Danny’s DNA Discoveries – Squamanitaceae of the PNW
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.
These white spored mushrooms, with attached gills, resemble the free gilled 'Lepiota', and were once thought to be closely related to them (having re-evolved attached gills), but now they appear to be further away in their own family in the dark spored Agaricineae, meaning there could have been an additional event where spore pigment was lost in that sub-order (along with the events where the 'Lepiotas' and Laccaria lost their spore pigment).
Like Lepiota and allies, these usually have scales on the cap that can't be removed. All species seem to have some sort of obvious partial veil. Most are saprophytic. These genera may be difficult to differentiate from other scaly genera, like Tricholoma (which more often has specifically notched gills and usually lacks a partial veil).
Cystodermella may be off by itself and not inside this family. To complicate things further, one genus re-evolved coloured spores (or never completely lost all their pigments) - Phaeolepiota (pale yellow or orange brown spores).
Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
Cystoderma and Cystodermella - click to expand
Cystoderma usually have amyloid spore walls (turning black in iodine) and Cystodermella don't. They look very much the same, but are not as closely related as you might expect. Cystodermella is not in the same family as the other "attached gilled" genera, and may be in its own family.
Species mentioned: Cystoderma amianthinum, Cystoderma jasonis, Cystoderma carcharias, Cystoderma fallax, Cystodermella granulosum, Cystodermella cinnabarina (terryi), Cystodermella adnatifolia
Cystoderma amianthinum group - This is the palest, brightest orange to yellow-orange coloured mushroom group in the section. The gills and cap flesh are almost pure white. It is thought to only leave a trace of a ring, but some specimens have had a fairly substantial ring and been mistaken for C. carcharius, below. There are two closely related genetic species in the EU (the second of which may be the real thing if an epitype sequence is to be believed). We may have 3 species in the group, none the real thing.
#1 - a sequence from WA matches EU species #1 quite well, and a sequence from BC differs by 4 bp in ITS1 only.
#2 - our one WA sequence is 6-8 bp from EU species #2 (the one that might be the real thing).
#3 - our most common species, that often has a reticulated cap like var. rugosoreticulatum, although that is a European variety and the DNA of our common species has only been found here in the UK so far.
Although some varieties of C. amianthinum were promoted to species level because they are genetically quite distinct (var. longisporum to C. jasonis, below, and var. sublongisporum to C. intermedia) our group members are closely related and could potentially remain as varieties. We need more collections to see how to tell these species apart.
Cystoderma intermedium's European type sequence is the same as Cystoderma carcharias below. Cystoderma amianthinum var. sublongisporum was reported from the PNW and declared a synonym, but I don't think it necessarily is, so it should be investigated what the genetics are of var. sublongisporum.
Cystoderma amianthinum #1 and #3 © Danny Miller
Cystoderma jasonis - Very similar, perhaps less likely to be found in moss, perhaps a little darker in colour and with a brown tint to the gills and cap flesh. It is so similar to the C. amianthinum group that it was never noticed to be present in the PNW until a UBC student found and sequenced one. The biggest difference is microscopic - longer spores. It used to be considered a variety of C. amianthinum (var. longispora) but it's not closely related enough to be a variety. Considered rare for now, but maybe not, as nobody has been looking for it. There is an albino form in Europe that should be looked for here.
Cystoderma jasonis © Noah Siegel
Cystoderma carcharias (Cystoderma carcharias var. fallax)
This species has the most well developed ring of any species in this group. Cystoderma carcharias is a rare white EU mushroom, and Cystoderma fallax is a common orange-brown WNA mushroom that was found to be identical except for colouration, so it was reduced to a variety, Cystoderma carcharias var. fallax. They can have identical ITS DNA, so thinking of it as only a variety might be the proper nomenclature, or perhaps even just a colour form. European sequences of both colours may vary from each other by as much as 3 bp, but most consider it all one species. Our local PNW var. fallax sequences are an additional 2-3 bp different. So far I am considering them all the same species. The ring can wear off making it resemble other species.
Cystoderma carcharias (var. fallax) © Steve Trudell, (var. carcharias) © Boleslaw Kuznik
'Cystoderma' gruberianum - This small (<3cm) clay-brown capped mushroom is found on Doug fir logs instead of on the ground, has a conic cap, and has very large amyloid spores compared to other species. It was described from Oregon, and is very rare (not found again since it was described in 1949). I'm not convinced it's actually a Cystoderma but somebody will need to find it again or sequence the type to figure out what it is.
Cystoderma gruberianum © University of Michigan
Cystodermella granulosa (Cystodermella granulosa var. ambrosii) - Cystodermella granulosa usually has a darker orange-brown to reddish-brown cap than Cystoderma jasonis. It has no extra sterile cells on the gill edges (cheilocystidia), unlike Cystodermella cinnabarina, below. It has inamyloid spores, like all Cystodermella. Our local sequences are 3bp and 3 indels different than European material, but so far considered to be the same species. This common species is also rarely found with a white cap, called Cystodermella ambrosii, but does not have any different ITS DNA when it does, so it has recently been redescribed as a variety, Cystodermella granulosa var. ambrosii, and may even more correctly be considered just a colour form.
Cystodermella granulosa © Andrew Parker
Cystodermella adnatifolia - This east coast species is also found out here (sequenced collections from near Mt. Rainier and near Stampede Pass in WA). It usually has a more brightly coloured cap than the very similar C. granulosa, and the same lack of cheilocystidia and inamyloid spores. It is probably uncommon, or very difficult to tell apart from C. granulosa in practice, since it has gone unnoticed.
Cystodermella adnatifolia © NAMA and the Field Museum of Natural History
Cystodermella cinnabarina - Also more brightly coloured like C. adnatifolia, this common species has large sterile cells on the gill edges, giving them a fringed appearance under a hand lens if you don't have a microscope. One OR sequence is an exact match except for a few ambiguous locations and a couple of WA sequences are 2 bp different from EU material, so there is strong genetic evidence of this species here.
Cystodermella cinnabarina © Lauren Ré
Cystodermella subpurpurea - Described from Oregon, so it's definitely here, but it's another rare species not found again since it was originally described in 1948. It has a very small (<1cm) grey cap with a purplish tinge. We need to find it again or sequence the type to find out what it really is.
Phaeolepiota - click to expand
Species mentioned: Phaeolepiota aurea
Phaeolepiota aff aurea - It is quite a wonder to find one, but it is fairly uncommon. It appears to be a sister genus to Cystoderma. Two Washington sequences are identical to each other, but 2% different than more than a dozen European sequences, so our species appears to be unique and needs a new name. Occasionally reported with free gills.
Phaeolepiota cf aurea © Steve Trudell
Floccularia - click to expand
Species mentioned: Floccularia albolanaripes, luteovirens
Floccularia albolanaripes - Described from Oregon, this mushroom somewhat resembles a veiled Tricholoma. It has bright yellow scaly cap often with brown on the disc, and a very shaggy stem.
Floccularia albolanaripes © Andrew Parker
Floccularia cf luteovirens - This European species is supposedly even scalier on the disc (bright yellow scales) and has been reported from the PNW, but perhaps only once since the 1950's. We need collections to prove whether or not it's actually here, or if these represent reports of F. albolanaripes.
Leucopholiota - click to expand
Species mentioned: Leucopholiota decorosa, lignicola
Leucopholiota sp. - Leucopholiota decorosa was described from NY, and is one candidate for what our species might be. Another is the boreal species Leucopholiota lignicola, described from Scandanavia, since all 4 collections found so far in the PNW are from about 51 degrees north or higher in BC. We need both local sequences and Scandanavian sequences to find out.
Leucopholiota sp. © Paul Kroeger
Squamanita/Dissoderma - click to expand
Squamanita has now been split into Squamanita and Dissoderma, which the larger species that have purple tones being moved into Dissoderma. If they were not split, Leucopholiota would have to move into Squamanita. They are very rare, very cool parasites on other mushrooms that seem to turn the mushroom into a hybrid, with the attacked mushroom on the bottom half and the purple-grey Dissoderma on the top. We do not have any known Squamanita in the PNW, only Dissoderma.
Species mentioned: Dissoderma paradoxum, contortipes, pearsonii, odoratum
Dissoderma odoratum EU - growing clustered with a notable sweet, grape odor. It parasitizes Hebeloma mesophaeum. We have one old sequenced collection from WA.
Dissoderma paradoxum OR - Greyish-purple parasite on Cystoderma amianthinum and Cystoderma carcharias. Note that the bottom half of the mushroom still looks like Cystoderma and the top half is a Dissoderma. The base is swollen and scurfy.
Dissoderma pearsonii UK - This species is very similar to D. pardoxum but lacks the scurfy swollen base and differs microscopically. It has been collected and sequenced once in WA in 1997.
Dissoderma stuntzii WA - differs by 7% in ITS, but otherwise cannot easily be distinguished from D. pearsonii. Known only from the type collection from 1975.
Dissoderma contortipes - Our smallest species (<1cm across) with almost round spores, mycenoid, greyish-purple on Galerina species. Known only from the type collection from 1993.
unsequenced Dissoderma paradoxa © Kit Scates Barnhart
Summary of Future Studies Needed