is, as his name would be a newer synonym of both.
Coprinopsis romagnesiana © Daniel Winkler
Coprinopsis picacea EU - A large, spectacularly mottled inky cap, with large, off white removable scales. Quite famous and common elsewhere in the world, but never reported here until found and sequenced recently in OR.
Coprinopsis picacea © Jonathan Frank
Coprinopsis lagopus EU group - These medium sized grey mushrooms, covered in shaggy white veil material, deliquesce in a matter of hours and are very short lived. They are so abundant, though, that they are still easy to find, even though any individual mushroom may not be around for longer than four hours or so (from sprouting to dissolving).
Coprinopsis lagopus F (#6) - there are seven genetic species that might be C. lagopus (numbered A through G by Wachter whose paper I reference on my Psathyrella page, and numbered 1 through 6 by Nagy). Two recent collections that have been sequenced (from Vancouver BC and Bridle Trails) were Coprinopsis lagopus F Wachter (Nagy's species #6). Perhaps that is our abundant species, but we need a lot more collections to find out which species occur here and their relative commonality.
Coprinopsis lagopides EU - reported from the PNW but we don't have a reliable type area sequence to know what this really is. If you think you find one, save it.
Coprinopsis lagopides var. trisporus WA - Fred Van de Bogart described this from WA but we don't have the type sequence or any other collections to know what it is and how it differs from the type variety. If you think you find one, save it.
Coprinopsis marcida MT - known only from the type sequence from Montana plus one Oregon collection that matches within 4 bp
Coprinopsis brunneistragulata WA/jonesii VT -these two species have basically the same ITS sequence. Various collections of C. jonesii sequence within 0.5%-1% of each other and the type sequence from WA of C. brunneistrangulata is in that clade. However, Wachter did not synonymize them because C. brunneistrangulata is the only species in this group with a perisporium in the spores (and it also was not found in a burned area like C. jonesii usually is). One recent collection in this clade was found in a burn.
Coprinopsis pachyderma WA - we have the type sequence.
Coprinopsis marcida © Bruce Newhouse, Coprinopsis jonesii © Alan Rockefeller
Coprinopsis cinerea EU (=Coprinus fimetaria EU) - Both Coprinopsis cinerea and Coprinopsis fimetarius are reported from the PNW, but they are now thought to be the same thing. C. cinerea is a short lived lookalike of C. lagopus but it grows on compost and dung and other very rich or decayed material. We have no local DNA to prove its presence here yet; we need collections.
Coprinopsis cinerea var. depressa WA - Not yet officially moved to Coprinopsis, this is a locally described (from WA) variety that differs by the shape of the spores. It definitely occurs here, but we don't have a type sequence to know what it is, or if it is just a variety or a valid species of its own.
Coprinopsis cf friesii EU - A small purplish-grey grass coprinoid with floccose veil material on the cap, that unlike the C. lagopus group, tatters more than it deliquesces. We have neither reliable EU sequences nor local sequences to know if that's what we have.
Coprinopsis friesii © Steve Trudell
Coprinopsis subdomestica FL - This mushroom is kind of a cross between Coprinopsis lagopus and Coprinellus flocculosus, with a warm brown and grey cap covered in thick veil material. It was not known from the PNW until sequenced in OR, matching the type sequence.
Coprinopsis subdomestica © Jonathan Frank
Coprinopsis nivea EU - This small to medium (<2.5 cm) sized white inky grows on dung. It has a bit variable ITS DNA (differing by a few positions even in the EU) also found in the PNW.
Coprinopsis nivea © Julie Jones
Coprinopsis cf stercorea EU - This miniscule (>=3 mm) white species may be what we find locally on dung, but we need a local sequence to find out if that's what our species is or if it's something else.
Coprinopsis cf stercorea © Danny Miller
Coprinopsis PNW05 - the first collection were tiny pale grey to pale tan fruitbodies on the cut edge of a trunk in WA. The second collection were larger and found on soil. It could be the same as one of the little known named species whose DNA is not yet known.
Coprinopsis PNW05 © Yi-Min Wang
Coprinopsis tectispora WA - Described from WA from a greenhouse, this smallish (1-2 cm) pale grey species (covered in shaggy veil material) has never been photographed in colour, so we need photos to understand it better. We have the type sequence.
'Coprinus' alnivorus WA (proposed new name: Coprinopsis alnivora) - This is one of the only species that was not recombined correctly by Scott Redhead et. al. when Coprinus was split into Parasola, Coprinellus and Coprinopsis. It has a ring (it is a small, pale species on wood) and that was thought to be an indication of a true Coprinus, but the more important features don't match (it doesn't have a thread running through the stem and it does have pleurocystidia). We have the WA type sequence, but it's never been found since.
Coprinopsis undulata WA - we have the type sequence, but no modern collections nor a photograph. C. undulata is said to be a compost species with thin white veil material on the cap that doesn't come off as easily by itself as some other species.
Coprinopsis sylvicola OR - a forest species said to have red-brown instead of white universal veil material when examined carefully, but perhaps best recognized microscopically. We have the OR ITS2 type sequence and now a recent WA collection with the whole ITS sequence and a photograph.
Coprinopsis sylvicola © Yi-Min Wang
Coprinopsis cf. kubickae EU/cf. phaeospora EU/cf. psychromorbida EU - Small whitish species (<1 cm or even <5 mm) on vegetation, with subtle brownish scales. These species were reported from the PNW, but no collections have been sequenced (or even photographed), so we need some collections to find out if we really do have them here. We have EU sequences of the first, but there are no reliable sequences of the other two yet to compare to.
Species that never deliquesce:
Psathyrella longipes CA -large, but not terribly stocky, with a smooth, solid brown cap and hanging veil remnants when young. Spores 10-13 x 7-10u. The unusually large spores >10u across was the first clue that they belonged in Coprinopsis instead of Psathyrella, where it still is now. We don't have the type sequence, but I have a handful of matching sequences from CA probably representing this species. It has also been sequenced from OR and WA.
Coprinopsis marcescibilis EU (=Psathyrella fragilissima OR, =Psathyrella elwhaensis WA) - with somewhat different spores, 12-15 x 6-8.5u. Psathyrella fragilissima has long been thought to be a newer synonym of C. marcescibilis. There are 3-5 bp differences between them, but even in the EU ITS can vary a bit, and my trees do not show support for separating the species. P. elwhaensis is differentiated by a slightly different shape of spore, but Smith's drawings of the two species' spores looked identical to me. The only other difference is that P. elwhaensis is said to never retain any veil material on the cap disc. P. elwhaensis is said to be much, much more common than P. fragilissima, yet we have many sequences of the supposedly rare P. fragilissima and no different sequences that might represent the supposedly common P. elwahensis. Therefore, I'm pretty sure that P. elwhaensis is not a distinct species.
Psathyrella huronensis MI - this has smaller spores than the others, 8-11 x 5-6u, so you might not recognize it as Coprinopsis (in fact it still needs moving to Coprinopsis from Psathyrella), as the spores barely exceed 10u. In one collection the average spore length was just 10.3u. It has a pale cap even when young (the others will get paler s they dry out). One EU sequence is 4 bp different than our one WA sequence. We still need ENA type area sequences to make sure ours line up.
Coprinopsis uliginicola WY (IDx4) recent: ORx1 - quite stocky, silky white fibrils on the cap. We have the type sequence and two local sequences that are a pretty good match to it except for 3 bp differences near the end, but both modern sequences have the same differences. California has a sister species.
Psathyrella subagraria NY (ORx1) is likely a Coprinopsis, probably another sister species of C. uliginicola, if not the same thing. It is said to have pleurocystidia while C. uliginicola does not.
Coprinopsis canoceps OR (WAx8 ORx9 IDx4) - white cap and stem veil fibrils with a pale brownish grey cap, like many Psathyrellas. The spores are not as large as other Coprinopsis, so it took DNA to show where this belonged. We don't have the OR type sequence, but we have some reliable recent local sequences from WA and CA. Europe may have 2 sister species to our true species.
Coprinopsis uliginicola © Kit Scates Barnhart, Coprinopsis canoceps © Alan Rockefeller
Narcissea - click to expand
Microscopically, these don't quite match Coprinopsis nor Coprinellus, but they look more like Coprinopsis, so that's where they used to be placed. Genetically, they turned out to be very closely related to Coprinellus, which is interesting. That definitely qualifies them for their own genus. The pileipellis is not a cutis, there are some spherical universal veil cells on the cap but there are no pileocystidia. The spores are strongly flattened with a polygonal outline. Sorry, but that's the definition of this genus. Luckily, we don't need to worry about that since we are not known to have any species here.
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