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Danny’s DNA Discoveries – Amanitaceae
by Danny Miller
Amanitaceae are cool.
These species have no partial veil (although the powdery grey A. 'farinosa' looks like it doesn't have a partial veil but belongs elsewhere). A. velosa is in this section but sometimes appears to have a partial veil. The volva is a sac. Brown or grey striate caps.
Our species in this group are not generally well understood. Sometimes the sac volva is white, sometimes it is grey (maybe only on the inside) and sometimes it has rust coloured stains. I have not yet been able to determine how any of these features indicate species yet, as they do not appear to be consistent. Even whether or not the cap is grey or brown does not always help indicate what species you have, as many species are found with both colours.
Amanita constricta © NAMA and the Field Museum of Natural History
This is a grey or brown capped species where the bottom of the volva is constricted against the stem, but other species may also have this feature, so I do not know how to identify it versus other undescribed local species mentioned below. It was described from California and its DNA has also been found in Washngton and BC so far.
Amanita pachycolea group
Amanita pachycolea group © NAMA and the Field Museum of Natural History
This is a large species, described with a large unconstricted sac volva, but other undescribed species have these features too, and specimens in this group have been found with constricted volvas as well. It was described from California. We have three genetic species in the west. The most common one, found so far in OR, WA and BC, is sometimes referred to as "NW04". A second species has been found in northern BC, Idaho + Wisconson (shown above). A third species has been found in Mexico. It is unclear to me if the "real" Californian A. pachycolea is NW04, the Mexican species, or something else, as we don't have any sequences of the type. But since all sequences of specimens identified as A. pachycolea were in the same clade of 3 species, I am going to assume that the "real" A. pachycolea is in this clade somewhere.
Similar Unnamed Species
We do not have the poorly understood Amanita vaginata in the PNW. But everytime a species is found that does not match one of the above two descriptions, and does not match the description of one of the distinctive, rare, species below, it is usually mis-labelled A. vaginata for lack of a better name. Here is a confusing look at some of the other unnamed and little known species we have genetic evidence of.
Amanita castellae n.p. © Buck McAdoo
This is described as possibly being paler than A. pachycolea with a dark ring around the cap near the inside of the striations. This photo shows neither of those characters. Found in Washington and outside the PNW.
Amanita "NAMA2018-OR1" © Danny Miller
One collection had a cap with a metallic colour and sheen, both collections had white stems. It was of modest proportions. Known from Oregon.
Amanita sp. "NAMA2018-OR2"
Amanita "NAMA2018-OR2" © Stephen Russell
This one had an especially brown cap, and an orange stem. The sequence was dirty in places so we'll want more sequences to be sure of its place in the tree. Found once in Oregon.
Amanita "NAMA2008-ID01" © NAMA and the Field Museum of Natural History
Found once in Idaho.
Amanita subnigra n.p.
Amanita subnigra n.p. © NAMA and the Field Museum of Natural History
Known from eastern North America and twice from Idaho. Only 2bp different than A. lindgreniana n.p. (next) but to be considered a separate species.
Amanita lindgreniana n.p. © Sava Krstic
This large species had a white, non-constricted volva. It is known from Washington, California and Oregon.
Amanita "sp-OR" © Buck McAdoo
These species are always brown or tan capped, at least once one was recorded with a white, partially constricted volva. The DNA is hard to read so I'm not sure if they represent distinct species or not. The DNA of the photographed collection is 4bp and 1 indel different than another Genbank Oregon collection, and other dirty sequences fit near these. Found in Oregon.
Amanita aff. fulva
Amanita aff fulva © NAMA and the Field Museum of Natural History
Found in Washington and BC, this brown species is somewhat related to the always brown Amanita fulva of Europe.
Rare but distinctive species
These species are distinctive enough that they might be recognizable.
Amanita 'velosa' © Daniel Winkler
This bright brown capped species often looks like it has a hint of a partial veil. It is very common in California with oak and can also be found in southern Oregon with oak. The Oregon sequence is 3bp and 1 indel different from California sequences I have so far. I am assuming there is only one species, but that needs to be verified.
Amanita populiphila © Michael Beug
This Kansas species with a tan cap and a stem that may widen is found in Idaho under Populus.
Amanita alaskensis n.p.
Amanita alaskensis n.p. © Buck McAdoo
This species, found in Alaska, Washington and probably BC has a dark grey cap and stem, with warts that remain on the cap, unlike most other species in this subgenus.
Amanita protecta © Noah Siegel
This very pale grey, stocky species from California may also leave warts on the cap. It is found with willow and oak in southern Oregon. Other whitish stocky species exist, though.
Other possible species: Amanita cf constricta from Northern BC known from Genbank, Amanita sp-OR03 a large white species with no ITS sequence yet and Amanita sp-NW05 easily confused with A. constricta and A. pachycolea.
None of the above characteristics. Universal veil may leave warts on the cap. Volva is often concentric circles or a collar. Rarely, a furry grey mushroom with no visible veils. Most mushrooms in this group are poisonous as well as hallucinogenic, in a way entirely different from "magic mushrooms" (Psilocybe). The most common symptoms after ingestion are vomiting and other gastrointestinal distress, delusions of grandeur, temporary coma-like state and amnesia. The toxins are water soluble.
Amanita muscaria and Amanita chrysoblema
Amanita muscaria © iNaturalist user naomimoriyama1, Amanita chrysoblema © Steve Trudell
Amanita chrysoblema orange, yellow and white forms, © Steve Trudell, Janet Lindgren and Steve Trudell
These are the famous Alice In Wonderland/Mario Brothers mushrooms with colourful caps (red, orange, yellow or white) with pale, removable warts on the caps from the universal veil, or "egg" that the mushroom appears to hatch out of.
The true Amanita muscaria is scarce in the PNW, usually found under introduced trees with almost pure white warts.
Amanita chrysoblema, our very common, spring and fall ubiquitous species found under native conifer and hardwood trees usually has yellowish warts (except the white form which usually has no colour in the warts either). What to call it was somewhat of a puzzle. It is too genetically distant from A. muscaria to be considered a variety, although we have some variety names for it. Traditionally, the different colour forms (red, orange, yellow and white) would each get their own variety or subspecies, but now we know there is no genetic difference between the colours, so that is not valid. (It would be valid to give each of them their own form, though. That simply means they look different and does not try to imply any genetic relationships).
First, we have to decide how many species are in the group. There are 4 clades of North American muscaria-like sequences that differ from each other by only 1 or 2 bp in ITS. That is usually not enough to be considered separate species, although that does happen if there are clear ecological, morphological or microscopic differences as well. If they are to be kept separate, here is how they work out:
Eastern North America probably has examples of three of the four clades, but the only clade found in the PNW so far is clade 4. That does have a valid name, Amanita chrysoblema. Two names were given to white forms, one of them at the variety level (var alba) and one at the species level (A. chrysoblema) so even though the variety name is older, since this needs to be a distinct species from A. muscaria, the newer species name A. chrysoblema is a valid name for our species. Remember, the colour turns out to not matter, so that is a valid name for all of our colour forms.
A. chrysoblema is also the oldest name in all 4 clades, so if they are considered the same, that name should apply to all of them. If they are to be considered different species, then things get more complicated. None of the existing names are distinct species names, they are all varieties and subspecies, so they will all need new names of their own or need to be designated varieties or subspecies of A. chrysoblema instead of A. muscaria. Luckily, here in the PNW, we don't need to worry about that.
We do have a tradition of calling our red forms here in the PNW by the name A. muscaria var flavivolvata and our yellow forms by the name A. muscaria var formosa. If all four clades are to be considered the same species, A. muscaria var flavivolvata is one technically correct name for all of our colour forms, just not the best name (the best name is A. chrysoblema because that name does not incorrectly imply that our species is a variety of A. muscaria. A. muscaria var formosa is a name for European yellow capped forms, but as I stated, likely represents the exact same species as A. muscaria, and is not a correct name for our common local species.
Amanita pantherina/gemmata group
These similar mushrooms can also be found in spring and fall, have yellow or brown caps (a slightly different shade of yellow than A. muscaria and A. chrysoblema sometimes have) and a collar volva instead of concentric rings. We do not have either of these European mushrooms in the PNW, but here's what I have been able to determine about what species we do have:
Amanita pantherindoides © iNaturalist users loather and cyndean
In Mycologia Volume 4 from 1912, Murrill listed all the known Amanitas on the west coast. Not a lot was known back then, so there are of course many species missing, but I imagine it is likely that the species he does include are common species here. Here are the species he described from Seattle that could represent A. pantherina/gemmata type species:
A. praegemmata - cap only to 6cm, persistent warts, honey-avellaneous (grey- to pinkish- brown) to dark honey on the margin, white partial veil, stem 1.5cm thick, volva attached rim.
A. pantherinoides - cap thick, small numerous warts, honey to dirty cream with brown center, white partial veil, stem 2cm thick, volva entire or undulate free rim
A. umbrinidisca - cap fleshy drying thin, large irregular patches, honey to straw w/umber disc, white partial veil, stem 1-2cm thick, volva sub-entire free rim
In 1977, Jenkins studied the types microscopically and synonymized A. pantherinoides and A. praegemmata, saying they were likely the same thing, our local panther species, with A. pantherinoides being the older name (it was published one paragraph earlier than A. praegemmata). He did not even consider A. umbrinidisca, because he thought it represented a very different kind of Amanita, but subsequently, others have figured it probably is similar to, if not the same species as the others. When we sequenced a couple dozen or so Seattle area A. pantherina like species, they were all the same species (we only have one common species).
What does this mean? If we assume that Murrill did not find something exceedingly rare that has not been found since (and at the same time somehow avoid finding the very common species), and if we assume that Jenkins was right and at least two of his species are the same, I believe Amanita pantherinoides is the correct name for our local panther species. But I cannot rule out the fact that one of Murrill's species might be our local gemmata-like species. That is a very common species, so why doesn't it appear in his list?
But for now the mycological community has decided to assume that all three are the same and they refer to our panther-like species.
Amanita ameripanthera is another name that is being put forward for our common local species, based on the fact that it is often dark brown instead of the "honey-avellaneous" and "honey to dirty cream" that was in the Murrill's descriptions. But there was a wide variety of cap colours in our sequences, from yellow to brown, and they all sequenced the same, so this appears to be another case of cap colour not being critically important. This would mean that A. ameripanthera does not represent a unique species and the name would not be needed.
Traditionally, it was thought that brown capped species were the panther species, and yellow capped species were the gemmata species (covered next), but that's not the way it works out for the two common species in the Seattle are. Amanita pantherinoides is yellow to brown, stocky, with persistent warts on the cap. Our gemmata species, Amanita pseudobreckonii n.p., below, is usually yellow, slender, and has warts that easily wash off.
Amanita pseudobreckonii n.p.
Amanita pseudobreckonii n.p. © Buck McAdoo
This very common species is always yellowish, usually slender and the warts wash off pretty easily. California mycologists have this tentative name for one of their local gemmata-like species. They have more than one common local species, but since our local species is a genetic sister species to A. breckonii, that particular name may also represent our species. I am looking for California sequences to verify this.
Alternatively, it is possible that one of Murrill's three species is not the same as the others, but represents our common gemmata-like species. If so, that name from 1912 would be the proper name for our species, not A. pseudobreckonii n.p. We will need type sequences of his original 1912 collections, if we can get them, to settle the matter.
Other, Southern "panther" and "gemmata" species
In Oregon and California, we have additional species not yet found in British Columbia or Washington.
Amanita "gemmata-NAMA2018" © Corinne Srsen, Amanita "stocky gemmata" © NAMA and the Field Museum of Natural History
Amanita "gemmata-NAMA2018" is another yellow, slender species with warts that easily wash off in the gemmata-panther group found in Oregon and California. I don't yet know how to tell it apart from Amanita pseudobreckonii, as you can tell from the fact that I described it in exactly the same way.
Amanita "stocky gemmata" is stocky like Amanita pantherinoides, but is more consistenly bright yellow. It appears to be more closely related to Amanita muscaria than it is to the panther-gemmata group, but it has a collar volva like the latter. It too is found in Oregon and California.
California likely has additional panther-gemmata species in their rich oak forests and other habitats that aren't present in the PNW.
Amanita aprica © Steve Trudell
This common, similar spring species is usually apricot yellow, with something between concentric circles and a collar at the base of the stem, and has a characteristic look like you ironed the warts into the cap. It was described from Washington.
Amanita alpinicola © Janet Lindgren
This rare species (at least outside of the interior) is stocky, pale yellow and grows in spring under 5-needle pines. It has a collar volva and also has fused warts that looked like they were ironed into the cap. It's often found partially buried. It was described from Montana.
Amanita 'farinosa' sensu Thiers
Amanita 'farinosa' sensu Theirs © Kit Scates Barnhart
This grey, powdery Amanita looks more like it belongs in subgenus Lepidella than it does subgenus Amanita, but consensus is strong that it belongs here. It is grey and powdery everywhere, with not much further evidence of either a volva or a veil besides the powdery universal veil (the lack of a ring also making it easily confused with subgenus Vaginatae). The real Amanita farinosa from Europe and/or Eastern North America is slender and the cap is striate. Our west coast species, which needs a new name, is stockier and therefore does not have many striations in the cap (striations usually come from thin flesh allowing the outline of the gills to be visible near the rim of the cap).
Summary of Future Studies Needed
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