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Danny’s DNA Discoveries – Hygrophoraceae (Waxy Caps)

by Danny Miller

 

Click here for my Pictorial Key to the Hygrophoraceae

 

Introduction

Waxy Caps (members of the Hygrophoraceae family) are many people's favourites, as they are often beautifully coloured 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) are especially long in this family, five times longer than the spores that grow on them, on average, which is what gives the gills their thick, waxy appearance. Not every species is equally colourful and waxy, sometimes the effect is more subtle and has to be learned to be recognized, but even the white and brown mushrooms in this clade are more of a pastel white or brown.

The gills are usually thick and widely spaced, as well as being waxy. Viscid caps are common, as are decurrent gills (but that's not always true). Rarely, there is a partial veil.

Colourful Mycenas might be mistaken for these mushrooms, but they are small, fragile and delicate and although they might be colourful, they are not at all waxy. Laccaria might also be mistaken as those do look sort of waxy, but they are always orange or purple with tough, fibrous stems.

Steve Ness is 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. Unfortunately, DNA is this family appears to mutate rapidly, and there is a greater distance between species than normal, meaning that ITS alone is very poor for figuring out the family tree and multiple gene sequences will be especially important here. Sometimes I have trouble placing a species in the correct genus or section with only ITS sequences. I wonder if it will turn out that ITS DNA will often need to vary by more than the typical/average 0.5% to actually indicate a unique species in this family.

 

Summary of Results from South Sound Mushroom Club Collections

Thanks to the collecting and sequencing done by the South Sound Mushroom club, I have been able to determine or confirm the following (more details further below):

Chrysomphalina - dry caps, growing on wood

Chrysomphalina aurantiaca and grossula © Kit Scates Barnhart, Chrysomphalina chrysophylla © A and O Ceska, Chromosera cf cyanophylla © Steve Trudell

Most waxy caps grow on the ground, but a few grow on wood. Chrysomphalina have dry caps, decurrent gills and may have a hygrophanous cap. Chromosera may be found on wood but is viscid all over and is discussed below.

Summary

Chrysomphalina aurantiaca - common, small, completely orange little waxy. This east coast species has never been sequenced from the east coast, so we need east coast DNA to confirm that ours is the same species.

Chrysomphalina grossula - uncommon yellow-green little waxy. One or two consecutive and equal chunks, four nucleotides long (GCAT) may be missing from our DNA compared to European DNA, but as a single event could drop all those characters, that probably doesn't represent enough of a change to constitute ours being a separate species. Otherwise, there is only 1 or 2bp difference.

Chrysomphalina chrysophylla - uncommon, brown capped, golden yellow little waxy. California DNA matches European DNA, so our local PNW DNA probably does too. We need local collections.

More Distant Relatives

Aphroditeola aff olida © Andrew Parker, Cantharocybe gruberi © Danny Miller, Pseudoomphalina angelesiana © Steve Trudell

Hodophilus paupertinus © A and O Ceska, Hodophilus sp. nov. © A and O Ceska

These species are either on the edge of the family tree or are not related to the waxy caps at all, but evolved to look waxy anyway. All are rare.

Aphroditeola aff olida - rare, small, pink capped, decurrent, smells strongly like tutti-frutti bubble gum! Our DNA is different than east coast DNA and if it turns out to also differ from European DNA (where A. olida is described) we will need a new name for ours. We need European samples. A possible second species was found in WA, differing by 1bp in ITS1 and 4bp and 2 indels in ITS2. There's a possible third species in Northern BC.

Cantharocybe gruberi - rare, large, yellow, decurrent, stem often off centre. We have DNA from CA, but it was described from ID, so we need specimens closer to there to verify it's all one species.

Pseudoomphalina angelesiana - this rare, violet-brown, viscid, decurrent small species was described from WA and while looking waxy, is not related (it's in the Tricholomatoid clade). It has previously been called Neohygrophorus.

Hodophilus paupertinus - this rare, drab yellow-tan-grey decurrent waxy-looking mushroom smells strongly of mothballs, but is not related to the other waxies. It was described from CA and we also have the DNA from BC.]

Hodophilus sp. nov. - this even rarer unnamed species from near Victoria, BC has darker yellow colouration (it reminds one of H. phaeoxanthus which means dark yellow, but it isn't that species) and also a strong mothball odor.

Cuphophyllus - click here for details

Cuphophyllus 'borealis' © Steve Trudell, C. subviolaceus © R Lebeuf, C. pratensis group © Steve Trudell

Formerly Camarophyllus, this genus is defined by the interwoven gill hyphae, but without a microscope, you may recognize them by dry stems, often dull colours and decurrent gills that are widely spaced apart. Some caps are <5cm across, but others can grow up to 10cm. Other species 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.

Summary

Cuphophyllus 'borealis' - common, small and white, dry or slightly viscid cap. It's unclear if we have more than one species and if the proper name is C. borealis or not, but the proper name is not C. virgineus. Some white species of Hygrophorus are easily confused 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.

C. burgdorfensis - this rare, small, yellow capped, viscid 'borealis' like species is only known from the type in Idaho. No DNA yet.

C. russocoriaceus/lawrencei -two rare, small, white mushrooms that smells strongly of cedar! C. lawrencei is a bit bigger and stockier.

C. aff lacmus - uncommon, purple-brown cap and gills, white stem. No odor, unpleasant taste that is either bitter, hot or nauseating. Needs a new name.

C. 'cinereus' - similar but with a mild taste, it was reported once from Mt. Rainier but we need samples from here and the EU to see if we really do have this species or not.

C. rainierensis/nordmanensis - rare, similar to C. aff lacmus, but they taste mild and smell of green corn, differing by spore size. These need DNA for study.

C. 'colemannianus' - uncommon, warm brown cap, pale gills and white stem. Ours might need a new name, we don't know yet.

C. recurvatus - rare, dark olive-brown cap, white gills, more slender slightly coloured stem. Similar to Gliophorus unguinosus below (but dry capped with more decurrent gills). DNA not verified yet.

C. pratensis group - not uncommon, two sister species, one of which is the real thing, it's just not clear which. Larger, orange to pink capped species (else pale).

C. graveolens - uncommon, similar, paler cap, more slender stem, smells sweet.

C. cremicolor - rare, smaller, pale cap, yellow gills, perhaps a slight fragrant odor. We need DNA to determine if this is different than C. graveolens.

Click here for more details of  Cuphophyllus

 

Hygrocybe s.l. and Hygrophorus

The rest of the mushrooms can be diviced into two clades, Hygrocybe and Hygrophorus, the two most famous genera in the family, differentiated by either parallel or divergent gill hyphae. Without a microscope, you can still often tell which of the two you have (after first eliminating the genera I have discussed above).

Hygrocybes are usually smaller and also 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 (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. But this is happening more and more as genetics force us to reckon with how species are truly related. You can still think of this group as Hygrocybe s.l. (sensu lato, in the broad sense).

Hygrophoruses are usually larger if they have a colourful stem, or are smaller with a paler stem. Sometimes they will have a partial veil. Mercifully, these are all still in the same genus. They have divergent gill hyphae. All of the Hygrocybe s.l. caldes shown below have parallel gill hyphae.

 

Hygrocybe s.l. - Chromosera

Chromosera cf cyanophylla © Steve Trudell, Chromosera citrinopallida © Christian Schwarz

Chromosera are viscid all over with bright colours that fade (like Gliophorus) with decurrent gills, and one grows on wood (like Chrysomphalina).

Chromosera cf cyanophylla - this is common, and for some reason it grows on wood so it is also keyed out above. 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. I believe east coast DNA, the type area for Mycena lilacifolia (now regarded as a synonym) is also distinct, so while they may be able to use a name like Chromosera lilacifolia for theirs, we need a brand new name for ours.

Chromosera citrinopallida - this rare 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. This is described from Washington so it definitely occurs here. We only have DNA from Iceland and Denmark, so we need local specimens to make sure their species is the same and to confirm the placement of our species. 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 irregularily yellow-white.

 

Hygrocybe s.l. - Gloioxanthomyces

Gloioxanthomyces nitidus © Andrew Parker

Gloioxanthomyces nitidus - we have one rare species in this genus here, and it is bright yellow and viscid everywhere, with an indented cap, decurrent gills and a very long, thin stem. It was described from eastern North America and we still need local DNA to confirm that it is the same species over here.

 

Hygrocybe s.l. - Humidicutis

Humidicutis marginata var olivacea © Bryce Kendrick

Humidicutis marginata - a rare, entirely orange species with a viscid cap. The gills are especially bright orange and stay orange, at least on the margins, long after the colour has faded elsewhere. This east coast species is rumoured from the PNW but we need specimens to confirm it is the same.

Humidicutis marginata var olivacea - this rare variety was described from Washington and has olive colours as well as orange colours.

 

Hygrocybe s.l. - Gliophorus

Gliophorus psittacinus group © Danny Miller, G. laetus group © Steve Trudell, G. cf irrigatus © Noah Siegel, G. flavifolius © Christian Schwarz

Gliophorus species are slimy-viscid all over and usually decurrent (but so are some others in Hygrocybe s.l.). A famous green mushroom is found here.

Gliophorus psittacinus group - common, sometimes green and sometimes orange (sometimes both!) and can change back and forth between those colours. One group member in California can be blue. It is a brighter green than the other famous green mushroom, the redwood yellow-green Hygrocybe virescens. The gills are not always decurrent and the cap may sometimes be pointy. So far we have DNA for 3 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.

Gliophorus laetus group - common, a different shade of orange, and smells 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. 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 cf irrigatus - rare European mushroom, dark grey-brown cap and stem, similar colours to Cuphophyllus recurvatus, above (a dry mushroom). In California they have named one local group member Hygrophorus subaromaticus which probably needs to be recombined into Gliophorus subaromaticus. We need samples of that and our local species to see which species we have. The also European G. unguinosus is thought to be a synonym for G. irrigatus.

Gliophorus flavifolius - rare California species 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 similar to Chromosera citrinopallida, above.

 

Hygrocybe s.s. - click here for details

Hygrocybe constans ('miniata') © Steve Ness, H. aff flavescens © Steve Trudell, H. aff singeri #1 and H. cf coccinea #1 © Steve Ness

These species are still properly in Hygrocybe itself.

Summary

Dry caps

H. constans (aff miniata) - this very common 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, with a dry cap that has small scales on it and gills that are not usually decurrent. The proper name for our species may not be H. miniata nor H. squamulosa, but H. constans.

H. substrangulata - similar and realted to H. constans, but usually found in wet moss and described as more scurfy than scaly on the cap. It is much rarer.

H. aff phaeococcinea - another similar and related undescribed species that may not have scales on the cap and usually has an orange-yellow cap margin even when the rest of the mushroom is still red and hasn't faded in colour. It is found in grass and probably rare. (One additional undescribed species in the miniata group may exist here).

H. reidii - small, bright orange with a dry, essentially smooth cap and often with an odor of honey gone bad. It is uncommon.

H. cantharellus - this species has a longer stem and more strongly decurrent gills than H. constans and the other dry capped species. The similar H. turunda is supposedly differentiated by scales that are darker than the rest of the cap, but we have not yet studied local specimens to see if this is the right name to use. H. coccineocrenata is similar, but not verified from here yet either.

H. virescens - from California can be found here under any introduced redwood trees, maybe bay. It is not viscid and is a beautiful, unique, lime-green to greenish-yellow colour.

 

Sticky caps

H. aff flavescens - this very common mushroom is yellow (or sometimes orange), 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.

H. ceracea - similar, but usually golden yellow with adnate to decurrent gills. Slightly sticky everywhere when wet. H. subceracea is rumoured to have a dry stem and different sized spores and needs study.

H. singeri group - the very famous witch's cap mushrooms are very common and identified by red, orange or yellow mushrooms that turn pitch black in age. They have pointy caps and are sticky, usually all over. We do not have H. conica here (that is a European species) but we do have H. singeri (described from OR) and at least two more cryptic species in the group so far.

H. coccinea group - larger and stockier than the H. miniata group, these mostly red mushrooms may have yellow on the stems. The cap may be slightly sticky and the gills are not deccurent. We have two species, neither the real thing.

H. cf aurantiosplendens - (not the real thing) is related and has a splotchy mix of red, orange and yellow in the cap.

H. 'punicea' - is related and has a darker red cap and a red to orange to yellow stem that is longitudinally fibrillose. No data yet on if it is the real thing.

H. laetissima - related California species with a bright red cap and white to yellow stem that is somewhat longitudinally fibrillose.

H. aff mucronella/reae - another undescribed, small, red to orange to yellow mushroom with a sticky cap and stem and gills that are not decurrent. It's distinctive feature is that it is supposed to taste very bitter!

H. cf minutula - I have no idea if ours is the real thing or not, we need more records of this species from its type area in New York. One of its synonyms is Gliophorus minutulus, because it is sticky all over like many Gliophorus, but it is actually a Hygrocybe. The cap is red fading to orange or yellow, the gills are yellow to orange and may or may not be decurrent, so this species is very difficult to tell apart from H. 'subminiata' and H. aff mucronella (although that is probably bitter).

H. 'subminiata' - is rumoured to exist here. It is quite small (less than an inch across) with a viscid red cap and yellowish stem and pale gills (that are decurrent). It is easily confused with H. minutula.

H. fenestrata np - small-ish, quite viscid everywhere, red fading rather quickly to orange and yellow (like many others) but usually with a milky translucent "window" on the cap disc.

We have genetic evidence of  at least 3 additional undescribed species that we know little about (more details on the detail page).

 

We also have locally described species that are probably legitimate but poorly understood:

H. californica - we don't have local sequences of H. acutoconica or H. persistens or H. cuspidata, but I suspect at least one of our local species that match these descriptions is H. californica, described, obviously, from California. They are red-orange-yellow and pointy capped, but do not turn black like the H. singeri group, at least not much.

H. americana - 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.

H. albicarnea - white to pale pink, decurrent gills, viscid cap. Snowbank species.

H. luteo-omphaloides - 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). This one is not viscid.

 

The following species have been reported (with fairly good microscopic evidence) but without genetic data yet:

H. 'atro-olivacea' - somewhat decurrent gills, a dry, somewhat scaly olive-brown cap that is almost black in the center.

H. 'parvula' - decurrent, bright yellow gills and a yellow-orange, sometimes slightly sticky cap, but the stem is sometimes even more highly coloured (pinkish or orange). Perhaps the stem can be pure yellow.

Click here for more details of Hygrocybe s.s.

 

Hygrophorus - click here for details

All of these species have divergent gill hyphae, unlike Hygrocybe s.l. with parallel hyphae and Cuphophyllus with interwoven hyphae. (Chrysomphalina and the more distant genera described up top have unknown hyphae, at least to me. This trick only works for the three main groups). Most have a viscid cap unless noted. A viscid stem will be noted. Hesler & Smith (see detail page) put a lot of weight on whether or not the stem was viscid, but that' s probably not as important to phylogeny as they thought.

Summary

H. caeruleus - a rare, beautiful baby blue stocky waxy, actually a strange form of Clitocybe odora.

 

Fibrillose partial veil

H. subalpinus - white and stocky, this is a common veiled conifer spring snowmelt species.

H. siccipes - a distinctive orange-brown and/or olive-brown veiled conifer common waxy that may prefer 2-3 needle pine that we have wrongly been calling H. hypothejus. We may also have the slightly more slender and duller coloured H. boyeri.

H. 'purpurascens' group - purple and spotting/staining more purple, common stocky spring and fall spruce and pine veiled mushroom. We may have two undescribed species. H. velatus is supposedly pale pink without spotting, but may be a proper name for one of our species and needs study.

H. olivaceoalbus group - mostly grey and stocky spruce species. The fibrillose veil is said to be black and the stem is viscid. Rare in the coastal PNW, perhaps we actually have H. korhonenii instead. H. olivaceoalbus var gracilis, also from the EU and possibly reported from hardwoods, is more slender and rarely reported from here but we don't have DNA yet.

H. 'fuscoalbus' - rare, reportedly has a white fibrillose veil instead of black. Nobody is sure what this European species is.

H. fuscoalboides similar, grey fibrillose veil and dry stem, described from Idaho, is likely the proper name for at least one of our local species in this group.

H. inocybiformis - similar grey, scaly, veiled waxy with spruce and fir that is dry everywhere (nowhere viscid). It is common in Idaho where it was described, but rare elsewhere.

 

Gills etc. spotting/turning pink/purple

H. 'purpurascens' group - the one with a partial veil, see above

H. cf erubescens group - common spruce/pine species, irregularly purple often with yellow. H. erubescens var gracilis (type OR) is a rare, more slender, more distant gilled mushroom that usually lacks the yellow. It is probably not a variety but needs a new species name.

H. cf capreolarius - this uncommon spruce species has more evenly distributed purple colours than the others.

H. cf russula - rare hardwood species with slightly closer gills than others.

 

Sweet Almond/Cherry Odor (specifically, not just sweet or fragrant)

H. agathosmus/odoratus - grey with spruce and pine/conifers, common/uncommon EU/Oregon species with different sized spores. H. odoratus is reportedly more slender.

H. bakerensis/variicolor - very common conifer species, brown disc with paler rim. H. variicolor, the older name, differs by a viscid stipe, but they may be the same.

H. secretani (monticola)/vinicolor - more evenly brownish cap flushed vinaceous, conifers. The Idaho type of H. monticola (where it is common, but rare elsewhere) appears to match the EU H. secretani, an older name. The rare H. vinicolor, also from Idaho, reportedly tastes bad with pinkish gills and a pinkish pruinose stem but might be the same.

 

White or Colourful Caps (not brown, grey or black)

H. piceae group - White, slender, viscid cap but dry stem, very similar to Cuphophyllus 'borealis', but not striate and the upper stem is very pruinose or fibrillose. It is common with spruce, as its name suggests. We have more than one unrelated species, this needs to be sorted out.

H. eburneus group - (as a group: white, very viscid, slimy stems, sometimes with yellow shades. No evidence yet that we have any of these species, some of ours may need new names). H. 'eburneus' - common, mid-sized, mostly conifers. H. cossus - aromatic, cap/stem aging pinkish buff, reported once from ID. H. glutinosus - ENA species, one record from OR, cap drying yellowish with red punctate dots on stem after drying. H. gliocyclus - spruce/pine, not uncommon in OR+ID, stocky yellowish cap when fresh, viscid stipe. H. flavodiscus - midsized, pine, yellow disk, not uncommon in OR+ID

H. melizeus forma minor (karstenii forma minor) - white with pale yellow gills, small, no tree info. This is presumably in Hygrophorus, but H. karstenii is a Cuphophyllus and nobody really knows what H. melizeus is supposed to be, so the one report of this variety of an EU species from Mt. Rainier is a mystery and should be looked for again.

H. aff chrysodon - this golden flecked, viscid stemmed white waxy is easily recognized, not uncommon, with conifers, and needs a new name.

H. pusillus - small, aromatic, white waxy, sometimes with a pink disc and a viscid cap. Not uncommon conifer species.

H. goetzii - small, pale pink, uncommon snowbank hemlock species from Oregon.

H. saxatillis - white cap, bright pink gills and a peachy odor, not uncommon, with conifers.

H. 'pudorinus'/fragrans group - a pale pink to orange viscid cap, punctate upper stem, may smell sweet. H. fragrans (or H. pudorinus var fragrans) is said to be more common, stain yellow-orange and have slightly coloured gills and a stronger odor.

H. speciosus - slimy everywhere, bright red-orange cap with white gills and stem. Favours larch.

H. sordidus/penarius - see discussion below under H. aff marzuolus.

 

Brown, Grey or Black on the Cap, at least in age on the disc

H. pacificus  -much like the much more common H. bakerensis and H. 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.

H. vernalis - brown capped spring snowmelt species (otherwise like the aromatic H. montanus but with a viscid stem), under hemlock. Fairly rare.

H. 'discoideus' - slender species with a bright, brown disc that is usually quite offset from the pale rim, viscid all over like H. occidentalis, next.Spruce? European species, ours possibly in need of a new name.

H. occidentalis - this CA species is found not uncommonly there and also in OR (under oak and pine). It is somewhat slender and has a dull grey-brown disc with a paler rim and is viscid all over, described like a browner H. eburneus (and also similar to H. 'discoideus' but without such a striking disc margin and with duller colours).

H. aff tephroleucus (aff pustulatus) - small, greyish cap, sometimes pointed, with notable dark scabers on the somewhat viscid stem. Uncommon, fir preference? H. morrisii is a 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, unconfirmed.

H. aff marzuolus - stocky spring species that is mostly white with a grey cap rim and dry stem. The cap may become more grey as it ages. Uncommon, unknown trees. This may represent some reports of H. sordidus and H. penarius, although one of those species may rarely occur here.

H. camarophyllus - fall species with blackish caps even when young, also stocky. Not uncommon under spruce and pine.

H. calophyllus - a similar blackish capped European mushrooms but with pink gills, uncommon under fir and pine.

 

LOCALLY DESCRIBED, rare, unknown DNA

H. albiflavus - a slender, white waxy with a fibrillose veil. Viscid stipe. Only known from the OR type. Unknown trees. Large spores.

H. amarus - yellowish gills that spot pink like other purple/pink spotting species, but with a buff to salmon to yellowish cap (not the usual purple cap). Notable bitter taste. Rare. May not be a unique species? Spruce and Douglas fir.

H. megasporus  -an uncommon conifer species, olive-grey brown with a viscid stem, (like H. olivaceoalbus, but no fibrillose veil), It is practically unknown, but with giant spores (even bigger than the olivaceoalbus grp) it should be easy to detect if found again to figure out what it is.

H. ellenae - white, cespitose species, not as stocky as H. sordidus (but a lot like ENA H. subsordidus). Stipe twisted striate, white tomentose at base. Only known from the type from Idaho. Pine and fir.

H. avellaneifolius - relatively slender stem, dark rusty brown to grey cap and darker avellaneous gills and a dry stem. Endemic to Idaho, conifers.

H. subpungens - 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.

Click here for more details of  Hygrophorus

 

Summary of Future Studies Needed

 

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