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Danny’s DNA Discoveries – White Spored Agaricaceae s.s. of the PNW
by Danny Miller

Click here for my Pictorial Key to all 'Lepiotaceae'

Introduction

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 of the Agaricaceae s.l., meaning that spore pigment may have gone away in several separate evolutionary events.

  • Free gills - some genera are very closely related to Agaricus (Chlorophyllum, Macrolepiota, and Leucocoprinus s.l. (Leucoagaricus, Leucocoprinus, and Macropsalliota)) and are considered inside the Agaricaceae s.s. This page covers these genera. In fact, the Agaricaceae s.s.'s ancestral trait may be white spores, with Agaricus and sister genera re-acquiring spore pigment.
  • Free gills - Some genera are a little more distant, and considered a sister family of their own, the Verrucosporaceae (or 'Lepiotaceae') (Lepiota, Cystolepiota, Pulverolepiota, Echinoderma and Melanophyllum). While it's tempting to call them the Lepiotaceae, that family was not legally described, and for now the proper family name is the Verrucosporaceae.

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. They probably did not evolve near Agaricus closely 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.

Leucocoprinus and Leucoagaricus used to be separated morphologically. Leucocoprinus had a striate cap margin, a clear ring that's often movable and granular-looking scale particles. Leucoagaricus were otherwise. However, they mixed together in the tree, and they have long been known to need revision. A revision has now been made, where Leucoagaricus americanus and allies were moved to Macropsalliota, Leucoagaricus rubrotinctoides and allies stayed in Leucoagaricus, and most everything else moved to Leucocoprinus.

If there are no scales and no warts, you'll have to decide between one of the Lepiota-like genera and Amanitaceae. Leucocoprinus 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.

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.

Summary of Interesting Results

Here are some of the newest, most interesting results of the study:

  • Leucocoprinus heinemannii has been found again and verified.
  • I've made an attempt to characterize macroscopically Lepiota vs Leucocoprinus s.l., which are easily confused without a microscope.
  • Leucocoprinus adelphicus and Leucocoprinus cupresseus from CA have been found in the PNW now, as has a sister species of the CA Leucocoprinus fuliginescens.
  • Leucoagaricus sp. 3, with greyish black scales uniformly distributed over the cap, is one of our most interesting newly discovered species.
  • Some species are actually species groups and some are sister species to their European counterparts that may need new names.

Chlorophyllum and Macrolepiota - click to expand
  • shaggy parasols, our largest 'Lepiotas', 20cm across or more, with a brown scaly cap and flesh that stains quickly orange when scratched (except in Macrolepiota)
  • white or green spore print (for one locally rare Chlorophyllum)
  • they have a distinctive look that can be learned.
  • Chlorophyllum includes one rare, local gastroid mushroom.

Species mentioned: Chlorophyllum molybdites, olivieri, brunneum, rhacodes (rachodes), agaricoides. Endoptychum agaricoides. Macrolepiota procera.

Macropsalliota - click to expand
  • many Leucoagaricus have been moved to Leucocoprinus, which now contains many morphological forms.
  • Macropsalliota is the new home for Leucoagaricus americanus and Leucoagaricus meleagris, somewhat clustered mushrooms that stain several possible colours.

Species mentioned: Leucoagaricus americanus, bresadolae, meleagris.

Leucoagaricus - click to expand
  • many Leucoagaricus have been moved to Leucocoprinus, which now contains many morphological forms.
  • true Leucoagaricus typically have red/brown radiating fibrels instead of the typical scales that break up into individual scales or in concentric patterns.
  • some are almost pure white and easily mistaken for some Leucocoprinus.
  • one has evenly distributed pure black scales (not just in the "eye").
  • one is yellow, but not bright yellow like some Leucocoprinus.

Species mentioned: Leucoagaricus rubrotinctus, pakistaniensis, glabridiscus, purpureolilacinus. Lepiota rubrotinctoides. Leucocoprinus flavescens.

Leucocoprinus s.l. - click to expand
  • SMALL, delicate fruit bodies usually a striate cap margin, a clear ring that's often movable and granular-looking scale particles (the classic definition of Leucocoprinus)
  • LARGE mushrooms (cap ~10cm, stem 1-2cm thick) that are not Chlorophyllum nor with erect scales like Echinoderma
  • SMALL mushrooms (cap <5cm, stem <0.5cm thick) with a smooth stem and a ring
  • try here first for small mushrooms that stain red, and for those with pink, black or mostly white scales
  • try Lepiota first for small mushrooms with a brownish eye of concentric cap scales that do not stain red appreciably

Species mentioned: Leucoagaricus leucothites, barssii, badhamii, adelphicus, cupresseus, erythrophaeus, georginae, roseilividus, ophthalmus. Lepiota naucina, fuliginescens, pulverapella, castanescens, flammeotincta, roseifolia, decorata, atrodisca, oculata, sequoiarum. Leucocoprinus brebissonii, birnbaumii, cepistipes, ianthinus, flavescens, cretaceus, heinemannii, straminella.

 

Summary of Future Studies Needed

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