Danny’s DNA Discoveries – 'Lepiotaceae'
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.
Click here to download the FASTA data of all my DNA sequences
Lepiota and allies are known for free gills, white spores and often scales on the cap that can't be removed (unlike the similarly defined Amanitaceae that have a universal veil that may leave warts, which unlike scales, are removable). Rarely, one may have a coloured spore print or be scale-less, but for the families covered on this page, they will all have free gills. All species seem to have some sort of obvious partial veil. Species in this family often have a coloured "eye" in the cap disc where the scales are especially dense. Like Amanita, some of them have evolved to produce the deadly Amatoxin and can kill you if you eat them (but not nearly as many species as we once thought). Others are eaten regularly, especially some Shaggy Parasols (Chlorophyllum) although one species of that can be dangerously poisonous as well. While most Amanita are mycorrhizal, most 'Lepiotaceae' are saprophytic.
The 'Lepiotaceae' have been thought to be a kind of sub-family of the Agaricaceae, not considered a family in their own right because they would be paraphyletic ("inside" Agaricaceae, not "beside" or sister to it). That family is known for dark chocolate spores and free gills. However, the 'Lepiotaceae' mushrooms lost their spore colour and mostly have white spores, something that also happened to Laccaria (that too is in the dark spored clade of agarics).
We now know the picture is not as simple as there being one group of 'Lepiotas' that evolved inside the Agaricaceae. The free gilled genera are in two groups fairly close to Agaricus, meaning that spore pigment may have gone away in either one or two separate evolutionary events.
There are also two similar looking groups with attached gills (think Cystoderma), once thought to be closely related to the free gilled genera but now appearing to be further away, meaning there may have been one or two additional events where spore pigment was lost in those groups. They probably did not evolve from Agaricus directly enough to have had an ancestor with free gills, so that explains the attached gilled 'Lepiotaceae' like Cystoderma.
To complicate things further, two genera re-evolved coloured spores (or never completely lost all their pigments), one in each clade - Chlorophyllum (one species has green spores) and Melanophyllum (green or red spores!). The attached gilled group also has one genus that did not completely lose its spore pigment, Phaeolepiota.
This family evolved one trufflized gastroid mushroom, Chlorophyllum (Endoptychum) agaricoides, that will not be recognizable to family. Another species, Endoptychum depressum, is actually an Agaricus and covered there as Agaricus inapertus.
If there are no scales and no warts, you'll have to decide between 'Lepiotaceae' and Amanitaceae. Leucoagaricus leucothites vs. Amanita smithiana/silvicola is an example. The two families have a slightly different look to their "free gills", a slightly different general stature (stocky vs stately), and the presence of a universal veil should be detectable in young Amanitas at least as a slight general shagginess.
Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
Chlorophyllum and Macrolepiota - click to expand
Species mentioned: Chlorophyllum molybdites, olivieri, brunneum, rhacodes (rachodes), agaricoides. Endoptychum agaricoides. Macrolepiota procera.
Family Agaricaceae. They have been well studied, and for once it is well known exactly which Chlorophyllum species occur in the PNW. Macrolepiota is a genus of similarly large mushrooms that does not occur in the PNW natively, although there are reports that cultivated Macrolepiota procera has escaped into the wild near Olympia, WA and can be found here.
Chlorophyllum molybdites - Dangerously poisonous, it is recognized by a green spore print in age, growth in grasses and gardens, and white areas between the scales. Young specimens may have white gills and not be identifiable. It is rare here, being a hot weather species, but it has made it all the way up north to Washington so far.
Chlorophyllum molybdites © Christian Schwarz
Chlorophyllum rhacodes (rachodes) - Most easily confused with the poisonous C. molybdites, but this has a white spore print (as do the rest of the species in this section). When young, the caps of some of these species start out solid dark brown before they start breaking up into scales (see photo).
Chlorophyllum rhacodes © Steve Trudell
Chlorophyllum brunneum - This species is known for a very abrupt bulb at the bottom of the stem, otherwise it is similar to the other two.
Chlorophyllum brunneum © Steve Trudell
Chlorophyllum olivieri - This species does not have the prominent white areas between the scales, but is more of a uniform brown.
Chlorophyllum oolivieri © Steve Trudell
Chlorophyllum agaricoides - Formerly Endoptychum agaricoides before it was realized that gastroid and truffle mushrooms don't usually deserve their own genus but are inside the genera they evolved from. It is round like a puffball. The spore mass turns yellowish brown (the DNA and the coloured spore print suggest a close relationship to C. molybdites), and it has a rudimentary stem inside, but the gills are not recognizable as such. It has evolved a strong odor of cabbage (many trufflized species need to evolve some sort of strong odor to enable them to be found by animals and dug up or opened). The ITS DNA of this species all over the world only differs by 1 or 2 bp. We have sequences of this eastern hemisphere mushroom from as close as New Mexico, so I expect our species is the same.
Chlorophyllum agaricoides © Steve Trudell
Macrolepiota procera - This species somewhat resembles Chlorophyllum rhacodes with scant brown scales on the cap (that is otherwise white), but it is not quite as large and does not stain orange when scratched. It is rare, with only a few reports of escaped cultivars around Olympia, WA so far.
Macrolepiota procera © Buck McAdoo
Echinoderma - click to expand
Species mentioned: Echinoderma asperum, eriophorum, perplexum, flavidoasperum. Lepiota acutesquamosa, aspera.
Family Verrucosporaceae. Echinoderma asperum (Lepiota aspera/Lepiota acutesquamosa) is a European species reported from the PNW, but that is not the species we have. Neither does it appear that our species is the same as the east coast Echinoderma eriophorum. We have at least 4 undescribed species in need of names.
True Echinoderma species have spores perhaps 6.5-9x3u and caps > 2.5cm across. There are similar species with erect cap scales and scaly stems (but they do not stain colour), with spores perhaps 5x3u, and although my ITS only trees sometimes show them near Echinoderma with weak support, a 4-gene study has shown that they belong in Lepiota. Some of those Lepiota are small, delicate mushrooms (easily distinguished from Echinoderma) but some are large and stocky, so you'll have to study the photos and look at the spores to tell them from Echinoderma.
Echinoderma PNW01 - This species, the first 2 times it was photographed, had a uniformly warm brown coloured cap, not showing any concentric rings of white between the scales. The third collection was mostly white with a few brown scales. I think a decade-old collection that was almost pure white may have been the same thing. In all cases, there were loose chunks of spines on the stem, especially near the bottom. It is uncommon, having been found twice in WA, once in Victoria, once on Texada Island (north of Nanaimo BC) and once in boreal BC. In one BC collection, it was with conifers. The white collection was in red cedar debris and had an unpleasant rubbery odor.
Uniformly brown Echinoderma PNW01 © Danny Miller & Marty Kranabetter. Mostly white PNW01 © Sharon Squazzo. Mostly white possible PNW01 © Danny Miller
Echinoderma PNW03 - This very similar species has a possibly colder brown cap than PNW01 and seems to show some white between the cap scales as it expands. It's unclear if the stem scales are more appressed than in PNW01 and fall off less easily. It is uncommon, having been sequenced more than a half dozen times in Vancouver BC, at least once in WA and also in Quebec and Arizona.
Echinoderma PNW03 © Danny Miller and Matthew Koons
Echinoderma 'favidoasperum PNW02' - This species has a warm yellow brown cap and a mostly smooth stem. E. flavidoasperum, from China, from which it differs by 6-7bp and 1-2 indels has a slightly reddening stem when handled. This rare sister species was found in the Cascade mountains in Manning Park BC and near Stampede Pass, WA.
Echinoderma PNW04 - Species PNW04 seems to be much like species PNW01 and PNW03, but with a rich milk chocolate coloured cap, and it appears to only have erect scales on the disc, with flattened scales everywhere else. It appears to have a fibrillose stem (not a scaly stem) and little white areas are showing between the brown scales. Also rare, it was found once SE of Salem, OR. It may be somewhat of a sister species of Echinoderma perplexum (differs by about 10%).
Echinoderma 'flavidoasperum PNW02' © Danny Miller (2 images), Echinoderma PNW04 © NAMA and the Field Museum of Natural History
Leucocoprinus - click to expand
Species mentioned: Leucocoprinus brebissonii, birnbaumii, cepistipes, ianthinus, flavescens, cretaceus, heinemannii.
Family Agaricaceae. Leucocoprinus does not deserve its own genus, according to strict rules, as it is parphyletic with Leucoagaricus ("inside" instead of "beside" or sister to it). But it seems to be a good monophyletic group of species with distinct characters so I am retaining the genus name here. This is usually a hot weather or tropical genus (mostly found in the summer), and except for one common "wild" species, others are only found locally in potted plants or greenhouses.
Leucocoprinus brebissonii - Black granular scales on the disc making an "eye". Very similar to the slightly stockier 'Lepiota' atrodisca, but never has any black on the ring. (That probably belonging in Leucoagaricus and described below in that section). It is now a wild species, found in forests. It was introduced to the PNW around 1994 and has become abundant. If it's found indoors, consider L. heinemannii, below.
Leucocoprinus brebissonii © Steve Trudell
Leucocoprinus cf birnbaumii - A bright yellow mushroom, covered in granules. Almost always found indoors or in artificially heated areas. This is a complex of species in Europe, so I don't know how many species we have or if the real thing is one of them. We need local samples.
Leucocoprinus cf birnbaumii © Michael Beug
Leucocoprinus 'cepistipes PNW01' - All white, or with pale brown scales. Found both indoors and outdoors. One outdoor (from woodchips) WA sequence is 5% different in ITS from EU sequences, implying ours is an unnamed sister species. We should test an indoor collection.
Leucocoprinus 'cepistipes PNW01' © Michael Beug
Leucocoprinus cf ianthinus UK - Purple. Found only indoors. Two asian sequences differ by 4 bp. I don't have any local or type area (UK) sequences. It seems plausible that we have the real species as it is never found in the wild.
Leucocoprinus cf ianthinus © Steve Trudell
Josh Birkeback did a study that found three additional rare species in western Washington - species that nobody else has ever found, to my knowledge, except for one. They should be looked for again.
Leucocoprinus cf flavescens - Also yellow, but not quite as bright, and often with a brownish disc and fewer granular scales on the cap. Found once in WA in a covered can outside a UW greenhouse. We need local DNA as well as type area DNA (Ohio) to compare to.
probable Leucocoprinus cf flavescens from eastern North America © Stephen Russell
Leucocoprinus cf cretaceus - All white with a more copious powdery covering than L. cepistipes. Found once in WA in wood chips with horse manure. We need local DNA to prove we have the real European species.
Leucocoprinus cf cretaceus from outside North America © Daniel Winkler
Leucocoprinus heinemannii - Black scaly disc, like L. brebissonii, but with larger, more fibrillose scales instead of tiny granules, and not quite as much of an "eye". It's a European species reported once in Washington in a greenhouse, and just recently found again by me in a potted plant just outside another greenhouse in Seattle.
Leucocoprinus heinemannii © Danny Miller
Cystolepiota - click to expand
Species mentioned: Cystolepiota seminuda, bucknallii, moelleri, oregonensis, hetieri, fumosifolia, petasiformis
Cystolepiota 'seminuda PNW01'/'seminuda PNW02' - Our least rare species in this genus are probably the most slender ones. They have a white cap with pinkish stem (the pink may intensify) and lots of shaggy material that disappears in age to become "half nude". There is a group of species matching this description all over the world; we have some east coast names but no west coast names that I know of for potential species, so we will probably need new names for ours. I have found 2 genetic species in the PNW, neither the likely real thing from Europe. Nor are they closely related to our best guess of the real thing. The second species was especially slender, but there was only one fruitbody.
Cystolepiota 'seminuda PNW01' © NAMA and the Field Museum of Natural History and Yi-Min Wang, C. 'seminuda PNW02' © iNaturalist user haunma
Cystolepiota 'bucknallii PNW03' - This beautiful, purple, cottony mushroom smells strongly and disagreeably like coal tar. This is a group of species in Europe and we have something distinct from all of them that likely needs a new name.
Cystolepiota 'bucknallii PNW03' © Lauren Ré
Cystolepiota cf moelleri EU - This very rare mushroom has pink velar material on the cap. We have potential European DNA but need local DNA to confirm if we have this species, or what the two BC and WA collections that resembled that species really are.
unsequenced Cystolepiota cf moelleri © A and O Ceska
Cystolepiota oregonensis OR - Copious velar material, staining rusty brown in age and where handled. Described from Oregon. We have one sequence that may be this, and unsequenced collections that are definitely this. We should sequence more collections to confirm that we now know what this DNA is. It strongly resembles the European Cystolepiota hetieri, which may be a group of species in the EU.
unsequenced Cystolepiota oregonensis © Danny Miller, sequenced possible C. oregonensis © Yi-Min Wang
Cystolepiota fumosifolia WA - Not petite (~3cm) white, cottony mushroom turning smoky coloured in cap and gills in age. Described from WA, and we have California DNA but need Washington DNA to confirm theirs is the same species.
Cystolepiota petasiformis WA - Copious cottony veil material, often forming a false conehead. Smells farinaceous. Described from Washington, we have California DNA and BC DNA (3 bp from each other in ITS2 only) but no Washington DNA yet. ITS doesn't place this species reliably inside Cystolepiota, but combined with LSU sequences, it does have support inside Cystolepiota.
Cystolepiota aff petasiformis - one WA sequence is different. No photo or description yet. We need more collections to confirm.
Cystolepiota petasiformis © Christian Schwarz
Melanophyllum - click to expand
Species mentioned: Melanophyllum haematospermum
Cystolepiota ('Melanophyllum') haematospermum group - Olive green spores when fresh, reddish brown spores when dried. This is a group of species, all called Melanophyllum haematospermum. Locally, we have sequenced 4 collections, and all 4 of them were different species, which was not surprising given that worldwide, there are over a dozen species concepts with ITS DNA >3% different from each other. I know of no study to try and determine the differences between them. They all seem to be found in conifer forests.
PNW01 had a convex to conical cap up to 5cm across. Stem base and mycelium blue-green. Spores 5.8-6.5x3.3-3.8u. Epithelium of pileipellis with sphaerocysts.
PNW02 has a small umbo, and the stems seemed especially thin for its size. Spores 5-5.5x3-3.5u
PNW03 has a very broad umbo.
PNW04 is easy to spot. It is much more delicate with much more distant gills than the other three. Spores 5-5.7x2.7-3.3u
Since they live "inside" Cystolepiota and not "beside" it, they technically don't deserve their own genus if you want to insist that Cystolepiota not be paraphyletic.
'Melanophyllum' PNW01, PNW02, PNW03 and PNW04 © Connor Dooley, Andrew Parker, Andrew Parker and Connor Dooley
Lepiota - click to expand
Species mentioned: Lepiota megnispora, ventriosospora, clypeolaria, erminea, alba, subincarnata, josserandii, helveola, castanea, pilodes, fuscosquamea, clypeolarioides, cortinarius, felina, fuscovinacea, cristata, kauffmanii, castaneidisca, subnivosa, subfelina, concentrica, amplifolia.
Leucoagaricus - click to expand
Species mentioned: Leucoagaricus leucothites, barssii, americanus, bresadolae, badhamii, adelphicus, cupresseus, erythrophaeus, georginae, roseilividus, rubrotinctus, glabridiscus, purpureolilacinus, opthalmus, pakistaniensis. Lepiota naucina, fuliginescens, pulverapella, castanescens, flammeotincta, roseifolia, decorata, rubrotinctoides, atrodisca, oculata, sequoiarum.
Family Agaricaceae. Some are still officially called Lepiota because they haven't been moved yet to Leucoagaricus , but make no mistake. They are not Lepiotas, which are in a different family (Verrucosporaceae). They are definitely members of Leucoagaricus.
Summary of Future Studies Needed
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