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Danny’s DNA Discoveries – 'Lepiotaceae' of the PNW
by Danny Miller

Click here for my Pictorial Key to the 'Lepiotaceae'


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 white spores, free gills, 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). Unfortunately, some species violate all three of those generalizations. 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. 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). Some even appeared to have lost their free gills, meaning that sometimes the gills are attached to the stem. That makes them a bit more difficult to identify as 'Lepiotaceae', but the scaly caps help. However, that's far from foolproof as other scaly genera exist that are similar but totally unrelated, like Tricholoma (which often will have specifically notched gills and seldom have a partial veil, unlike the mushrooms on this page).

We now know the picture is not as simple as there being one group of 'Lepiotas' that evolved inside the Agaricaceae. It is more complicated than that, and there are probably up to 4 groups of Lepiotas. The free gilled genera are in two groups fairly close to Agaricus, meaning that spore pigment may have went away in either one or two separate evolutionary events. The attached gilled genera are further away, and may have lost their spore pigment in one or two separate evolutionary events, but they did not "lose" their free gills, they probably did not evolve from Agaricus directly enough to have had an ancestor with free gills, so that explains the attached gilled 'Lepiotaceae'.

  • Free gills - some genera are very closely related to Agaricus (Chlorophyllum, Macrolepiota, Leucoagaricus and Leucocoprinus) and considered inside the Agaricaceae s.s.
  • Free gills - Some genera are a little more distant, and considered a sister family of their own, the Verrucosporaceae (Cystolepiota, Echinoderma and Melanophyllum).
  • Attached gills - Some genera are surprisingly far away, in the family Squamanitaceae (Squamanita, Lepiota, Cystoderma, Phaeolepiota, Floccularia and Leucopholiota).
  • Attached gills - Cystodermella is kind of by itself, at least a bit. No family name for it yet.

To complicate things further, three genera re-evolved coloured spores (or never completely lost all their pigments), one in each of three of the clades - Phaeolepiota (pale yellow or orange brown), Chlorophyllum (one has green spores) and Melanophyllum (green or red!).

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:

  • We have at least 4 species of Echinoderma, all probably undescribed. We do not have E. asperum.
  • Leucocoprinus heinemannii has been found again and verified.
  • I've made an attempt to characterize macroscopically Lepiota vs Leucoagaricus, which are easily confused without a microscope.
  • Leucoagaricus adelphicus and Leucoagaricus cupresseus from CA have been found in the PNW now, as has a sister species of the CA Leucoagaricus fuliginescens.
  • Leucoagaricus sp. 3, with greyish black scales uniformly distributed over the cap, is one of our most interesting newly discovered species.
  • Cystoderma jasonis occurs in the PNW and has always been mistaken for C. amianthinum.
  • Cystodermella adnatifolia occurs in the PNW and has always been mistaken for C. granulosa.
  • Some species are actually species groups and some are sister species to their European counterparts that may need new names.

Free Gills (Agaricaceae s.s. and Verrucosporaceae) - click to expand

Genera mentioned: Chlorophyllum, Macrolepiota, Echinoderma, Cystolepiota, Melanophyllum, Leucocoprinus, Leucoagaricus, Lepiota

Attached Gills (Squamanitaceae and Cystodermella) - click to expand

These genera have re-evolved attached gills, making them difficult to differentiate from other scaly genera, like Tricholoma (which more often has specifically notched gills and usually lacks a partial veil).

Genera mentioned: Cystoderma, Cystodermella, Phaeolepiota, Floccularia, Squamanita, Leucopholiota


Summary of Future Studies Needed

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