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

Click here for my Pictorial Key to the 'Lepiotaceae'


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.

  • 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, Pulverolepiota, Echinoderma and Melanophyllum).

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.

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:

  • 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.
  • 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)
  • a distinctive look that separates them from the erect scaled Echinoderma and the not quite as large Leucoagaricus, and far more common than both lookalikes
  • Chlorophyllum includes one rare, local gastroid mushroom.

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

Echinoderma - click to expand
  • medium to large species with erect scales on the cap and staining colours, uncommon to rare. Similar species with smaller spores that have erect scales but flesh that does not stain are covered under Lepiota. Echinoderma are closely related to Cystolepiota.

Species mentioned: Echinoderma asperum, eriophorum, perplexum, flavidoasperum. Lepiota acutesquamosa, aspera.

Leucocoprinus - click to expand
  • small, delicate fruit bodies
  • usually a striate cap margin, a clear ring that's often movable and granular-looking scale particles

Species mentioned: Leucocoprinus brebissonii, birnbaumii, cepistipes, ianthinus, flavescens, cretaceus, heinemannii, straminella, flavescens.

Cystolepiota and Pulverolepiota - click to expand
  • small, delicate fruitbodies
  • a shaggy appearance overall from a cottony universal veil material, no ring on the stem. Uncommon to rare.

Species mentioned: Cystolepiota seminuda, bucknallii, moelleri, oregonensis, hetieri, fumosifolia, cystidiosa, petasiformis, pulverulenta

Melanophyllum - click to expand
  • this genus evolved from within Cystolepiota, so perhaps these species should go by that genus (it is paraphyletic inside Cystolepiota).
  • a small (<5cm) amazing mushroom that evolved coloured red and/or green spores
  • greyish-brown cap, bright red free gills and scales all over the cap and stem.
  • membranous veil material seems to often be left on the cap margin as well as on the stem in a ring zone. This is very much unlike the cottony universal veil material of Cystolepiota.

Species mentioned: Melanophyllum haematospermum

Lepiota - click to expand
  • small to medium mushrooms (cap <10cm, stem <1cm thick)
  • stems may be smooth, scaly or shaggy
  • caps smooth or scaly, but not shaggy, granular or striate, fruitbodies not staining red
  • smooth stemmed species will usually have a brownish "eye" of scales on the disc that break up concentrically and not stain appreciably red (although rarely such a mushroom will be a Leucoagaricus).
  • (see Leucoagaricus for small mushrooms that stain red, have pink, black or pure white scales or have reddish radiating fibrils instead of scales that break up individually or concentrically)

Species mentioned: Lepiota megnispora, ventriosospora, clypeolaria, spheniscispora, erminea, alba, subincarnata, josserandii, helveola, castanea, pilodes, fuscosquamea, clypeolarioides, cortinarius, felina, fuscovinacea, cristata, kauffmanii, castaneidisca, subnivosa, subfelina, concentrica, amplifolia, thiersii

Leucoagaricus - click to expand
  • 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, have reddish fibrils that radiate out from the cap centre instead of breaking up individually or concentrically, 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, although rarely such a mushroom could be a Leucoagaricus).

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. Leucocoprinus cretaceus.


Summary of Future Studies Needed

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