Summary of Interesting Results Here are some of the newest, most interesting results of the study: Pholiota gummosa was found in WA

Pyrrhulomyces - click to expand One brightly coloured pinkish-orange species Species mentioned: Pyrrhulomyces astragalinus. Gymnopilus laeticolor.   Pyrrhulomyces astragalinus EU (Gymnopilus laeticolor WA) - BC, WA, OR and ID sequences all match EU sequences. Pyrrhulomyces astragalinus © Daniel Winkler

Kuehneromyces - click to expand Some of  the only hygrophanous species in this group. Usually clustered and dull brown coloured. Species mentioned: Kuenheromyces mutabilis, lignicola, vernalis. Pholiota atripes, conica, nigripes, obscura.   Kuehneromyces mutabilis EU - this fall species has a ring and scales on the stem. BC sequences are 2 bp and possibly a couple of indels different from an EU sequence, but I am assuming ours is the same species. Kuehneromyces lignicola NY (vernalis) (Pholiota atripes WY, Pholiota conica ID) - this spring species has less of a veil (seldom leaving a ring) and no scales. You will learn to recognize the changing caramel cap colour and distinctive dark stem. A BC sequence matches east coast sequences. The type sequence of P. atripes matches. There is no type sequence of P. conica yet, but studies have shown it to very likely also be the same. Kuehneromyces obscurus ID (Pholiota nigripes ID) - This very similar spring species has broad gills (K. lignicola's are narrow) and this has much larger spores, but as this species has been all but forgotten, it is probably identified as K. lignicola when found.  We should try to find modern collections and sequence them to find out how common this is, and to get a colour photograph (none are known to exist). The type sequences of P. obscura and P. nigripes matched each other so the species will now be known by the epithet that is older by 2 pages, K. obscurus. For P. nigripes, the subtle veil was not detected at all in 5 collections and it was described as a different species on that basis. Kuehneromyes mutabilis © Michael Beug,     K. lignicola © Steve Trudell

Flammula - click to expand A few yellowish, thinly viscid species, possibly hygrophanous and umbonate, that are somewhat tall and slender, but they will have learned to be recognized. Most likely to be confused with Pholiota spumosa, which is more viscid and more likely to have a brown disc. Species mentioned: Flammula alnicola, malicola, malicola var. macropoda, flavida. Pholiota oregonensis, subvelutina.   Flammula alnicola EU (Pholiota subvelutina OR) - Tian and Matheny have formally accepted PNW sequences as official representatives of this EU species, so it does occur here, even though there's quite a lot of genetic variation among sequences, 3-5 bp as well as another half a dozen ambiguous locations. They also synonymize P. subvelutina based on the type sequence. They don't mention Pholiota malicola EU nor Pholiota flavida EU, both often synonymized with this, but no plausible sequences of either are around, and they probably would have mentioned them if there weren't synonyms, so they probably are. I think the older name P. flavida isn't being used over the newer name P. alnicola because P. flavida is ambiguous and so not used anymore. Where does this leave the locally described Pholiota malicola var. macropoda ID? It's probably a synonym too. Smith's Pholiota study found that the PNW's common species were this variety, and occasionally P. flavida. Since sequences are showing that our common species is F. alnicola, that is probably what Smith was finding and calling F. malicola var. macropoda. Flammula abieticola NC (Pholiota flavida var. graveolens ID) - Tian and Matheny also showed that this could be a distinct sister species, only differing by a very few bp in ITS, which wouldn't seem to matter given how F. alnicola varies more than that intra-species, but sequences of it did clade together for them and one study implied they were not biologically compatible with the others. Unfortunately, some local sequences fall in between this clade and F. alnicola, so to me, ITS DNA does not seem to separate these two alleged species. We should sequence more collections in this group to see how they all fall out in the tree. Pholiota oregonensis OR - Smith placed this in the group of that became Flammula, so we need sequences to find out if it's distinct or a duplicate of one of the above. It is said to have more distant gills than the others, so it could very well be distinct. It's even possible it's not actually a Flammula. Flammula alnicola complex member © Steve Trudell

Hemipholiota - click to expand One large white, scaly species (the cap somewhat discolouring to brown). Species mentioned: Hemipholiota populnea, Pholiota destruens.   Hemipholiota populnea (Pholiota destruens) EU - WA and OR sequences only differ in ITS2 by 1 bp and 1 indel or so from a European sequence. It would be nice to get an ITS1 sequence from Europe to make sure our species also matches in that region. Hemipholiota populnea © Michael Beug

Hemistropharia - click to expand One large red, scaly species. Species mentioned: Hemistropharia albocrenulata.   Hemistropharia albocrenulata NY - this east coast species is found across Europe, so although we don't have any local DNA yet, I'd bet it's the real species that's found here. It usually has very dark spores like Stropharia, but is found on wood like Pholiota. Hemistropharia albocrenulata © Buck McAdoo

Pholiota - click to expand Everything else. Species mentioned: Pholiota highlandensis, carbonaria, fulvozonata, molesta, subsaponacea, brunnescens, luteobadia, subangularis, flammans, kauffmaniana, aurivella, adiposa, limonella, abietis, connata, filamentosa, hiemalis, terrestris, gummosa, melliodora, squarrosoides, squarrosa, kodiakensis, flavopallida, lurida, lenta, scamba, spumosa, baptistii, baptistiae, velaglutinosa, rubronigra, humii, decorata, lubrica, lubrica var. luteifolia, rufodisca, avellaneifolia, fulviconica, ferrugineolutescens, alniphila, occidentalis, mixta, agglutinata, fulvodisca, nubigena, abruptibulba, acutoconica, aurantioflava, bakerensis, brunnea, canescens, crassipedes, curcuma, ferruginea, galerinoides, gruberi, iterata, lactea, luteola, macrocystis, milleri, multifolia, olympiana, pallida, paludosella, mutans, polychroa, populicola, proximans, pseudopulchella, pulchella var. pulchella, pulchella var. brevipes, rivulosa, scabella, scamboides, sienna, sipei, subangularis, subechinata, subflavida, subfulva, subgelatinosa, sublubrica, subminor, subochracea, subpapillata, tuberculosa, P. curvipes, umbilicata, velata, verna, vialis, vinaceobrunnea   (Burned ground/wood species) Pholiota highlandensis NY (P. carbonaria EU/CA, P. carbonicola CA, P. fulvozonata ID) - usually slender, stem <4 mm thick (but some are thicker). No recent local sequences yet, but we have all three type sequences and an old, original WA sequence, and they are officially synonymized. Pholiota molesta ID (P. subsaponacea ID) - usually squat, stem >4 mm thick (but some are thinner). We have the type sequences and 4 WA sequences used in the study that synonymized them. Pholiota brunnescens OR (P. luteobadia MI) - the only one where the stem stains brown. All 4 recent PNW burn sequences are this species, so maybe it's our most common species locally. There are a handful of ambiguous locations in some sequences, including the type sequence, as well as some run length differences (indels) but besides that there's very little variation among the sequences. An ecological and morphological study showed them all to be the same species. They are common as a group but only recently did we learn to reliably tell them apart so some are undoubtedly less common than others. Crassisporium funariophilum EU  ('Pholiota' subangularis ID) - is in a different family. It has a dry cap and reddish, thick walled spores. It is usually restricted to the spring whereas the Pholiotas can be found spring or fall. Pholiota highlandensis © Brandon Matheny,     P. molesta © Brandon Matheny,     P. brunnescens © Steve Trudell Here's a (hopefully) interesting story about how the correct name of Pholiota highlandensis/carbonaria/carbonicola/fulvozonata got chosen among the four competing names that turned out to all refer to the same thing. In 1818 Elias Fries, by some accounts only the world's second mycologist, found it, presumably somewhere near Sweden, and named it Agaricus carbonarius, because all gilled mushrooms were called Agaricus at the time and because the epithet means "burn associated". Then in 1872, Charles Peck, America's most prominent early mycologist, found it in Highland Falls, NY, and named it Agaricus highlandensis. Then in 1944, Alexander Smith, one of America's most prominent mycologists at the time, found it in California and called it, well... he accidentally referred to it as Pholiota carbonaria half the time and Pholiota carbonicola the other half of the time in the paragraph he wrote about it. Then in 1951 Singer got around to changing the name of the European Agaricus carbonarius to Pholiota when he realized that nobody had moved that one yet to its proper genus (moving the thousands of Agaricus species to their new proper genus is something that had slowly been going on for many decades already). But he didn't know that Smith had already used the name Pholiota carbonaria, so most people think he erred in reusing the name Pholiota carbonaria and that he should have picked a new name. In 1952, Singer got around to changing Agaricus highlandensis to Pholiota highlandensis, but he didn't provide enough information about what he was doing, so the name change didn't stick until 1990 when somebody made it official. Then in 1962, Singer saw the confusion about the name that Smith had chosen and selected Pholiota carbonicola as the official name of Smith's mushroom to try and avoid the problem that would make his 1951 renaming invalid, since that name was not in conflict with anything else, like Pholiota carbonaria was. Finally, in 1968 Smith found and named Pholiota fulvozonata in Idaho, which turned out to also be the same thing, but that name is clearly not the oldest name among them so it can be discounted. Normally, as Agaricus carbonaria was found and described way sooner than any of the others (1818), you would expect the official name chosen from among all the synonyms would be Pholiota carbonaria, which might be a valid choice if we accept Singer's view that we should consider Smith to have named his 1944 mushroom Pholiota carbonicola and not Pholiota carbonaria. But in "Pyrophilus taxa of Pholiota" in 2018, the authors, who tested the DNA of all of the above as well as microscopy and ecology, and concluded they were all the same, seemed to take the view that the bold type faced name in larger font at the top of the page, Pholiota carbonaria, was the name Smith really meant to use, and that Pholiota carbonicola was a typo, and you can't just pretend that Smith meant to use Pholiota carbonicola instead of Pholiota carbonaria because that would conveniently avoid a duplicate name. So therefore, Pholiota carbonaria only dates back to 1944. Pholiota highlandensis dates back to 1872 through its synonym Agaricus highlandensis. True, Agaricus carbonarius dates back to 1818, but it's not an Agaricus, is it? It's a Pholiota. They declared the official name Pholiota highlandensis. But the story doesn't stop there. Since then, it has been argued that P. carbonaria and P. carbonicola are not two competing names for the same mushroom, but instead simply orthographic variants of each other (minor spelling differences of the same thing). If you accept this view, then Singer was allowed to choose either one of them as the official spelling, it would not matter which name Smith intended: the clarifying author, Singer, gets to decide, and you have to follow his lead. The other name disappears and is no longer around to conflict with anything. So now it is accepted by most that there was no Pholiota carbonaria Smith in existence when Agaricus carbonaria Fries was recombined into Pholiota carbonaria. Now the official name has gone back to being Pholiota carbonaria, and even the authors of the 2018 paper have conceded. I am bucking the trend and have listed Pholiota highlandensis as the official name on this page for now. Now you have a bit of insight into what it's like to be a nomenclaturist (somebody who studies the proper names of things) as opposed to just a taxonomist (someone who studies how organisms are actually related to each other but doesn't necessarily worry too much about what they're called). If you actually did really enjoy this discussion, and you find that you have a strong opinion one way or the other right now, perhaps being a nomenclaturist is for you.   (brightly coloured orange-yellow species, scales not prominently erect and not tightly clustered) Pholiota flammans EU (=Pholiota kauffmaniana WA?) - a striking bright yellow to orange species covered in bright yellow scales. EU sequences differ by 2 bp and 6 somewhat suspicious indels from each other. A WA and OR sequence match each other but are between 2 and 5 bp and a half dozen to a dozen suspicious indels from the EU sequences. I'm not quite sure what to make of that, but for now I'm assuming it's one species worldwide with some variability in ITS, although it could be investigated if the colour forms show any genetic distinction. Pholiota flammans © Steve Trudell and Ben Woo   Pholiota adiposa EU (=Pholiota aurivella EU, =P. limonella NY?) - a similar bright yellow mushroom with brown scales that remind me of onion skin. One old pre-genetics study suggested that what we have locally is usually the east coast P. limonella (90% of the time) with most of the rest being P. aurivella and rarely, P. adiposa. However, worldwide sequences vary in a bunch of ambiguous locations as well as other places, with no rhyme or reason to which of the 3 names are attached to the sequence, so they may all be the same thing. The three BC, WA and OR sequences we have differ from each other in 4-5 ambiguous locations and 3-4 other places that don't seem ambiguous yet, definitely within the noise of the worldwide sequences, so my guess is ours (and Europe's) most common species is just one species, Pholiota adiposa. It is now commonly accepted that P. aurivella is the same thing. We need more collections from back east and here to determine if P. limonella is the same thing too, and if not, if we have both here. Pholiota abietis WA/P. connata MI/P. filamentosa EU/P. hiemalis ID - other supposed group members that have been reported from the PNW without any genetic information known. We need more collections and more studies to figure out the exact identity of the species found here, but it's reasonable to assume we have more than one species here since the spore sizes of some are reported to be quite different. Pholiota cf aurivella © Michael Beug     (Unusual for Pholiota, mostly found on the ground) Pholiota terrestris OR - found in clusters on the ground, often prominent erect fibrillose scales on a brown cap and stem. Hard to recognize as a Pholiota because the wood it is associated with is always buried. We have a number of PNW sequences of this native which all match, but as it is a species with variable colours and scaliness, this could be investigated to make sure it's all the same species. Pholiota gummosa EU (P. melliodora OR) - this is a sister species of Pholiota terrestris and also often found in clusters on the ground, but it is pale yellow-brown and the scales might tend to wash off more easily. The type sequence of P. melliodora differs from reliable EU sequences mostly in ambiguous locations, and is probably a synonym. We have 2 recent WA collections as well that were pale yellow-brown, one with a greenish tint. Smith placed P. gummosa, P. melliodora and P. terrestris in three different subgenera. Pholiota terrestris © Leah Bendlin,     Pholiota gummosa © Paul Dawson     (Prominently erect scales, tight clusters) Pholiota squarrosoides NY - clustered on wood, with prominent erect scales. We have a half dozen east coast sequences, and a few local sequences that match, one of ours is 4 bp different. This species is recognized by a viscid cap and sweet odor. Pholiota cf squarrosa EU/kodiakensis AK - Very similar, yet not closely related, is P. squarrosa with a dry cap, garlic odor, and yellow young gills that turn greenish in age. We have over a hundred EU sequences, so we know what this is, but that sequence is not known from WNA (although a similar sequence has been found in ENA). What are our local sightings of P. squarrosa? One possibility is Pholiota kodiakensis, a similar sister species from Alaska with a short type sequence and a matching recent sequence. This appears to be the same as the newer EU species Pholiota lundbergii. We need local collections to find out what species we actually have. Pholiota cf squarrosa and cf squarrosoides (unsequenced) © Danny Miller     (pale, almost white cap colours, perhaps small) Pholiota lenta EU - an all white Pholiota with some scales on the cap and especially the stem. We have reliable EU sequences and one short but matching sequence of an original Smith collection from 1946 from OR. This is an especially pale member of the large Pholiota lubrica clade, detailed below, and easier to recognize than most of them. Pholiota scamba EU - one of our smallest Pholiotas, pale capped and scaly mostly on the stem. Confusable with other LBMs like Galerina. One WA sequences matches a bunch of reliable EU sequences. This species is on a branch of its own.   probable P. lenta © Andrew Parker,     P. scamba © Daniel Winkler     (bright yellowish-brown 'spumosa' cap colour - indistinctly scaly) Pholiota spumosa EU (P. baptistii/baptistiae ID, P. subflavida WA?, P. vialis WA?) - yellowish-orange-brown cap usually with a darker plain brown disc, usually indistinctly scaly. Similar colours to Flammula, above, but a different stature. Sequences from all over the PNW match many EU sequences quite well. The Tian/Matheny paper sequenced the local type of P. baptistii (formerly incorrectly spelled P. baptistiae) and found it is a synonym. The colour difference between this and members of the Pholiota lubrica group, below, can be subtle, but this species has larger spores than any of those. P. subflavida is thought by some to be the same thing. P. vialis was even thought by Smith himself to perhaps be a duplicate of this too. This should be confirmed. probable P. spumosa © Steve Trudell     (cap colour not bright yellowish like P. spumosa - the large P. lubrica clade) Pholiota lenta EU - described above, is a scaly, almost white member of this clade. Pholiota flavopallida ID (=P. lurida MI) - pale whitish rimmed Pholiotas often with a yellow-brown disc and no scales anywhere. The rim seems to be more well defined than in some of the below species and the colouring seems somewhat irregular (although a two-toned cap is quite common). We have the type sequence of P. lurida, and a couple sequences that Tian identified as P. flavopallida from China, but that is far away from Idaho so the identifications might not have been correct. However, that does show that sequences of collections matching the microscopic characters of P. flavopallida (pleurocystidia sometimes have thickened walls) match sequences of P. lurida (without thickened walls) and that is a very slight microscopic difference, the kind which usually doesn't have any genetic significance. Therefore, I think they probably are synonyms, with P. flavopallida being the older name. Pholiota flavopallida © Yi-Min Wang   Pholiota velaglutinosa OR (=P. rubronigra CA?) - the type sequences match, and they are described as differing by one having a glutinous partial veil, although that seems like a significant thing to be different or be overlooked if they are the same species. Also, P. rubronigra was supposed to be fragrant. Smith also found P. rubronigra in Idaho, so if they are not the same, we might have both species here, but it's going to be hard to recognize. It's more common in the south of our region and California.   probable Pholiota velaglutinosa © Dimitar Bojantchev   Pholiota humii ID - one of many species that has a dark orange-brown disc and a paler margin (but the disc may not as defined as in P. flavopallida above and the colours seem more regular). It has no scales anywhere and is is differentiated from similar species below by supposedly subtly different shades of brown, but may need microscopy for confirmation. Collections of this species have been long going by the name P. decorata (described next) because the guide books have been confused about whether or not that species has scales (it does). The type sequence of P. humii is dirty in one spot, but we have other sequences to fill in what the missing part is (that match exactly everywhere else). Pholiota humii © Yi-Min Wang   Pholiota decorata WA - has scales on both the cap and stem that may disappear in age, a less fragrant odor, and pleurocystidia with thinner walls. We don't have the type sequence, but we do an original Smith collection and other trustworthy sequences. Its sister species is the less common and paler Pholiota lenta, above. P. decorata © NAMA and the Field Museum of Natural History   Pholiota lubrica EU (lubrica var. luteifolia ID? P. rufodisca ID?) - we have some reliable EU sequences of P. lubrica and a paratype sequence of P. rufodisca from NM is very close, differing in at least 3 locations (one of them a run) from the closest EU sequences (and in a couple extra places from another EU sequence). We don't have the ID holotype sequence to know that the paratype matches it. If the species are the same, P. lubrica is by far the older name. We have no data for whether or not P. lubrica var. luteifolia from ID and WA is the same species or not. P. lubrica is said to have a somewhat scaly stem and white veil and gills when young. P. lubrica var. luteifolia and P. rufodisca are said to have yellow veil and gills when young, but differ in the fact that the latter has pleurocystidia with thickened walls (and we've already seen that does not necessarily indicate a separate genetic species). Pholiota avellaneifolia ID - similar somewhat scaly stem, with a yellow veil but avellaneous (smoky) gills when young, this does seem to be a separate genetic species. We have the type sequence and a good match with a recent WA collection, differing in only 2 locations near the beginning. Until this collection this species was little known and in these photos the veil is still intact so I believe it is a young specimen showing the young smoky gills even before the spores mature. Pholiota avellaneifolia © Leah Bendlin   Pholiota fulviconica FL (=P. ferrugineolutescens CA?) - the holotype sequence of P. ferrugineolutescens is the same as an isotype sequence from Idaho of P. fulviconica, but we don't have the holotype sequence from Florida, so it's possible they aren't the same species. But assuming they are, the CA name is a newer synonym. However, they are not described as being the same, with P. fulviconica being described as almost identical to P. spumosa and P. ferrugineolutescens being described as very similar to P. lubrica but with a stem that stains yellow. This is suspicious. I don't know that either have ever been photographed in colour.   Pholiota alniphila OR (Pholiota occidentalis OR?) - these are differentiated from P. decorata etc. and P. lubrica etc. above by yet more subtly different shades of brown. We have a paratype sequence from ID of P. occidentalis that I assume is correctly identified, and we have sequences of modern collections from OR and BC. Some authors synonymize it with the older Pholiota alniphila, if that's not true, then we have 2 species here. Conceivably even three, since there is also a Pholiota occidentalis var. luteifolia that we don't have sequences of to know if it is genetically different.   Pholiota mixta EU (P. agglutinata ID, P. fulvodisca ID) - Another species with a red-brown disc and pale rim, somewhat indistinctly scaly. The stem base stains brown in age. Both local type sequences turned out to be the same as the EU species P. mixta, and they have been officially synonymized. They seem to only have ever been separated by subtly different shades of brown on the disc. Pholiota aff mixta - one OR sequence differs by 3% from all the others and could be a unique sister species, it seems (at least in our only photo) with more pale colours showing than for P. mixta itself. Pholiota aff mixta © Jonathan Frank     Pholiota nubigena CA - a gastroid Pholiota from CA. We don't have the type sequence, but we have a reliable CA sequence of this distinctive mushroom also reported from the drier areas of the PNW. This appears to be in the large Pholiota lubrica clade with the above species. probable Pholiota nubigena © Steve Trudell   Little known species In order to learn more about these, we'll need either a type sequence or sequences of new collections that key out to one of these species in Smith's Pholiota monograph so we can get an idea of what these species might be. It's possible that some may not even belong in Pholiota. Here is a copy of Smith's monograph you can use to ID your unusual Pholiota collections that don't exactly match one of the above (and I didn't give very good descriptions, so I admit it's going to be difficult to feel confident of a perfect match). Pholiota abruptibulba OR - with an abrupt stem bulb and other microscopic differences, said to be related to P. occidentalis, P. verna and P. fulvodisca. Pholiota acutoconica OR - a very conical mushrooms said to be related to P. fulviconica and P. lenta. Pholiota aurantioflava ID - described as somewhat like P. gummosa. Pholiota bakerensis WA - described as somewhat like P. lubrica. Pholiota brunnea ID - Pholiota canescens OR - this is similar to Tubaria confragosa and might be a Tubaria. The real T. confragosa probably does not occur here, and our species needs a new name, but Tubaria canescens is already taken so we cannot use this epithet for our local species. Pholiota crassipedes WA - Pholiota curcuma SC - Smith reported this from ID too, and it might be related to Phaeomarasmius. Pholiota ferruginea WA - said to be somewhat like P. lubrica. Pholiota galerinoides WA - this could be related to Kuehneromyces. Pholiota gruberi ID - Pholiota iterata OR - Pholiota lactea WA - possibly related to Phaeomarasmius. Pholiota luteola ID - Pholiota macrocystis ID - Pholiota milleri ID - Pholiota multifolia MO - reported by Smith from WA Pholiota olympiana WA - Pholiota pallida ID - probably a Kuehneromyces Pholiota paludosella OH - reported by the Key Council from the PNW Pholiota mutans MI - Smith wasn't clear if this was present in the PNW or not. It might be a Meottomyces, perhaps the same as Pholiota pattersoniae CA. Pholiota polychroa OH - reported once from BC Pholiota populicola WA - probably a Kuehneromyces Pholiota proximans MI - reported by Smith from ID, found recently in WA. Now officially Phaeomarasmius proximans, see that page. Pholiota pseudopulchella OR - Pholiota pulchella var. pulchella WA and var. brevipes ID - Pholiota rivulosa ID - Pholiota scabella OR - Tian and Matheny sequenced the type and it is now Stropharia scabella Pholiota scamboides ID - Pholiota sienna OR - Pholiota sipei OR - Pholiota subangularis ID - now officially Crassisporium funariophyllum Pholiota subechinata WA - possibly a Phaeomarasmius Pholiota subfulva NY - reported by Smith from ID Pholiota subgelatinosa OR - Pholiota sublubrica ID - said to resemble P. lubrica (photo in MM) Pholiota subminor OR - Pholiota subnigra WA - now known to be the local name for at least one of our species in the Cyclocybe erebia group. Pholiota subochracea WA - Tian and Mathaney sequenced an old OR Smith collection (but not the type) and discovered it belongs in Hypholoma, where it was originally placed before Smith changed his mind and moved it to Pholiota. Pholiota subpapillata ID - may belong in Keuhneromyces Pholiota tuberculosa EU (P. curvipes EU) - not found by Smith, but reported in the PNW since then, assumed to be the same species as P. curvipes. Probably not a Pholiota, one possibility is Simocybe, another is Phaeomarasmius. Pholiota umbilicata ID - Pholiota velata ID - Pholiota verna WA - said to resemble P. decorata (photo in MM) Pholiota vinaceobrunnea ID - also said to be close to P. decorata.     Also, Stropharia scabella OR

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