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Danny’s DNA Discoveries – Hygrophoraceae (Waxy Caps)
by Danny Miller
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 (from now on referred to as H&S), in the most important book on this family in North America, North American Species of Hygrophorus from 1963, 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.
'Hygrophorus' caeruleus - Clitocybe odora
There was always something strange about this rare, beautiful, pale blue conifer mushroom, described from ID but also found in WA and OR. The gill hyphae was parallel like Hygrocybe, which it clearly wasn't, and the basidia were a little short, always <45u, but based on its appearance as a waxy cap of some kind it was named Hygrophorus caeruleus without knowing at the time what genus of waxies it really belonged to, as Hygrophorus used to be the catch all genus for all waxies. DNA results surprisingly came back with both the ITS and the LSU DNA regions practically identical to Clitocybe odora (maybe one base pair substitution out of over 1300, or 99.93% identical). Perhaps some environmental phenomenon sometimes affects Clitocybe odora to make it waxy and lose most of its black licorice smell. (Although it is still sometimes faintly fragrant and the original description was that it smelled disagreeable, nobody has reported the famous black licorice odor in one). Or maybe this is one of those rare cases where significant changes in other DNA that affects how the mushroom looks and smells occurred without any changes to the supposedly faster changing ITS and LSU regions. (Another example is Agaricus mesocarpus and Agaricus inapertus, where one is secotioid, but I will get to that in my Agaricus report). It is uncommon but not rare that enough other DNA changes to make a new species (without ITS and LSU changing), but that usually affects things like habitat and they usually still look almost identical. So it might be a unique species, but we'll still have to call it a waxy Clitocybe. The name Clitocybe caerulea is not taken, so that is available for a recombination, if necessary, but it may be decided that it will continue to be called an odd-looking Clitocybe odora.
'Hygrophorus' caeruleus (Clitocybe odora) © Noah Siegel
Fibrillose partial veil - these species have an actual fibrillose partial veil leaving fibrillose strands around the stem almost like a ring.
Hygrophorus subalpinus group
White and stocky, this is a common veiled conifer spring snowmelt species (found near melting snow at high elevations, but also apparently found in fall), described from WA. We have Oregon DNA from the fall and California DNA from the spring, 3bp apart, of a first species, but no Washington DNA yet.
A second species from WA in fall and boreal Quebec in September has 10% different DNA, the one specimen photographed seemed less stocky without as obvious a veil, but let's not read too much into one photograph yet.
Hygrophorus albiflavus - a slender, white waxy with a fibrillose veil. Viscid stipe. Only known from the OR type. Unknown trees. Large spores.
Reports of Hygrophorus sordidus/penarius from the PNW may represent finds in the Hygrophorus subalpinus group if the fleeting veil is not noticed. Another theory is Hygrophorus marzuolus, below. Any reported finds should be sequenced to verify which species occur here.
Hygrophorus subalpinus #1 and #2 © Jonathan Frank and Danny Miller
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 many 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 officially all one species.
Hygrophorus siccipes © Michael Beug
A species from Quebec is very similar and also with pines, but more consistently slender and with duller colours that tend to fade, and its DNA has been found in CA and boreal BC. It may occur here, as H&S reported both robust and slender forms from the PNW, so we need to look for it.
possible Hygrophorus boyeri © Kit Scates Barnhart
Hygrophorus 'purpurascens' group
Unevenly purple and somewhat stocky, the gills and other parts spot and stain even more purplish over time. It is a common spruce and pine mushroom, notably found often in spring as well as fall (we need spring collections to see if they match the DNA of fall collections). Several similar species are purple and 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 and is probably our common species, but it doesn't match DNA from Europe, where it was described, so our species needs a new name. We appear to also have a second undescribed species, found once in OR (>10% difference in ITS). It definitely spotted purple on the gills.
Hygrophorus velatus - this supposedly rare mushroom is only known from the type in Idaho, collected in August. 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' that hadn't stained yet and one of the above unnamed species should be going by this name. We need specimens to determine how many additional new names we need for our H. purpurascens mushrooms and to understand how to recognize this species better if it isn't one of the above.
Hygrophorus aff purpurascens © NAMA and the Field Museum of Natural History, Hygrophorus aff purpurascens #2 © Jonathan Frank
Hygrophorus cf fuscoalboides/aff korhonenii
Hygrophorus olivaceoalbus is the name being used for most collections in this group, a mostly grey and stocky spruce species with big spores (>10u). The fibrillose veil is said to be black and the stem is viscid. It is fairly common in ID, CA and CO, but rare in the coastal PNW. However, we have not found the DNA of this European mushroom in the PNW (except for one possible record in boreal BC), so we are not using the correct name for our collections. 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 from Europe or any local specimens sequenced, so I don't know what it is and if it is actually distinct. Hygrophorus fuscoalbus is a rare species that reportedly has a white fibrillose veil instead of black. Nobody is sure what this European species is, even in Europe, so the name may not be used going forward. Something that might be it is sequenced in France and Quebec, so that could possibly be the species, but we still haven't found that DNA in the PNW.
So what species are actually here? Hygrophorus fuscoalboides is a similar species with a grey fibrillose veil and dry stem was described from Idaho (only 2 records including the type, no trees noted), so it is actually here. Given that stem viscidity in other species has turned out to not be that important a defining character, this could be the proper name for many of our local collections with viscid stems. One collection from Michigan and one from somewhere in Canada were identified as H. fuscoalboides (they may not be identified correctly), and those sequences match a sequence from boreal BC. That DNA is 4bp and 2 indels from the newer European species Hygrophorus korhonenii. The one western WA sequence I have of a mushroom in this grouip is 3bp and 1 indel from the purported Hygrophorus fuscoalboides sequence in northern BC.
In summary, if we only have one genetic species, it is likely Hygrophorus fuscoalboides. If I am wrong and do not understand what that species is, then we do have whatever it really is, at least in Idaho where it was described, but we also have Hygrophorus aff. korhonenii (it may or may not be the actual species as it only differs by 5 bp or so, not a whole lot for this genus). We need additional samples with stocky vs slender statures and white vs grey partial veils and different amounts of stipe viscidity to clinch the matter.
Hygrophorus cf fuscoalboides © Danny Miller
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. Interestingly, we have DNA from Europe, but no local DNA to see if they have our species or an undescribed different species.
Hygrophorus inocybiformis © Ben Woo
Purple species with Gills etc. spotting pink/purple
Hygrophorus 'purpurascens' group has already been discussed in the veiled section above.
Hygrophorus cf erubescens group
This common irregularly coloured purple spruce and pine species seems to often have yellow tones. We need local specimens to see how our species compares to the European species.
Hygrophorus erubescens var gracilis from Oregon is a locally described variety, but the type DNA is not close to the European species so it is probably not a variety and needs a new name. It is rarer, more slender, has more distant gills and usually lacks the yellow tones. These two species don't appear to necessarily be related to the other purple staining species.
Hygrophorus cf erubescens group © Buck McAdoo
Hygrophorus aff capreolarius
This not uncommon spruce species has more evenly distributed purple colours than the others (less splotchy, so to speak). One WA specimen matched DNA of a Quebec specimen and was different that a European sequence, so we appear to have a sister species in North America that needs a new name.
Hygrophorus aff capreolarius © Danny Miller
A buff to salmon to yellow tinged cap instead of the usual purple cap, bitter tasting species with yellow gills that spot pink. The type is from WA and this may represent a duplicate of one of the above species, so we need specimens. It is rare, only known from the type, plus 1 specimen in ID and 1 in WY. Spruce and Douglas fir. Never photographed in colour.
This is a rare hardwood species with slightly closer gills than others. We also need specimens to see if this is the same as the European species, although there may be more than 1 European species going by that name, so that makes it a little less likely that ours is the real thing. H&S found it only once in WA, and there have been a few reports since then.
Hygrophorus cf russula © Noah Siegel
Sweet Almond/Cherry Odor (specifically, not just sweet or fragrant)
Hygrophorus agathosmus group
Uniformly grey cap and white to pale grey stem, somewhat stocky (although the pictured collections are not) spruce and pine species (and maybe Douglas fir, but it should be checked if those collections represent H. odoratus, below). It has regular sized spores. The DNA of this European species has shown up in BC and WA. DNA of a sister species, about 8 bp and 2 indels different than that found in the EU has also shown up once in WA and twice in southern OR, so we may have a cryptic sister species as well. DNA throughout Europe varies by as much as 2bp and 7 or more indels, but not quite as much as 8bp and 2 indels.
Hygrophorus agathosmus and aff. agathosmus © Danny Miller
This is a very similar, also uniformly grey capped mushroom with conifers, with a hint of yellow in the cap and a whiter to pale yellow stem. It is supposedly usually a bit more slender. It was described as being more uncommon than H. agathosmus, but they are easily confused, so we will have to find out. It has large spores. 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 © NAMA and the Field Museum of Natural History
This very common conifer waxy has a brown disc with a paler rim. It was described from WA. We don't have the type sequence, but sequences from WA and BC probably represent this species. It has regular sized spores.
Hygrophorus variicolor - a rare (except in California) waxy, 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. This is the older name, but H. bakerensis is the name commonly used.
Hygrophorus bakerensis (variicolor?) © Steve Trudell
Hygrophorus secretanii (monticola)
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 - very similar (with the same large spores) but supposedly tastes bad and has somewhat pink gills and a pink pruinose stem. It was 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 - similar but said to have a different non-almond sweet odor and is described below.
Hygrophorus secretanii (monticola) © Andrew Parker
White or Colourful Caps (not brown, grey or black)
Hygrophorus piceae group
White, usually 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. There are at least 4 competing concepts in Europe for which species is the real thing, so we don't know for sure.
What's interesting is that two species have been found in WA, and although they look identical, they do not appear to be 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 Hygrophorous tree. One one the two might be the real thing (or is at least somewhat related), but the other one isn't. We just don't know which is which. Both might require new names.
Hygrophorus piceae #1 and #2 © 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 with yellow shades. They are not all related to each other.
Hygrophorus eburneus - this common, white, mid-sized, mostly conifer species is a European mushroom and we need local specimens to see if our species is the same.
Hygrophorus cossus - 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 - 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 - 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 - European species with spruce/pine, not uncommonly reported in Oregon and Idaho, stocky, yellowish cap when fresh. We have European DNA but need local specimens to compare. H&S say that our species seems a little different than in Europe.
Hygrophorus flavodiscus - eastern species, midsized, yellow disc, with pine, also not uncommonly reported in Oregon and Idaho. Sister species to H. gliocyclus. We have ENA DNA but need local specimens to compare.
So with a half dozen similar looking species reported in the PNW, which are there DNA evidence for? None. So far we have 2 genetic species from local collections, not matching anything else at the moment, so possibly unnamed, 1 from WA and OR and a second only from an environmental sample in OR.
Hygrophorus cf eburnius © NAMA and the Field Museum of Natural History, H. cf gliocyclus © Kit Scates Barnhart, H. cf flavodiscus © Noah Siegel
Hygrophorus 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.
See the discussion below under Hygrophorus aff marzuolus.
Hygrophorus aff chrysodon
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 not uncommon and found with conifers, but Washington DNA is clearly a different species than the EU, so ours needs a new name. Interestingly, California DNA appears to be >3% different from WA DNA so far, and may represent a third species.
Hygrophorus aff chrysodon © Steve Trudell
A white, cespitose species, not as stocky as H. sordidus (below, but a lot like the ENA H. subsordidus). Stipe twisted striate, white tomentose at base. Only known from the type from Idaho. Pine and fir.
Small, aromatic, white waxy, sometimes with a pink disc and a viscid cap. Not uncommon, with conifers. Our local DNA seems to match the LSU region of a reliable eastern North American sequence (where this species was described) so it looks like we have the real thing.
Hygrophorus 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. Known only from the Oregon type, no sequence yet.
Hygrophorus pusillus © Kit Scates Barnhart
A small, pale pink snowbank, uncommon species with large spores like H. vernalis, and also with hemlock. Described from Oregon.
Hygrophorus goetzii © Steve Trudell
White cap, bright pink gills and a peachy odor, described from Oregon, not uncommon, with conifers. We don't have the type sequence, but sequences from Victoria and boreal BC match and probably represent this species.
Hygrophorus saxatilis © Michael Beug
Hygrophorus 'pudorinus'/fragrans group
These have a pale pink to orange viscid cap, punctate upper stem, may smell sweet and are larger than H. goetzii (below) with closer spaced gills than the dry capped Cuphophyllus pratensis.
There seems to be consensus what the common H. pudorinus var fragrans is. It is said to bruise yellow-orange and have slightly coloured gills and a stronger odor. It is sometimes considered a variety of H. pudorinus and sometimes considered its own species as H. fragrans. The type is from Oregon. We have sequences from near Victoria, BC.
We have a sister species in WA and boreal BC (8bp and 2 indels different), but I have no idea what the real European H. pudorinus var pudorinus is to know if that's it or not. They seem to prefer spruce or maybe pine. H&S reports that var fragrans is our more common species, and var pudorinus is a Rocky Mountain species (found mostly in Idaho and other Rocky states). We shall see.
Hygrophorus cf pudorinus (sister to var fragrans) © NAMA and the Field Museum of Natural History, H. pudorinus group © Andrew Parker
Bright red-orange cap, but white gills and stem. Very slimy everywhere and a fleeting slime partial veil. It favours larch.
The type of H. speciosus var kauffmanii (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
Brown, Grey or Black on the Cap (at least in age on the disc)
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 but not H. bakerensis. It is not uncommon in California but rare here (reported once each from WA and ID).
possible Hygrophorus pacificus © Ben Woo
A brown capped spring snowmelt species (otherwise like the aromatic H. secretanii/monticola and also with large spores, but with a viscid stem), under hemlock. This WA collection is the first DNA documentation of this WA species. BC DNA matches it fairly closely (2/0-5). Fairly rare.
Hygrophorus vernalis © Sava Krstic
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. Rumour has it our DNA is different than in Europe so our species may need a new name but I need specimens to confirm. (H&S noted our specimens have shorter spores than in Europe). The same DNA as Europe was found in Quebec, though, although that specimen resembled H. morrisii, below.
Hygrophorus occidentalis - this California species, found not uncommonly there and also 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, if the same species as out west, is related to H. eburneus. We don't have any local DNA to tell and no known colour photographs.
Hygrophorus 'discoideus' © Andrew Parker
Hygrophorus aff tephroleucus (aff pustulatus)
Small, greyish cap, sometimes pointed, with notable dark scabers on the somewhat viscid stem. Formerly considered two species based on just how slender it is, they are now considered just one species, and perhaps H. tephroleucus, being described 3 pages earlier in the same 1801 publication will get priority. Uncommon, preference for fir? Our species seems to be 6 bp and 6 indels different from European specimens, so could very well be an undescribed sister species.
Hygrophorus morrisii - 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), but we don't have good sequences from either area to know what it is or if we really have that species here (one Quebec sequence was really H. discoideus).
Hygrophorus aff tephroleucus (aff pustulatus) © Andrew Parker
Hygrophorus 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. Our species is a sister species to the true European species, at an 83% match.
Hygrophorus sordidus/penarius - 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. aff marzuolus 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 aff marzuolus © Josh Powell
Fall species with blackish caps even when young, also stocky. Not uncommon 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 camarophyllus © Andrew Parker
Hygrophorus cf calophyllus
A similar blackish capped European mushrooms but with pink gills, uncommon under fir and pine. No DNA from Europe nor here yet to say if we have the same species.
Hygrophorus cf calophyllus © Danny Miller
An uncommon conifer species, olive-grey brown with a viscid stem, described from Washington (like H. olivaceoalbus, but no fibrillose veil), It is practically unknown, but with giant spores (even bigger than the H. olivaceoalbus grp) it should be easy to detect if found again to figure out what it is.
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 not uncommon, but only found there. Conifers.
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