Danny’s DNA Discoveries – boletes of the PNW
Boletes look like regular gilled mushrooms in that they have a cap and a stem, but they have a soft fleshy pore surface under their cap instead of gills. The pore layer is easily separated from the flesh of the cap, and the whole mushroom will feel soft enough to easily chew. Unrelated polypore mushrooms (and others) may have a pore surface, but they are usually tougher - at least as tough as leather if not as hard as a chunk of wood, and the pores cannot usually be easily removed from the rest of the mushroom. Boletes usually grow out of the ground (they are mostly mycorrhizal), while polypores more often grow out of the sides of trees without stems, or are simply a pored surface lying flat without a cap on a piece of wood, but occasionally they will have a stem too. Bolete spores are often long and thin, routinely three times longer than they are wide.
This page covers the Boletales families Boletaceae, Gyroporaceae (which I can't confirm deserves to be a separate family from Boletaceae) and Suillaceae, which include all the boletes plus some gastroid and trufflized boletes.
For the gilled mushrooms (and their gastroids) that are in other families of the Boletales, see my page on Gilled Boletes.
abundant common uncommon rare - colour codes match my Pictorial Key and are my opinions and probably reflect my bias of living in W WA. Rare species may be locally common in certain places at certain times.
Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
Suillus - click to expand
Either viscid or with a partial veil, often both. Stem may have dots that look painted on. (The rare Aureoboletus and Pulveroboletus may be viscid or possess a veil, respectively, but will have extremely vivid yellow pores and never any stem dots).
Family: Suillaceae. Genera mentioned: Suillus, Truncocolumella
We do know a lot about Suillus now, thanks to the 2016 paper "Global Suillus ITS phylogeny" by Nyugen et. al. In general, North America will sometimes have distinct species from Europe, but the same species are usually found on the east and west coast here. I have a few single sequences of potential new species, but they could also be bad sequences, so until I see the sequence a second time, I'm not going to report anything.
'Truncocolumella' citrina OR - this false truffle is recognized by yellow colours inside and out and an interior that is not entirely homogenous (there is a false columella inside). Species in this genus need to be moved to the genus Suillus. Smith described a 'Truncocolumella' citrina var. separabilis ID that is assumed to be the same thing, but we should find and sequence it to make sure.
'Truncocolumella' citrina © Steve Trudell
Suillus suilloides CA - a truffle Suillus from CA that has been reported from OR. I have no OR sequences to compare with my CA sequences, but I'm sure the reports are true.
Two sequences from Oregon don't match anything yet. KT800307 seems to be related to Suillus tomentosus and Suillus suilloides. KX260627 may be related to Suillus quiescens. We don't know if there's something wrong with the sequences, or if they are regular species or trufflized species.
First, the species that lack a partial veil. They do not blue, except for S. tomentosus and S. discolor. Check young specimens, as some veiled species don't leave much of a trace except for along the cap margin when young.
Suillus brevipes ENA - dark cap (rarely pale), no dots on the stem (thus a clean look), often a short stem, under 2-3 needle pine. This east coast species is found all over the PNW. Suillus pallidiceps from ID is a pale capped version that may or may not be a separate species. We need collections to find out.
Suillus quiescens CA - very similar 2-3 needle pine species to S. brevipes, perhaps with a paler cap, a hint of yellow at the stem apex and a few obscure glandular dots, found genetically in the soil in OR but never proven to have fruited up here until finally found in WA recently. It is so similar to S. brevipes that it may be more common than we think and just overlooked.
Suillus punctatipes WA - larger pores, many dots. Douglas fir. Suillus imbellus OR from Oregon is a gastroid, misshapen mushroom whose relation to S. punctatipes should be investigated. It may be the same thing. I don't have any DNA yet.
Suillus subalpinus WY ('granulatus'/flavogranulatus?) - smaller pores, many dots, high elevations with 5-needle pine. Described from WY and also sequenced from WA, OR, MT and CA. Suillus granulatus (EU) and/or Suillus weaverae (ENA) have been reported from here but I think they represent findings of S. subalpinus. It is already accepted that S. granulatus, a 2-3 needle pine species, is not in North America. Suillus flavogranulatus, described from Idaho, had its type sequenced and it turned out to be a Suillus lakei, a very different species, so perhaps there was some error. One theory is that it is an older synonym for S. subalpinus. If you think you find S. weaverae or S. flavogranulatus in the PNW, save it.
Suillus brevipes © Steve Trudell, S. quiescens © Noah Siegel, S. punctatipes © Michael Beug, S. subalpinus © Jonathan Frank
Suillus placidus EU - whitish with many stem dots. 5-needle pine. EU species. ENA sequences match pretty well, but the one WNA sequence I have from AZ is 3 bp and 4 indels different in ITS. The 2016 Nyugen paper does call S. placidus a member of a species complex, so it can't be ruled out yet that our local species may be different. We need local collections.
Suillus tomentosus ENA? - rough, almost dry cap and no veil, orange colours, including in the pores. Slowly blues. 2-3 needle pine. Described from North America, probably from back east, and confirmed from here too.
Suillus discolor ID (Suillus tomentosus var. discolor) - perhaps a less yellow cap than S. tomentosus, otherwise similar. It is found under 5-needle pine, which is perhaps the best way to distinguish it. Described from ID as a variety of S. tomentosus, which it is a sister species of, so it could be thought of as a variety.
Suillus fuscotomentosus CA - described from CA under 2-3 needle pine, it has a dark brown cap and does not blue at all. It is rumoured from OR but we need collections to match to my CA sequences to prove it.
Suillus placidus © Noah Siegel, S. tomentosus © Noah Siegel, S. discolor © Jonathan Frank, S. fusctotomentosus © Noah Siegel
Rough-capped Suillus: These do not appear very Suillus-like because the caps are not very sticky, in fact they are pretty rough and scaly, but they do have an obvious veil. Some exhibit a slight bluing: S. lakei in the base of the stem and S. ochraceoroseus throughout.
S. lakei OR (amabilis?/flavogranulatus?) - a pinkish brown distinctively scaly cap with large pores that stain brown. Described from OR. Douglas fir. It is not currently thought that Suillus lakei var. pseudopictus (especially pink and scaly) is a different organism, but we don't have a type sequence. Suillus amabilis is a much older CO spruce species that seems very similar so we should get a type sequence or type area sequences to see if we should be using that older name instead. Some authors are already saying we should (Læssøe and Petersen of Fungi of Temperate Europe). The type collection of Suillus flavogranulatus from Idaho turned out to be S. lakei, but they are quite distinct species, so there must have been some mix-up. We need more modern collections of S. flavogranulatus to see what it is. So far, one collection turned out to be Suillus subalpinus, above.
Suillus ampliporus NY ('cavipes') - dark brown with large pores. Easily recognized by its hollow stem. Common with larch, although larch is uncommon here. It had been going by the European name S. cavipes until it was discovered that all of North America has a sister species, S. ampliporus, described from NY. It might even be considered a variety. We don't have any PNW DNA to confirm that ours is the same species, but we have CA DNA, and all over North America where it has been sequenced it's the same species, so I'm sure we have it too.
Suillus ochraceoroseus ID - large, very rosy (more than S. lakei) with even larger pores. Larch. Described from Idaho.
Suillus lakei © Steve Trudell, S. ampliporus © Ben Woo, S. ochraceoroseus © Ben Woo
Next, species with glandular dots on the stem (they do not blue). First, with a weak partial veil, that usually does not stay on the stem to form a ring, you may only see some remnants along the edge of the cap.
Suillus brunnescens OR (='borealis' ID) - young white cap and stem staining chocolate brown, weak purple- to red-brown veil, with stem dots. 5-needle pine. Closely related to S. luteus and quite similar (but that is consistently brown with a strong ring and under 2-3 needle pine). Described by Smith from Oregon in 1964. Suillus borealis, described by him from Idaho a year later, turned out to be the same mushroom after sequencing the types, which is probably a good thing as the name S. borealis had already been taken by Peck for a Labrador mushroom in 1895. Now there is no confusion.
Suillus americanus NY (=sibiricus) - a very yellow Suillus with reddish-brown patches near the cap margin. Sometimes with large pores. Glandular dots. 5-needle pine. Suillus sibiricus was described from Siberia thinking that it was different than the east coast Suillus americanus, but it really isn't (it's the same species all around the world), so we don't need that name. We have a sequence from CA, but we should get a local sequence to verify for sure that ours is the same species as the rest of the world.
Suillus glandulosipes CA (=neoalbidipes) - duller yellow with smaller pores and many stem dots. 2-needle pine. The east coast S. neoalbidipes turned out to be the same as the older California mushroom S. glandulosipes. As explained in the introduction, for some reason we seem to share species between the 2 coasts, which doesn't always happen.
possible Suillus brunnescens © Michael Beug, S. americanus © Andrew Parker, S. glandulosipes © Kit Scates Barnhart
These species with glandular dots usually do have a conspicuous ring on the stem (they also don't blue).
Suillus luteus EU - the Slippery Jack itself. Dark cap colour and purple sheathing veil (splays up). Lots of glandular dots. The closely related S. brunnescens, above, can look similar. 2-needle pine. This EU species is found worldwide.
Suillus acidus NY (=subolivaceus WA) - the Slippery Jill. Olive-brown with a wide band around the stem for a veil. Not bluing. Lots of glandular dots. 5-needle pine. S. subolivaceus was described from WA thinking it was different than the east coast version, S. acidus, but once again, it is not.
Suillus luteus © Michael Beug, S. acidus © Andrew Parker
Finally, the sometimes difficult to recognize species without conspicuous glandular dots on the stem (there may be a few). The ring may or may not be conspicuous. Usually not bluing.
Suillus caerulescens WA (=imitatus var. imitatus) - streaky yellow-brown cap, sometimes fibrillose and possibly even scaly like its sister species S. lakei. Pores bruise brown and only the bottom of the stem bruises blue. Douglas fir. Described from WA.
Suillus ponderosus OR (=imitatus var. viridescens) - a little darker brown than S. caerulescens, occasional green patches, often bigger and stockier, with yellow-orange slime on the veil when young. Douglas fir. Described from OR.
Suillus clintonianus NY (=S. grevillei var. clintonianus) - usually chestnut coloured with a yellow rim and a yellow to chestnut veil, but harder to recognize when the colours fade. Also staining brown, but sometimes blue in the stem base. There is often a reticulated pattern of brown spots at the top of the stem from decurrent pores that looks like glandular dots. Larch. This NY species, also found throughout the PNW, is sometimes considered a variety of the EU S. grevillei, and sometimes distinct. We now know that the EU variety is not found here.
Suillus caerulescens © Steve Trudell, S. ponderosus (normal and green) © Noah Siegel and Michael Beug, S. clintonianus © Steve Trudell
Suillus elbensis NY (='viscidus'/'laricinus'/'aeruginascens') - the grey Suillus, large pores, Slowly blues. Stem may be reticulated like Boletus. Larch. This NY species has been called a few incorrect European names in the past, but they are not our species. One ID sequence is 3 bp different than some east coast sequences, but even type area east coast sequences vary by as much as 5 bp or more, so we very likely have the same species here.
Suillus flavidus EU (=umbonatus UT/helenae OR) - yellowish with an umbo (pointed cap). A brownish stainer. Not bluing. Large pores, sticky brown sometimes fleeting veil. 2-needle pine. This EU species has some newer synonyms from the west coast, but it appears that they are not distinct genetically in ITS and that the EU species may be found worldwide. Coastal S. helenae may be more slender than montane S. flavidus/umbonatus and may represent a more slender form. S. helenae was described without a veil, but the black and white photo of the type collection appears to show that at least one specimen had ample veil material hanging from the cap margin.
Suillus pseudobrevipes ID/albivelatus ID - fairly nondescript 2-3 needle pine species with a white veil (like many others), not bluing (like many others). S. albivelatus is said to have flesh that can stain pale vinaceous and less veil material (S. pseudobrevipes' more ample veil may sheath). S. albivelatus (from Idaho) seems to have identical ITS DNA as the older (by one year, also from Idaho) S. pseudobrevipes except for a few ambiguous locations. The 2016 Nyugen paper shows it as a slightly distinct genetic species, but I can't reproduce that. They are so similar and difficult to differentiate, we should consider that there really might be only one species, S. pseudobrevipes.
Suillus elbensis © Steve Trudell, S. flavidus © Kit Scates Barnhart, S. pseudobrevipes © Noah Siegel
Leccinum - click to expand
Leccinum are the scaber-stalked boletes, not to be confused with Suillus which has glandular dots that look painted on compared to scabers that can be removed. Most Leccinum can be recognized by whiter pores than many Boletus (not as yellow). Very young specimens might not have developed the scabers yet and are easily confused with the Boletus edulis group (unless you note blue/grey/red staining).
Family: Boletaceae. Genera mentioned: Leccinum
Although they were well studied long ago and we have many species named locally, how to differentiate them is a lost art and we will have to await a more modern study using type sequences and colour photography before I will be able to give you a definitive ID on your Leccinum. One character to note is if it stains red before staining bluish-grey, or if it stains directly to bluish-grey (although it has not been proven that can differentiate species). Another thing to note is the host tree, which is thought to be very indicative of what species you might have. Unfortunately, the ITS region of Leccinum is a little weird, with copies of the same pattern over and over again with slight differences, which makes them hard to align. Any tree you make is not going to reflect reality, you pretty much have to look at every sequence by hand to figure out what it is. Therefore I do not include a tree for Leccinum in my files.
Leccinum scabrum EU - brown cap, under urban birch, said to have flesh that will not turn blue. This EU species is verified from Monmouth, OR but we should get more collections to see what lookalikes are also here. ITS DNA can vary by 2% in this EU species. Many sequences of L. rotundifoliae, which varies even more in ITS, can have the same ITS DNA as L. scabrum, but that is an alpine species.
Leccinum rotundifoliae Siberia - very similar brown birch species (perhaps paler when young), but in the wild interior alpine instead of the urban lowlands. This species was reported from Idaho but we need modern collections to confirm its presence here. A recent soil sample from interior BC was this species, so it probably is here. Its ITS DNA is very much like that of L. scabrum, so you may not be able to tell these two definitely distinct species apart by only sequencing ITS, except it varies even more than L. scabrum.
Leccinum schistophilum EU - one interesting surprise was finding out that a sequence of a brown urban birch collection from Vancouver turned out to be this lookalike instead of L. scabrum. It is said to have flesh that turns blue. We don't have enough local urban birch collections to know what all species we have nor how common each one is, or how the blue staining actually works.
Leccinum cf idahoense ID - brown cap, in the wild. Found in Bellingham with alder (and some conifers) and in the southern interior of BC on birch root tips. L. idahoense is said to be a conifer species from the interior that slowly greys, so this may not be that. If so, we don't know what L. idahoense really is, nor do we know what this species that we've been finding is.
Leccinum 'cf idahoensis' © Sharon Squazzo
Leccinum cf mazanitae CA - This must be our most common reddish capped species, as every sequence from the PNW of a red capped Leccinum (almost a dozen so far) seems to be the same genetic species in ITS. Alternatively, there is more than one species that can't be distinguished by ITS, more study is needed. Mostly, the collections have been called L. manzanitae, said to have a reddish-brown cap, be found under Madrone and Manzanita and stain directly blue-grey, but the names on the collections don't mean much and some of the collections were found under conifers (although at least one was near Madrone). We need more collections with good data about the nearby trees and colour staining, and non-ITS sequences. Perhaps this is the really the supposedly common conifer species described below, Leccinum ponderosum.
Leccinum versipelle EU (atrostipitatum?)- many black scabers even when young, hardwoods like birch. Stains red then blue-grey. This EU species is verified in AK but we need local collections to see if the synonymy is correct.
Leccinum insigne MI - orange to reddish-brown cap with flesh staining directly to bluish-grey. We don't have a type sequence, but we have possible sequences from the east coast, and a find in boreal BC matches it. We need more collections from the type area and from here.
Leccinum 'cf manzanitae' © Jonathan Frank
Species described from the PNW without DNA or colour photographs. This means these species are here by definition, although there may be more than one name for the same species, or some may be the same as other species not listed here.
Leccinum brunneum CA - described from CA by Thiers who said it's also found in Idaho, this is a brown capped aspen species with flesh mostly turning "fuscous". We don't know anything about it (no DNA).
Leccinum clavatum ID - a whitish to pale tan capped conifer species described from Idaho staining directly to blue-grey.
Leccinum fibrillosum ID - a liver brown spruce and pine species staining red then blue-grey.
Leccinum discolor ID - described from Idaho under aspen and pine. Stains red then blue-grey.
Leccinum fallax ID - a dark red spruce species that stains red then blue-grey.
Leccinum ponderosum OR - orange to reddish brown capped species staining blue only in the stem base, found under pine. Described from OR. Supposedly very common so lets start collecting it. I wonder if this is the species already sequenced but usually labeled L. manzanitae?
Other species described from the EU and reported from the PNW with no genetic proof yet.
Leccinum holopus EU - a European birch bolete with a very pale tan cap, said to stain red. Reported from here but no DNA yet to prove it. We do have reliable EU DNA to compare to. The ITS DNA is difficult to differentiate from Leccinum cyaneobasileucum, another L. scabrum look-alike with a brown cap, next.
Leccinum cyaneobasileucum EU - a tan to brown capped birch species that turns blue in the base of the stem (but others may as well), with ITS DNA that can't be distinguished from L. holopus. This EU species is reported from CA and OR, but now that we have found a sequence of the blue staining Leccinum schistophilum in the PNW, a species that is indistinguishable except by spore size, reports of L. cyaneobasileucum may represent L. schistophilum.
Leccinum aurantiacum EU - although every red capped collection is usually called by this EU name, we know what its DNA looks like and it has never been found here, nor is it expected to be.
Leccinum vulpinum EU - although some authors have suggested that perhaps this is an EU species that actually occurs here instead of L. aurantiacum, we don't know what its DNA looks like and there's no convincing reason yet to believe it is here.
Boletus s.l. - click to expand
Everything else that isn't a Suillus or Leccinum. The stem may have a net-like reticulation at the apex, but won't have scabers nor glandular dots. The cap is usually dry and their is usually no veil (rarely, Aureoboletus and Pulveroboletus are found with vivid yellow pores, with either a viscid cap or a partial veil).
Familes: Boletaceae, Gyroporaceae. Genera mentioned: Boletus, Pulchroboletus, Hemileccinum, Chalciporus, Caloboletus, Rubroboletus, Neoboletus, Suillellus, Hortiboletus, Porphyrellus, Butyriboletus, Xerocomus, Xerocomellus, Gyroporus, Aureoboletus, Buchwaldoboletus, Pulveroboletus, Gastroboletus, Chamonixia, Octaviania
Chamonixia and Octaviania are false truffles in the Boletaceae family that will be covered here in more detail later. Chamonixia appears to be a distinct genus somewhat related to Leccinum. I don't know enough about Octaviania to know if it's monophyletic and/or distinct or not.
The porcini and its relatives are now the only species left in Boletus. Everything else is being moved to a new genus. They are large, brownish, have white to pale yellow pores, a reticulated stem top, bulbous stem, mild taste, and do not usually appreciably turn blue (but occasionally they might). The odd one out is Boletus fibrillosus, with a slightly felty cap, streaky brown stem that is not bulbous, and whitish pores, most easily mistaken for 'Boletus' mirabilis, with a much more velvety cap.
Boletus edulis EU - the king bolete or porcini with a baked bread coloured cap and stem is found worldwide.
Boletus edulis var. grandedulis CA - This variety has a darker reddish-brown cap and stem, with brown pores at maturity. It differs in ITS in only a single location. Sequences of Boletus edulis typically vary in several locations from each other, but var. grandedulis seems to have a particular single mutation.
Boletus rex-veris CA - the "spring king" is our spring species with a reddish-brown cap and whitish to yellowish pores in age.
Boletus edulis © Michael Beug, B. edulis var. grandedulis © Daniel Winkler, B. rex-veris © Michael Beug
Boletus barrowsii AZ - this almost white hardwood species has sequences 2 clades of sequences, with almost six differences between them plus another half dozen ambiguous locations. It is being investigated if there are 2 species here. #1 is found in WA, CA and NM. #2 is found in WA and CA.
Boletus regineus CA - our "California queen" bolete has an almost black cap with a white bloom and is most often found with oak. It is squat. While we do have the EU King B. edulis in the west, we do not have the EU Queen Boletus aereus, so our species with a different cap cuticle was given its own name. It has been sequenced from OR. Boletus mottiae CA, with a very retculated cap, may be the same species with very similar sequences, and I can verify that one WA collection had a somewhat reticulated cap and did sequence to be B. regineus. Our Queen is rarely reported all over the PNW.
Boletus fibrillosus CA - a dark brown, almost felty cap and a dark brown streaky stem. KOH turns the cap red. The similar Boletus pinophilus EU, which does not stain red in KOH, has been reported from here, but I believe they are all mistaken reports of B. fibrillosus. If you find a mushroom like this that does not stain red in KOH, let us know! This mushroom could also be partially responsible for the purported Boletus mottiae CA, as very dark capped reticulated specimens have been found of it (see here for one that even turned appreciably blue), although as explained above, the type specimen was probably a B. regineus.
Boletus subalpinus OR - a secotioid sister species to Boletus regineus. It is pale brown with whitish to olive-brown pores and does not blue.
Boletus barrowsii © Steve Trudell, B. regineus © Michael Beug, B. fibrillosus © Steve Trudell, B. subalpinus © J M Trappe
Butyriboletus - the "butter boletes", similar to Boletus (large, mild taste and reticulated stem tops), but they do stain blue and the pores are usually more yellowish. The stems aren't quite as distinctively bulbous, though. Often there is some pink/red in the cap.
Butyriboletus abieticola CA - a conifer species found at high elevations with a pinkish tan scaly or cracking cap, sequenced from OR, matching the CA type sequence very well.
Butyriboletus primiregius CA - a red capped spring conifer mountain species. OR and ID sequences closely match the CA type sequence.
Butyriboletus autumniregius CA - a red capped fall conifer mountain species. OR sequences closely match the CA type sequence.
Butyriboletus querciregius CA - a yellow to pinkish capped oak species not as brightly coloured as the previous two species (perhaps even mostly yellow), that fades to yellow-brown. The stem is often bulbous. We have OR and WA sequences. Unfortunately, the sequence of the chosen CA type collection is dirty, so it's not the best sequence to match against to see if you have this species.
Butyriboletus persolidus CA - a similar oak species with a brownish cap, and a more cylindrical stem has been reported up here but not yet sequenced from north of CA. It is very difficult to tell apart from older B. querciregius after that red/yellow cap has turned dingy brown; we need collections to verify it is here. I suspect it is.
Butyriboletus persolidus var. nov. OR - A variety of B. persolidus from OR with the same ITS sequence but different morphology is being described.
Butyriboletus abieticola © Dimitar Bojantchev, B. primiregius © Richard Morrison, B. autumniregius © Daniel Winkler, B. querciregius © Michael Beug, B. persolidus © Jonathan Frank
'Boletus' mirabilis WA ('Aureoboletus' mirabilis) - the admiral bolete, with a suede-like velvety cap and tree-trunk like stem may get its own genus. It has yellow pores, unlike the similar, not quite as velvety Boletus fibrillosus. It is in a sister relationship to Aureoboletus, so one could either consider it an Aureoboletus or give it a different genus name. Since Aureoboletus are viscid capped boletes, and the suede-like velvety cap of this species could not be any more different, it makes sense to place this mushroom in its own genus. It was described as Cerioporus mirabilis, but that genus was already in use for some polypores, so there is no good synonym to use for a genus name for it, and one would likely have to be created.
'Boletus' mirabilis © Fred Rhoades
Caloboletus - known for their very bitter taste and large size. They usually strongly turn blue.
Caloboletus frustosus CA (= Caloboletus conifericola AK) - rough brown cap, reticulated stem top, no red in the stem. The Alaskan species C. conifericola is probably the same thing, as the older C. frustosus has been sequenced from CA all the way up through BC, so they are being synonymized.
Caloboletus rubripes CA - red stem bottom, stem not reticulated, paler young cap, not quite as bitter. The EU Caloboletus calopus with red on the stem but a reticulated stem top has been reported from the PNW, but so far, reports are turning out to be the non-reticulated species C. rubripes. We need collections with a reticulated stem top to see if that species is here.
Caloboletus marshii CA - this CA oak species has a paler brown cap and non-reticulated bulbous stem. We have the type sequence and it is also sequenced in southern OR.
Caloboletus frustosus © Michael Beug, C. rubripes © Steve Trudell, C. marshii © Christian Schwarz
Chalciporus - known for their hot and/or bitter taste, small size, cinnamon brown pores, and yellow stem base and mycelium.
Chalciporus piperatus #1 - This species does not turn blue. Europe has 3 genetic species, one 2% different and one 5% different from the first one. Our most common species matches #1 (our first 3 sequences matched #1), but it's not necessarily the real one, as we don't have a type sequence.
It should be noted that one collection of this was somewhat gastroid, so there may be a gastroid species in this genus with ITS DNA matching a non-gastroid species as well.
Chalciporus piperatus #2 - Our fourth sequence of Chalciporus piperatus, from WA, turned out to be EU species #2, so we do have more than one species in this complex here. From our only collection so far, it also does not turn blue, and it appears it has a stockier stem and is more brightly coloured.
Chalciporus cf piperatoides MI - something close to this rare east coast species is found here. It does turn blue and is also somewhat brightly coloured, with pores that are almost yellow when young. Sequences from OR and CA differ by 5 bp in ITS, so perhaps this is a species complex as well, like the more common C. piperatus. I don't have any east coast sequences to compare to to verify that ours is a match. More local and east coast collections would be helpful.
Chalciporus piperatus #1 © NAMA and the Field Museum of Natural History, C. piperatus #2 © Yi-Min Wang, C. cf piperatoides © Jonathan Frank
Porphyrellus aff. porphyrosporus EU - a dark grey bolete with dark grey pores, bluing. Possibly bitter. A half dozen EU sequences are almost identical with only one outlier, and ours differ from those by 4-5 bp, so we may have an unnamed sister species. AZ may have a 3rd species.
Porphyrellus aff. porphyrosporus © Michael Beug
Rubroboletus - known for red pores and reticulation at the top of the stem. They turn blue. Very poisonous, probably the most dangerous genus of boletes.
Rubroboletus eastwoodiae CA - a very bulbous stem. We have CA sequences but no sequences from the PNW, but the reports from the PNW (WA and OR) of this very distinct species are undoubtedly true.
Rubroboletus pulcherrimus CA - a clavate stem. Red-brown cap and stem. See below.
Rubroboletus haematinus CA - also a clavate stem, but with a paler brown cap and a yellow stem. The pores start out yellow, and only turn red in age, except for the margin where they may stay yellow, so be careful not to misidentify this very poisonous species. Keep reading.
For the last two species, we have two purported morphological species and two genetic species, sequenced from CA and OR, but without type sequences, it's not proven which is which or that these are the only species, but I am pretty sure. One genetic species has always looked like R. pulcherrimus, and could very well be. The second genetic species was labeled R. haematinus nine times and R. pulcherrimus three times. Given that old R. haematinus can look like R. pulcherrimus, I am assuming these are the sequences of R. haematinus. They differ a bit amongst each other by up to 1%, but most of the differences are accounted for by 4 ambiguous locations in one sequence, with only 1-2 differences usually remaining, so it is all plausibly one species. Therefore, absent type sequences of the two, I believe it's fairly clear what the sequences of these species look like and that we have both of them in the PNW.
'Rubroboletus eastwoodiae © Daniel Winkler, R. pulcherrimus © Noah Siegel, R. haematinus © Wendy Boes USDA Forest Service
Neoboletus - known for red pores, no stem reticulation and very strong bluing. Most easily confused with Suillellus, but separated by dark colours and a cynlindrical stem with yellow flesh in the stem base. There is a possibility that Sutorius may be an older name for this genus.
Neoboletus 'luridiformis' group EU - this is a complex of two species in Europe, and we probably don't have either of them, but 2 unique species of our own, soon to get names. I do not yet know how to tell them apart, and I only have DNA of one of them, from OR. We need more local collections.
Gastroboletus turbinatus var. flammeus ID - our most common gastroid bolete blues quickly, has yellow flesh, red-yellow pores and a blackish-brown cap, for quite a colour combination. It is a variety of the east coast Gastroboletus turbinatus. Our G. turbinatus var. flammeus is described from ID. It's not yet clear to me if our variety will also be promoted to species. I don't yet know how much it differs genetically from our local epigeous species. These gastroids are similar to 'Pulchroboletus' vividus and amyloideus.
Gastroboletus ruber WA - our most trufflized bolete, looking more like a false truffle. Reddish exterior that wears away exposing yellow pores and a whitish stem, bluing. We have some OR sequences of this WA mushroom.
Neoboletus 'luridiformis' group © Michael Beug, 'Neoboletus' (Gastroboletus) turbinatus var. flammeus group © Michael Beug, Gastroboletus ruber © Sava Krstic
Suillellus - also known for red pores, no stem reticulation and very strong bluing. Most easily confused with Neoboletus, but separated by paler colours, a spindle-shaped to cylindrical stem and red flesh in the stem base.
Suillellus amygdalinus CA - we have sequenced this CA mushroom under OR oaks as well.
Suillellus aff. amygdalinus - different by >3% in ITS, this species is getting its own name. I believe the sequence I have is from OR.
Suillellus amygdalinus group © Michael Beug
Pulchroboletus - red on the stem, most easily confused with Xerocomellus and Hortiboletus.
'Boletus' smithii AZ - this interesting bolete has a reddish brown cap with a bloom that when rubbed off makes the cap get even more red. Bluing. Only the top of the stem is red, as opposed to other species that have red stem bottoms only. It has not been moved to Pulchroboletus yet, but that is likely where it belongs, based on DNA and its close resemblance to other species in that genus.
'Gastroboletus' vividus OR and 'Gastroboletus' amyloideus CA - secotioid versions, with yellow and red caps, yellow flesh and pores that turn red, and a red-topped stems. G. vividus only varies by 1 to 2 bp in ITS DNA from B. smithii. It has been sequenced in CA but is reported from southern OR too. I have a couple of sequences from OR that might be G. amyloideus and it is a distinct genetic species in ITS. They resemble the secotioid 'Neoboletus' turbinatus var. flammeus.
'Pulchroboletus' smithii © Ben Woo, 'Gastroboletus' vividus © J M Trappe
Hortiboletus - bright red colours on the cap, most easily confused with Pulchroboletus and Xerocomellus. The flesh of the stem base has orange grains in it, supposedly unlike those other two genera.
Hortiboletus coccyginus CA - bright pinkish red cap, but the orange brown stem is not red. Found with cottonwoods and some non-native trees. Sequenced from WA and OR.
Hortiboletus cf 'rubellus' EU - deep red cap and stem base, bright yellow stem top and pores. Similar to Xerocomellus dryophilus. I need EU sequences and local sequences to prove it (I have neither) but studies have shown that our species is different and in need of a new name.
Hortiboletus "willow" - this seems to be a fat stemmed species growing under willow in Seattle. The DNA was also found once in NY and once in Germany.
'Hortiboletus coccyginus © Noah Siegel, H. cf 'rubellus' © Steve Trudell, H. "willow" © Shannon Adams
Xerocomellus - small to medium boletes with red stems, bright yellow pores and brown to black caps that may also have some red. Similar to Pulchroboletus and Hortiboletus. The species can be difficult to tell apart.
Xerocomellus diffractus OR - cracked cap, perhaps with pink in the cracks. Reddish stem. Blues slowly. Formerly misidentified as the EU Xerocomellus chrysenteron.
Xerocomellus amylosporus ID - very similar, darker cap with irregular cracks, staining inky blue, stem stains dingy brown. Also sequenced from WA.
Xerocomellus mendocinensis CA - very similar, very red stem, darker cap, less cracking (may resemble X. zelleri), but bluing quickly. The spores are truncated. Formerly misidentified as the EU Xerocomellus truncatus. Also sequenced from WA and OR.
Xerocomellus diffractus © Michael Beug, X. amylosporus © Yi-Min Wang, X. mendocinensis © Noah Siegel
Xerocomellus zelleri WA - Beautiful black cap contrasted with yellow pores and red stem. Some may blue slightly. Slightly velvety but not very bumpy cap, with a pale rim. Slender and old growth forest only.
Xerocomellus atropurpureus OR - very similar and previously confused with X. zelleri, bumpy cap not velvety, stockier.
Xerocomellus rainisiae WA - paler, velvety cap, red stem base, turns rapidly greenish-blue.
Xerocomellus zelleri © Noah Siegel, X. atropurpureus © Steve Trudell, X. rainisiae © A and O Ceska
Xerocomellus salicicola CA - pink cap when young, red stem apex with some reticulation, cracking. Hardwoods. Also sequenced from OR.
Xerocomellus dryophilus CA - red cap, red stem bottom, with oak. Rumoured from Oregon, but we still need genetic proof.
There are secotioid/trufflized species as well. 'Gastroboletus' xerocomoides CA has the same ITS DNA as X. amylosporus. Xerocomellus macmurhphyii CA and Xerocomellus behrii CA are two distinct oak truffle species also found in OR.
Xerocomellus salicicola © Christian Schwarz, X. dryophilus © Noah Siegel
Aureoboletus citriniporus CA - unusual for having a viscid cap and stem, which will make it key out to Suillus. This CA species has a brownish-black cap, bright yellow pores, and a pale whitish stem that is orange at the apex, and it has been sequenced from WA.
Aureoboletus flaviporus CA - The Aureoboletus species that was said to occur in the PNW was the similar CA species Aureoboletus flaviporus, but that species has a cinnamon-red brown cap, and a red-brown stem that is white on top. At least some PNW reports of it were a case of mistaken identity, so I am awaiting proof that A. flaviporus actually occurs in the PNW. It is rumoured to be at least as far north as Eugune, OR, so we need collections.
If 'Boletus' mirabilis (above) does not get its own genus, it will stay here, as an Aureoboletus that looks unlike all the others.
Aureoboletus citriniporus © Michael Beug
Pulveroboletus aff. ravenelii - this SC mushroom has a bright yellow veil, which will make it key out to Suillus as well. It has been reported once from Oregon (being so distinctive it's hard to image it was a misidentification), and occasionally in California, but our west coast species is getting a new name. I don't have the sequence yet. We need local collections.
Pulveroboletus ravenelii (from the east coast) © Noah Siegel
Buchwaldoboletus - overhanging cap margin, stem sometimes eccentric, sometimes growing on wood. Usually staining blue.
Buchwaldoboletus lignicola EU - brown cap with overhanging margin and brown eccentric stem, large yellow pores, often growing on wood. One Oregon sequence was 3 bp and 1 indel different than reliable EU sequences, so ours is probably the same species.
Buchwaldoboletus cf sphaerocephalus EU - yellow cap, gills and stem, cap with overhanging margin. Rumoured from Idaho and WA, we need local and EU sequences to verify what was found.
Buchwaldoboletus lignicola © Christian Schwarz, B. sphaerocephalus © Noah Siegel
Xerocomus - Phylloporus gilled boletes are related.
Xerocomus oregonensis OR - the cap is slightly felty like Boletus fibrillosus, but with larger yellow (sometimes brightly so) pores. Not usually bluing. It's a little tricky to characterize and identify. Three species have been reported from the PNW, Xerocomus subtomentosus EU, Xerocomus ferrugineus (=spadiceus) EU and our local Xerocomus oregonensis. So far, only one genetic species is found in the PNW, from everywhere from the coast through the Cascades and into Arizona, and it's quite variable looking, but always with similar DNA, so our one variable species probably explains reports of multiple species (X. subtomentosus has not been found here yet, and neither has X. ferrugineus although that also occurs in Alaska). Our sequences are usually within 1 bp of each other plus a couple of ambiguous locations.
Xerocomus oregonensis © Michael Beug
Hemileccinum subglabripes NY - fairly non descript and sometimes hard to identify. Yellow pored, almost like a Leccinum with scabers that do not develop very visibly, but this feature is not very obvious. Does not blue. This east coast species is also found on the west coast. Sequenced from OR and reported from WA and ID too.
Hemileccinum subglabripes © Noah Siegel
Gyroporus - placed in its own family right now, Gyroporaceae. My own ITS trees do not show that it deserves to be a family separate from the Boletaceae, but as ITS is not great for relationships above the species level, I await further multi-gene studies to answer the question for sure.
Gyroporus borealis VT - A small bolete with hollow chambers in the stem, and what look like notched white pores, a concept usually reserved for gilled mushrooms. It does not blue. Found in the PNW under urban landscaped hardwood trees like chestnut (and formerly thought to be the EU Gyroporus castaneus until this NA species was recently described from VT). They did not sequence ITS for the type, nor do a large study of the PNW, but for now it appears that our only species is G. borealis and that is what we are assuming until we get an ITS sequence of the type and more local collections to study.
Gyroporus borealis © Danny Miller
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