Danny’s DNA Discoveries – Amanitaceae of the
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.
The Amanitaceae family is defined by white spores, free gills (although they don't always look as free as other free gilled families) and the presence of a universal veil. That universal veil is often composed of material that makes Amanitas look like they "hatch" out of eggs, leaving warts all over the cap and other areas of the mushroom after the mushroom grows. Warts are defined as removable pieces of material that are not part of the mushroom itself. The similarly defined Lepiota "family" is often scaly, but those scales are part of the mushroom body and can't be removed.
Most mushrooms you will find in this family are Amanita. The rare Limacella genus is related, but the universal veil is a layer of slime, so they are entirely slippery (cap and stem) white spored mushrooms with free gills. But as the free gills can be subtle, Limacella are hard to recognize without practice.
Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
Amanita section Vaginatae - click to expand
The so-called "grisettes". 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.
Species mentioned: Amanita constricta, pachycolea, sp-NW04, sp-NW11, sp-OR01, sp-OR02, castellae, velosa, populiphila, alaskensis, protecta, sp-OR03, obconiobasis, subnigra, lindgreniana, vaginata, vaginata var alba, sp-NW05, sp-NW09
Some of the species that occur in the PNW 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.
We do not have the poorly understood Amanita vaginata in the PNW. But everytime a species is found that does not exactly match A. constricta or the A. pachycolea group (and does not match the description of one of the more distinctive, rare, species below) it is usually mislabeled A. vaginata for lack of a better name.
Click on the name of the species, when possible, to bring up the page for that species on Rod Tulloss' excellent world-wide Amanitaceae site, www.Amanitaceae.org
Amanita section Phalloideae - click to expand
This group contains mostly deadly poisonous species. Nothing you can do to these mushrooms will remove the poison (a cyclopeptide called amatoxin) and prevent them from killing you if you eat them. They are recognized by the sac volva (as if the mushroom is sitting in a sac), usually bulbous stem, usually a non-striate cap and partial veil. (But both the volva and the partial veil can fall off).
Species mentioned: Amanita phalloides, ocreata
Amanita phalloides - The "death cap" has a yellow to yellow-green cap, although albino versions of this and many other Amanitas have been found. You are more likely to encounter this introduced European species in urban areas, like someone's yard or a city park with exotic trees (oak, chestnut, hazelnut) or perhaps under our native Garry oak. It's becoming more common in our cities as the years go by, but fortunately still rare in the wilderness of the PNW (although it has escaped to the wilderness of California, where their native trees are closer to its preferred European trees that it is used to).
This one species is responsible for about 80% of the serious poisonings and deaths from mushrooms in North America. It should be the first mushroom everybody here learns.
Amanita phalloides © Danny Miller
Amanita ocreata group - These species are pure white, giving them the name the "Destroying Angel". Amanita ocreata is a California species, and we seem to have additional species too: the genetic species best matching the type collection has been found once in California, during the 1998 NAMA foray. "sp-ocreata01" has been found in Oregon and California, and "sp-ocreata02" has been found in California. These collections were pure white but are more closely related to Amanita phalloides than they are to Amanita ocreata. More study is needed, but they are usually rare species here, found in the lowlands with hardwoods.
Amanita 'sp-ocreata01' © Joanee Schwartz
Amanita section Caesareae - click to expand
The so-called "coccoras". These Amanitas are considered edible, but can very much resemble deadly poisonous species. Recognized by a sac volva and a partial veil (just like the deadly poisonous A. phalloides) but typically with a more striate cap, non-bulbous stem and thicker sac material that tends to leave one giant eggy patch on the cap. They can get quite large. These are fairly rare in the PNW, unlike the deadly poisonous A. phalloides which is becoming more and more common in urban areas.
Species mentioned: Amanita calyptroderma, lanei, calyptrata, vernicoccora, calyptratoides
This section is named after the famous Caesar's Amanita, the bright orange Amanita that was allegedly one of Caesar's favourite foods, so he decreed that only he was allowed to eat it and anybody else caught doing so was put to death. Locally, they mostly prefer oak but occasionally conifers.
Amanita calyptroderma - This quite large fall species has a bright yellow-orange cap. It is a California species that grows with oak but possibly many other trees. It is found rarely in Oregon, Washington and BC under Douglas fir. There have been attempts to call it Amanita lanei or Amanita calyptrata, but those names were never legally published for one reason or another so the correct name for it is Amanita calyptroderma.
Amanita calyptroderma © Steve Trudell
Amanita vernicoccora - This quite large spring species has a dull yellow cap. It is also a California oak species rarely found in Oregon and Washington with oak and perhaps Douglas fir.
Amanita vernicoccora © Michael Beug
Amanita calyptratoides - This species is also described from California under oak, but has been found a couple of times in southern Washington under oak too. It has a more plain brown cap than the other species, is not quite as large, and has a stem that sort of appears water soaked, even when it isn't, and a ring that tends to "dissolve" and so older specimens may appear to be a grisette and belong in section Vaginatae.
Amanita calyptratoides © Cindy Trubovitz
Amanita section Validae - click to expand
Continuing the reputation of bizarre properties in Amanita, this section contains species with hemolytic properties when raw - in other words, they make red blood cells explode, at least in a Petri dish. Recognized by an abrupt bulb at the stem base and usually a coloured partial veil. Parts of these species usually turn red when handled (blush).
Species mentioned: Amanita augusta, franchetii, aspera, porphyria, novinupta, cruentilemurum, rubescens, orsonii, spissa, excelsa, citrina, sponsus, sp-NW10
Amanita augusta - Somewhat resembling a "Panther" Amanita in section Amanita (brown cap with warts) but the warts and the ring on the stem are bright yellow, the stem base is either bulbous or with scales like A. muscaria instead of a collar, and the base of the stem may stain a bit reddish in places. This mushroom has often been incorrectly called Amanita franchetii and Amanita aspera, similar mushrooms from Europe, but our common species was described from California.
Amanita augusta © Steve Trudell
Amanita porphyria - Grey or brown cap, grey partial veil, chevron pattern on the stem and subtle blushing on an abrupt stem bulb. Our species is the same as the species described from Europe.
Amanita porphyria © Steve Trudell
Amanita novinupta - A small, mostly white, strongly blushing species found under oak and conifers. Described from Oregon and relatively rare. Found in spring. Its name means "blushing bride".
probable Amanita novinupta © Noah Siegel
Amanita cruentilemurum n.p. - Practically identical to A. novinupta, but usually found in the fall with more elongated spores. Also found with oak and conifers. Its name means "bloody ghost". It has been found in California and New Mexico, and turned up once on southern Vancouver Island, so it's very rare up here so far.
Amanita cruentilemurum © Bob Chapman
Amanita sponsus n.p. - this California oak mushroom was recently found once under Oregon oak. It is a stocky club shaped mushroom with a very subtle volva, stains slowly brownish but not red, but most distinctively, has coarse brown warts textured into the white cap.
Amanita sponsus n.p. © Bruce Newhouse
Amanita rubescens - Another strong blusher, but this time usually with a brownish cap when young instead of pure white. Its warts are usually a little more pronounced, whereas A. novinupta's warts may appear as soft, cottony patches. Reported under various confiers and hardwoods.
Our local DNA is sometimes an exact match but sometimes 1-2 bp from European sequences that could be Amanita rubescens, which probably means it is the same species, but a unique species was described from India (Amanita orsonii) with only 3bp differences so I can't say for certain if we have the real thing or only 1 species. We do need confirmation that the European sequences I'm comparing to, which Rod calls Amanita sp-rubescens02, are in fact A. rubescens. This collection has not been sequenced, but seems to match other collections that have been sequenced.
Amanita cf rubescens © Paul Kroeger
Amanita 'spissa' - This brownish-grey capped species does not readily blush. There will be red stains around insect tunnels, but the stem won't strongly stain from handling nor will the flesh stain strongly after cutting like A. rubescens will. It can be told apart from Amanita augusta, another species that does not stain strongly, by the greyish white veils without any yellow. It is more closely related to A. augusta than it is to A. rubescens. It was found once with exotic trees in Vancouver BC (reported under conifers in Europe) and is assumed to be rare here.
As to its identity, other collections with the same DNA as ours have been identified in Europe as the European Amanita spissa, but in the absense of a type sequence that is not a guarantee. The older European Amanita excelsa is said to be a synonym, but collections identified as A. excelsa in Europe have different DNA than our collections, so perhaps there really are two species, and the one found here is actually Amanita spissa. If not, it is a closely related species. For now, Rod is calling it Amanita sp-rubescens01.
Amanita 'spissa' © Paul Kroeger
Amanita sp-NW10 - A very pale yellow capped species with a prominent bulb, resembling Amanita citrina of Europe. Known only from one collection in the Olympic Mountains of WA, but this photo of a similar collection from a spruce-hemlock forest in Haida Gwaii, northern BC seems to resemble it. We will see if that is what this is. If not, then what is it? No DNA yet.
Amanita cf 'sp-NW10' © Paul Kroeger
Amanita sections Lepidella and Saproamanita - click to expand
Sometimes not easily recognized as Amanitas, they may represent old lineages before strong, obvious volvas evolved. The universal veil has not developed to leave the same type of material on the cap or bottom of stem. It may simply be a mass of cottony tissues that makes the mushrooms look fluffy, until it washes off. Saproamanita can be recognized by being found far away from trees (in grasses, marshes or desert areas), as they are the only Amanita that are not mycorrhizal, another possible artifact of them being an old lineage. They are mostly white mushrooms, and at least Lepidella can contain deadly poisonous species (Don't confuse with Matsutake! That could be a deadly mistake).
Species mentioned: Amanita smithiana, silvicola, magniverrucata, pruittii, armillariiformis, prairiicola
The toxins found in Lepidella are not the same as in the Death Cap, these toxins may "only" destroy your kidneys, not both your liver and your kidneys. Some have created a new genus for Saproamanita, but the necessity of them having their own genus is controversial, so I will refer to them as a section of Amanita. It may be hard to tell Lepidella apart from Saproamanita by looking at the mushroom, they are best separated by habitat - forest versus prairie grassland.
Amanita section Amanita - click to expand
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.
Species mentioned: Amanita muscaria, muscaria var. formosa, muscaria var. guessowii, muscaria subsp. flavivolvata, muscaria var. alba, chrysoblema, aprica, pantherina, gemmata, junquillea, pantherinoides, praegemmata, umbrinidisca, ameripanthera, breckonii, pseudobreckonii n.p., alpinicola, farinosa, aurantisquamosa
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 uncommon in the PNW, usually found under introduced trees with almost pure white warts.
Amanita chrysoblema, our abundant, 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 aprica - This 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 aprica © Steve Trudell
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 (neither A. pantherina nor A. gemmata), but here's what I have been able to determine about what species we do have (it should be noted that a recent European study showed that A. gemmata may actually just refer to A. muscaria, so even in Europe they may start calling their yellow 'gemmata' species A. junquillea). Amanita pantherina-like mushrooms have a lookalike in subgenus Validae - Amanita augusta, that among other things, will not have a single collar volva and may blush red.
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 most common species in the Seattle area. Amanita pantherinoides is yellow to brown, stocky, with persistent warts on the cap. Our most common "gemmata" species, Amanita aff breckonii, below, is usually yellow, slender, and has warts that easily wash off, although some of our less common "gemmata" species might be confused with yellowish A. pantherinoides.
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 common 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.
Amanita aff breckonii - Our most common species in WA, this species is always yellowish, usually slender and the warts wash off pretty easily, although occasionally it can be stockier like A. pantherinoides. It is a sister species to a California species that has not yet been found in the PNW, Amanita breckonii.
Amanita pseudobreckonii n.p. - another yellow, slender species with warts that easily wash off in the gemmata-panther group found in Oregon and California. It is very difficult to tell apart from A. aff breckonii in Oregon where they are both common.
Amanita "stocky gemmata" is sometimes stocky like Amanita pantherinoides but consistenly bright yellow. Unfortunately, it can also be relatively slender and bright yellow and has now been found in Olympia WA, where it will be easily confused with the other species. 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 is found in Oregon and California as well.
Amanita aff breckonii © Buck McAdoo, Amanita pseudobreckonii n.p. © Corinne Srsen, Amanita "stocky gemmata" © NAMA and the Field Museum of Natural History
It is possible that one of Murrill's three species is not the same as the other two, but represents one of the above 3 gemmata-like species. If so, that name from 1912 would be the proper name for it. We will need type sequences of his original 1912 collections, if we can get them, to settle the matter. California has additional "panther-gemmata" species in their rich oak forests and other habitats that aren't present in the PNW, making identification for them even more difficult than for us.
Amanita alpinicola - This 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 and the DNA has been found in Idaho, as well as in WA north of Spokane at 5500' in July. There are reports that it may be in BC and OR as well, presumably inland and at elevation.
Amanita alpinicola © Janet Lindgren
Amanita 'farinosa' sensu Thiers - This grey, powdery Amanita looks more like it belongs in section Lepidella than it does section 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 section 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).
Amanita 'farinosa' sensu Theirs © Kit Scates Barnhart
Amanita aurantisquamosa - This tan coloured species has hints of orange in the scales, irregular patches of veil on the cap and a volva that is somewhat sac-like at first but soon breaking up. It is rare, only known from Southern Idaho and we do not have any sequences of it yet.
Amanita aurantisquamosa © Ellen Trueblood
Limacella - click to expand
Hard to recognize, as the universal veil is a layer of slime, but they can be recognized with practice by the free gills (although not always as obviously free as in other families) and slimy nature, especially on the caps. The species fall into three groups, which some consider separate genera, but I will treat them as subgenera. All are rare.
Genera mentioned: Limacella, Zhuliangomyces, Limacelloides
Summary of Future Studies Needed
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