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Danny’s DNA Discoveries – Russula

by Danny Miller


Click here for my Pictorial Key to Russula


Introduction - read this for a background on Russula


Brevipes and Compactae Clades (click one for details)

R. nigricans © Ben Woo and Danny Miller

These two clades of Russulas are easy to recognize, and used to be thought of as one group, but they form two separate genetic clades.


Brevipes - click for details

R. cascadensis - small (<9cm), hot tasting with a creamy yellow spore print.

R. brevipes group - will grow larger than 10cm, mild or hot tasting, almost white spore print. We have about 6 species in this group, none with a name, and none are actually R. brevipes itself.


Compactae - click for details

R. nigricans and R. albonigra - do occur here as thought

R. adusta group - we have 3 species in this group, perhaps including the real R. adusta

R. acrifolia - what we have been calling R. densifolia is actually closer to R. acrifolia, if not the same thing, so use that name for now instead.


Click here for more details of the Brevipes and Compactae.

Fetid Russulas – Ingratula and Crassotunicata Clades (click one for details)

R. laurocerasi group member © Joy Spurr, R. cerolens © Ben Woo and R. crassotunicata © Kit Scates Barnhart

These similar mushrooms used to be grouped together in one clade, Ingratula, until it was discovered that the similar R. pallescens belonged in a separate clade with the unique R. crassotunicata. They taste somewhat peppery, which is bad enough, but even worse than that, they often have this putrid, nauseating odor and flavour that kind of makes you want to throw up. So, they win the worst tasting Russulas award. Although, I once had a professional chef prepare one of these (sp. Woo 3) to see if it could be made palatable, and a few of us ate some, and it didn’t suck. But definitely not recommended.


Ingratula - plain brown - click for details

R. cerolens  - very common, mistakenly called R. sororia, R. pectinata or R. pectinatoides.

R. recondita group - 3 rare, closely related species best differentiated from R. cerolens microscopically

Ingratula - yellow brown - click for details

R. 'foetens' - unnamed species often reported as unpleasantly nauseatingly sweet like vomit

R. 'laurocerasi'/'fragrantissima' group - 2 unnamed species often reported as pleasantly sweet like almond or maraschino cherries


Crassotunicata - click for details

R. pallescens - rare, almost yellow, small, slender species with white spores (the others have cream spores), no odor. Mistakenly called R. farinipes

R. crassotunicata - common, all white, browns significantly, cap cuticle is a thick, gelatinous, peelable layer.


Click here for more details of the Ingratula and Crassotunicata clades.

Heterophyllidia I and II Clades - click for details

R. sp. Woo 13, R. 'aeruginea', R. 'brunneola' © Ben Woo

This group needs the most study. They are difficult to identify and not much is known about them. They just aren’t found often enough to study and most of the time when one is found, it isn’t recognized. That’s unfortunate, because most if not all of the group members are mild tasting and especially dense and crunchy textured making this group potentially among the better tasting Russulas (although none have been tested enough to be sure of their edibility). There at least a dozen species, and few of them have been named yet or even have a good description. Here is a not-at-all foolproof way of trying to identify this section:

So... a Russula with a matte, pruinose or powdery cap (left photo) might be one of these (except the very common purple capped Russulas in the R. murrillii group). Any green Russula, even with a shiny cap (middle photo), might be one of these. That can be really subtle, a wet cap can hide the matte-ness and any dry cap might start to look matte. Brown, pruinose caps are common in this group (right photo).


R. 'aeruginea'  - our least uncommon member, a shiny green capped species. Other Russulas are very similar, like R. graminea.

R. sp. Woo 13 - a beautiful matte, green capped species, other species are not entirely green but with a lot of brown.

R. 'brunneola' - a matte, brown capped species with white spores (almost all others have cream coloured spores).

R. 'mustelina' and 'medullata' - two matte, brown (maybe mixed with other colours), cream spored European species, but a half dozen undescribed species are similar.

R. sp. Woo 16 - a beautiful purple, matte capped Russula, but with a purple flushed stem (unlike the much more common R. murrillii group).


R. malva np - mistakenly called R. cyanoxantha, this large, matte capped southern Russula has mottled purple colours (mostly found in California) has a provisional name. Some analyses show it in a separate clade, which I call Heterophyllidia II. FeSO4 does not turn the flesh of this species pink, unlike the other Heterophyllidia.


Click here for more details of the Heterophyllia clades.

Amoenula, Archaeae, Malodora and Glutinosae Clades

We do not appear to have any Russulas from these sections in the PNW, so we’re not going to learn anything about them today. If you can find one, you’ll be famous. Russula mariae from the east coast is in clade Amoenula, but reports of that species in the west have not been proven yet.

Core Clade

R. emetica, R. phoenicea, R. americana © Ben Woo

The “core” species (most closely related to the official first Russula, Russula emetica), are not recognized so much by how they look, but by the fact that they are often hot tasting and don't belong to one of the previous groups. A surprising number of species come in either poppy red or yellow (or a mixture, left photo). Some species have a lot of stem flushing (right photo). Unfortunately, the chemical that makes them taste hot can go away or wash out, so sometimes they taste mild. A drop of sulfo vanillin will usually turn the flesh of a “hot” species purple (in this clade or elsewhere), even if a certain specimen tastes mild. This was one of the most interesting results of Anna’s study of Ben’s chemical tests on his Russula collection.


White Spored Members - click for details

Most species are together in one clade, but a few white spored species live in the cream spored clade (R. stuntzi, R. parapallens, R. aff. viscida). These three won't necessarily have pure white spores, but could have a little tint to them.

R. emetica/montana  - two very common similar mushrooms that can be poppy red, yellow, or both. R. montana's stem may grey in age, but best distinguished microscopically. At least two other rare species are almost identical. The local R. crenulata exists but not much is known about what it really is. These have been mistakenly called R. bicolor or R. silvicola in the past.

R. parapallens - also common and bright red and/or yellow but usually with a dark splotch of colour on the disc. Two rare species (R. laccata and R. 'viscida') are similar but perhaps closer to wine-red, with the stem of R. 'viscida' browning considerably in age.

R. phoenicea/hypofragilis - have purple tones. R. phoenicea also has green/olive tones. These have mistakenly been called R. fragilis in the past.

R. stuntzii - grey cap that may have purple tones.


Cream Spored Members - click for details

R. americana/rhodocephala - poppy red caps with poppy red flushed stems, differentiated by their tree associates. Mistakenly called R. rosacea or R. sanguinea in the past.

R. queletii - purple caps with purple flushed stems. Mistakenly called R. sardonia or R. torulosa in the past.

R. pseudopelargonia/salishensis - reddish-purple caps with a bit of stem flushing, difficult to differentiate from each other. Mistakenly called R. pelargonia before.

A handful of rare species have red, yellow or white caps. They are unnamed, poorly understood, or both, and resemble and/or are close relatives of R. alcalinicola, R. albidula/persicina and R. renidens.


Click here for more details of the White Spored and Cream Spored Core Clade members.

Crown Clade

The remaining Russulas have not been definitively separated out into smaller groups yet, unfortunately, and there are a lot of them. Around half of the Russulas fall into this large clade. There have been many smaller groupings made over the years, but they have not proven to be genetically valid groupings, so I’m going to treat them all as one large group. It gets its name from the fact that they may represent the newest branches, at the “crown” of the genetic tree. I also call them the “garbage” clade, since you can think of these as species that did not end up fitting into any smaller, well defined group, but instead got thrown in to the large garbage bin of clades. They tend to have relatively dark, yellow spores (usually darker than the hot “core” clade Russulas) and are mostly mild, with a few notable exceptions. The many mild species vary a lot in their density, texture and flavour, so some make better edibles than others. Many people indiscriminatly eat all mild Russulas with relatively few reported poisonings, but most cannot be identified reliably so the edibility of most species is still officially unknown. A few species, like the shrimp Russula aff xerampelina are popular edibles. Let's break them down into sub-clades:


Hot Species - click for details

R. mordax © Ben Woo


Although most of these species are mild, unfortunately a few of them are hot, making them easily confused with the “core” clade.

Click here for details about the hot species.


Shrimp Russulas - click for details

R. sp. Woo 61 (aff xerampelina, R. viridofusca, R. favrei © Ben Woo

These are some of the most popular edibles in Russula, probably because they are easier to identify than most, not because they are the best tasting. They are not as dense and well-textured as some other species, but the shrimpy-ness disappears in cooking. The cap can be many different colours, but the following combination of characters identifies them:

Also in this section I discuss our local species that resembles R. olivacea, which differs in significant ways.


R. sp. Woo 61 (aff xerampelina) - usually a wine-red cap (left photo) with significant stem flushing

R. viridofusca - usually a yellow-brown cap (centre photo) without much stem flushing. Sometimes referred to as R. isabelliniceps np

R. favrei - usually a brown cap with a dark brown disc (right photo), also without much stem flushing. Mistakenly referred to as R. elaeodes


R. sp. Woo 58 (cf olivacea) - described above. We do not actually have the real R. olivacea.

Click here for details about the shrimp/olivacea species.


Greying Russulas - click for details

R. vinosa, R. 'vinososordida' © Ben Woo

Some Russulas with brightly coloured caps will turn grey after you scratch the stem (as the white Compactae group can). They might also turn reddish before turning grey. Two separate sub-clades have species that behave this way (although not every species in the clades exhibit this feature). The two most common species are not that closely related (one is in each sub-clade).


R. vinosa (occidentalis) - purple/green cap (left photo)

R. 'vinososordida' - orange cap (right photo)

R. 'californiensis' or something close, which may occur this far north (rosy-red cap) as well as a rare species with a dirty yellow-brown cap are to be looked for.

Click here for details about the greying species.


Yellowing Russulas - click for details

R. sp. Woo 45 (cf abietina) © Ben Woo

Some Russula stems and other fleshy parts will turn a fairly bright yellow in age. They seem to make up a good sub-clade of related species, although not every species in the clade may exhibit the yellowing staining very easily.


R. puellaris group - we have been incorrectly calling these R. puellaris. We have two common conifer species, R. sapinea and something we are calling R. 'abietina' but may or may not be that, and a third rare unnamed species.

R. versicolor - with birch, the only reported acrid tasting species. The colours are, as the name implies, variable and the yellowing on the stem is subtle.

Click here for details about the yellowing species.


Encrusted Cap Hyphae and Yellow Russulas - click for details

R. murrillii group member, encrusted primordial hyphae © Danny Miller, R. 'turci' and R. postiana © Ben Woo


R. murrillii group - we have the local R. murrillii, the European R. 'turci' and a rarer unnamed third species, with purple to reddish-purple matte/pruinose caps. They somewhat resemble the rare, unrelated R. sp. Woo 16, but that has a purple flushed stem.

R. postiana - rare, shiny yellow capped species with the encrusted cap hyphae, but not enough to make the cap matte. The spores are especially dark yellow. Mistaken for R. lutea.

R. olivina - rare, shiny yellow capped species without the encrusted cap hyphae.

Click here for details about the pruinose purple species and our yellow Russulas.


Russula zelleri group - click for details

R. pseudotsugarum © Ben Woo

This is a group of at least 5 related species, very hard to tell apart. For the most part, they are somewhat fragile, mild tasting, purple (sometimes with red), yellow spored Russulas with normal shiny caps. Five of them definitely match that description; two other mystery Russulas are somewhat related but look different. They are much like the R. turci group except they don’t have the encrusted pileal primordial cystidia, so they do not have a matte or pruinose textured cap.


R. zelleri group - mistaken for R. cessans, this group of purple to reddish-purple smooth/shiny capped species (in contrast to the matte/pruinose caps of the R. murrillii group) includes the common R. zelleri, R. obscurozelleri and R. pseudotsugarum. Two rarer species do not yet have names nor descriptions.

Click here for details about the R. zelleri group.


Other Russulas - click for details

© Ben Woo - R. graminea, R. sierrensis, R. benwooii, R. benwooii


R. graminea - (first photo) a not uncommon smooth, green capped Russula easily mistaken for the Heterophyllidia R. aeruginea.

Two rare species, sp. Woo 48 and sp. Woo 49, of which little is known.

R. sierrensis - (second photo) a not uncommon highly variable mushroom with a red, purple, brown (or all of the above) cap, easily mistaken for the R. zelleri group and others.

R. benwooii - (last two photos) a very common large Russula mistaken for R. 'xerampelina' shrimp Russulas or R. vinosa. It is often (but not always) brownish tan, perhaps with areas of olive, and a stem that may be slightly flushed grey or vinaceous.

Once, under introduced birch in Seattle, something close to R. velenovskyi was found.

Click here for details about the other Russulas.


Summary of Future Studies Needed


Brevipes and Compactae

Figure out how to distinguish the half dozen or so members of the Russula brevipes group and name those that are distinct species.

Determine if R. atrata is distinct from R. albonigra.

Determine how to distinguish the three members of the R. adusta group and which constitute new species.

Determine if hot taste reliably separates R. aff acrifolia and if it is a unique species needing a new name.


Ingratula and Crassotunicata

Investigate reports of R. granulata

Determine how to differentiate the three R. recondita group species and put names on them.

Determine how to better separate sweet smelling species 3, and 4 (and if 4a is a unique species or not), and put names on those and sp. 5.



Study the entire Heterophyllidia group, there’s not much that doesn’t need study here.


Core Clade

Determine what R. crenulata is and if it’s sp. 24 or 24a.

Figure out if our species is the real R. laccata

Figure out if R. sp. 29 is R. viscida or not

Better ways to distinguish R. salishensis and R. pseudopelargonia, including how to recognize sp. 38.

A better understanding of sp. 31, sp. 33 and R. alcalinicola.

Figure out what R. flava var pacifica is


Crown Clade

Distinguishing R. firmula and R. ‘punicea’ including getting a proper name for R. ‘punicea’.

Give a new name to our R. ‘olivacea’

Investigate how to recognize greying species 56 and 65 and give them names

Sort out the yellowing group – Name #42. Figure out if we really have R. sapinea. Figure out what R. abietina really is and is it #45 or does that need a name?

Why are R. murrillii and R. turci sometimes described backwards from each other? How to tell sp. 71 apart from those two.

Sort out how to recoginze the various members of the R. zelleri group, especially how to recoginze sp. 51 and sp. 51a.

Learn something more about sp. 48 and sp. 49, of which really nothing is known yet.

Get a good description of the many faces of R. sierrensis and how to recognize it.

Figure out if we have R. velenovskyi or if ours needs a new name, and how to recognize it better.

Solve the mysteries of what are R. placita, R. marina, R. inconstans and R. maxima

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