Danny’s DNA Discoveries – Hygrophoraceae (Waxy Caps) of the PNW
Introduction - click to expand
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.
Warning: BLAST will not be able to make any kind of a useful tree out of ITS sequence data for this family! You will need to use better alignment software, like 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.
Summary of Interesting Results - click to expand
Here are some of the newest, most interesting results of the study:
Summary of Results from South Sound Mushroom Club Collections - click to expand
Steve Ness, Lauren Ré, Luca Hickey and Eric Chandler are leading a study of waxy caps with the South Sound Mycological Society and lots of the information we know about this family is coming from the collections they have been making and sequencing.
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.
Species mentioned: Cuphophyllus borealis, virgineus, niveus, burgdorfensis, russocoriaceus, lawrencei, lacmus, subviolaceus, rainierensis, nordmanensis, cinereus, cinerellus, colemannianus, recurvatus, subradiatus, pratensis, graveolens, cremicolor
Cuphophyllus 'borealis' group NY - 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, 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.
'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.
Cuphophyllus russocoriaceus UK and '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 sp. iNat68420842 - this unique, probably unnamed species, doesn't match any known sequences yet. It appears to be odorless and hygrophanous with some stem nattering. The previous species are believed to either have a cedar odor or not be hygrophanous, but more collections are needed to find out.
Cuphophyllus sp. iNat68420842 © Lauren Ré
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. Collections from WA and northern BC were very close to European sequences, 1 bp and 1 long indel different in the WA collection. Another WA collection was 5 bp different, but as the waxy caps tend to show a little more genetic variation than usual, and as Voitk said to expect some more genetic variation than normal, I don't think we have a second species.
Cuphophyllus rainierensis WA and 'Hygrophorus' nordmanensis ID - said to be similar mushrooms that taste mild and smell of green corn, differing from each other by spore size. However, I do not believe them to be separate species. 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 basidium). Recently, a collection from Mt. Rainier (the area the original C. rainierensis was found in) that smelled of green corn was sequenced and found to be only 1bp and 1 long indel different than the European C. lacmus, so odor and taste may vary and it may not be correct to consider different smelling collections different species. Cuphophyllus nordmanensis has not yet been renamed from Hygrophorus nordmanensis and would need to be, if it was determined to be a separate species. But I don't think that will happen either.
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 (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.
probable Cuphophyllus colemannianus © Christian Schwarz
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.
Cuphophyllus recurvatus © Luca Hickey and Yi-Min Wang
Cuphophyllus pratensis EU group - Probably four or so larger species, orange to pink dry capped (else pale). 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 three others, between 4% and 8% different from each other. Also see a lookalike below.
Cuphophyllus pratensis group member © Steve Trudell
'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 graveolens © Christian Schwarz
Cuphophyllus aff aurantius - 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 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 aff aurantius © Luca Hickey
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).
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, persicolor, capreolarius, amarus, russula, piceae, eburneus, cossus, chrysaspis, glutinosus, gliocyclus, flavodiscus, melizeus forma minor, karstenii forma minor, sordidus, penarius, chrysodon, ellenae, pusillus, subpungens, goetzii, saxatilis, karstenii, pudorinus, fragrans, speciosus, speciosus var. kauffmanii, pacificus, vernalis, discoideus, occidentalis, tephroleucus, pustulatus, pustulatoides, morrisii, marzuolus, camarophyllus, calophyllus, megasporus, avellaneifolius. Clitocybe odora.
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.
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 aff cyanophylla - 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 WA and we have a WA sequence that represents the real thing. Greenland and the EU seem to have a sister species, 2% different.
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
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.
Hygrocybe s.l. - Gliophorus s.l. - click to expand
Gliophorus species are slimy-viscid all over, sometimes with decurrent gills. To distinguish those with decurrent gills from similar Chromosera, Gloioxanthomyces and Hygrocybe, these have gills with duller colours than the cap and stem, and 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.
This section also includes the mysterious 'Gliophorus' flavifolia (white with bright yellow gills) from CA whose DNA is too distant from anything else to properly place it. It could be in Gliophorus, or elsewhere, or deserve a new genus. If it is to be found in southern OR, as suspected, then we will have to worry about it and figure it out.
Species mentioned: Gliophorus psittacinus, laetus, sciophanus, irrigatus, unguinosus, flavifolius. 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 4 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.
#1 seems to have a dark orange phase not found in the others species (see photos). It's the first clue we have about how to tell them apart. #2 appears to be the most common and can be blue, although blue is the rarest colour reported. #4 has not been photographed yet (well, it has, but as part of a mixed collection where we don't know which one is #4).
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 psittacinus group #2 © Steve Ness, Steve Ness and Luca Hickey
Gliophorus psittacinus group #1 and #3 © Steve Ness
Gliophorus laetus EU group - these are possibly different shades of orange, and they usually smell strongly fishy. The gills should always be decurrent and the cap should not be pointy. So far we have 2 PNW species, neither of them the real thing, and a sister species of Gliophorus sciophanus, which looks like it belongs to this group even though G. sciophanus is supposedly a synonym of G. psittacina (it isn't). One of the three G. aff. sciophanus collections had the fish odor. One WA find 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 #1 © Steve Ness, #2 © Yi-Min Wang, G. aff. sciophanus © Yi-Min Wang and Steve Ness
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' flavifolius CA - a California species that may also be found under Douglas fir in southern Oregon. Very white stem, very bright yellow gills, and a white cap disc surrounded by yellow. Adnate gills. It is stockier than Chromosera citrinopallida, above. The DNA is not close enough to anything for us to know yet what genus this is.
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.
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).H. chloochlora, H. nigrescens, H. occidentalis, H. papillata, H. caespitosa, H. glutinipes, H. noninquinans
Summary of Future Studies Needed
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