Brevipes and Compactae Clades - click to expand

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.

• completely white when young (other Russulas have a coloured cap cuticle and may be whitish, but not usually this white)
• stain brown in age, sometimes a lot
• quite dense (thus the term "compact") and heavy
• have subgills (shorter gills that don't go all the way from stem to cap, most other Russulas don't)
• there is no top cap layer to peel off easily (there is a coloured cuticle on most other Russulas that peels, at least somewhat)
  • Compactae: these turn black in age, sometimes with an intermediate red phase. It can be quite striking!
  • Brevipes: do not, but they brown considerably in age.

Species mentioned: Russula cascadensis, brevipes, nigricans, dissimulans, albonigra, sordida, subsordida, atrata, adusta, densifolia, acrifolia, anthracina

Brevipes - click to expand Species mentioned: Russula cascadensis, brevipes These are the mushrooms that are “underneath” the famous, popular edible Lobster mushroom. An orange mold called Hypomyces lactifluorum, covered in pimples, grows all over the mushroom and throughout the flesh (turning the gills into shallow ridges somewhat like Chanterelles), but unlike most molds, improves the flavour and texture of the food it is growing on. They are already crunchy to begin with, but the Lobster mushroom gets rid of the “grainy” texture that they sometimes have and enhances that crunchiness, and adds a complexity of flavour that they otherwise lack. Some of these Russulas are hot, like hot peppers, but I don’t know of anybody reporting a hot taste in a Lobster mushroom, so perhaps they only grow on the mild tasting ones, or, perhaps more likely, the hot taste that is known to be somewhat volatile is taken away by the mold. The Lobster mushroom is fairly easy to recognize because of its bright orange colour in the outline of the shape of a Russula and by the tiny pimples that cover the entire surface. (As an interesting side note, the spores of the mold only grow within the pimples, not everywhere. They sacrifice quantity for the quality of being able to eject them more forcibly like a volcanic eruption. This is a quality shared by the sexual stage of most of the “ascos” in the Sordariomycetes class, including the famous “zombie ant” fungus, Cordyceps, which is also covered in pimples). Lobster mold growing on a Russula ‘brevipes’ group member, uninfected and infected – Photos © Steve Trudell You can also use a Lobster mushroom to dye wool and other fabrics. This makes the lobster mushroom an incredible wild mushroom to learn for both culinary and crafting purposes.   Russula cascadensis OR - The smallest mushroom in this clade, with the cap usually <9cm across. The spore print is cream coloured (somewhat yellowish), darker than the other members. It is almost always hot tasting, at least raw, and is not recommended as an edible. I imagine it has been eaten “by mistake” so we would probably know if it was universally poisonous, but we just don’t know for sure. This mushroom was first discovered and described from Oregon. It appears to be a conifer species. R. cascadensis © Ben Woo   Russula brevipes group - Larger, usually >9cm across, and sometimes 20cm or more (one of our largest Russulas). The spore print is mostly white, with at most a hint of yellow. The taste may be mild or hot. Some specimens have a really mesmerizing blue tinge to the gills, or even a thin blue line running around the top of the stem at the base of the gills. Russula brevipes was discovered and described from the east coast in 1890, because Peck recognized that the mushrooms he saw were ever so slightly different than the ones in Europe, so he thought we might have a different species here. Many people disagreed at the time, and this kind of debate is happening within almost every group of mushrooms to this day. DNA studies are helping to sort out the answers. Nobody had yet formally suggested that our west coast mushrooms might be different from all of the above, the current theory being that we have the same R. brevipes they have on the east coast (not the species found in Europe). But our studies are showing that our local mushrooms are unique. In fact, we have found up to a half a dozen different species may grow in the PNW that for now, cannot easily be told apart. It was thought that the hot ones with the blue tinge represented one variety, “variety acrior”. But more than one of the species can exhibit these characters, so the truth is not that simple. For now, none of our half dozen or so species have names, so the best we can do is to say that we have mushrooms in the Russula brevipes group. Russula brevipes var. megaspora CA - one variety has larger spores than the others, so patient mycologists that look at the spores of every specimen will be able to identify this variety. It most likely needs promotion to species. Russula 'brevipes PNW24' - one fairly consistently “hot” species grows with oak (at least most of the others grow with conifers) so habitat may an indicator of this species. Russula 'brevipes Woo 21'/'brevipes PNW25' - two conifer species seem to be the most common, but we are still working out how to tell them apart. Unfortunately, both of them have been observed mild or acrid, with or without the blue tinge to the gills and blue line around the stem apex. Russula 'brevipes US3' has been found on Vancouver Island (both normal and as a lobster), Bridle Trails State Park in WA with conifers, as well as other parts of North America and the world, but it's not clear yet if it's as common as the first two species. Russula 'brevipes MX2' - yet another species has been found in Mexico and northern BC near Alaska, but not yet in the PNW. That range makes me think that it could grow here too. California and the east coast also appear to have additional species that have not yet been found here. Russula 'brevipes Woo 21', one of two common conifer species © Ben Woo Some group members have a blue line around the base of the gills (photo © Paul Kroeger) or a bluish tint to the gills themselves (photo © Danny Miller, subtlely shown in the way the light is hitting the small Russula on the right). This is often accompanied by an acrid taste.

Compactae - click to expand Species mentioned: Russula nigricans, dissimulans, albonigra, atrata, adusta, densifolia, acrifolia, anthracina These mushrooms can go through quite a transformation. One member from Japan, Russula subnigricans, is deadly poisonous and its close relative that grows in California, Russula cantharellicula, may also be poisonous, so we do not eat any of these, just in case, even though none of the ones that grow here have been proven to be poisonous. The enzymes that make the Compactae turn black seem to deteriorate the DNA quickly, as it is proving very difficult to sequence ones that have been kept for any length of time. We have had to find new material to attempt to solve the mysteries of this group, as none of Ben's old collections were able to be successfully sequenced.   Russula nigricans EU - probably our most common clade member, can get quite large (20cm), has gills spaced the most widely apart, turns distinctly red when damaged and then very black and has a dry cap. On the east coast they have the related species R. dissimulans, but although some have thought our species looks more like that than the European R. nigricans, our more than half dozen specimens tested so far are all proving to be R. nigricans. It seems to associate with many different trees in Europe so maybe that’s why it was able to migrate over here successfully, where it is commonly found with Doug fir and hemlock, even though those are different genera than the conifers found over there. Since R. albonigra, below, was split into many species, there is a possibility that will happen with this species too. Russula nigricans starts out completely white, turns red where damaged and eventually, completely pitch black – Photos © Ben Woo and Danny Miller   Russula atrata complex ('atrata PNW10'/'atrata PNW10a'/'atrata PNW10b'/'atrata PNW10c') (R. albonigra EU misapplied) - these are also large and dry capped, but goe straight to black without the red, have their gills a little closer together and may taste of menthol. A recent paper clarified that there were 5 species going by the name R. albonigra in the EU, and none of them occur in NA. Our two "species" that they knew about were left unclarified, and subsequently I have found 2 additional clades of local sequences. The ENA names Russula sordida NY and Russula subsordida NY could very well be available names. So is Russula atrata OR. It was described from Oregon near oak and pine (and said to be common down through central California), as having an unusually thick cap cuticle. Don't expect to be able to tell these species apart. The way the 5 EU species are told apart is by looking at tiny appendages of certain cystidia. But since all the worldwide collections fell into more than a half dozen clades 0.5% apart in ITS, they decided to make them all distinct species, something that may or may not have been the right decision. They may have opened up a can of worms requiring dozens of more species to be described worldwide if their pattern is followed. Perhaps it's best to refer to all of our species as the Russula atrata complex as they are all closely related within 1% or so in ITS. Russula albonigra is actually quite distant in ITS, and definitely the wrong name for any of our species. Russula 'atrata PNW10' and R. 'atrata PNW10a' © Danny Miller,     R. 'atrata PNW10c' © Drew Henderson (2 images)   Russula adusta group - The Russula adusta group is a common (as a group) bunch of more regular-sized mushrooms (up to ~10cm) with greasy or sticky caps instead of dry caps that may smell of wine barrels. The stem bases may get wrinkly. They go at least a little red before turning grey, but not always as pitch black as the previous two species can. Our studies show that we have three species here. One of them may be the real thing (or is at least close, 1-2 base pair differences in ITS2 but 3-5 differences in ITS1 which may or may not indicate a different species). Russula 'adusta PNW04' - was found near Victoria, BC with old growth Doug fir and hemlock, near Snoqualmie Pass in WA and near Goose Lake in southern WA. Russula 'adusta PNW05' - has the same description, and found so far in BC, WA and OR. Russula adusta EU - was found at Mt. Rainier at the Dalles under unspecified trees, and in Alaska with spruce and aspen and birch, and is the closest genetic match to the real species. It’s still >=1% different, and may or may not be the “real thing” or a third local species. Russula 'adusta PNW04' © Steve Trudell,     Russula 'adusta PNW05' © Marty Kranabetter,     Russula adusta © NAMA and the Field Museum of Natural History   Russula 'acrifolia IN85' - is a Compactae species that is consistently hot tasting. It turns red where scratched, then eventually to grey or black. It's probably a little greasy capped like R. adusta instead of dry capped. Tree association reports are vague. ITS is somewhat variable in this species (between 3 and 7 differences between them), and there could be more than one species hiding in here, especially considering how R. albonigra was split based on small ITS differences. One possible identity of Russula acrifolia itself is in this clade of species. Russula 'acrifolia PNW11' - four OR collections have a very large indel difference, plus 3 bp differences, plus 2 ambiguous differences. This seems distinct enough for me to separate it from IN85. We need to verify it is acrid too. Until now it has been assumed that our hot tasting Compactae was Russula densifolia, since some mycologists think that can also be a hot tasting Russula. This indicates that the Europeans should also take a closer look at their species to see if the “real” R. densifolia is mild or hot or possibly both.  Russula anthracina is another hot tasting Russula that usually does not turn red before turning black. We're not sure what it is. Most sequences show it as a unique species, but at least one collection turned out to be near this Russula acrifolia group. Until that is sorted out, I can't be sure of the proper name of our species. Russula 'acrifolia IN85' © Danny Miller,     R. 'acrifolia PNW11' © Jonathan Frank and Bitty Roy (2 images)

Fetid Russulas - Ingratula and Crassotunicata Clades - click to expand

They taste somewhat peppery, which is bad enough, but even worse than that, they often have a putrid or nauseating odor and flavour.

• most of them have a yellow-brown to brown cap cuticle with a strongly striate margin
• often taste somewhat peppery. Ingratula have a putrid, nauseating odor and flavour
• some brighter yellow-brown ones smell nauseatingly sweet like marashino cherries, the plain brown ones do not.
• if you cut a fresh stem lengthwise, you may find hollow diamond shaped chambers inside (true for many species in both clades).

• Crassotunicata has a small, white spored, bright yellow member (which is easily mistaken for core clade species) and an odd one out that is white (but browns considerably) with a very thick, peelable gelatinous cap cuticle.

Species mentioned:  Russula cerolens, sororia, pectinata, pectinatoides, granulata, recondita, laurocerasi, fragrantissima, foetens, pallescens, farinipes, crassotunicata

Ingratula - plain brown - click to expand Species mentioned: Russula cerolens, sororia, pectinata, pectinatoides, granulata, recondita, amerorecondita, garyensis   Russula cerolens OR - Our most common member is R. cerolens, mostly dark brown, often with an even darker center, <10 cm across, with the usual striate margin for this section. The spore warts are strongly connected almost into a reticulum. It’s found with spruce, pine, Doug fir, etc. and maybe some oaks too. For some reason, we had been using the EU and ENA names R. sororia, R. pectinata and R. pectinatoides for this and the following mushroom group, instead of the local name. Some have reported R. granulata with red-brown granules on the cap, so I do want collections of that to figure out what it really is. Only Christine Roberts, in her PhD dissertation, seems to have recognized that R. cerolens was not just a rare endemic, but actually the most common mushroom in this clade found here. Russula cerolens © Ben Woo   Russla recondita group - Perhaps without a dark stain in the center, but the only reliable way to tell you don't have R. cerolens is to find spore warts that are mostly isolated (they are strongly connected to each other in R. cerolens). In 2016 Russula recondita was described from Europe giving me a name I can use for our three species that are closely related to it. No word yet on how to tell the three apart. Russula 'garyensis Woo 6' - a sister species to an east species from Gary, Indiana. Found once in WA (no trees noted) and once in OR (near alder and hemlock). Russula amerorecondita (Woo 6a) - more common back east but also occurs here, found once at UBC under oak. Formerly referred to as “Russula recondita complex clade I” in the R. recondita paper until it got an official name recently. Russula 'recondita Woo 6b' - found once on Vancouver Island under Garry oak, once in Oregon under Garry oak, and twice in California under different oak.   Russula 'garyensis Woo 6' © Ben Woo, unfortunately showing that these species may have a dark center as well. Russula amerorecondita © Stephen Russell, an Indiana collection, rather bright yellow in youth (aging more brown) Russula 'recondita Woo 6b' © Christian Schwarz

Ingratula - yellow brown, sweet odor - click to expand Species mentioned: Russula laurocerasi, fragrantissima, foetens, fluvialis   Russula laurocerasi/fragrantissima/foetens group - We appear to have 3 or 4 species that are brighter yellow-brown and smell strongly sweet. They can get quite large (20cm)! These are the European names we’ve been incorrectly using for them. Russula 'laurocerasi Woo 3' - with a specific almond or maraschino cherry odor that is usually pleasant but may be unpleasant to some, was found in mixed live oak and conifers in California but also in Oregon, perhaps near Eugene. It is most closely related to R. laurocerasi (whose proper name may be R. grata). Russula 'laurocerasi Woo 4' was found near Snoqualmie Pass WA, in Idaho with cedar, hemlock and maybe Doug fir, and somewhere in the Seattle area. Russula 'laurocerasi Woo 4a' was found in the Olympics, near Mt. Rainier, near Stampede Pass in Washington, and in Idaho where it was under spruce, pine, and fir. It has 4 differences and 2 indels in ITS2 and only 1 difference and 1 indel in ITS1, but there’s not enough evidence yet to definitively separate it from Woo 4.  All these species have “wings” on the spore warts that Woo 5 does not possess. The spore size of Woo 3 is larger than those in Woo 4, but all have smaller spores than Woo 5. R. fluvialis EU (Woo 5) (R. foetens misapplied) is NOT usually reported as having an almond or marashino cherry smell, but instead a more generically sweet odor. Its odor is typically reported as nauseating or sickeningly sweet like vomit, whereas more people seem to find the other two species’ odors more pleasant than this one. This species is more closely related to R. foetens than the others, and the description of local collections very much matches that description. It has been found in the interior (the Blue Mountains of WA, in Idaho and in Northern BC). The one collection with nearby trees noted was hemlock and fir. This species has the largest spores. When R. fluvialis was described recently, a PNW collection of Russula Woo 5 was included in the official clade of the new species. It only differs by 1 bp in ITS2 and 2 chunks of indels in ITS1 right next to each other from EU collections. All the differences could be explained by 2 or 3 mutations. But it should be noted that R. fluvialis was described without a distinct odor, which is odd. unsequenced Russula laurocerasi group member © Joy Spurr, showing the cool hollow diamonds of most mushrooms in section Ingratula. Russula 'laurocerasi Woo 3' © Ben Woo,     R. 'laurocerasi Woo 4' © Danny Miller,     R. 'laurocerasi Woo 4a' © NAMA and the Field Museum of Natural History R. fluvialis (Woo 5) © Ben Woo

Crassotunicata - click to expand Species mentioned: Russula pallescens, farinipes, crassotunicata   Russula pallescens EU (R. farinipes misapplied) - These next species have been separated out into a separate section called Crassotunicata. This smaller, more slender mostly yellow species has whiter spores and gills than the preceding species. It’s more hot and less nauseating in its odor and taste. It’s a fairly rare species probably associated with conifers. This species and R. farinipes are very similar and both from Europe. Most people thought ours more closely matched R. farinipes and only Christine Roberts (that I know of) in her PhD dissertation thought correctly that ours was actually R. pallescens. Russula emetica and others in the core clade can be yellow and taste hot with white spores, and they are more common, but they are probably less strongly striate and less likely to have the hollow diamonds in the stem. Russula crassotunicata WA - described from Washington under conifers. It may be found all over North America and also Japan. Easy to identify with its white colours that brown considerably with a unique thick, peelable gelatinous cap cuticle. It too may have hollow diamond shape chambers in the stem. Russula pallescens © Ben Woo,     R. crassotunicata © Kit Scates Barnhart

Heterophyllidia I and II Clades - click to expand

This group needs the most study. They are difficult to identify and not much is known about them. 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:

• spores white to cream (not usually that dark of a yellow)
• mild tasting and often dense
• sometimes the caps are matte or even pruinose (but not always) instead of shiny and sticky, which is quite beautiful.
• gills might be flexible and buttery instead of brittle.
• more often in brown, green and grey colours, only rarely the red wine colours you see in other Russulas
• the flesh may or may not brown in age and usually there is no coloured stem flush, but that’s not always the case.
• the microscopic spot where the spore attaches to the basidium is small and doesn’t darken in iodine as much as it does for other Russulas

Species mentioned: Russula brunneola, heterophylla, virescens, smithii, aeruginea, mustelina, medullata, vesca, parazurea, atroglauca, grisea, modesta, malva, cyanoxantha

Russula smithii WA - a greyish green and/or brown mystery, described from the Olympics. Nobody knows which Russula Smith found. Some think it might be a Heterophyllidia. We need a type sequence or new collections.

Russula brunneola VT - This has white spores, with no real hint of yellow ever appearing in the gills. The cap is a beautifully textured dark brown. It’s described from the east coast under spruce and probably other conifers. Here, it is usually, but not always, found with spruce. One Alaskan collection had green and grey colours mixed in, and a stipe that greyed. One WA collection had lots of brown on the stem. All these characters are said to occur uncommonly, but that was the only collection that showed variation. We need a sequence of a definitive R. brunneola from the east coast to prove that our species is the same, but all of our sequences closely match a sequence of a “not quite proven” specimen from Quebec, with 3 differences in ITS1 and 2 differences in ITS2 between coasts. For now, I’ll keep using the name R. brunneola. There are rumours of green capped white spored species like R. heterophylla or R. virescens that are probably false (perhaps the specimens found did not really have white spores and are actually species described later in this section).

Rusula brunneola © Ben Woo

Russula sp. Woo 11 - This species has a cool pinkish brown matte to suede-like capped mushroom and was only found twice in coastal WA and OR, with hemlock but where spruce and true fir were also present in Oregon. It appears to have very pale spores like R. brunneola.

Russula . sp. Woo 11 © Ben Woo

Russula aeruginea EU - This pretty yellow-green to olive-green Russula has cream spores (most of the rest of the species on this page have cream spores). It has a shiny cap, unfortunately, so it can be mistaken for unrelated green Russulas, like the darker yellow-orange spored R. graminea (see the Crown clades) and others. One collection showed a fairly pure white spore print, so that may cause confusion too. It’s the, shall we say, least uncommon mushroom in this entire section. Our collections, like those in Europe, are found near various trees including Doug fir, hemlock, pine and alder (both conifers and hardwoods).

PNW collections from Washington, Oregon and Idaho are very close to the European species with only 2-3 different nucleotides in ITS2 and a perfectly matching ITS1. But European collections, our PNW collections, Alaska collections and Montana/boreal BC collections form 4 different clades all about 3 bp from each other. For now, let's assume that it is a variable species and not that there are a whole bunch of sister species that need to be described individually.

Russula sp. Woo 13 - This even prettier green Russula has a beautiful matte, not shiny cap, and cream spores. It is one of my favourite mushrooms. It doesn’t have a name yet, just lucky #13. It seems to be found with the usual conifers (probably Doug fir and hemlock). It has probably been mistaken for R. heterophylla, which does not seem to occur here,  by those who did not check that the spore print is not white.

Russula sp. Woo 15 - Another Russula, #15, has a matte cap that can sometimes be green (see the edge of the cap of the smaller stemmed specimen) but most of them might be (or usually show some) brown as well. It has cream spores. It’s also a Doug fir and hemlock species.

Russula PNW12 - known from one WA collection, a root tip sequence on spruce and hemlock from Vancouver Island BC and one collection in China. It is green, but as it was waterlogged, the cap texture is not known. The tree position doesn't indicate that. It was very rusty in age, unusual for a Heterophyllidia.

Russula aeruginea © Ben Woo,     Russula sp. Woo 13 © Ben Woo,     Russula sp. Woo 15 © Ben Woo,     Russula PNW12 © Danny Miller

Russula mustelina EU - This European mushroom has a dull matte brown cap, usually without any green tones but maybe a pinkish or vinaceous tint. It has cream spores. It is reported to not always poke out of the ground very far and is found under conifers.

It averages about 3 bp differences in ITS2 from the European species but for now I will assume that is the same thing, as European sequences differ by around 4 bp themselves. One local sequence had an extra two pairs of bp reversed from the others. Christine Roberts and Darryl Grund both suspected this species might occur here in their respective dissertations, but it wasn’t proven until the study of Ben’s collections.

Russula ‘medullata Woo 18’ - Also matte brown, but usually with some olive and greyish tones, unlike R. mustelina. It has cream spores. It is 9 bp different from the European species. In Europe, it’s found under hardwoods, but here, it’s found under conifers (both Doug fir/hemlock and spruce/pine forests). That is another reason to consider it a separate species.

Russula mustelina © Ben Woo,     Russula 'medullata Woo 18' © Ben Woo

Russula modesta NY - a fragile Russula usually with a mixture of green, grey and purple colours. It is found here so far only under landscaped oaks in Idaho with Amanita phalloides, matching east coast sequences from Indiana.

Russula modesta © Joe Matanzas,     R. modesta from IN © John Plischke and Brian Hunt

Other rare, unnamed Russulas

Unfortunately, there are at least 3 other species probably with pale, creamy yellow spores that are brownish (with perhaps a hint of pinkish or violet) like R. Woo 15, R. mustelina and R. aff medullata. They are not as common, but will be easily confused with these species. The Heterophyllidia have the biggest need of future study.

Woo 9 is a dull (matte capped?) brown colour and found once near Gold Bar, WA (trees not noted). It appeared to be staining red then brown, but that seems odd, so more collections matching this DNA will have to be found before anything tangible can be determined. It was not photographed fresh.

Woo 10 is another dull (matte capped?) brown mushroom with reddish violet tones when young. It was found once in the North Cascades, WA under Doug fir, hemlock and cedar. It was not photographed fresh.

Woo 14 is a smooth to shiny capped species that is tan coloured with some green tints (much like sp. #15, but without the matte cap). It was found twice, on the Olympic coast and in Idaho (and also in Alaska). The nearby trees were not noted, and there are 1-2 base pair differences in the DNA between them. The Idaho sequence oddly has a chunk of 8 nucleotides missing that is present in the other two.

Woo 16 is unusual. It has dark purple pruinose cap (that fades to cream), perhaps with some purple stem flushing. The Heterophyllidia are usually not purple and don't have stem flushing. The cap is matte and therefore can be mistaken for the Russula murrillii group (see the Crown clades), except those species do not typically have any stem flushing, and are more fragile (not nearly as dense as this one is) with darker yellow spores than this mushroom (which has cream spores), and the caps do not seem to fade as readily. It was found under conifers like fir, Doug fir and hemlock.

Russula sp. Woo 14 © Ben Woo,     Russula sp. Woo 16 © Marty Kranabetter and Michelle C. Torres-Grant (2 images from CA)

Russula PNW07 and CA01 - these rare species are only known from 2 collections and 1 collection respectively from an oak savannah in southern Oregon. Nothing much is known about them yet.

Russula PNW07 and CA01 (2 images) © Jonathan Frank

Other very pretty (and not brown) Russulas in this section that are rumoured to be in the PNW but without evidence are R. vesca, R. parazurea (powdery blue, truly beautiful!), R. ‘atroglauca’ (greyish-green darkening to a truly beautiful almost blackish green) and R. grisea. There’s genetic evidence they may all be in California, so keep an eye out! R. ‘atroglauca’ has also been found in Alaska with spruce, birch and aspen (probably associated with the hardwoods, and in this case, it may or may not be the actual European species found here on the west coast) so it may also be found in the PNW which is between those locations.

Russula malva n.p. – Heterophyllidia II - A famously beautiful, large species with many variegated shades of purple mottled together, with whitish spores, is found in California and rarely in southern Oregon, usually with hardwoods like oak. It has gone by the name of R. cyanoxantha, but ours is a new species that may get the new name R. malva soon (n.p. means the name isn’t official yet). It is in a separate clade I call Heterophyllidia II. FeSO4 won’t turn this mushroom’s flesh pink like it will the rest of the mushrooms in this section (and many others).

Russula malva n.p. © Christian Schwarz

Core Clade - click to expand

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.

• any white spored Russula not found in previous sections is probably in this section, whether or not you can detect the hot taste.
• hot Russulas with cream spores are usually found here. Make sure to rule out the very common, darker yellow spored R. mordax (and the rarer purple/red, spruce Russulas R. 'firmula' and R. ‘punicea’, and R. versicolor), found in the Crown clades below
• a core clade cream spored Russula that doesn't taste hot is going to be difficult to identify; you'll have to learn their individual characters

Species mentioned: Russula emetica, silvicola, bicolor, hydrophila, griseascens, montana, crenulata, betularum, nana, cremoricolor, raoultii, parapallens, fragilis, phoenicea, hypofragilis, atropurpurea, lilacea, stuntzii, laccata, viscida, americana, rosacea, rosacea var macropseudocystidiata, rhodocephala, sanguinea, sanguinaria, queletii, sardonia, torulosa, salishensis, pelargonia, pseudopelargonia, renidens, albidula, persicina, alcalinicola, exalbicans, gracillima, flava var. pacifica

Crown Clade - click to expand

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 olympiana are popular edibles. Let's break them down into sub-clades:

Species mentioned: Russula mordax, firmula, punicea, versicolor, veternosa, tenuiceps, xerampelina, olympiana, viridofusca, isabelliniceps, favrei, serissima, subrubens, elaeodes, olivacea, vinosa, occidentalis, vinososordida, decolorans, mendocinensis, claroflava, flava, californiensis, versicolor, odorata, puellaris, cremeirosea, sapinea, nauseosa, abietina, blackfordiae, sphagnophila, brunneoviolacea, turci, murrillii, postiana, lutea, risigallina, chlorineolens, chamaeleontina, olivobrunnea, olivina, intermedia, cessans, lilacea, zelleri, obscurozelleri, pseudotsugarum, graminea, urens, cuprea, nitida, sierrensis, velenovskyi, basifurcata, benwooi, placita, murina, inconstans, maxima, persobria, citrina forma separata

Hot Species - click to expand Although most of these species are mild, unfortunately a few of them are hot, making them easily confused with the “core” clade. R. mordax - perhaps our most common Russula, often reddish with a paler disc (see photo). Mistakenly called R. veternosa. R. 'firmula'/'punicea' - two rare probably hot, purple, coastal spruce Russulas Russula versicolor - found under birch, may be hot tasting in the gills and is described below under the yellowing Russulas. Species mentioned: Russula mordax, firmula, punicea, versicolor, veternosa, tenuiceps   Russula mordax WA - Arguably our most abundant Russula, found with doug fir and hemlock, our most common trees. It is usually very acrid. The cap is reddish, sometimes pinkish to orange, and there is usually a paler splotch of yellow-orange in the center, unlike other species that have a darker center. It may have a slight stem flush but doesn’t brown very much. The odor is often floral, also causing confusion with the “core” clade. As you can see from the second photo, it is of course susceptible to colour variations and unfortunately won't always have its typical colouration. We have been calling this R. veternosa for years, a similar European species that is actually a relative of Russula 'firmula' and Russula 'punicea', described next. In 1936, Burlingham found and described this species from Seattle, but it appears that nobody noticed until the study of Ben’s Russulas revealed that more local Russulas matched this species than any other single species, and they were all a match to the type of R. mordax, often an exact DNA match. We wouldn’t have needed DNA studies to figure this out if we had paid closer attention to her publications. Russula tenuiceps, an east coast oak species, is said to be very similar (and if the same, that older name would take precedence), but although Russula mordax has been found under oak here, others think the species do show enough differences that they are not likely the same. Somebody please study east coast oak specimens so we can find out for sure. This mushroom is in the same general clade as the red-grey staining Russula vinosa, along with a few others, but it does not demonstrate that property, nor is it mild like the rest of them. R. mordax © Ben Woo and Steve Trudell   Russula ‘firmula Woo 55’ - This usually hot purple Russula was found by Ben 7 times with coastal spruce, so it very much resembles R. queletii but usually without the prominent stem flushing. It has much darker yellow spores. Then it was found once with spruce in Idaho (so much for its identity as a coastal species) The ID collection was unfortunately mild tasting and pale capped, further complicating its identification. Our local species is about 2% different in DNA than the European R. firmula. Russula ‘punicea’ CA - This fairly rare (up north in the PNW) related wine coloured Russula (from the same hot tasting clade in the Crown clade as R. firmula and R. veternosa, the species that R. mordax was long mistaken for) is also found with spruce on the coast and usually has a prominent stem flush like R. queletii, but it usually is redder than it and R. firmula (again, with darker spores than R. queletii). It was described from California in 1997, but unfortunately the name R. punicea was already taken back in 1945, so it needs a new name. An attempt to sequence the type failed, but collections from the type area of coastal CA match a recent coastal spruce OR sequence and two Ben Woo sequences from coastal WA and OR. R. ‘firmula Woo 55’ © Ben Woo and Krista Willmorth (2 images),     R. ‘punicea’ © Ben Woo and Ann Goddard

Shrimp/olivacea Russulas - click to expand First, the shrimp Russulas. 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: large, mild tasting Russulas. In the “crown” clade; the only other Russulas to regularly get >15cm across are R. vinosa< and R. benwooi. they smell like shrimp, at least in age a drop of FeSO4 turns them green, which doesn’t usually happen in any other group. the flesh is especially susceptible to browning the stems are sometimes flushed pink the spores are rather yellow so the gills are quite yellow in age Also in this section I discuss our local species that resembles R. olivacea, which differs in significant ways. It is quite dense and is probably one of the best edibles in the genus. does not smell like shrimp, does not turn green in FeSO4, does not brown much, and is much denser. It's cap is usually purple and/or green. It is also large with significant stem flushing. Species mentioned: Russula xerampelina, olympiana, viridofusca, isabelliniceps, favrei, serissima, elaeodes, olivacea, subrubens   Russula olympiana WA (Russula xerampelina EU misapplied) - Woo 61. Our abundant species is significantly different genetically than the European Russula xerampelina EU, so it finally got its own name, although I do not yet know of a way to separate them other than by country of origin and genetics. It is usually wine coloured (some shade of purple, red or both). There is often a lot of pink stem flushing. It is usually found with the typical Doug fir and hemlock and other conifers. Since this can be one of the easier groups of Russulas to identify, it is a popular edible. Unfortunately, sometimes it can be almost any colour, as seen in the second brown photo, where it could easily be confused with our 2 brown species. Russula olympiana © Ben Woo   Russula viridofusca WA - This shrimp Russula is also found with different conifers but is usually pale, pinkish-, yellowish- or greenish-tan and has little stem flushing. It can be green or even reddish and so probably can’t be unmistakenly differentiated from the others. It was originally proposed as Russula xerampelina var. isabelliniceps in 1965 by Grund who ended up describing it as R. viridofusca, from Washington in 1979. It is very common, collected by Ben more often than R. olympiana (31 vs 13 times), but that may be because he didn’t feel the need to sample the latter very often. We have the ITS1 sequence of the type, and longer sequences matching it. Russula viridofusca © Ben Woo (2 images)   Russula serissima MA (=Russula favrei EU, Russula 'elaeodes' EU misapplied) - We have been calling this species Russula elaeodes, but that species is probably the same as R. pseudoolivascens and R. clavipes, which turns out to be something different. This is another case where there were similar lookalike European species, and somebody chose the wrong one as our local species. Our species matches a pretty unanimous consensus of what the European R. favrei is, found with conifers. Then it was discovered that the older Russula serissima MA was the same thing, so that is the proper name. Here it might prefer hemlock and spruce. Its colour scheme is usually brown with an especially dark brown center. It has little stem flushing just like R. viridofusca. Perhaps it can have purple tones like R. olympiana so it also may not always be easy to distinguish. Russula serissima © Ben Woo   Russula subrubens EU - one single OR coast collection with a yellow cap matches dozens of EU sequence of R. subrubens. It is a willow species from dunes near the coast or from arctic/alpine habitats in the EU. It is described with a red cap, although ours had no red in it, and was found under California bayberry. If it does occur here with a red cap too, it might look like Russula olympiana, but this species seems to have no stem flushing. It has also been found in CO and MT. Russula subrubens © Buck McAdoo   Russula 'olivacea Woo 58' - This is not a shrimp Russula, but it shares some features. It’s large and has a pink flushed stem with fairly dark yellow spores. But this is an especially dense Russula (so some say it is one of the best edibles but others caution about possible poisonings), with a somewhat velvety purple and/or green cap that has a very interesting texture when fresh. It does not smell of shrimp. It does not usually brown as much as the shrimp Russula and FeSO4 will not usually turn it green, but both of those are variable features as I have found one collection that browned significantly and did turn green in FeSO4. It turns out we do not have the real R. olivacea here, but a sister species. It will need a new name. It is uncommonly found with conifers like Doug fir, hemlock and spruce.   Russula 'olivacea Woo 58' © Ben Woo (2 images) and Fernando Macias-Jimenez

Greying Russulas - click to expand 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. Three separate clades have species that behave this way (although not every species in the clades exhibit this feature). Species mentioned: Russula vinosa, occidentalis, vinososordida, decolorans, mendocinensis, claroflava, flava, californiensis   Clade 1 Russula vinosa EU (=R. occidentalis) - another large Russula with olive colours in the center and purple colours on the edges (or perhaps either one of those colours). The spores are a only faint creamy yellow, not as dark as many in the crown clade. Scratch it and wait a few minutes to a few hours) and it will turn reddish and then grey. Russula vinosa was described from Europe and then our west coast species was later considered a variety of that mushroom, eventually getting a promotion and getting its own name, Russula occidentalis. Anna’s study showed that there is very litle genetic difference between our species and the original R. vinosa (perhaps 2 differences or so on average) so if that and the macroscopic differences are not considered enough to justify a separate species, ours would just be considered R. vinosa. It is found with various conifers. Russula vinosa (occidentalis) © Ben Woo   Clade 2 Russula 'vinososordia Woo 57' - This large Russula (and a couple other species) are not in the same clade as R. vinosa, and represent a separate genetic lineage that stain similarly red/grey. It is uncommonly found with various conifers, but differs by usually having orange colours and a dark yellow spore print. When bruising is noted, sometimes a red stain is noticed before a grey stain, but sometimes only the grey staining is noticed. There are two European Russulas that match this description, R. decolorans (in clade 3, a Fries species from 1838) and R. vinososordida (from 2000, recently discovered to be a lookalike). Before 2000, it was assumed, of course, that all our specimens were R. decolorans (the only known species at the time). After 2000, nobody thought to check to see which species ours was. This study has shown that all 10 of Ben's PNW specimens matching this description are 1% different than the type of R. vinososorda, but not at all close to R. decolorans. This is probably a little too reliably different genetically to be considered to be R. vinososordida - a similar 1% difference separates the very different looking R. mendocinensis from both species. Also, our species is a conifer associate whereas R. vinososordida is a hardwood associate. It should be noted that R. decolorans has since been found in Alaska, and now recently in Vancouver BC, so both species do occur here, supposedly easily separable microscopically. Perhaps the confusion in the past was due to our local species being a conifer species like R. decolorans instead of a hardwood species like R. vinososordida. Russula mendocinensis, as I stated, is a very closely related mixed conifer-hardwood forest species (I’m not sure which trees it actually grows with) in California that can look and act surprisingly diverse. The type was purple, staining orange (but greying was not noticed), and was supposedly quite acrid tasting. However, a mushroom very much like R. vinososordida from CA sequenced to be R. mendocinensis instead, so it may not always look different than R. vinososordida. This will have to be looked into. R. claroflava (=R. flava), a bright yellow mushroom that stains reddish then grey, not yet known from the PNW, has been reported from here before but could represent sightings that were actually “aff vinososordida” or perhaps Russula sp. Woo 65. Russula flava var pacifica was described from Oregon, but it is clearly not a variety of R. flava, but very likely actually a “core clade” member, as stated on those pages. Save any yellow specimens you find that stain this way. Russula 'vinososordia Woo 57' © Ben Woo   Russula 'californiensis Woo 56' - Russula californiensis is second species closely related to R. vinososordida, a California pine species that is rosy red with a greying stem and mild taste. Our species is different and probably needs a different name (at least 6 substitutions and 2 indels different in ITS2, or >1.5%). It is rare, having only been found once with conifers (not pine) near Snoqualmie Pass WA, and was pinkish/rosy red with some greyish yellow tones. Although the greying in age or when damaged wasn’t specifically noted, it probably does. It was mild with creamy spores. Russula 'californiensis Woo 56' © Ben Woo   Clade 3 Russula decolorans EU - while all of Ben's orange Russulas that eventually stained grey (often turning red/orange first) turned out to be Russula aff. vinososordida and not Russula decolorans, we did later find R. decolorans in Alaska, northern BC and now finally in Vancouver BC and Oregon, so it is here too. It is probably also a conifer species. The two are supposedly easily separable microscopically. Perhaps the confusion in the past was due to both local species being conifer species like R. decolorans instead of hardwood species like R. vinososordida. Russula decolorans © Marty Kranabetter

Yellowing Russulas - click to expand 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. Species mentioned: Russula versicolor, odorata, puellaris, cremeirosea, sapinea, nauseosa, abietina, blackfordiae, sphagnophila, brunneoviolacea   Russula versicolor EU - The colours can be quite variable as the name suggests (usually with purple, red or maybe green), but careful attention to an admittedly slight stem yellowing can help identify it. Also, this is the only yellowing species group that is acrid tasting, especially in the gills. What is probably the real R. versicolor, under birch, is uncommon here. Russula 'versicolor Woo 44' - Less uncommon is the sister species, Woo 44, usually found under cottonwoods but also under birch. Two Seattle specimens of the "sister" species were found under cottonwood instead of birch, one Seattle collection was found with birch, and one northern BC collection was found with conifers. This species has 2-3 differences and 1 indel in ITS2 and a further 4 differences in ITS1 from sequences of R. versicolor itself. DNA matching this sister species is found all over Europe with the "real" R. versicolor as well, so the species are not distinguished by geography. Russula versicolor © Danny Miller,     Russula 'versicolor Woo 44' © Ben Woo   Russula 'puellaris Woo 42' - This is the rarest of three similar species in the yellowing clade, found once on Washington’s Olympic peninsula near conifers (spruce, Doug fir, hemlock and cedar), once on the Oregon coast west of Eugene and once in OR near Mt. Hood. Yellowing was not noted in the notes, but you can see yellowing in the first photo. This group of mushrooms have previously gone under the name Russula puellaris. That similar, closely related European Russula was thought to be our local species until these studies have revealed differently. Woo 42 is about 3% different from R. puellaris, probably enough to justify it being a different species. It is also about 3% different from R. cremeirosea. This first collection appeared to have more red tones than purple tones (although the colour was bad), but the second collection was purple with a darker disc just like the other 2 species. Russula 'puellaris Woo 42' © Ben Woo and Ann Goddard   Russula ‘sapinea Woo 43’ - Most yellowing Russulas in the PNW are this common one and the next one. This may be Russula sapinea EU, which is not known to turn yellow over there, but it does over here, so it should be more closely examined in Europe. The genetic variation is a bit high in sequences of this species (up to a full %), but without forming any groups, so I think it may just be all one species. It’s possible somebody will find a reason to split it into more than one species in the future. It is usually purple, but can be red-purple or closer to red like the previous species. Many different conifers were around when it has been found. Rumours of R. nauseosa being in the PNW are probably not true. I don’t even know if that European species is in the yellowing clade or in the R. zelleri group (found on the "zelleri group" page), but whatever it is, it’s not actually here. Russula 'sapinea Woo43' © Ben Woo   Russula 'abietina Woo 45' - This is the other common species. R. abietina was described from NY as being like R. puellaris except for not staining yellow. There is some confusion over what R. abietina really is. A study of Peck’s type collection in 1947 gave Singer the impression that it was just a mixture of the existing species R. blackfordiae and R. sphagnophila. If R. abietina truly was a new species, the Russulas that seem to be what Peck described have turned out to be one of two things. Firstly, this mushroom, that perhaps doesn’t always turn yellow in age or maybe the yellowing was not always noticed. When collected here, it’s usually noticeably yellowing. When collected in Tennessee, yellowing has mysteriously not been noted. You might expect to find the same species in New York as you do in Tennessee, so perhaps this mushroom is R. abietina. Secondly, it might be a mushroom in the Russula zelleri group (see that page). These were collected in California, I believe, which may or may not have the same species as what is found in New York. The aforementioned R. blackfordiae, R. sphagnophila and also R. brunneoviolacea are three poorly understood Russulas with dubious reports from the PNW, which this study can somewhat debunk. It is not even certain if those species belong in this clade, the R. zelleri clade, or the shrimp Russula clade. I’m going to take a guess that perhaps R. abietina is this mushroom in this yellowing clade, but all we can say for now is that it looks like it superficially, so thus my calling it “cf” abietina. To prove it, somebody will need to study the NY type, if available, and/or a number of matching collections from the New York area. Unfortunately, as far as I know, this mushroom looks identical to R. sapinea. The microscopy should be examined to see not only if this species could in fact be R. abietina, but if it can be told apart from R. sapinea and R. sp. Woo 42 (aff puellaris). This species is another conifer species. Russula 'abietina Woo 45' © Ben Woo

Encrusted Cap Hyphae (matte purple caps) and Cream/Yellow Russulas - click to expand These purple (or reddish-purple) members have pruinose or matte purple caps, the texture you often see in the Heterophyllidia clade, but they are more fragile and less dense, with darker yellow spores. Microscopically, you can identify the reason for the pruinose cap by finding “encrusted pileal primordial hyphae”. Pileal means in the pileus cuticle, the top layer of the cap. They are crusty bumps on the walls of otherwise cylindrical cells, a few of which are pointed out in this micro photo: In this section I also cover yellow Russulas that may or may not have the encrusted cap hyphae - but even if they do, they do not demonstrate it with a matte/pruinose cap texture; they usually have regular, somewhat shiny caps. Species mentioned: Russula turci, murrillii, postiana, lutea, risigallina, chlorineolens, chamaeleontina, olivobrunnea, olivina, intermedia, basifurcata   Note that the similar Woo 16 has a pruinose purple cap that seems to be more likely to fade, and perhaps with some stem flushing and denser flesh. Russula ‘turci’ (Woo 72) EU - This is the most abundant of the three local species, a European species found under various conifers that sometimes has an odor of iodine in the base of the crushed stem. The caps are most often dark purple, but may occasionally have some red tones or even have faded to yellow-brown. If any of the three normally purple species have faded to yellow, they could be confusable with the unrelated conifer species R. olivina, below. But if you look carefully at the caps of a fresh specimen (or under a microscope), there is something special about that “matte” appearance, an almost powdery look that makes this group especially beautiful, especially when dry, due to encrustations on certain cells in the cap cuticle. Species in the EU labelled R. turci make up as many as three different species, and without a type sequence, we don't know for sure which species is the real thing, but ours most closely matches a consensus of what probably is the real one. The genetic variation found in our local species (about 5 bp differences in ITS between us and Europe) is not necessarily enough to justify considering ours a separate species. An unusual amount of genetic variation in this species and R. murrillii has been found, possibly indicating that some kind of evolutionary pressure is currently causing speciation. Perhaps we are witnessing the formation of new species, and eventually, they will sort themselves out into distinct clades with further distance between them. Russula ‘turci Woo 72’ © Ben Woo, uncommon yellow form © Ben Woo   Russula sp. Woo 71 - This very closely related species cannot yet be told apart from R. turci. It is rare, found by Ben 3 times in WA near Glacier Peak with fir, Doug fir, hemlock and cedar nearby. Twice it was deep purple like R. turci, but once it had red tones like R. murrillii more often does, below. It has since been found once near Vancouver, BC. Russula sp. 'Woo 71’ © Ben Woo   Russula ‘murrillii’ (Woo 70) OR - In 1913, Burlingham described this fairly common mushroom collected by W. Murrill in Corvallis, OR under conifers. She didn’t directly compare it to R. turci, but others soon did, and recognized it as a related but separate species. This species is more often reddish-purple than R. turci is (but might also be deep purple or yellow-brown), and reportedly lacks the iodine odor that R. turci sometimes has. The caps may not be as obviously pruinose as R. turci, and the stature is usually a bit smaller. I don’t have type sequences of either species, but many sequences of specimens from the type areas of both species are consistent with each other so there is a consensus of what represent these species. However, the descriptions of this species and R. turci seem somewhat switched with each other which I can’t quite explain. The description for R. turci says it’s often wine coloured and the description for R. murrillii is supposedly almost always pure violet, but dozens of collections I've examined show the opposite is true. I await type sequences or some official decisions being made to make sure I have our species names correct. Russula ‘murrillii Woo 70’ © Ben Woo, encrustations on the pileal primordial hyphae © Danny Miller     Yellow Russulas – encrusted hyphae or not   Russula 'postiana Woo 69' - This rare, often bright yellow Russula has been going by the name R. lutea (synonym R. risigallina) over here for many years. It’s not always yellow, and specimens with other colours have been called R. chamaeleontina, for its chameleon-like properties, but that species is now assumed to be the same as R. lutea. Although our species DNA is twice as close to R. postiana as it is to R. lutea, the one collection with spore measurements better matched R. lutea. Hmm. Although in Europe the true R. postiana can have olive or brown tones (and maybe even a hint of pink or lilac), the only documented case of our species in the PNW is being bright yellow in Seattle under Madrone, although it’s famous for its spruce habitat in Europe. Like the R. turci group, R. postiana is in the clade with encrusted primordial hyphae, but not to the same extent, because although you can find them under a microscope, they do not seem to be sufficient in this species to make the cap matte or pruinose; it’s still relatively shiny. The spores are especially dark yellow, almost orange, possibly the darkest colour of any of our Russulas. This is another case of there being two similar European Russulas, in this case R. lutea (risigallina) and R. postiana, with mycologists deciding that our bright yellow Russula was probably R. lutea, but DNA evidence showing it is closer to R. postiana. (A similar thing happened as described on previous pages with R. pallescens vs. R. farinipes, with R. vinososordida vs. R. decolorans, with R. cerolens vs. Russula sororia and pectinata and also with R. sapinea vs R. puellaris). Ours may not be the same species, though. Ben's sequences of ITS2 showed 1-3 differences, not necessarily enough to justify a new species, but more recent sequences of ITS1 have shown that there are an additional 3-4 differences in ITS1. Russula chlorineolens OR - this is a trufflized sister species, with a strong chlorine odor and distorted gills that can turn deep yellow/ochre. The cap is cream colour and not nearly as yellow as its epigeous relatives. It is from spruce and hemlock. Russula 'postiana Woo 69' © Ben Woo,     R. chlorineolens © Michael Beug   Russula 'olivobrunnea Woo 46' - This rare slender lookalike of R. postiana can also be bright yellow, but it is not closely related to any of our other local Russulas and does not have the encrusted pileal hyphae. The spores are dark yellow, but not quite as dark as R. aff postiana, and a little larger. It is said to grow with spruce and fir, instead of hardwoods, but the three times it was found in the Blue Mountains of Eastern Washington and Idaho, the trees were not noted (although they are largely conifer forests). Recently, it was just collected near Bellingham under Doug fir and alder so it is on the west side as well which gave us an ITS1 sequence and shows that there are a total of 6 bp differences in ITS from R. olivobrunnea, pretty clearly enough to make this a separate species in need of its own name. Some related species are R. olivobrunnea which is a wine coloured species with a very dark centre and Russula olivina, a yellowish species with olive or brown tones, both from Europe. Russula Woo46a - a close relative to the yellow Woo46, but like R. olivobrunnea, it is purple coloured and illustrated in the zelleri group section. Russula PNW16 - another relative of R. olivobrunnea, but also purple and illustrated in the zelleri group section. Russula 'olivobrunnea Woo 46' © Ben Woo and Sharon Squazzo   Russula sp. Woo 65 - This sometimes yellow species also doesn't have encrusted cap hyphae and is probably most easily mistaken for Woo46, but is is a stockier mushroom with paler yellow spores. It was only found by Ben twice, once in Idaho and once near Glacier Peak, WA (near Doug fir and hemlock). In the two collections the cap colours were yellow (flesh did not stain) for the ID collection, and dirty brown (flesh stained orange then grey), for the WA collection. But there may have been a mixup and only one of those may represent this species. This species is in a group with Russula sierrensis and R. integra. Subsequently it has been found on Vancouver Island and Arizona (all with conifers, varying from sea level to high elevation) and we have a full ITS sequence, and it whitish or yellow, and the flesh does not seem to appreciably stain, so the dark brown collection that had flesh that stained orange then grey may have been a mixup. I don't know what it was. Russula Woo65 (incorrect, mix-up?) © Ben Woo,     R. Woo65 © Marty Kranabetter (2 images) Russula Woo65 (from AZ) © Jonathan Frank (2 images)   Russula 'basifurcata PNW01' - this white oak Russula has a cap tinged with yellow or reddish-yellow, and a pale yellow spore print. This meets the description of R. basifurcata, but as that is a poorly understood species, I'm giving it a code for now. It has been sequenced twice, once under oak in WA and once in CA. Ben never found it. Russula 'basifurcata PNW01' © Michael Beug   Russula 'intermedia PNW02' - This is a very pale yellow capped Russula (almost white), with very dark ocher spores notable by the fact that the gills are much brighter than the cap. It is rare, only found once so far near Mt. Rainier (and a memory that I might have seen one 10 years ago near Leavenworth). The nearby trees were not noted, but the closely related R. intermedia is a birch species with a bright red cap that might fade to yellow in the centre. Our species is 8bp and 1 indel different in ITS. Russula 'ocher oaks PNW03' - a very similar almost white cap and very dark ocher spores, but this species is with oak and is found in a different part of the genetic tree. One collection showed pink around the rims. Rumour has it that California has 2 similar oak species, this is probably one of them. We have one sequence from OR and one from CA. Russula 'intermedia PNW02' © Danny Miller (2 images),      R. 'ocher oaks PNW03' © Valerie Engle (2 images)

Russula zelleri group - click to expand This is a group of at least 5 related species, very hard to tell apart, and one similar species outside the group. 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. Species mentioned: Russula cessans, lilacea, zelleri, obscurozelleri, pseudotsugarum The first 5 species have been going under the name R. cessans, a European Russula that may or may not be in this group (another possibility is the yellowing clade), but that does not actually occur here. Nobody had studied our R. 'cessans' closely before the study of Ben’s collections to notice that there are really multiple species, and even if they had, they are very difficult to tell apart from each other. Russula zelleri was described, you guessed it, by Burlingham in 1936 from Oregon, in the same publication where she described R. mordax, and similarly, not much attention was paid to it, but if it had been, we might have noticed long ago that what we have long called R. cessans over here was actually R. zelleri and other related species. Here is an interesting story about a mushroom that does not occur in the PNW, R. lilacea, and how the very different R. zelleri group members were mistaken for it. In 1965, Darryl Grund’s UW PhD dissertation stated that the mild, dark yellow spored, purple European Russula lilacea was found very commonly in the PNW. He failed to notice that the description of R. lilacea was of a white spored mushroom, and he mistakenly stated that it was a yellow spored mushroom. He was actually finding members of the R. zelleri group. When Ben Woo made his Russula key in 1989, he followed Grund and stated that R. lilacea was very common in the PNW, but he did not copy Grund’s description of it, he went back to the original texts and wrote down the actual description of R. lilacea, not noticing the discrepency. So Ben’s key states that the mild tasting, white spored purple Russula lilacea is very common here. Then, many people happened to find a mild tasting R. phoenicea, a white spored Core clade member, and mistook that species for R. lilacea, not looking at the microscopy.   Russula zelleri OR - Russula zelleri itself is the third most common species in the group, occurring with spruce and pine. While usually purple, it has been noted to come in red or green colours too. There is a single ITS1 difference between recent collections and the 1936 Oregon type, and among the collections there may be 1 or 2 ITS2 differences. Russula subloculata ID - This may be one of those common cases where the secotioid mushroom evolves so quickly, ITS doesn't change. You cannot tell this apart from R. zelleri by ITS, so that is the species that is trufflizing. Russula zelleri © Ben Woo    Russula pseudotsugarum WA - The second most abundant species in the group, it can be found with spruce and pine as well, but also Doug fir and hemlock. It was recently named by Anna for its ability to associate with Doug fir (Pseudotsuga). It can also occasionally come in red or other colours like brown and green, making it very hard to recognize. It is almost identical to the next species. One collection had a very, very grey stipe, but that may have just been unusual waterlogging.   Russula pseudotsugarum © Ben Woo   Russula obscurozelleri WA - The most abundant species in the group, almost identical to R. pseudotsugarum and found near the same trees. It is named for its difficulty to tell apart from the other zelleri members. It can also fade to brownish yellow colours. One Washington collection tasted acrid, which confuses me. Russula obscurozelleri © Ben Woo   Rare relatives of R. zelleri   Russula sp. Woo 51 - This rare species is very similar, purple or purple-red, found only twice with spruce and pine on the Oregon Coast. We only have ITS2 of this species. There’s no way yet to tell it apart from R. zelleri. There is a hypogeous species in California, CA17, with one collection having the same ITS2 as Woo51. Russula sp. Woo 51 © Ben Woo   Russula sp. Woo 51a - from a coastal Victoria BC Douglas fir forest and from a CA pine forest. It has a quite deep purple cap. Russula Woo51a © Marty Kranabetter,     R. Woo51a © Taye Bright (from CA)   Russula sp. Woo 52a - two sequences from WA (different by 3-4 bp from each other) and one from Smithers, BC that matches one of the WA sequences. Our one good photo shows faded purple. Russula Woo52a © Michael Beug   Russula Woo46a - a close relative of the yellow Woo46, illustrated in the yellow Russula section, but like another close relative, R. olivobrunnea, it is purple coloured and therefore resembles all of the mushrooms in this section, although it is not closely related to them. This is currently very difficult to ID. It is known from a single Victoria BC collection. I wonder if it has very dark yellow spores like R. olivobrunnea. Russula PNW16 - a similar purple capped close relative of R. olivobrunnea, which does appear to have dark yellow spores like that does, but it's unclear how much darker than the other R. zelleri relatives in this section those spores are, and if that can help identify it. We have one collection from ID and one from OR. Russula Woo46a © Marty Kranabetter,     Russula PNW16 © Ed Barge

Other Russulas - click to expand Species mentioned: Russula graminea, urens, cuprea, nitida, sierrensis, velenovskyi, benwooi, placita, murina, inconstans, maxima, persobria, citrina forma separata   Russula graminea EU - This European Russula is fairly closely related to the R. zelleri group, described on the "zelleri group" page. It is green or greenish-yellow, so it could be mistaken for R. aeruginea and possibly others. Four Alaskan collections, however, only have some green, usually on the disc, with the rest of the cap being purple (the most common colour of other R. zelleri group members). It was not known from the PNW until Anna’s study of Ben’s collections. Our DNA is basically an exact match to R. graminea in Europe, with the exception of some confusion about how long a string of identical letters might be in a couple of places. This likely represents not being able to reliably tell the length of the string, not actual differences. Everybody probably mistook it for R. aeruginea and never looked at it too closely, but it is just as common (or not uncommon) as R. aeruginea is. It is known as a spruce species in Europe, but here it can also be found with our common conifers like Doug fir and hemlock. In Alaska, it was always found in a spruce-hemlock forest. The hot tasting yellow-green European Russula urens/cuprea that has been reported as being commonly found here has no evidence of actually existing in the PNW. R. graminea was once recorded as hot tasting, so this is probably the basis of those reports. If you find a hot green Russula with dark yellow spores, save it so we can double check. Russula graminea © Ben Woo   Russula sp. Woo 48 - These next two species are also somewhat related to R. zelleri, but not closely enough to look like it nor necessarily share many of its properties. R. sp. Woo 48 is rare, found once in Shelton near Olympia, WA near unspecified conifers. It had some purple in the center but the rest was pale to medium brown or had faded to brown. We don’t know what its usual colour scheme would be and if it can mimic the other purple species or not. For now, I can't say anything about how to recognize it. Russula sp. Woo 48 © Ben Woo   Russula sp. Woo 49 - A rare sister species to sp. Woo 48, it was found once near Satus Pass in S. Central WA near pine and Garry oak, it was entirely cream coloured with little pigment. Also found near Santa Cruz, CA, where it was reported as red and yellow, and near Sly Park CA. It is not known what its usual colour scheme is. Very little is known yet about how to recognize either of these species. Russula sp. Woo 49 © Ben Woo   Russula nitida - This is a European birch species with a wine-brown cap and a flushed stem. Our single collection had neither of those properties, but it was found in a Seattle park with birch as one of the many types of nearby trees. Little is known about how to recognize it over here.   Russula sierrensis CA (=Russula pacifica CA) - Both type sequences match and are from the same publication, so no name has priority, but I've been using the name R. sierrensis, as that was sequenced first, so I will continue to do so. It is fairly common at times. It can be found all over the PNW (from the Idaho interior to the Washington Cascades to the Oregon coast) under conifers (either Doug Fir or spruce seem to always be present, although in California it’s reported with oak and pine). The reason we never noticed it north of the California border before is that this study showed it doesn’t often look like it did when it was first found, it’s a much more variable-looking species than ever thought, so it was never recognized. Unfortunately, I have no good means of identifying it yet, but this species should be studied in more detail since it is not uncommon (Ben found it 14 times). It’s supposed to be a mild tasting purple-red Russula with a pale yellow spore print, but the spore print can often be dark yellow and the colour can be almost anything. Oh look upon the many faces of Russula sierrensis and despair. It’s possible one of these specimens could have gotten mixed up and not be this mushroom, but not all of them. Any Russula not confidently identified as something else (and that might be most of them) should be microscopically examined carefully for a match to this species, with separate spore warts that rarely connect and pileocystidia (special cells in the cap cuticle not always present in other species). It would be really helpful to get a reliable description of this species. The type of Russula pacifica seemed to stain grey, and this species is close to Woo65, which in one collection stained grey, although R. sierrensis was never reported to stain grey. This is somewhat mysterious. The many faces of Russula sierrensis © Ben Woo   Russula 'velenovskyi Woo 64' - This rare mushroom, related to R. sierrensis and others in the R. vinosa group, does not stain and was only found once under an introduced birch tree in Seattle. It is about 5 bp different in ITS2 (~0.7%) than the European species and may represent a unique species. In Europe, it has been found with other kinds of trees as well including conifers. It has a peachy-orange cap and creamy yellow spores that are not as dark as most of the others. It kind of resembles R. vinososordida but does not stain.   R. 'velenovski Woo 64' © Ben Woo   Russula benwooi WA - I have saved one of my favourites for last. Normally, when we discover a new species in the PNW, all we are really doing is admitting that our species isn’t really the same as the European mushroom whose name we’ve been using for it, but that enough differences were found to justify separating it out and giving it a different name. That’s not the case here. Arguably the most interesting result to come out of Ben’s study was that one very abundant Russula, found at least 60 times (second only to how often R. mordax was found) was something completely different than expected. This Russula was large, and kind of looked like our R. 'xerampelina' and R. vinosa, and was always mistaken for one of those two. Did it smell like shrimp or turn green in FeSO4? No, and anybody spending just a few seconds to try those things during the hundreds of times it has been collected would have noticed it was not a shrimp Russula like they thought. But when you’re finding them by the hundreds, who has time to test every one? You just start assuming you’re looking at yet another one. Anybody who thought it was R. vinosa could have dispelled that myth quickly by scratching it and noting that it did not change colour, but sometimes it has a faint grey tinge to the stem and if you don’t look too closely, you could imagine it was probably a greying reaction happening, and not bother to spend any more time on it. Thus this Russula, not closely related to any known Russula that was ever rumoured to exist in the PNW, went unnoticed for decades. I’m just as guilty as everyone else for not looking closely at it but dismissing it as one of the two other unrelated species. It may be closely related and is similar to R. paludosa from Europe, but nobody ever suspected that mushroom might be here. As soon as I heard of this cool discovery from Anna’s study of Ben’s collection, I advocated for naming it after him, which finally happened in late 2017. It is a large Russula, easily reaching 10cm. This fact alone might help make it possible to identify (as opposed to the similarly chameleon-like but normal-sized R. sierrensis. The cap colours can be quite variable, but are often brownish-tan, perhaps with areas of olive and darker spots of red or purple (or sometimes entirely olive or red or purple). The stem may be purple flushed (like our R. 'xerampelina') perhaps with a faint greyish tint (as if it is greying like R. vinosa), both also large species, which it has traditionally been mistaken for. In one Alaskan collection, the purple flush on the stem was significant. In one collection, it had yellow rhizomorphs! Russula benwooi © Ben Woo R. viridofusca (in the xerampelina group of shrimp Russulas) on the left, with the stem turning green in FeSO4. R. vinosa in the middle, with the stem turning grey where scratched. R. benwooi on the right, with the stem not changing colour when scratched and turning the ordinary rust colour of iron in FeSO4 instead of green. Photos © Danny Miller   Unsolved Russula mysteries If you are thinking that we should pay close attention to other Russula species described by Burlingham in 1936, you’re right. There are others, whose identities are not yet understood, as well as a few Russulas described by others that are not yet understood. R. placita WA - (a hot purple Russula) R. murina OR - (slowly slightly hot olive-grey Russula) R. inconstans FL - (slowly slightly hot Russula with a red margin and yellow-brown center) R. maxima WA - (a large, mild, purple Russula with a rose coloured stem) described by her in 1915. R. persobria OR - (no idea what it is except the cap appears to be green in the dried type collection) described from Oregon in 1939. R. citrina f. separata WA - (a hot white spored yellow capped Russula in clade Russula core). The European Russula citrina is now thought to be a synonym of the European Russula ochroleuca. We likely don't have that species here, so this form described by Singer from Washington is likely a completely different species that what Singer thought it was so it can't keep its name that incorrectly indicates it is a form of Russula citrina. But what is it? No idea. If it doesn't have another name already, it could possibly be called Russula separata to honour his discovery of it. Very little is known about these species except all but the last one are likely somewhere in this “crown clade” and microscopically, do not match any known species enough to suggest what they might be. Absent sequences of the types and further study, they will remain a mystery.