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Danny’s DNA Discoveries – Hygrophoraceae (Waxy Caps)
by Danny Miller
First, the dry capped species.
Hygrocybe 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 (~2.5cm across) with a dry cap that has small scales on it and gills that are not usually decurrent. 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). Ours is not the true H. miniata of Europe. H. squamulosa may not be here either, the only east coast sequence purporting to be that species (type from NJ) 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. H. constans is an Oregon species from 1912 given its own name because 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.
Hygrocybe constans ('miniata') © Steve Ness
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 (and with slightly larger spores). It is much rarer. The presence of this UK species in the PNW was never suspected until the South Sound Mushroom Club DNA project found it.
Hygrocybe substrangulata © Steve Ness
Hygrocybe aff phaeococcinea
Another similar and related undescribed species that may not have scales on the dry 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 (unlike the others?) and probably rare. Since our species is not the true European H. phaeococcinea, we really don't have a good description of it yet and it will be hard to recognize. The presence of this species here was also a discovery by the South Sound Mushroom Club.
Hygrocybe aff phaeococcinea © Steve Ness
Hygrocybe sp. MP3955113 - somewhat closely related to H. phaeococcinea in the H. miniata group, it appears to have started out red and faded to orange and yellow. It has a noisy chromatogram in places (but clean 5.8s region) so 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 sp. MP3955113 © NAMA and the Field Museum of Natural History
This is not 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 uncommon, and also a newly discovered European species by the South Sound club. It was found in the forest.
Hygrocybe reidii © Steve Ness
This species has a longer stem and more strongly decurrent gills than the dry capped H. constans and the others. It is an east coast species also found here.
Hygrocybe turunda is supposedly differentiated by scales that are darker than the rest of the cap. In Europe, there are 4 genetic species that appear to be able to go either way, so I'm not sure what dark scales say about species, since we haven't sequenced any dark scaled specimens from here yet. To confuse things, there has been some debate as to whether or not our local species with dark scales should be called H. turunda or H. coccineocrenata (both European species), but the answer could be neither and we may need new names for any additional species that turn up in the PNW. Hygrocybe turunda var sphagnophila is an east coast name. We have no type sequences of it, but it could turn out to a valid unique species or variety and also be found here.
Hygrocybe aff 'cf minutula'
A species that is 5% different in ITS than H. 'cf minutula', below, but looks more like Hygrocybe cantharellus than H. minutula, was found once at 3000' in the Washington cascades. It is not viscid at all like its closest relative and does not seem to belong in the H. minutula group. Clearly more than one specimen will need to be found to verify anything at all about this new species.
Hygrocybe cantharellus © Steve Ness, unsequenced H. cf turunda © Andrew Parker, H. aff 'cf minutula' © Steve Ness
From California and can be found here under introduced redwood trees, maybe also bay laurel. It is not viscid and is a beautiful, unique, lime-green to greenish-yellow colour.
Hygrocybe virescens © Steve Trudell
Most species discussed next will now be viscid, at least on the cap.
Hygrocybe aff flavescens
This very common 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. It could be difficult to tell apart from H. cf minutula, but even when that species fades there should still be some red somewhere, unlike this species, and this species is much more common.
Hygrocybe aff flavescens © Steve Trudell
This similar mushroom is golden yellow everywhere with adnate to decurrent gills and is slightly sticky everywhere when wet. Our DNA differs by 2-3 substitutions in ITS and a couple to few indels in ITS2 from European material and is likely the same species.
Hygrocybe subceracea, 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.
Hygrocybe ceracea © Steve Ness
Hygrocybe 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 potentially three more cryptic species in the group so far (#1 once from WA, #2 once from BC and three times from WA, #3 once from WA). California may have additional species. Work will need to be done to figure out how many species we have and how to differentiate them. Traditionally they've been separated by the viscidity of the stipe and colour of the young gills, but that may not hold.
Hygrocybe aff singeri #1 © Steve Ness, Hygrocybe aff singeri #2 © NAMA and the Field Museum of Natural History, H. aff singeri #3 © Eric Chandler. H. singeri not photographed.
Hygrocybe acutoconica is a rarely reported pointy capped orange-yellow waxy like H. singeri, but that does not turn black. It has also been known as H. persistens and H. cuspidata (the latter when it has red tones).
Sequences from Europe labeled H. persistens, where it was described, show at least 3 different sequences, at least 5 bp apart from each other and sometimes more. The type of H. acutoconica var microspora from the east coast has been sequenced and is similarly different. The few collections from the west examined by Hesler and Smith did not have the smaller spores of that variety. One local sequence from Washington (below on the left) has been sequenced and is about 5 bp from all of the above, but it is only a short sequence of a portion of ITS2 so it is expected that when we get more of the region sequenced there are probably going to end up being more differences than that. Although it is true that waxy cap species may show greater variety than usual within a species, it is probably best to assume that this is a group of species. Hygrocybe californica was described from California and could be the proper name for our species, but more local and California sequences are going to be needed to confirm that. The state of affairs up until now, before DNA scrutiny, was that all of the above mentioned species were the same, with H. acutoconica being the oldest and best name, but as there is evidence of genetic difference, I am going to resurrect the name Hygrocybe californica as the best name to use for our species for now.
probable Hygrocybe californica © Danny Miller, Andrew Parker
Hygrocybe 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 European species nor even that closely related (not sister species). The second, less common species, has an all-yellow stem in the one photograph I have.
Hygrocybe cf coccinea #1 © Steve Ness, Hygrocybe cf coccinea #2 © Steve Ness
Hygrocybe aff aurantiosplendens
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 but isn't (with splotches of red, orange and yellow on the cap).
Hygrocybe aff aurantiosplendens © Steve Ness
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 laetissima © Lauren Ré
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, so it's still up in the air if ours is the same species. East coast specimens seem to be clearly distinct.
Hygrocybe 'punicea', California © Christian Schwarz
Hygrocybe aff 'mucronella/reae'
It is not looking like ours is the same as the European H. mucronella (also called H. reae), but instead a sister species that needs a new name. It is yet another 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!
Hygrocybe aff mucronella © Steve Ness
Hygrocybe cf minutula
The type area of H. minutula is New York. One of its synonyms is Gliophorus minutulus, because it is reported to be 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. Something that looks somewhat similar (but we could be very wrong about this, and it might not be related at all) was found twice in Washington at sea level. It was quite small and looked wet, but was not necessarily viscid all over. More specimens need to be studied to determine the viscidity of our species, and if reports of H. minutula represent this species or something completely different. If not, very little is known about this species.
H. 'subminiata' has been reported a few times from Washington, but we don't have DNA or even a photo of a local collection of that Jamaican species to know if that's one of the species we have here. It is rumoured to be related to the mushrooms in the H. coccinea group, but that can't be confirmed. It is quite small (less than an inch across) with a viscid red cap and yellowish stem and pale gills (that are decurrent). It would be easily confused with H. cf minutula.
Hygrocybe cf minutula © Steve Ness
Hygrocybe fenestrata np
A smallish mushroom which is 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. 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, but for now it is "np", or a proposed name only. It was thought to belong in Gliophorus because of it's extremely viscid nature, but it too belongs in Hygrocybe.
Hygrocybe fenestrata np © Steve Ness
Poorly understood other species
Hygrocybe sp. MO336328 - this matches the DNA of a mushroom that was identified back east as H. parvula, where H. parvula is described from. I don't know how much that one looked like H. parvula, but I know that our specimen certainly does not. It's a yellow, decurrent waxy and not much more is known about it (at first glance it looked like Glioxanthomyces nitidus).
Hygrocybe sp. MO336328 © Steve Ness
Hygrocybe sp. iNat19620764 - yet another red-orange small species. We need to know its gill attachment, viscidity, taste, etc to even begin to tell it apart from the many similar waxies.
Hygrocybe sp. iNat19620764 © Lauren Ré
Hygrocybe aff ceracea - fairly closely related to H. ceracea, but red instead of orange. It has been suggested that it might resemble the European mushroom Hygrocybe moseri in some ways, but at this point everything is just a guess. I know of no way yet to recognize it.
Hygrocybe aff ceracea (red) © Steve Ness
Locally described species that are probably legitimate but poorly understood
Hygrocybe 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. No DNA is available of anything resembling it yet.
Hygrocybe 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). We need samples.
Hygrocybe albicarnea - 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 yet.
Species reports without genetic data yet
Hygrocybe 'atro-olivacea' - 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.
Hygrocybe cf atro-olivacea © Renée Lebeuf
Hygrocybe 'parvula' - Unlike sp. MO336328 above, we do have reports that look exactly like this east coast species. It has 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 and H. sp MO336328 could be related, we'll have to see.
Hygrocybe cf parvula © Buck McAdoo
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