Hygrophorineae (waxy caps)

Warning: BLAST will not be able to make any kind of a useful tree out of ITS sequence data for this family! (It usually doesn't, but it really doesn't, in this case). You will need to use better alignment software, like those found in MEGA. Perhaps in part because of the age of this lineage (or perhaps their DNA mutates faster) there is a greater genetic distance between species than normal, meaning that BLAST's primitive alignment algorithm is only useful for determining if two collections are the same species or not.

Waxy Caps (members of the Hygrophoraceae family) are many people's favourites, as they are often not only beautifully coloured but with a special look about them that can make them look like fake mushrooms made of wax, especially in the gills. The basidia (where the spores are grown) can be especially long in this family, perhaps five times longer than the spores that grow on them, which is what gives the gills their thick, waxy appearance. Even the white and brown mushrooms in this clade can have especially brilliant shades of white and brown. Not every species is obviously waxy, sometimes the effect is more subtle and has to be learned to be recognized. It may be most obvious in the gills.

They all have white spores and the gills are usually thick and widely spaced, as well as being waxy. Viscid caps and stems are common, as are decurrent gills (but none of that is always true). Rarely, there is a partial veil. Lookalikes: other colourful genera in this family that aren't quite as waxy, colourful Mycenas that are smaller, fragile and delicate, and Laccaria that also look waxy, but they are always orange or purple with tough, fibrous stems.

Not every genus in this family looks waxy. This page only covers the ones that do, and they don't all clade together. Cuphophyllus, Hygrophorus and Hygrocybe s.l. either evolved their waxiness independently in 3 or more separate events, or the other genera lost their waxiness in multiple events.

Hesler & Smith (from now on referred to as H&S) is the most important book on this family in North America, North American Species of Hygrophorus from 1963.

abundant common uncommon rare - colour codes match my Pictorial Key and are my opinions and probably reflect my bias of living in W WA. Rare species may be locally common in certain places at certain times.

Summary of Interesting Results - click to expand

Here are some of the newest, most interesting results of the study:

Cuphophyllus - click to expand

Formerly Camarophyllus, this genus is defined by interwoven gill hyphae, but without a microscope, you may recognize them by dry stems, pastel colours and decurrent gills that are widely spaced apart. Some caps are <5cm across, but others can grow up to 10cm. Other genera in the family may also have these characters, so it's best to browse this section first before moving on to find your mushroom elsewhere. Note that this genus of waxy caps is separate from the other two clades.

Species mentioned: Cuphophyllus borealis, virgineus, niveus, russocoriaceus, lacmus, subviolaceus, rainierensis, cinereus, cinerellus, colemannianus, recurvatus, subradiatus, pratensis. Hygrophorus burgdorfensis, lawrencei, nordmanensis, graveolens, subaustraligus. Camarophyllus cremicolor.

Cuphophyllus 'borealis PNW09' - A small and white, dry or slightly viscid capped waxy with no odor. Several white species of Hygrophorus are very, very similar but are either quite viscid everywhere, not as striate, have a pruinose stem apex, or some colour to them on the disc and a non-mothball odor. It's unclear if we have more than one species and if the proper name is C. borealis or not. The east coast, where the species was described (NY specifically), has at least 3 genetic species, and ours matches a common one, so it could possibly be the real thing. There are about a dozen species worldwide known so far, and that's with limited sampling. Four or so species in Europe could be the "real" C. virgineus (probably not found here, despite reports to the contrary). The European name C. niveus is available for one of the species in Europe, since the genetic diversity we've found in this group shows that it may not be the same thing as C. virgineus as currently assumed. Since other locations have more than one species, it is to be expected that once more sequencing is done here we'll find ours is a species group. We could use more collections.

'Hygrophorus' burgdorfensis ID - this small, yellow capped, viscid 'borealis' like species is only known from the type in Idaho. No DNA yet to know if it is a unique species or how it fits in. Its official name is still Hygrophorus burgdorfensis, but it most likely is a Cuphophyllus. It is possible, if the yellowish cap does not necessarily indicate a separate species, that this name could end up being used for (one of) our common white local species. We need the type sequence.

Cuphophyllus PNW08 - our one WA collection had yellow tones in the in situ shot. It should be investigated if this could be H. burgdorfensis.

Cuphophyllus 'borealis PNW09' © Steve Trudell, Cuphophyllus PNW08 © Yi-Min Wang (3 images)

Cuphophyllus russocoriaceus UK

'Hygrophorus' lawrencei OR - Two almost identical small and white waxy caps, that smells strongly of cedar! It's an amazing odor to experience. C. lawrencei is a little stockier, with the cap usually between 1 and 3 cm, and the stem more than 0.5cm thick. C. russocoriaceus' cap is reported as being between 0.5cm and 2.5cm and the stem usually 0.25cm thick or so.

They are not sister species, but rather the cedar smelling species are sprinkled in the tree amongst the dozen C. virgineus/borealis species around the world discussed above, which is interesting. C. russocoriaceus is a UK species and DNA from the UK and from WA differ by only a couple of nucleotides and indels. C. lawrencei is described from Oregon, and while we have no sequences from Oregon, we have California and Washington sequences that differ by 2bp and a handful of indels that probably represent this species. It was thought that all of our local PNW specimens were likely to be C. lawrencei before C. russocoriaceus DNA was found in Washington, so perhaps C. lawrencei is less rare here. (Technically, Hygrophorus lawrencei is the current name as it has not been recombined into Cuphophyllus yet, but will be).

Cuphophyllus russocoriaceus © Steve Ness, Cuphophyllus lawrencei © Alan Rockefeller

Cuphophyllus PNW01 (=Hygrophorus subaustraligus CA?) - this white species species, found once in WA and twice in CA, appears to be odorless and hygrophanous with some stem nattering. It may have a yellow disc. This could be Hygrophorus subaustraligus CA, and if so, that species probably needs to be moved to Cuphophyllus.

Cuphophyllus PNW01 © Lauren Ré (3 images) and Yi-Min Wang

Cuphophyllus lacmus EU - Our purple-brown cap and gilled mushroom with a viscid cap, white stem and no odor has an unpleasant taste that can be either bitter, hot or nauseating. We appear to have this European species here, according to Voitk's 2020 Cuphophyllus paper, which shows that there is more genetic variation than usual in this species. We do not appear to have the eastern North American C. subviolaceus, which Voitk shows is a sister species that is about 3% different, and not necessarily a synonym as is currently thought. The supposed yellow stem base of C. lacmus does not appear to be a reliable way to distinguish the two, as our local material does not have it.

Cuphophyllus rainierensis WA and 'Hygrophorus' nordmanensis ID - are said to be similar mushrooms that taste mild and smell of green corn, differing from each other by spore size. However, it does not appear that they are distinct. For almost 50 years, they were each known from only one single collection, separated by spore size, even though H&S admitted that C. lacmus had quite a variable spore size, depending on if the basidia held 2 spores each or 4 spores (2 spored versions have larger spores since fewer need to fit on the basidia). Recently, a collection of C. rainierensis from Mt. Rainier (the type area) that smelled of green corn was sequenced and found to be almost exactly the same as some EU sequences of C. lacmus. Then, a collection of C. nordmanensis with the large spores from near the ID type area also sequenced to be almost exactly the same as the other two. So there is no need to rename Hygrophorus nordmanensis to Cuphophyllus nordmanensis, since it is probably not a distinct species.

Cuphophyllus cinerellus EU - we recently found a sequence in WA matching pretty closely to the type sequence, and pretty much exactly to a sequence quoted in the paper that designated the type. It is found in mossy bogs, and is not supposed to have an odor (but may). It might be differentiated from the above species by a dry cap that is slightly scaly with subtler purple tones (mostly smoky brownish grey with a hint of purple).

Cuphophyllus cinereus EU - a European species described as similar to C. lacmus but with dry cap, a mild taste and no odor. It was reported once from Mt. Rainier, but given that the special odor of C. rainierensis does not necessarily indicate a different species, and that viscidity has already been shown to not necessarily indicate a different species, I hesitate to believe that this one collection from well over 50 years ago actually represents a unique species found in the PNW. It is probably one of the above two species, perhaps the dry capped C. cinerellus. We have no DNA from anywhere yet.

Cuphophyllus lacmus (as rainierensis) © Noah Siegel, Cuphophyllus cinerellus © Connor Dooley

Cuphophyllus cf colemannianus EU - a beech/oak species with a tan cap and a thick, white stem. There are 2 or 3 possible groups of sequences representing that species in Europe, and we're not sure which is the real one. California has a species matching that description, and it's been reported up here too. We need local samples to see if it might match any European sequences, and a better concept of which European mushroom is the real thing.

Cuphophyllus recurvatus NY - A somewhat brown capped mushroom with white gills and a more slender, slightly coloured stem growing in grass. It is similar to Gliophorus unguinosus (but dry capped with more decurrent gills). Pale capped and dark olive-brown capped collections from WA have the same sequence. We need sequences from eastern North America (where it was described) to make sure ours is really the same species. One east coast collection misidentified as C. colemanianus differs by 4 bp, 1 indel and 3 ambiguous nucleotides from our C. recurvatus sequences. Plausibly, that collectionis C. recurvatus showing that our sequences are very similar and may represent the real thing. It should also be compared to the very similar, European 'Hygrophorus' subradiatus, which some have wondered might be the same species, but nobody is really sure what that species is and rather than worry about it longer, we may just give up on using that name so it won't matter.

unsequenced Cuphophyllus colemannianus © Christian Schwarz, Cuphophyllus recurvatus © Sadie Hickey and Yi-Min Wang

Cuphophyllus pratensis EU -
Cuphophyllus pratensis PNW04 -
Cuphophyllus pratensis PNW05 -
Cuphophyllus pratensis PNW06 -
Cuphophyllus pratensis PNW07 -

Possibly five largish species, pale orange, dry capped (else pale). C. pratensis and PNW04 seem the most highly pigmented. PNW04 and PNW06 seem more slender stemmed, like H. graveolens, described next. PNW06 has a stem bulb. They can be confused with Hygrophorus pudorinus, which has a viscid cap and more closely spaced gills. Most EU sequences by far are one particular species, which is probably the real one. We have that species here as well as four others, between 4% and 8% different from each other. Normally that would imply five distinct species are here, but as ITS DNA varies considerably in this family, it's also possible that some lumping will occur.

'Hygrophorus' graveolens OR - A similar mushroom described from Oregon with a paler cap, more slender stem and a sweet odor. It's official name is still Hygrophorus graveolens as it has yet to be formally renamed. It is a sister species to the entire complex of Cuphophyllus pratensis group members.

'Camarophyllus' cremicolor WA (still called Camarophyllus cremicolor because nobody has renamed this one to Cuphophyllus yet either) may be the same thing. It was described from Washington and said to be smaller, with an even paler cap, yellower gills, and perhaps but not always a slight fragrant odor. We need DNA to determine if this is different than C. graveolens. If they are the same, this is the older name that would have priority, although exceptions can be made if the older name describes an unusual or aberrant form of the mushroom.

Cuphophyllus pratensis © Lauren Ré (2 images), C. 'pratensis PNW04' (from AZ) © Mike Dechter, C. 'pratensis PNW04' (from MI) © MO user Huafang, C. 'pratensis PNW05' © Danny Miller

Cuphophyllus 'pratensis PNW06' © Bruce Newhouse (2 images), C. 'pratensis PNW07' © Eric Chandler, C. graveolens © Jordan Gates (2 images)

Cuphophyllus 'aurantius PNW02' - yet another amazing find by the South Sound Club of a species nobody has ever reported from the PNW before. This completely orange Cuphophyllus is not quite as bright as similar Hygrocbyes, and has a fairly uniform colour, completely dry cap and decurrent gills. It is about 7% different than the Jamaican species Cuphophyllus aurantius, there seem to be several distinct similar species around the world, including a different one in the Rockies. This species may not have the interwoven gill hyphae typically found in Cuphophyllus.

Cuphophyllus 'aurantius PNW02' © Sadie Hickey (2 images) and Warren Cardimona

Hygrophorus - click to expand

All of these species have divergent gill hyphae. Most have a viscid cap unless noted. A viscid stem will be noted. The stems are often less colourful than the caps, but if they have a colourful stem they will usually be larger than a Hygrocybe s.l. Some have a partial veil (that never happens in the other two groups). Note that this genus is a separate clade from Cuphophyllus and all the others.

Species mentioned: Hygrophorus caeruleus, subalpinus, albiflavus, sordidus, penarius, siccipes, hypothejus, boyeri, purpurascens, velatus, olivaceoalbus, olivaceoalbus var. gracilis, fuscoalboides, whitei, fuscoalbus, inocybiformis, agathosmus, agathosmoides, albofloccosus, odoratus, bakerensis, variicolor, secretanii, monticola, vinicolor, erubescens, erubescens var. gracilis, neoerubescens, persicolor, capreolarius, amarus, russula, parvirussula, piceae, eburneus, cossus, chrysaspis, glutinosus, gliocyclus, flavodiscus, melizeus forma minor, sordidus, penarius, chrysodon, ellenae, pusillus, subpungens, goetzei, goetzii, saxatilis, karstenii, karstenii forma minor, pudorinus, fragrans, speciosus, speciosus var. kauffmanii, pacificus, vernalis, discoideus, occidentalis, tephroleucus, pustulatus, pustulatoides, morrisii, marzuolus, camarophyllus, calophyllus, megasporus, avellaneifolius. Clitocybe odora.

Hygrophorus caeruleus - Clitocybe odora - click to expand

This blue waxy cap turns out to not be a waxy cap after all, but an aberrant Clitocybe odora.

Species mentioned: Hygrophorus caeruleus. Clitocybe odora.

Fibrillose partial veil - click to expand

These species have an actual fibrillose partial veil leaving fibrillose strands around the stem almost like a ring.

Species mentioned: Hygrophorus subalpinus, albiflavus, sordidus, penarius, siccipes, hypothejus, boyeri, purpurascens, velatus, olivaceoalbus, olivaceoalbus var. gracilis, fuscoalboides, whitei, fuscoalbus, inocybiformis

These are not all related to each other. It seems several groups have independently evolved a fibrillose veil. We used to think that evolving a veil was an unusual event that wouldn't be that common, and even placed more importance on this than the ecology of growing on ground or on wood. It turns out we had that backwards.

Hygrophorus subalpinus WA - White and stocky, this is a veiled conifer spring snowmelt species (found near melting snow at high elevations), but unfortunately it can also be found in the fall, making it hard to distinguish from PNW03, next.

Hygrophorus 'subalpinus PNW03' - a second species may be fall only, from WA, northern Alberta, and boreal Quebec, which has 10% different DNA, but cannot yet be told apart morphologically. The first known photo below from WA looked more slender than the first species, but a subsequent find in Alberta was quite stocky just like H. subalpinus.

Hygrophorus albiflavus OR - a slender, white waxy with a fibrillose veil. Viscid stipe, and larger spores than H. subalpinus. It is only known from the OR type collected on Sept. 29th, 1946. Unknown trees. No sequence yet. We need a type sequence or modern collections. It is just possible that this could be the name of PNW03. We should check for a viscid stipe on PNW03 and measure those spores.

Reports of Hygrophorus sordidus/penarius from the PNW may represent finds in the Hygrophorus subalpinus group if the fleeting veil is not noticed (but they have much smaller spores than H. subalpinus). Another theory is that they are Hygrophorus 'marzuolus PNW10', below, which has spores the size of H. sordidus, but not H. penarius. So what are collections with the spore size of H. penarius? Any reported finds should be sequenced.

Hygrophorus subalpinus © Jonathan Frank, H. 'subalpinus PNW03' © Danny Miller

Hygrophorus siccipes OR - A distinctive orange-brown and/or olive-brown veiled supposedly stocky (but sometimes slender) conifer common waxy that may prefer 2-3 needle pine. This is our local species that has been going by the name H. hypothejus. H&S found one example of the common H. 'hypothejus' without a viscid stem and described it as H. siccipes from Oregon, putting a lot of weight on the viscidity of the stem as an indicator of not only species, but what would now be subgenus. It turns out that whether viscid stemmed or not, they are the same species, and ours is not the same as the European H. hypothejus, so now we have a proper name for our species. Ironically, most of them are viscid stemmed but now the name of our species means "dry stemmed".

This species has, like perhaps some other Hygrophoraceae will turn out to have, a lot of genetic variation (~2% or so), but as no ecological, morphologic or microscopic differences have been found, it is still officially all one species in the paper on this section. I have found 3 clades of species, as much as 5% apart, but if you treat an indel of a large chunk as one difference, the amount of difference between the clades falls to about 2%. If all sequences end up falling into one of the 3 clades, and not in between, it should be investigated if in fact this is more than one species.

Hygrophorus boyeri PQ - A species from Quebec is very similar and also with pines, but more consistently slender and with duller colours that tend to fade. The study describing it only found it in CA and boreal BC. We expected to find it here, in between, as H&S reported both robust and slender forms from the PNW, and we finally found a WA collection maching the type.

Hygrophorus siccipes © Michael Beug, Hygrophorus boyeri © Michael Beug

Hygrophorus 'purpurascens PNW04' Fall - Unevenly purple (starting out pale and staining/turning purplish pink) and somewhat stocky and large, the gills and other parts spot and stain even more purplish over time. It is a spruce and pine mushroom, notably found in fall. It may be a denser species than our spring species, PNW05. Several similar species are purplish and/or spot and stain purple (described in the next section), but this one has a fibrillose partial veil. So far, DNA from BC, WA, and twice from OR all match each other, but it doesn't match DNA from Europe, where it was described, so our species needs a new name.

Hygrophorus 'purpurascens PNW05' Spring - this snowbank species looks the same and is also stocky but may be less dense. It differs in ITS by >10%. It has been sequenced twice in OR.

Hygrophorus velatus ID - this supposedly rare mushroom is only known from the type in Idaho, collected in August, which was never sequenced. It was a veiled, slender specimen that was pale pink and did not appear to further spot or stain. Some think it was a young H. 'purpurascens PNW04' that hadn't stained yet. We need specimens to find out.

Hygrophorus 'purpurascens PNW04' Fall © Kathleen Dobson and Jonathan Frank, H. 'purpurascens PNW05' Spring © Jonathan Frank

This next species group is recognized by a coloured fibrillose partial veil, and grey to olive-grey colours.

Hygrophorus olivaceoalbus EU - the name we give to most collections in this group, but actually rare here, so far only sequenced locally from the wet, west coast spruce forests of Vancouver Island. No good way to recognize this species yet, except it may be the only one obligated with spruce. It is said to have a black fibrillose veil and a viscid stem, but modern studies note that the amount of viscidity is not a good indicator of species in this group, unlike what is found in the H&S key.

Hygrophorus fuscoalboides ID - our least uncommon species, found with conifers, said to have a grey fibrillose veil and dry stem, but we now know the stem viscidity is not important. That leaves very few known differences so far. We have the ID type sequence and sequences from BC and WA that match. This has been synonymized with Hygrophorus olivaceoalbus var. intermedius from CO.

Hygrophorus whitei CA - described by H&S as a yellowish clone of H. eburneus, the type sequence showed this name belongs to an H. olivaceoalbus clone instead. It is said to be slightly more slender than H. fuscoalboides. We have the CA type sequence and one matching WA sequence.

H. olivaceoalbus var gracilis EU, rumoured from the PNW under hardwoods, is slender. It has now been synonymized with Hygrophorus mesotephrus, which they have not discovered in North America yet, so I do not believe the rumours that it is found here. If you think you find it, save it.

Hygrophorus fuscoalbus EU is a rumoured species that reportedly has a white fibrillose veil instead of black. Nobody is sure what this species is, so the name may not be used going forward. A recent study supposes that H&S' report of this in the PNW probably represented Hygrophorus agathosmoides, so it is no longer believed to be found here.

Hygrophorus fuscoalboides © Richard Morrison, Hygrophorus whitei © Danny Miller

Hygrophorus inocybiformis ID - This similar grey, scaly, veiled waxy with spruce and fir is dry everywhere (nowhere viscid). It is common in Idaho where it was described, but rare elsewhere. EU sequences are about 6bp different than our one WA/ID border sequence, so they investigate if they have the real thing or not.

Hygrophorus inocybiformis © Andrew Parker, unsequenced H. inocybiformis © Ben Woo

Sweet Almond/Cherry Odor - click to expand

These have a sweet almond/cherry odor (specifically, not just a sweet or fragrant odor).

Species mentioned: Hygrophorus agathosmus, agathosmoides, albofloccosus, odoratus, bakerensis, variicolor, amarus, secretanii, monticola, vinicolor, pacificus

Hygrophorus agathosmoides PQ - Uniformly grey cap and white to pale grey stem. Spruce species. Sequenced from BC, WA and OR.

Hygrophorus albofloccosus CA - very similar Douglas fir species, but with stem fibers that stay white and don't turn brown (thus the name), a perhaps slightly more robust fruitbody (but not true in the examples shown) and slightly shorter spores than H. agathosmoides. Also sequenced from BC, WA and OR. It can easily be confused with another conifer species with much larger spores, H. odoratus, below.

Both these species have mistakenly been called Hygrophorus agathosmus, but that only seems to occur in Europe (and has similar DNA to H. agathosmoides, only differing by 1-2 bp and 5-7 indels). They both have "regular" sized spores compared to the larger spores of H. odoratus.

Hygrophorus agathosmoides © Sadie Hickey, H. albofloccosus © Danny Miller

Hygrophorus odoratus OR - This is a very similar, also uniformly greyish capped mushroom with conifers, said to be either more slender, or more yellowish than the Hygrocybe agathosmus group, and it has much larger spores. It was described as being more uncommon than the species above, but they are easily confused, so I'm assuming the spores were checked before they made that statement. We have the type sequence from Oregon and the same DNA from Washington. These species appear to be related to H. pustulatus/tephroleucus, below.

Hygrophorus odoratus © Kathleen Dobson, NAMA and the Field Museum of Natural History

Hygrophorus bakerensis WA - this conifer waxy has a brown disc with a paler rim. We have an isotype sequence, and many other sequences from WA and BC. It has "regular" sized spores. Conifers.

Hygrophorus variicolor OR - a rare (except in California) waxy, this is almost identical except reportedly with a viscid lower stem, said to be often overlooked, and with similar spores. This makes one wonder if this is the same species as H. bakerensis. California specimens (where H. bakerensis is supposedly rare) and/or the type need to be examined to see if the viscid stem indicates a separate species. Most people think they are the same. This is the older name, but H. bakerensis is the name commonly used.

Hygrophorus amarus WA - this is probably another synonym. It also has a darker disc and a paler margin, but was described from an odd collection with slightly different colouration, gills that appeared to spot pink, no odor and a very bitter taste. There were only 3 collections ever made, and the type collection has almost the same sequence as H. bakerensis. Perhaps the other two are something different, but even if so, it wouldn't matter. This name is in between age of the other two. Even though H. bakerensis is the newest name, its popular usage up here means it will continue to be the name used, at least for a while.

Hygrophorus bakerensis (variicolor?) © Steve Trudell

Hygrophorus secretanii EU (=monticola ID) - A more evenly brownish ("pale alutaceous") cap flushed vinaceous, with conifers. The type of H. monticola is from Idaho and it is relatively common there, but rare elsewhere. The older European species H. secretanii seems to match our type so that older name may prevail. It has larger spores than H. bakerensis.

Hygrophorus vinicolor ID - very similar (with the same large spores) but supposedly tastes bad and has somewhat pink gills and a pink pruinose stem. It is rare, only reported by H&S twice from Idaho, including the type, and never since, and may represent an aberrant H. secretanii/monticola. Specimens are needed.

Hygrophorus pacificus CA - similar mushroom but said to have a different non-almond sweet odor and is described below. We have no sequences yet.

Hygrophorus secretanii (monticola) © Andrew Parker

Pink/Purple/Orange stocky species - click to expand

Some of the purplish-pink ones become more coloured in age because the gills or other areas spot irregularly. The colour may be mostly in the gills. This section does not include some slender species that are mostly white with hints of pink or orange.

Species mentioned: Hygrophorus purpurascens, erubescens, erubescens var. gracilis, neoerubescens, persicolor, pudorinus, fragrans, capreolarius, russula, parvirussula, saxatilis, karstenii, speciosus, speciosus var. kauffmanii

Hygrophorus 'purpurascens' EU group has already been discussed in the veiled section above.

Hygrophorus neoerubescens EU (H. erubescens EU misapplied) - starts out whitish and develops pinkish/purple splotches, found under spruce and pine. It may have yellow tones. This used to be called H. erubescens but the 200 year old drawings were sometimes of a Rhodocollybia, so that name can't be used anymore. We have a sequence from northern BC, but no more local collections yet. We need some to prove it's here too.

Hygrophorus 'erubescens var gracilis' OR - is a locally described variety, but the type sequence shows it needs to be raised to species level. It is rarer, more slender, has more distant gills and usually lacks the yellow tones. We have no modern collections nor photographs of this species yet (and need some).

Hygrophorus 'parvirussula PNW11' (H. russula EU misapplied) - This is a similar hardwood species always lacking yellow tones and supposedly with slightly closer gills than H. neoerubescens and H. capreolarius (described next). Two WA sequences under oak are closer to the chinese species H. parvirussula than to the EU species H. russula. They look 4% different, mainly in ITS2, but I can explain all the differences by 13 indels, which is not a lot of difference for this family. Perhaps this is H. parvirussula. H&S reported H. russula only once in WA, and there have been a few reports since then.

Hygrophorus 'capreolarius PNW06' - This spruce species has more evenly distributed purple colours than the former two (less splotchy, so to speak) and also lacks yellow tones. One WA specimen matched DNA of a Quebec specimen and was >6% different than a European sequence, so we appear to have a sister species in North America that needs a new name.

unsequenced Hygrophorus cf neoeerubescens © Buck McAdoo, Hygrophorus 'parvirussula PNW11' © Michael Beug, Hygrophorus 'capreolarius PNW06' © Danny Miller

Hygrophorus pudorinus EU (=Hygrophorus persicolor EU) - starts out more uniformly pinkish/purple, and fades (kind of the opposite of H. neoerubescens and others above). It may also have yellow tones after fading. It is very closely related to H. neoerubescens. A recent study showed that H. pudorinus and H. persicolor are synonyms and provided sequences that match the type, which also match some local sequences.

Hygrophorus pudorinus © Mary McCallum (2 images)

Hygrophorus fragrans OR (=H. pudorinus var. fragrans) - with a pale orange (perhaps pinkish) viscid cap, punctate upper stem, and a sweet odor. It has more closely spaced gills than the dry capped Cuphophyllus pratensis, above. It is said to bruise yellow-orange and have slightly coloured gills and a stronger odor compared to H. pudorinus. We have the type sequence and other matching local sequences. Some sequences are 8 bp and 2 indels different than others, but a recent study showed no ecological nor morphological differences, so this is a species that can vary somewhat in ITS.

Hygrophorus fragrans © Danny Miller (2 images) and the Field Museum of Natural History

Hygrophorus karstenii EU (=Hygrophorus saxatilis OR) - White cap, bright pink to peachy gills and a peachy odor. H. saxatilis was described from Oregon in 1942 with conifers, on the basis of it having gills more towards pink than the yellowish gills of H. karstenii, but EU sequences of Hygrophorus karstenii described from the EU in 1887 are practically identical, so it looks like our name is a newer synonym. We have local sequences from WA, Victoria and boreal BC. The Victoria sequence is the longest and cleanest, most of the rest are shorter and somewhat dirty, but in the clean areas they all match almost precisely. Most sources still list H. karstenii as a newer synonym of Cuphophyllus pratensis, but that appears to be a propagation of an older mistake, and the DNA is certainly not bearing that out.

Hygrophorus karstenii forma minor EU - known from one single collection from WA near Mt. Rainier decades ago, this is probably the same thing. H&S separated it from H. saxatilis by a thinner gelatinous cuticle on the cap, lacking the odor and having pale yellow gills instead of pink, but all these traits are known to be variable and they themselves weren't sure it deserved to be considered a separate species. They are identical microscopically. We sequenced one collection with orange gills (photo below) and it has the same sequence as the usual pink gilled collections from the rest of the world, so with the known colour variation of the gills, I'm convinced that all our collections are properly called H. karstenii.

Hygrophorus ellenae ID - A cespitose species, with white gills that slowly turn peachy. It is also only known from a single type collection and was separated from H. saxatilis on the basis that that species should have coloured gills even when young, but it seems likely that this too may be the same thing as the above two species. Pine and fir.

Hygrophorus karstenii (sequenced and unsequenced) © Eric Chandler and Michael Beug

Hygrophorus speciosus NY - Bright red-orange cap, but white gills and stem. Very slimy everywhere and a fleeting slime partial veil. It favours larch, but common in those habitats.

The type of H. speciosus var kauffmanii MI (said to be stockier) is 3 bp and 1 indel from the consensus DNA of the type variety. Our local WA DNA is 1 bp from var kauffmanii and 2 bp and 1 indel from var speciosus. For now I am going to assume it is all one species. Both varieties are described from eastern NA, but only collections visually matching the type variety have been formally reported from the PNW.

Hygrophorus speciosus © Steve Trudell

Mostly white species - click to expand

The caps may have yellow, orange or pink tones, but the stocky ones with pink and orange tones are covered above. Caps with a significant amount of brown, grey or black are covered next.

Species mentioned: Hygrophorus piceae, eburneus, cossus, chrysaspis, glutinosus, gliocyclus, flavodiscus, melizeus forma minor, chrysodon, ellenae, pusillus, subpungens, goetzei, goetzii, vernalis.

Hygrophorus subalpinus group - is described above in the "Fibrillose Partial Veil" section.

Hygrophorus 'marzuolus PNW10' - a stocky spring species that is mostly white with a grey streaks on the cap and with a dry stem. This is described below in the grey capped section.

Hygrophorus sordidus DC/penarius EU - white species, are also discussed below with Hygrophorus 'marzuolus PNW10'.

Hygrophorus piceae EU group (PNW07 and PNW08) - White, viscid cap but dry stem, very similar to Cuphophyllus 'borealis', but not striate and the upper stem is very pruinose or fibrillose. PNW07 is usually slender. PNW08 may be somewhat stockier. It is common with spruce, as its name suggests. There are at least 4 competing concepts in Europe for which species is the real thing, so we don't know for sure, but PNW07 is the most common EU species, so perhaps that is the real thing. This is thought of as a somewhat slender mushroom, so that makes sense.

What's interesting is that two species have been found in WA, and although they look identical, they do not appear to be as closely genetically related as is usually the case with species groups. There could be two completely distinct species that evolved to look the same as each other, although in different parts of the Hygrophorus tree.

Hygrophorus piceae PNW07 and PNW08 © Danny Miller, NAMA and the Field Museum of Natural History

Hygrophorus eburneus group

As a group, these species are white, have very viscid, slimy stems, and sometimes have yellow shades. They are not all closely related. Here are the species that have been reported from here:

Hygrophorus eburneus EU - this white, mid-sized, mostly conifer species is a European mushroom and so far our sequences of this species differ by >5%.

Hygrophorus cossus EU - very similar and probably related, from Europe. It is aromatic with a cap and stem aging pinkish buff, reported only once from Idaho.

Hygrophorus chrysaspis EU - from Europe, gills browning in in age, is only rumoured from the PNW, so unless it can actually be found, this is a dubious report.

Hygrophorus glutinosos NY - from eastern North America, one possible record from Oregon, with cap drying yellowish with red punctate dots on stem after drying. No sequences from anywhere on the planet of anything resembling this are available yet.

Hygrophorus gliocyclus EU - with spruce/pine, a stocky mushroom with yellow tones in the cap.

Hygrophorus flavodiscus ENA - midsized, yellow disc, with pine, also rarely reported in Oregon and Idaho. Sister species to H. gliocyclus. We have ENA DNA but need local specimens to compare.

Here are the species that have been sequenced from here:

Hygrophorus 'eburneus PNW01' - none of the above have been confirmed from the PNW through DNA. Our collections from WA, OR and CA that resemble H. eburneus differ by 5% from EU sequences, so it is probably an undescribed species. DNA of a second species is only known from an environmental sample in OR, so I will ignore it until it is found to fruit.

Hygrophorus gliocyclus PNW12 - this sister species (7 bp different) of the EU H. gliocyclus, found in BC and OR, resembled H. gliocyclus in the BC collection and is yellowish in most collections, but had grey caps in one OR collection (this would make it hard to ID). H&S did say that our collections seemed a little different than in Europe, but grey was never reported.

Hygrophorus 'eburneus PNW01' © Danny Miller, H. 'gliocyclus PNW12' © Mitch Milgram (yellowish) and Jonathan Frank (greyish), unsequenced H. cf flavodiscus © Noah Siegel

Hygrophorus 'chrysodon PNW09' - This golden flecked, viscid stemmed white waxy is easily recognized when the flecks have not rubbed off. It is probably not related to anything in the supposed H. eburneus group, although it is mostly white with bits of yellow and a viscid stem. It is found with conifers, but Washington DNA is clearly a different species than the EU, so ours needs a new name.

Hygrophorus 'chrysodon CA01' - a second species in the group, found in WA, OR, and CA. A third species has been found in CA. Although sometimes stocky, it can be slender like PNW09.

Hygrophorus 'chrysodon PNW09' © Danny Miller, and NAMA and the Field Museum of Natural History, H. 'chrysodon CA01' © Jonathan Frank (2 images)

Hygrophorus pusillus ID - a small, aromatic, white waxy with a viscid cap and a pink disc when fresh. With conifers. The odor is probably the best way to recognize it as the pink tones is not always obvious (only visible in the 2nd photo). Our local DNA seems to match the LSU region of a reliable eastern North American sequence, so this Idaho species may occur there too.

Hygrophorus subpungens OR - a small, aromatic white species that ages avellaneous in the cap and clay colour in the gills. Dry cap and stem. It may look and smell like a duller/darker H. pusillus. Known only from the Oregon type, no sequence yet.

Hygrophorus goetzei OR (=H. goetzii) - A small, pale pink snowbank, species with large spores like H. vernalis and sister to it, also with hemlock. Described from Oregon. We have one OR collection whose sequence probably represents this.

Hygrophorus 'vernalis PNW14' - a sister species to H. vernalis (which is found under the brown capped section) but with caps more pinkish, but not quite as pink as H. goetzei. It also seems to lack the dark brown disc of H. vernalis. Known from one OR collection.

Brown/Grey/Black on cap - click to expand

Brown, grey or black somewhere on the Cap (at least in age on the disc).

Species mentioned: Hygrophorus pacificus, vernalis, discoideus, occidentalis, tephroleucus, pustulatus, pustulatoides, morrisii, marzuolus, sordidus, penarius, camarophyllus, calophyllus, megasporus, avellaneifolius

Hygrophorus pacificus CA - Much like the much more common H. bakerensis and H. secretanii/monticola, but a different odor (strong and penetratingly aromatic, but not specifically of sweet almonds). It has more distant, yellowish gills and a more rusty-brown cap. Found with spruce and with larger spores like H. monticola (not like H. bakerensis). It is not uncommon in California but rare here (reported once each from WA and ID). We don't have any DNA yet.

Hygrophorus vernalis WA - A brown capped spring snowmelt species with a viscid stem (similar to the more aromatic H. secretanii/monticola which also has large spores and lacks the viscid stem), under hemlock. It also resembles H. goetzii, which is almost identical but has a pale pink cap. This WA collection is the first DNA documentation of this WA species. BC DNA matches it fairly closely (2 bp and 0-5 indels).

Hygrophorus 'vernalis PNW14' - (also described under the mostly white capped section) is a sister species but with caps more pinkish, but not quite as pink as H. goetzei. It also seems to lack the dark brown disc of H. vernalis but may have a uniformly pinkish brown cap. Known from one OR collection.

Hygrophorus 'discoideus PNW13' - A slender species with a bright, brown disc that is usually quite offset from the pale rim, viscid all over like H. occidentalis, described next. Under spruce? Reported from CA, OR and ID. Our one WA sequence is 5 bp and 4 indels different than most EU sequences, so our species may need a new name. (H&S noted our specimens have shorter spores than in Europe).

Hygrophorus occidentalis CA - this California species, found not uncommonly there and rarely in Oregon (under oak and pine) is also somewhat slender and has a dull grey-brown disc with a somewhat paler rim and is viscid all over, like a browner H. eburneus. The Tennessee isotype shows that it is related to H. eburneus, but now I need local DNA to see if theirs is a sister species back east or the real thing. There are unfortunately no known colour photographs.

Hygrophorus pustulatoides PQ - Small, greyish cap, sometimes pointed, with notable dark scabers on the somewhat viscid stem. Our species was recently described under spruce from Quebec (and also sequenced from BC, WA and OR). Until now we have been assuming our species was the European Hygrophorus pustulatus, but now we know that doesn't seem to occur here, and our species differs slightly by usually not having a scaly disc and differs by up to 6 bp and 6 indels in ITS. It was also formally declared that the EU Hygrophorus tephroleucus is a synonym of H. pustulatus, (formerly differentiated by being more slender).

Hygrophorus morrisii MA - similar dry stemmed species with brown to grey-brown cap and stem without scabers. It is an east coast conifer species rarely reported in the west (ID + CA). We don't have sequences from here to see if we really do have this species, but we do have a reliable sequence from Quebec to match it to.

Hygrophorus 'marzuolus PNW10' - Stocky spring species that is mostly white with grey streaks on the cap and with a dry stem Unknown trees. Our species is a sister species to the true European species, at an 83% match.

Hygrophorus sordidus DC/penarius EU - Some reports of Hygrophorus sordidus, an all dirty-white species from eastern North America may represent this species, as may some reports of the very similar European Hygrophorus penarius, with folks not realizing how pale capped H. 'marzuolus PNW10' can be (just a little grey on the margins, as shown). There is one single report of H. penarius from WA in 1915 that may be real, but it may be pretty rare. Any reported finds of either species should be sequenced to verify which species occur here.

Hygrophorus camarophyllus EU - a fall species with blackish caps even when young, also stocky. Slightly viscid, streaky cap. Under spruce and pine. There's not total agreement on what the European species DNA is, but we match many European sequences and could have the actual species.

Hygrophorus cf calophyllus EU - A similar blackish capped European mushroom, but with a very viscid, more uniformly coloured cap. It may have pink gills. Under fir and pine. There's no DNA from Europe yet, but one EWA sequence may represent this species. We should also sequence a pink gilled collection to see if it really is the same species.

Hygrophorus camarophyllus © Matthew Koons, H. calophyllus © Andrew Parker, unsequenced H. calophyllus (pink gills) © Danny Miller

Hygrophorus megasporus WA - A conifer species with an olive-grey brown disc with a paler margin and a viscid stem, described from Washington (like H. olivaceoalbus, but no fibrillose veil). The gills appear more distant than described. It is practically unknown, but with giant spores (even bigger than the H. olivaceoalbus grp). We don't have the WA type sequence, but we have a sequence from an old Smith collection from OR that probably represents this species. Recently, a matching OR collection was found and it was photographed in colour for the first time.

Hygrophorus avellaneifolius ID - A relatively slender stem, dark rusty brown to grey cap and darker avellaneous gills and a dry stem. The type is from Idaho where it is said to be not uncommon, but only found there. We have no sequences yet to really understand what it is. We need collections. Found with conifers.

Hygrocybe s.l. have parallel gill hyphae, and both the smaller and the larger species have bright colours on the stems. They never have a partial veil. It has been decided to split this genus up into a number of other genera (including Chromosera, Gliophorus, Gloioxanthomyces and Humidicutis as well as Hygrocybe), although they are all closely related so you could argue that the split didn't need to happen. This becomes especially true because the individual genera are hard to characterize and recognize, which is less than ideal for a genus. This is the third and largest clade of waxy caps.

Note that Chrysomphalina is in this clade, but as they are semi-waxy and not typically thought of as waxy caps (and grow on wood, which is unusual for mushrooms on this page except for Chromosera) I treat them on the non-waxy Hygrophorineae page.

Hygrocybe s.l. - Chromosera - click to expand
Chromosera are viscid all over with decurrent gills. To distinguish between similar Gliophorus, Gloioxanthomyces and Hygrocybe you are looking for gills at least as brightly coloured as the stem and cap, only yellow or lilac colouration, and a stem that is not long and thin. One species grows on wood (the only species on this page that does).
Species mentioned: Chromosera cyanophylla, citrinopallida. Mycena lilacifolia.

Chromosera 'cyanophylla CA01' - growing on wood. It is viscid/slimy all over, has decurrent gills and is purple when young, fading to yellowish. The purple persists longest in the gills. Our species is 5% distinct in ITS from this European species. On the East coast, the type area for Mycena lilacifolia, DNA is also distinct, so while they will be able to use a name like Chromosera lilacifolia for their species, we are getting a brand new name for ours.

Chromosera citrinopallida WA - this species is also viscid all over with decurrent gills, but grows on the ground. It is bright yellow fading on the cap and stem to white. The irregular yellow colouring helps differentiate it from similar yellow species of Gloioxanthomyces, Gliophorus and Hygrocybe. It is also really small, usually coming in at <1cm wide. It is similar to the California species 'Gliophorus' flavifolius below, but that species has brighter, adnate gills, an even whiter stem, and the cap is white in the center and yellow on the outside instead of irregularly yellow-white. This is described from high elevation WA but we have low elevation WA sequences that also match. Greenland and the EU seem to have a sister species, 2% different.

Chromosera 'cyanophylla CA01' © Steve Trudell, C. citrinopallida © Connor Dooley, Matthew Koons and Kendra Dedinsky

Hygrocybe s.l. - Gloioxanthomyces - click to expand
We have one rare species in this genus here, and it is viscid and bright yellow everywhere, including in the gills, with an indented cap, decurrent gills and a very long, thin stem.
Species mentioned: Gloioxanthomyces nitidus

Gloioxanthomyces cf. nitidus SC - described from eastern North America and we still need local DNA to confirm that it is the same species over here.

probable Gloioxanthomyces nitidus © Andrew Parker

Hygrocybe s.l. - Humidicutis - click to expand
A rare, entirely orange (with olive colours when fresh) species with a dry cap. The gills are especially bright orange and stay orange, at least on the margins, long after the colour has faded elsewhere.
Species mentioned: Humidicutis marginata var. marginata and var. olivacea

Humidicutis marginata var. olivacea WA - this beautiful variety was described from Washington and has olive colours as well as orange colours as well as bright orange marginate gills. We have a few WA sequences.

Humidicutis marginata var. marginata NY - this entirely orange east coast type variety is rumoured from the PNW but we need specimens to confirm that. Perhaps people are finding var. olivacea where the green has faded and are mistaking it for this.

Humidicutis marginata var olivacea © Steve Ness

Hygrocybe s.l. - Gliophorus s.l. - click to expand

Gliophorus species are slimy-viscid all over, sometimes with decurrent gills. Similar mushrooms are found in Chromosera, Gloioxanthomyces and Hygrocybe, but those are usually not quite as slimy-viscid. Usually. Gliophorus are never red like Hygrocybe can be (they probably lack the ability to produce betalain pigments). A famous green mushroom is found here. Gliophorus does not appear to be monophyletic, so perhaps some of the three groups will be split into additional genera.

Species mentioned: Gliophorus psittacinus, laetus, sciophanus, irrigatus, unguinosus. Hygrophorus subaromaticus. Hygrocybe hondurensis, flavifolia.

Gliophorus psittacinus EU group - sometimes with green (or even blue) but sometimes pure orange (sometimes both!) and they can even change back and forth between those colours. The green is a brighter green than the other famous green mushroom, the redwood yellow-green Hygrocybe virescens, which has a completely dry cap. The gills are supposedly not decurrent and the cap is often pointy, but since we have several species in this group, we have yet to show that is always true. So far we have DNA for 5 PNW species (none of them the real thing) and we don't even know for sure if they are a closely related group or scattered throughout the genus.

PNW01 seems to have a dark orange phase not found in the others species (see photos) and is seldom observed green. It's the first clue we have about how to tell them apart. PNW02 appears to be the most common and can be blue, although blue is the rarest colour reported. PNW04 has not been photographed yet (well, it has, but as part of a mixed collection where we don't know which one is PNW04). There is also PNW03. PNW08 is from peaty soil under sedges/spiraea in a wet area on edge of poor fen. It is very small, translucent, with yellow-green tones.

Gliphorus psittacanus var. californicus CA is an available name for one of our species, assuming that California species is not endemic to California only. I just don't know which one it might be. Gliophorus perplexus MI from back east has had its type sequenced, and it only occurs back east and does not appear to be one of our species.

Gliophorus laetus EU group - these are different shades of orange, never have a green stage, and they usually smell strongly fishy. The gills should always be decurrent and the cap should not be pointy. So far we have 3 PNW species: PNW05 and PNW06, and a sister species of Gliophorus sciophanus, which seems to belong to this group even though G. sciophanus is supposedly a synonym of G. psittacina (it isn't). One WA collection in this group looked intermediate between G. laeta and Hygrocybe hondurensis, but doubtless we do not have H. hondurensis here and it makes perfect sense that our undescribed local species would not fit into any one description.

Gliophorus 'laetus PNW05' - most collections are bright orange.

Gliophorus 'laetus PNW06' - one collection was more dully coloured, but another looked the same as the others.

Gliophorus 'sciophanus PNW07' - usually red-orange, although one outlying collection was yellowish. In one of the first 3 collections the fish odor was detected.

Gliophorus cf irrigatus EU - dark grey-brown cap and stem, similar colours to Cuphophyllus recurvatus, above (a dry mushroom), with gills that are not always decurrent. In California they have named one local group member Hygrophorus subaromaticus which probably needs to be recombined as Gliophorus subaromaticus. We need samples of that and our local species to compare to the EU DNA that I do have to see what our species are. The also European G. unguinosus is thought to be a synonym for G. irrigatus.

Gliophorus 'irrigatus PNW09' - a greyish-tan cap and stemmed species that is closest to G. irrigatus. The more they dry out, the more silver-metallic coloured they get. It is not what they are calling Hygrophorus subaromaticus in CA, that is white capped and not yet known from the PNW.

unsequenced Gliophorus irrigatus group member © Noah Siegel, 'Gliophorus' (Hygrophorus) 'irrigatus PNW09' © Yi-Min Wang (3 images)

'Gliophorus' flavifolius CA - a California species that has been found in OR and WA (but not yet sequenced from there). Very white stem, very bright yellow gills, and a white cap disc surrounded by yellow. Adnate gills. It is stockier than Chromosera citrinopallida, above. It is very viscid on the cap and stem but was overlooked for transferring to Gliophorus, where it belongs. It is currently still going by the name Hygrocybe flavifolia.

Hygrocybe s.s. - click to expand
The rest of the species are still properly in Hygrocybe itself.
Species mentioned: Hygrocybe virescens, miniata, constans, squamulosa, substrangulata, phaeococcinea, reidii, cantharellus, turunda, turunda var. sphagnophila, coccineocrenata, singeri, conica, californica, persistens, cuspidata, acutoconica, acutoconica var. microspora, flavescens, chlorophana, ceracea, subceracea, coccinea, marchii, aurantiosplendens, laetissima, punicea, mucronella, reae, parvula, minutula, subminiata, fenestrata, glutinipes, constrictospora, americana, luteo-omphaloides, atro-olivacea. Gliophorus minutulus, fenestratus. Omphalina occidentalis. Hygrophorus albicarneus.

Dry caps - click to expand
First, we discuss the dry capped species, some of which can even be scaly. Most of them are red, fading to orange and yellow, but first, one rare green species. None are conical or umbonate. Some species in the viscid capped section, next, may not have viscid caps so check there too.
Species mentioned: Hygrocybe virescens, miniata, constans, squamulosa, substrangulata, phaeococcinea, reidii, cantharellus, turunda, turunda var. sphagnophila, coccineocrenata

Hygrocybe virescens CA - Described from California, this can be found here rarely under introduced redwood trees (and red cedar, which makes me wonder why it is so rare). We have a sequence from California found under bay laurel, which matches a WA sequence under red cedar quite well. It is not viscid (unlike the slimy green Gliophorus psittacinus group), but it is related to the viscid Hygrocybe flavescens group. It is a beautiful, unique, lime-green to greenish-yellow colour.

Hygrocybe virescens © Steve Trudell

Hygrocybe constans OR (H. 'miniata' misapplied) - This species is reddish (but as so many others do, may fade to orange or yellow). It is found in soil, moss or rotting debris in forests. It is small (~2.5cm across) with a dry cap that has small scales on it and gills that are not usually decurrent (but might be)? We have many local sequences, most very good matches for each other. There are a some alleles that can make them look different, so the few differences between them might be accounted for by other undiscovered alleles. Or perhaps a multi-gene study will show there is more than one species not distinguishable by ITS. Ours is not the true H. miniata of Europe. H. constans is an Oregon species from 1912 given its own name because Murrill thought the red colouring didn't fade to orange or yellow (the red was more constant), than usually happens with this species group. It has long been thought to be the same as H. miniata, since the amount of fading can be quite variable, but now that name is a good candidate for the proper name of our local species, although it has yet to be proven.

H. squamulosa NJ - may not be here, although it was reported one single time from WA by H&S, and has been described as "impossible to separate" from H. miniata by Stuntz. The only east coast sequence purporting to be that species was unlike any of our local DNA, and it wasn't much different from DNA of regular east coast H. 'miniata', so it remains to be seen if H. squamulosa is a true distinct species or not, and if so, if it actually occurs here too. If you think you find it here, save it.

Hygrocybe constans © Steve Ness and unusual yellow collection © Connor Dooley.

Hygrocybe substrangulata UK - This is very similar and related to H. constans, but usually found in wet moss and described as more scurfy than scaly on the cap, slightly decurrent gills and with slightly larger spores. It is rarer, but how rare, we're not sure. The presence of this UK species in the PNW was never suspected until the South Sound Mushroom Club DNA project found it. Our local collections are about 3 bp and 1 indel different than a UK reference sequence. They are also brighter red instead of orange and maybe a little more slender than in the UK. Is that enough to make ours a distinct species?

Hygrocybe substrangulata © Connor Dooley

Hygrocybe 'phaeococcinea PNW02' - Another similar and related red species with a smooth, dry cap, hints of orange-yellow on the cap rim and stem, decurrent gills and usually found in grass. It is probably rare, but might just be overlooked.

Since our species is about 4% different from European sequences, we really don't know if ours is the same or how closely it will match the description. In the EU, it is also known from dunes and lake sediments and it is said to develop a blackish cap covering in age and turn dark brown on drying, whereas the samples collected here turned darker on drying but retained the red colours. Perhaps that is one sign that our species is indeed different. The presence of this species here was one of the many discoveries by the South Sound Mushroom Club.

Hygrocybe 'phaeococcinea PNW02' © Steve Ness, Eric Chandler and Connor Dooley

Hygrocybe reidii EU - This is not necessarily related to the previous species in the H. miniata group. It is small, pretty consistently bright orange with a dry, essentially smooth cap, adnate gills and often with an odor of honey gone bad. It is also a newly discovered European species by the South Sound club, and may or may not actually be rare. It was found here in the forest, although in the EU it is reported from grasslands.

Hygrocybe reidii © Steve Ness

Hygrocybe cantharellus NC (two forms) - This species has a rough cap, but a much longer stem and more strongly decurrent gills than the scurfy capped H. constans and the others. The cap often is bright (reddish to orange, once pure yellow!) and may or may not have black scales on the cap. Until now those with black scales were considered a separate species, but now we see the ITS DNA is the same for both of them (except for a possible couple of ambiguous locations). This species, like many others, can start out all red but parts can fade to yellow or orange (which has happened to the stem in the first photo). If your collection has duller colours and black scales, you probably have H. cf turunda, described below. If it has brighter colours (although the difference might be subtle), with or without black scales, it might H. cantharellus but it also might be Hygrocybe cf cantharellus #2 (described next).

Hygrocybe glacialis EU - one WA sequence of something that looked like H. cantharellus turned out to match Swiss type area collections of Hygrocybe glacialis made by the person who described the species. One was almost pure red with a stem as brightly coloured as the cap, but the other was orange. The scales are the same colour as the cap. Other identical sequences were labeled H. turunda by others, which tells me it may sometimes have black scales. How do you tell H. glacialis apart from H. cantharellus? The author suggests that the cystidia on the stem are shorter in H. glacialis. Hopefully there is a better way to tell them apart. Hygrocybe coccineocrenata UK with scales that only slightly darken, has been reported from the PNW. We need to find out what reports of that species really are, but we don't know for sure what that species is.

Hygrocybe 'turunda PNW03' - this is what we had been calling all the collections with black scales on the cap. Red-orange species with black scales seem to be a colour form of Hygrocybe cantharellus, but yellow-orange-brown collections sometimes sequence to be a unique species, 5% different in ITS than both H. cantharellus and the EU species H. turunda. This species may be more likely to be found in wet areas than the other two. It is probably undescribed. However, the colouration does not seem to be a fool proof way to distinguish the two black scaled species. The photos show this and H. cantharellus can look very similar. Hygrocybe turunda var sphagnophila is an east coast name that could be considered for our collections, but given that different geographic areas often have unique genetic species in this group, we'll need east coast sequences of that to find out. Some say it's the same as Hygrocybe coccineocrenata.

Hygrocybe cantharellus (red and yellow without black scales, orange with black scales) © Steve Ness, Steve Ness and Yi-Min Wang,     H. glacialis © Steve Ness and Jordan Gates

Hygrocybe 'turunda PNW03' © Yi-Min Wang, Steve Ness, and Warren Cardimona

Hygrocybe PNW16 - somewhat closely related to H. phaeococcinea in the H. miniata group, it appears to have started out red and faded to orange and yellow. Based on its close genetic relationship to other dry capped species, I'm going to guess that it has a dry cap. More specimens are needed to see if this is a legitimate species and if so, how to recognize it. It has only been found once near Mt. Rainier, WA.

Hygrocybe PNW16 © NAMA and the Field Museum of Natural History

Viscid caps - click to expand
Most species discussed next will now be viscid, at least on the cap.
Species mentioned: Hygrocybe singeri, conica, californica, persistens, cuspidata, acutoconica, acutoconica var. microspora, flavescens, chlorophana, ceracea, subceracea, coccinea, marchii, aurantiosplendens, laetissima, punicea, mucronella, reae, parvula, fenestrata, glutinipes, constrictospora. Gliophorus fenestratus.

I cover them in 4 groupings:

very conical species

yellow/orange species without red

stocky red species (>=5cm across, the stems are a little thicker than the slender species)

slender red species, <5cm across

1. CONICAL SPECIES

Hygrocybe singeri OR group - The very famous witch's cap mushrooms are abundant and identified by red, orange or yellow mushrooms that turn pitch black in age. They have pointy caps and are viscid, usually all over. We have H. singeri (described from OR) and potentially four more cryptic species in the group so far. Almost every new sequence has yielded a new potential species, except for #2. (#1 found once from WA, #2 found seven times, #3 once from WA, and #4 once from OR). However, the "species" are only about 3-4% apart in ITS from each other, so barring the discovery of morphologic or ecologic differences, I reserve the right to consider Hygrocybe singeri just one species with unusually variable ITS, especially since California and Quebec have their own "species" 3-4% different as well. I sincerely hope these are all not 7 different species. We do have a varietal name we can raise to species if necessary, Hygrocybe singeri var. albifolia. Traditionally they've been separated by the viscidity of the stipe and colour of the young gills, but that may not hold.

Hygrocybe 'conica PNW22' - We have not yet found Hygrocybe conica EU here (that also appears to be a complex, and is so far known from Europe, northern BC and California), but we did find a sister species in BC. It has an orange-red somewhat metallic cap, is somewhat umbonate but not as conical, and the cap and stem are not really viscid, at least not as much as in H. singeri.

Hygrocybe 'conica PNW25' - one WA collection is a unique species in this complex, not in the H. singeri clade, but in the greater conica/singeri clade. There appears to be debris stuck to the cap and not the stem so I am going to surmise that this too is a species with a viscid cap but not a viscid stem.

Hygrocybe aff singeri #1 © Steve Ness,     H. aff singeri #2 © Sharon Squazzo,     H. aff singeri #3 © Eric Chandler,     H. aff singeri #4 © Connor Dooley.     H. singeri not photographed.

Hygrocybe 'conica PNW22' © Sarandeep Khella,     H. 'conica PNW25' © Steve Ness

Hygrocybe acutoconica group (CA01 and PNW05) - Hygrocybe acutoconica is a rarely reported pointy capped orange-yellow species from NE back east, like H. singeri but without red tones and not turning black significantly. We have two genetic species in this group, but I don't know what to call them yet. CA01 is more common and from WA and CA, and PNW05 is from OR and CA (so far). PNW05 has been found pure yellow in CA and one collection had a somewhat blackening stem.

H. acutoconica NE - might be one of our two species, but I don't have reliable east coast sequences of that to know what it is.

H. persistens EU - probably not one of them. It is considered the same thing as H. acutoconica, but probably isn't, as the EU seems to have a whole group of genetically distinct species, with none of those sequences matching ours.

H. cuspidata ON - might be one of them. It is an east coast species with a red cap. We don't have sequences of it to know what it is, but we need red capped local collections to sequence to see if they are different than the two genetic species we already know about.

H. californica CA - most likely to be one of them, since it is locally described from the west coast. It is currently thought to be a synonym of H. acutoconica but probably isn't. The spore size of #1 didn't seem to match, so perhaps this is our #2.

Hygrophorus albinellus CA - might be one of them. This is a long forgotten west coast name, we need sequences to see what it is.

Hygrocybe 'acutoconica CA01' © Sadie Hickey,     H. 'acutoconica PNW05' (one showing a blackening stem and one pure yellow) © Jonathan Frank and Josh Walker

2. YELLOW-ORANGE SPECIES (usually never red)
Hygrocybe 'flavescens PNW06' - This mushroom is yellow (or sometimes orange), with adnexed gills and has a sticky cap and maybe stem too. H. chlorophana is a European species, H. flavescens is an east coast species, and ours is neither but closer to eastern NA material (but still >7% different).

Hygrocybe 'flavescens PNW06' © Steve Trudell

Hygrocbye ceracea EU - This similar mushroom is golden yellow everywhere with adnate to decurrent gills and is slightly sticky everywhere when wet. Some collections are dark orange and almost red, making them easily confused with many other species. Other collections are bright yellow, making them easily confused with H. 'flavescens PNW06'. Our DNA differs by 2-3 substitutions in ITS1 and a couple to few indels in ITS2 from most European sequences but is likely the same species. (A few EU sequences are a cryptic second species but I'm assuming the more common sequence is the real thing). One WA sequence stands a bit apart from the other PNW sequences and the type area sequences.

Hygrocybe subceracea FL - from Florida, has a dry stem and different sized spores. A sequence of a mushroom that seemed to meet that description from Quebec is close to but distinct from sequences of H. ceracea. It has been reported locally, but we need specimens to examine to prove whether or not it is here. Considering that some EU literature lists H. ceracea with a dry stem and spore size in between North American reports of H. ceracea and H. subceracea, it's not clear that H. subceracea will really be confirmed from the PNW. We need collections.

Also see Hygrocybe 'parvula PNW20' ("red chunk"), below in section 4, for a species that sometimes is red but often is found only with orange/yellow tones. This species is one theory for what reports of Hygrocybe subceracea actually are.

Hygrocybe ceracea © Steve Ness (2 images) and Bryce Edward Englehardt

Hygrocybe 'ceracea PNW07' - Fairly closely related to H. ceracea, (about 11 bp and 8 indels different) but more towards orange instead of golden yellow, if the two WA collections are any indication. For this family, that's not necessarily enough genetic variation to indicate a separate species, but our specimens fall out matching one of the two sequences fairly well, without anything in between. That plus the colour differences make me want to keep them separate for now.

Hygrocybe 'ceracea PNW07' © Steve Ness (picture comes with note: "more yellow than picture suggests")

Hygrocybe PNW01 - this species has only been found once in OR but that collection was orange with yellow tones on the cap edge, gills and stem bottom. The gills were decurrent, and at least the cap viscid. We'll need more collections to see if these colours are representative. It was found in a swampy oak woodland.

Hygrocybe PNW01 © Connor Dooley

Hygrocybe parvula NY - one variety doesn't have any red in it, but another variety does, so it is described below in section "4. Slender Red Species". It has a yellow cap, yellow decurrent gills, and most notably an orange stem that is slightly brighter than the cap.

Hygrocybe 'constrictospora PNW08' - This is orange to yellow and decurrent with a long stem, somewhat resembling Gloioxanthomyces, but this one is barely viscid if at all. I know I have placed it in the viscid capped section, because it is related to viscid capped species, but the few collections we have don't seem to be viscid. It has been found and sequenced three times from WA. It is related to Hygrocybe constrictospora (about 4% different in ITS) but doesn't really look like it. However, a microscopic analysis of one specimen did find constricted spores, just a little wider than expected for H. constrictospora itself. For the real Hygrocybe constrictospora, see the red section.

Hygrocybe 'constrictospora PNW08' © Steve Ness

Hygrocybe 'constrictospora PNW09' - This sequence is only 4% different from Hygrocybe 'constrictospora PNW08', but interestingly, mostly in ITS2. ITS1 only differs by 2 bp and 1 indel. It's ITS2 that differs in about 15 places. This is interesting because ITS1 usually varies more than ITS2. The cap and gills seem very pale or have perhaps faded to white, at least at the margins, in both WA collections, unlike the first species. The notes from one collection from WA say that the stem appeared viscid, and it was growing on a lot of moss on a rotten log. It is also related to Hygrocybe constrictospora (about 3% different in ITS) but doesn't look like it.

Hygrocybe 'constrictospora PNW09' © Steve Ness

Hygrocybe PNW23 - sequenced from OR and CA, so far no red was noted in the collections. It was slightly viscid and probably found with redwood and sequoia. It may clade near H. constrictospora, PNW18 and PNW19.

Hygrocybe PNW23 © Damon Tighe (2 images) and Jordan Gates (2 images)

Hygrocybe 'minutula PNW24' - dark orange caps fading to pale orange, the stems seemed pale orange. Our one WA collection sequences about 5% different in ITS from a couple ENA sequences purporting to be Hygrocybe minutula NY. However, that species starts out scarlet. Maybe this one will too if found fresh, but it seems like this is not what people have been finding and mistaking for H. minutula, that could be any number of our local species.

Hygrocybe 'minutula PNW24' © Matthew Koons (2 images)

3. STOCKY RED SPECIES - (most are fairly closely related to each other)

Hygrocybe 'marchii PNW10' and 'marchii PNW11' (H. 'coccinea' misapplied) - Larger and stockier than those covered previously, these mostly red mushrooms may have yellow on the stems (unlike what the photos suggest, PNW10 is not more likely to have yellow on the stem). The cap may be slightly sticky and the gills are not deccurent. We have two species, both more closely related to H. marchii than to H. coccinea, but as the former is a poorly known species, we've been calling both of our species H. coccinea. The more distantly related PNW11 is more common. Both can have quite a variable amount of red.

Hygrocybe 'marchii PNW10' © Steve Ness,     Hygrocybe 'marchii PNW11' © Steve Ness

Hygrocybe 'aurantiosplendens PNW12' - Related species (that are also stocky without decurrent gills) are not well understood yet. We have a sister species in Washington of the European H. aurantiosplendens (it differs by more than 10%) with splotches of red, orange and yellow on the cap.

Hygrocybe laetissima CA - Also very similar and related, described from California, it has now been found in Washington by the South Sound Mushroom club. It has a brighter red cap, and a white to yellow stem that is longitudinally fibrillose. In California it is reported from Bay Laurel and Redwood, but in Washington it was found with red cedar that had some bigleaf maple, red alder and hemlocks.

Hygrocybe punicea EU - Another similar, H. coccinea group relative, with a darker red cap and yellow to orange to red stem that is also longitudinally fibrillose. California sequences are 4bp and 1 indel different than European sequences, which for some families can be enough to be a distinct species, but for this family, is not a lot of variation, so ours may be the same species. East coast specimens seem to be clearly distinct. We need PNW collections.

Hygrocybe 'aurantiosplendens PNW12' © Steve Ness,     Hygrocybe laetissima © Lauren Ré,     Hygrocybe punicea, California © Christian Schwarz

4. SLENDER RED (when young) SPECIES

Hygrocybe 'mucronella PNW13' - Very bitter tasting. It is a small, red to orange to yellow mushroom with a sticky cap and stem described with adnate gills (but appearing sinuate in some of our WA collections). It looks like we have a sister species to the European H. mucronella (also called H. reae) as local ITS DNA (found in WA and OR) is more than 10% different, so ours probably needs a new name.

Hygrocybe 'mucronella PNW21' - another very bitter, entirely viscid sister species more recently discovered once in OR, almost 10% different in ITS from H. mucronella and more than 10% different than PNW13.

Hygrocybe 'mucronella PNW13' © Steve Ness

Hygrocybe 'mucronella PNW21' © Connor Dooley

Hygrocybe PNW14 - this slender species with a long stem is entirely viscid and tasted very bitter, like PNW13 and PNW21. It seems to have a much longer stem than that species. Sequenced only once from WA, whereas we have 4 sequences of the previous species. This species is not even closely related to H. mucronella or our bitter relative, above, so it is indeed mysterious.

Hygrocybe PNW14 © Connor Dooley

Hygrocybe PNW15 ("opposite") - This one is easy to identify with a bright yellow cap and gills, which are strongly decurrent, but a bright red stem (the opposite colouration of many others). It has been mistakenly called Hygrocybe parvula (described next), because that species can have a brighter stem than cap, but never as strikingly as this species. One of the two collections (Metaline Falls) appeared viscid (and its closest relative is viscid), and one probable CA coastal collection reported the surfaces were "tacky to subviscid". It was found in the PNW three times, twice in the WA Olympics at about 3,000' and once near Metaline Falls near the Idaho border at 2500'.

Hygrocybe parvula NY - viscid yellow cap and gills, gills usually strongly decurrent, with an orange-yellow stem. The notable thing about this species is how the stem is a bit more brightly coloured than the cap, but not nearly as much so as Hygrocybe PNW15 ("opposite"). The one discrepancy is that it was described as moist to subviscid, but at least in our area, appears to be distinctly viscid. Sequenced from WA, matching two NY type area sequences that seem to be the real thing.

Hygrocybe 'parvula PNW20' (var. "red chunk") - these sequences are only 2 bp different from H. parvula, plus the insertion of one long chunk of 9 characters, but morphologically it is quite distinct, with a red to orange cap (the other features being a viscid cap, decurrent gills, and orange-yellow stem). This is quite a common colour pattern, unfortunately, so it will easily be confused with all five of the poorly understood species below. The DNA is close enough to consider it a variety of H. parvula. One microscopic analysis showed that the spores of this variety were wider than those of the type variety (Q=2 vs. Q=3) although that is an odd result as the type variety is not described as having spores that narrow.

Hygrocybe cf subceracea FL - the only other species reported from here to have small spores (in fact even smaller than H. parvula), so it needs to be investigated which of the species on this page is what H&S called H. subceracea, and if that is in fact what they are. It is somewhat morphologically similar to "red chunk".

Hygrocybe PNW15 "opposite" © Steve Ness,     H. parvula © Steve Ness,     H. 'parvula PNW20' (var. "red chunk") © Steve Ness and Michael Beug

Hygrocybe fenestrata n.p. - A smallish mushroom which is quite viscid everywhere, red fading rather quickly to orange and yellow (like many others do) but usually with a milky translucent "window" on the cap disc. The gills are usually notched with a decurrent tooth. It is a fairly new discovery from California now known to occur in Washington, and when it gets a name, that name may be Hygrocybe fenestrata ("windowed"), but for now it is an "n.p.", or a provisional name only.

Until it was sequenced, it was originally thought to belong in Gliophorus because of it's extremely viscid nature. Even though we don't have photos of young enough specimens to show the red colour, it had been noted to be red when very young, and as we now know, Gliophorus probably lacks the ability to create betalain red pigments like Hygrocybe can, so that was a clue that this was a slimy Hygrocybe and not a Gliophorus. This is likely a sister species of the similar EU Hygrocybe glutinipes (another slimy Hygrocybe that isn't known from the PNW), a complex differing by up to 5% in ITS. This species is >10% different in ITS from that complex.

Hygrocybe fenestrata np © Steve Ness,    probable faded specimens w/o red © Maria Morrow

Hygrocybe PNW17 - this appears to be mostly red with a yellow-orange rim in places, very viscid all over and with adnate gills. "Growth from underneath Thuja plicata roots. Pressed tightly against soil and roots, densely clustered". It has been sequenced from California and Washington.

Hygrocybe PNW17 © Peter Russell, Lauren Ré and Lauren Ré

Hygrocybe PNW18 - some of these were quite young, and showed red, orange and yellow colours, but interestingly, the youngest ones were the palest colours, so this doesn't seem to follow the usual pattern of starting off red and fading to orange or yellow. They appeared to have adnate gills. It seems shiny enough to be viscid. This seems to be a lookalike of Hygrocybe ceracea and its relatives, when lacking in red tones. Found in Washington in 2018 (older, redder, specimens) and then again in 2020 (young ones).

Hygrocybe PNW18 © Lauren Ré

Hygrocybe PNW19 - again, mostly red with bits of orange, viscid all over, but clearly decurrent gills. Closely resembles many others. The next step is to get spore characteristics from each of these species and try to definitely determine what H&S would have called each of them. This probably mostly resembles the "red chunk" variety of H. parvula, but hopefully has larger spores.

Hygrocybe constrictospora EU - one ID collection matches EU sequences. Perhaps it is best recognized microscopically by its constricted spores, but our one collection had an orange cap with a red rim, and an orange stem.

Hygrocybe PNW19 © Sadie Hickey,     H. constrictospora © Ed Barge

Poorly understood other species - click to expand
Species mentioned: Hygrocybe americana, luteo-omphaloides, atro-olivacea, minutula, subminiata. Omphalina occidentalis. Hygrophorus albicarneus. Gliophorus minutulus.

1. Locally described species that are probably legitimate but poorly understood

Hygrocybe americana OR - This was called H. acuta until they realized that name was already taken and it had to be renamed. It is described from Oregon. It has a viscid, dark grey-brown very pointy cap and a dry white stem. No DNA is available of anything resembling it yet.

Hygrocybe luteo-omphaloides WA - Otherwise known as Omphalina occidentalis, this little yellow moss waxy with decurrent gills, was described from Washington. It may be one of the above species or something new, and it may not even be a Hygrocybe, that has not been proven (the basidia are pretty short). We need samples.

Hygrophorus albicarneus OR - white to pale pink, decurrent gills, viscid cap. Snowbank species. Described from Oregon, it is so rare, it is technically still called Hygrophorus albicarneus because it hasn't been moved to Hygrocybe albicarnea yet, where it more likely belongs based on gill trama.

2. Species reports without genetic data yet

Hygrocybe atro-olivacea MI - described from Michigan, but reported once from Washington by Hesler & Smith in their iconic monograph. We need to see if it really is the same species. It has somewhat decurrent gills, a dry, somewhat scaly olive-brown cap that is almost black in the center. No east coast nor west coast sequences yet.

Hygrocybe cf minutula NY - a reddish orange capped species described from NY with variably attached gills. It is described as very viscid all over and has been placed in Gliophorus as Gliophorus minutulus as a result, but the only east coast sequences purporting to be this species are in Hygrocybe, but we don't know for sure they were identified correctly. It is likely that one or more of the many unnamed species on this page represent reports of H. minutula, but so far none of these genetic species match purported sequences of H. minutula back east. So what were they finding and calling by this name?

Hygrocybe cf subminiata Jamaica - also quite small (less than an inch across) with a slightly viscid red cap and dry yellowish stem, and with whitish decurrent gills. We also need to figure out what reports of this species in the PNW really are. As for H. minutula, one or more of the unnamed species on this page may have gone by this name. We have no idea what this species is genetically, it has never knowingly been sequenced.

Hygrocybe cf atro-olivacea © Renée Lebeuf

One study showed that the DNA of the following Hygrocybe species were found inside seeds in either Washington or Oregon, far away from their normal range, and never known to have fruited (so the mushrooms may never be found here).