Danny’s DNA Discoveries – gilled boletes of the PNW
Gilled boletes are in the order Boletales, more closely related to boletes than they are to most other gilled mushrooms which are found in the Agaricales. In fact, Phylloporus, the original gilled bolete, was one of the first mushrooms to teach mycologists that their morphologic classification system might not reflect their natural relationships. It was easy to assume that gilled and pored and spiny mushrooms belonged together with others like them. But when they first looked at the "gilled bolete", shown above, with its colour and stature exactly matching a number of boletes with a dark cap and bright yellow pores, and they turned it upside down, they found gills instead of pores. When the microscope was invented, they found it produced spores over twice as long as they are wide, just likes boletes do. Then they knew something was up and that mushrooms must have evolved back and forth between gills and pores more than once. As it turns out, it's not that big a change. Imagine pinching two gills together at regular intervals and you are creating pores. As as the DNA era has shown, convergent evolution (the same look coming about independently) is quite common. But Phylloporus was one the first clues.
Since then the microscope discovered that other mushrooms seemed to be bolete relatives, and DNA has confirmed it. You'll just have to learn how to recognize the various genera, but there is often a clue - gills that (especially near the stem) do something weird (after all, they didn't evolve with other gilled mushrooms) - either fork, wriggle, or become interveined almost to the point that they might start to look like pores where they attach to the stem.
I will also cover false truffles in the gilled bolete families.
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:
Phylloporus - click to expand
Boletaceae - These are very closely related to Xerocomus boletes, in fact it's not certain if they belong in Xerocomus or deserve their own genus. Their ITS sequences can be strange, with multiple copies of some portions, slightly modified, making them hard to align. Therefore, I am not certain as to their relationship to Xerocomus or each other in the ITS only trees I make.
Dark cap and bright yellow... gills, looking like so many true boletes with the same colour and stature.
Species mentioned: Phylloporus rhodoxantha, arenicola
Phylloporus rhodoxanthus EU - brown cap sometimes with red tones, decurrent gills that might stain blue. This species is reported from here, but even though I don't have EU DNA of this species to prove it one way or another, it is strongly suspected that it does not occur here but that reports represent one or more of the undescribed species below, for instance #1. We need more collections both here and from the EU to prove it, especially local collection with red in the cap to see what they really are.
Phylloporus arenicola OR - This locally described fall species is definitely here, described as having notched gills and no blue staining. We don't have a type sequence, but based on the description (but not the time of year), the best match so far might be #4. I'm not sure, because it apparently has lookalikes we didn't know about.
Phylloporus PNW01 - fall, brown cap with fairly dark areas, decurrent gills or notched gills with a decurrent tooth, slow, slight blue staining. Perhaps this unnamed species has been mistaken for P. rhodoxanthus, although no red was noted in the cap. It has been found in BC and WA so far.
Phylloporus PNW02 - fall, olive-brown cap, notched gills, no bluing noted even after cutting it in half. Stem may have red fibrils. Only known so far from CA and AZ.
Phylloporus PNW03 - spring and fall, known from a sequence from OR (spring) and BC (fall), notched gills, bluing distinctly.
Phylloporus PNW04 - spring, known from a single sequence from OR, notched gills and no bluing noted. Perhaps the best match to Phylloporus arenicola so far, except that is supposed to be a fall species.
Phylloporus PNW01 © Yi-Min Wang
Phylloporus PNW02 from AZ © Terri Clements and Donna Fulton
Phylloporus PNW03 and PNW04 © Jonathan Frank
Hygrophoropsis - click to expand
Hygrophoropsidaceae - Commonly mistaken for a chanterelle by beginners, so therefore one of our "false chanterelles" with an orange cap and strongly decurrent bright orange forked gills. The caps on some species may be dark brown or even pale (almost white).
Species mentioned: Hygrophoropsis aurantiaca, rufa, pallida
Orange capped species
Hygrophoropsis aurantiaca EU - all sorts of shades of orange seem possible in this species. Our local sequences are 2 bp and 2 indels and 1 ambiguous location different than EU sequences in ITS1, and ITS2 is identical. For now, I'm going to consider that our collections are the real thing.
Hygrophoropsis PNW05 - another orange species, found once near Seattle, but I don't know how to tell it apart from H. aurantiaca.
Very pale capped local specimens should be collected to see if they are a separate species, as there is a pale capped east coast species, Hygrophoropsis pallida MA, but we don't have DNA of that to know what it is, nor if our own pale capped collections have unique DNA. We need collections.
Hygrophorus aurantiaca © Stephen Russell and Crysta Lehmann, H. sp. #5 © Danny Miller
Dark brown capped species
Hygrophoropsis PNW02 - no photo, found once in BC, a "robust dark form", resembling Hygrophoropsis rufa EU, but we have DNA of that EU species and none of our species match it.
Hygrophoropsis PNW03 - our most common dark brown capped and stemmed species, found in BC, WA and OR, perhaps not as robust as #2, although some collections are robust.
Hygrophoropsis PNW04 - another dark brown capped and stemmed species found in WA and OR, perhaps not as robust as #2, but no idea how to differentiate it yet from #3 or #2.
Hygrophoropsis PNW03 © NAMA and the Field Museum of Natural History (2 images), H. PNW04 © Stephen Russell (2 images)
Gomphidius and Chroogomphus - click to expand
Gomphidiaceae - Easily recognized fairly large, fleshy mushrooms with strongly decurrent gills and grey to black spores. Gomphidius are very slimy with white flesh that blackens. Chroogomphus are not slimy and have orange coloured flesh with water soluble pigments that can turn your urine red like beets.
Chroogomphus albipes CA and Chroogomphus loculatus OR are secotioids. Gomphogaster leucosarx is a trufflized mushroom in this family.
Species mentioned: Gomphidius glutinosus, largus, oregonensis, maculatus, pseudomaculatus, smithii, subroseus. Chroogomphus tomentosus, vinicolor subsp. californicus, ochraceus, pseudovinicolor, leptocystis, albipes, loculatus. Gomphogaster leucosarx.
Mushrooms in this family are oddly specific about which Suillus they parasitize. Since the EU generally has different Suillus species than we do, you'll notice that out of a dozen and a half or so species in the Gomphidiaceae found in the PNW, not a single species that wasn't described from here has been proven to be here (yet). I've never seen that in any other mushroom group.
Gomphidius oregonensis OR -drab brown cap, bright yellow stem base, often clustered growth. Supposedly parasitic on Suillus caerulescens under Doug fir.
Gomphidius pseudomaculatus ID is a single collection from Idaho said lack the glutinous veil of G. oregonensis, but Idaho is dry and I'm not sure it's really a unique species. We need collections to find out.
Gomphidius cf glutinosus EU - perhaps a purplish cap and solitary growth. Bright yellow stem base. There are many reports of this in the PNW, but no genetic proof. We already suspect that eastern North America has an undescribed species and not the real thing, so we need local collections to find out what is really here. Whatever it is supposedly parasitizes Suillus ponderosus under Doug fir and perhaps a second species grows with coastal spruce.
Gomphidius largus ID may be a synonym of a very large G. glutinosus, so we need the type sequence or new very large collections.
Gomphidius oregonensis © Buck McAdoo, G. cf glutinosus (what is it really?) © Michael Beug
Gomphidius subroseus OR - red cap fading to pinkish tan, bright yellow stem base, smaller size than G. oregonensis and usually a bit more brightly coloured even when faded. Parasitic on Suillus lakei under Doug fir. One sequence varies by 4 bp in ITS1 only, but I think there's only one species here.
Gomphidius smithii OR - pinkish tan cap, a little stem base yellowing. Difficult to differentiate from faded G. subroseus, its closest genetic relative, when the stem base is yellow, as shown in the two very similar pictures. Even microscopically they are similar. Parasitic on Suillus lakei under Doug fir as well. A BC sequence differs by 2-4 bp from several AZ sequences. Are there two species on the west coast? If so, which is the type from OR? I suspect there's only one species here, but we need OR collections.
G. subroseus © NAMA and the Field Museum of Natural History, G. smithii © Tajinder Sidhu
Gomphidius cf maculatus EU - drab cap, little stem yellowing. It's the only species said to lack any kind of fibrillose veil underneath the slime veil. With larch? We don't have an EU nor local sequences, but we have China and Quebec sequences that differ from each other by 2 bp and 3 ambiguities. Perhaps that's close enough to mean that they and we will have the real species, but we need local and EU sequences to find out.
Some of my ITS only trees show it is likely more closely related to Chroogomphus than to Gomphidius. At least a couple of papers show it inside Gomphidius with weak support. I am assuming it belongs in Gomphidius, but I thought I would point out that it would be nice to have a multi-gene study confirm that it isn't outside of both genera.
Gomphidius cf maculatus © Kit Scates Barnhart
Gomphogaster leucosarx ID - a blackening secotioid (almost a false truffle) probably more related to Gomphidius than Chroogomphus. No photo nor DNA yet to see if it deserves its own genus. It may not.
Chroogomphus tomentosus WA - an orange woolly dry capped species, often mistaken for a chanterelle by beginners. Parasitic on 'Boletus' mirabilis under hemlock.
Chroogomphus loculatus OR - this misshapen secotioid's type sequence is identical to all of our C. tomentosus ITS sequences, so it may only be a secotioid variety.
Chroogomphus leptocystis BC - a very similar, sister species perhaps with some grey in the cap, perhaps best differentiated microscopically. We only have 1 purported sequence of this from BC, but it sequences sister to C. tomentosus like it should, so it's probably identified correctly. We need more collections and a photograph.
Chroogomphus tomentosus © Michael Beug, probable C. loculatus © Thom O'Dell
Chroogomphus pseudovinicolor ID - a reddish-orange to burgundy-orange very fat-stemmed species with a smooth, dry cap that is never umbonate. The confusing group of species below are not as stocky, often have a viscid cap and may be pointy. The only sequence we have is an unusually purple collection. I'd like sequences of more "normal" orange collections like the second photo below, or a type collection, to see if there's more than one species here. Parasitic on Suillus pseudobrevipes under pine.
Chroogomphus pseudovinicolor #1 (sequenced) © Andrew Parker, C. cf pseudovinicolor #2 (unsequenced) © Danny Miller
Chroogomphus aff. vinicolor - may be viscid, but more pointy capped and darker red-brown than the following confusing species. So far the sequence we have from OR and the one from boreal BC do not match east coast type area sequences of C. vinicolor, nor do they match a CA sequence that might be subsp. californicus, so we may have our own variety in need of its own name. This theory is born out by ecology, as subsp. californicus is suspected to be parasitic on Suillus pungens, which is not proven north of CA, so ours is suspected to have a different host. We should get more collections to see if more varieties are present here as well.
Chroogomphus ochraceus group OR - said to be viscid as well but less pointy and less dark than C. aff. vinicolor, but in practice very similar, except for microscopically having thinner walled cystidia, but even that may not be reliable. Two sister species are found in OR, one of them probably the real thing and one of them a lookalike sister species. We don't know how to tell them apart yet, nor even reliably from C. aff. vinicolor. #1 is found in OR and BC so far, #2 just in OR. We need more collections of all of them. Perhaps one of them may be parasitic on Suillus tomentosus under pine and perhaps there's a 3rd California species that may be parasitic on Suillus fuscotomentosus under pine.
Chroogomphus aff. vinicolor, ochraceus #1 and ochraceus #2 © NAMA and the Field Museum of Natural History
Chroogomphus 'ochraceus PNW03' - it may resemble C. ochraceus, but it sequences outside the group of the first two, closer to the secotioid C. albipes than it does C. vinicolor or C. ochraceus. This one does seem to have brighter orange colours and is not umbonate.
Chroogomphus 'ochraceus PNW03' © Buck McAdoo
Chroogomphus albipes group CA - even more secotioid than C. loculatus. we appear to have two genetic species here, not one, probably one of them the real thing and one of them in need of a new name. #1 is found in ID and OR. #2 is found in OR. No photos of sequenced collections yet.
possible Chroogomphus albipes group member © Ben Woo
Paxillus, Alpova and Melanogaster - click to expand
Paxillaceae - Large but thin-fleshed mushrooms with indented caps, a strongly inrolled margin, strongly decurrent gills and brown spores. The caps are large, yellow- to olive-brown discs and the mushrooms strongly stain brown.
Alpova and Melanogaster are trufflized mushrooms in the same family. Some ITS only trees show that Alpova is indeed a separate genus from Melanogaster, but it doesn't always show up that way. However, a 3-gene study was able to confirm with high confidence that Alpova is monophyletic and distinct.
Species mentioned: Paxillus involutus, cuprinus, ammoniavirescens, obscurisporus. Alpova diplophloeus, concolor, trappei. Melanogaster tuberiformis, macrocarpus, euryspermus, natsii, ambiguus, intermedius, broomianus, vittadinii.
While guidebooks have long said Paxillus involutus is the species here, under urban birch, it has recently thought that Paxillus cuprinus might be our species instead, but these DNA studies show that it is not an either/or, they are both here and so are two more. Telling them apart and figuring out which trees they each go with is the next step. Do not eat any of them, they contain a deadly toxin that can slowly kill you. The only bona fide report of a mycologist dying from mushroom poisoning was from this genus, before we realized the cumulative effects of the toxin.
Paxillus involutus EU - slender species said to be with urban birch, verified from Vancouver BC a number of times but trees not noted. Supposedly the DNA was found on a root tip of a Douglas fir.
Paxillus cuprinus EU -also slender, verified both with and without birch from BC, WA and OR. One collection was under native hazelnut, Corylus cornuta.
Paxillus obscurisporus EU - perhaps stockier and probably with hardwoods, found once in WA and once in boreal BC.
Paxillus aff. ammoniavirescens - a similar stocky hardwood species with a more reddish-brown spore print (the others may be more yellow-olive-brown). Our collections sequence 3-6 bp different than EU sequences and may represent an undescribed species. Found in BC, WA and OR, trees not noted (possibly cottonwood).
Paxillus involutus © Hiba Mezerreg, P. cuprinus © Daniel Winkler, P. obscurisporus © Danny Miller, P. aff. ammoniavirescens © Danny Miller
Alpova and Melanogaster - false truffles with a uniform dark, jelly-like oily marbled centre. Alpova are found with alder and sometimes not as pure black as Melanogaster.
Alpova diplophloeus WA - under alder, fruity/garlicky odor, yellow-brown exterior bruising red, brown interior.
Alpova concolor OR - best differentiated microscopically.
Alpova concolor © Yi-Min Wang
'Alpova' trappei OR - brighter yellow exterior when young, covered with some yellow rhizomorphs and not staining red. It belongs in Melanogaster with the darker species, which should have been predictable as it is with conifers not alder and ecology is always more important than colour for determining relationships. A southern species known only from OR and CA for now. It is not the same as Melanogaster luteus, which is a European false truffle with a very different sequence.
Alpova trappei © Matt Trappe
Melanogaster macrocarpus OR - a dark orange-brown false truffle perhaps with some dark rhizomorphs, with a black jelly marbled interior. Smells of onions and motor oil. It (and probably all local Melanogasters) have long incorrectly gone by the name Melanogaster tuberiformis EU, a closely related European false truffle not actually found here. M. macrocarpus is a >1% ITS variation species complex, but it's all the same species.
Melanogaster euryspermus OR - our second most common Melanogaster, with an odor of citrus and sewer gas. It is a 3% ITS variation complex (definitely on the higher end of ITS variation seen within one species), but all of them represent the same species.
Melanogaster natsii OR - we have the type sequence and one other, but it appears to be rare. I don't have a description yet.
Melanogaster PNW06 OR - unique DNA from any known species, supposedly found with Garry oak but the same DNA was supposedly also found on madrone and Doug fir roots. Found twice in southern OR. It's possible this represents misidentified reports of one of the below species, but either way, it probably needs its own name.
Melanogaster ambiguus EU - we have EU DNA thanks to Alvarado 2021, and it has been reported recently from all over the PNW, but we don't have any local sequences to see if they really represent this species or not. M. euryspermus used to be considered a variety of this species, so it's possible reports of this actually represent that species. We need collections.
Melanogaster intermedius UK - we have likely UK DNA, and it has been reported from southern OR, CA and BC, but we don't have any local sequences to see if they really represent this species or not. This used to be considered another variety of M. ambiguus, so it's possible reports of this actually represent M. euryspermus, another former variety of M. ambiguus known to be common here. We need collections.
Melanogaster broomianus EU - we have EU DNA thanks to Alvarado 2021 to compare to, and it has been reported from OR almost a hundred years ago, but as it has not been found again in the extensive truffle research done by the Trappes, I highly doubt it's actually here. We could use some collections that key out to this to see what they actually are.
Melanogaster vittadinii EU - was reported once in OR, but nobody really knows what this species is supposed to be as it isn't a legally published name, so I highly doubt it was identified correctly. We do have one reported EU sequence to compare to, but that's probably not correctly identified either.
probable Melanogaster macrocarpus © Matt Trappe, M. euryspermus © Jim and Matt Trappe
Tapinella - click to expand
Tapinella atrotomentosa EU - very large dark brown cap with strongly decurrent yellowish gills and a large, black-velvet eccentric stem on conifer stumps. Brown spores.
Tapinella panuoides EU - lacks a stem, on conifer stumps with brown spores. Yellowish brown cap and gills. Hard to differentiate from Crepidotus except for the odd forked, crimped (wrinkled) or interveined gills near the point of attachment).
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