Danny’s DNA Discoveries – Hydnellum, Sarcodon
and Phellodon of the PNW
Click here for my Pictorial Key to the Sordid Toothed Fungi.
These medium to large toothed fungi either have a vague cap and stem form or are top shaped like the kids toy (becoming wide on top from a single point of attachment to the ground). The colours are more sordid than the clean orange-white look of the hedgehogs (Hydnum). All are thought to be mycorrhizal. There are also unrelated toothed fungi that look like earpicks, jellies, conks or simply bundles of teeth (Search for "toothed" on the main page).
Sarcodon - large, brown spored, cap and stem - although most species (but not the type) turned out to be inside Hydnellum, and one may need a new genus, for now Neosarcodon. Perhaps a simpler solution would be to move all of these into Hydnellum, the older genus, but a multi-gene study will have to confirm that Neosarcodon can move too, as it is placed on the basis of ITS only.
Hydnellum - large, brown spored, top shaped - but now including many former Sarcodon that are cap and stem shaped. The two forms are mixed in throughout the tree. The forms seem to have evolved back and forth regularly. Rather than recognizing three genera, perhaps we should study if we can recognize one expanded Hydnellum genus, as with Bankera/Phellodon. This genus is poorly studied in the PNW and much work is still to be done to figure out what species are here.
Phellodon - medium sized, white spored, thin top shapes, fragrant maple odor when dried (smaller and thinner than the similar shaped Hyndellums).
Bankera - large, white spored, cap and stem, fragrant maple odor when dried - basically a white spored Sarcodon. While practically all Phellodon/Bankera have a sweet/spicy odor, some Sarcodon/Hynellum do too, so the spore print is the best differentiating factor. Bankera lives paraphyletic inside Phellodon, so most people considered these part of Phellodon now.
Note that the traditional families of the Thelephorales do not hold up, as many genera currently placed in the Theleophoraceae are closer to the Bankeraceae. There seem to be two major clades in a 4 gene study, with Thelephora and Tomentella in the Thelephoraceae, and everything else found in the PNW in an expanded Bankeraceae.
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.
Click here to download the FASTA data of all my DNA sequences
Hydnellum s.l. - click to expand
Large, brown spored mushrooms, either top shaped or vaguely cap and stem shaped.
'Neosarcodon' atroviridis AL - greyish brown like so many others but discoloring purplish and greenish. One single report in the PNW of this neotropical species has never been confirmed, so it is not known if this occurs here or not. It would be far out of its accepted range. My guess is something in the Sarcodon leucopus group was mistaken for this. It can discolour purplish, and sometimes, greenish too. Sarcodon atroviridis either needs a new genus, such as Neosarcodon, or perhaps it could be moved to Hydnellum along with all the rest of the Sarcodon, although we'll need a mutli-gene study to see if that works as Neosarcodon is only placed using ITS.
Sarcodon (vaguely cap and stem shaped) - true Sarcodon usually have spore lengths in the 7.5 - 9u range.
Scaly cap - (note that Hydnellum scabrosum and Hydnellum underwoodii also have a distinctly scaly cap, but also a dark greenish stem base, at least in H. scabrosum. H. underwoodii has much smaller spores).
Sarcodon squamosus EU - a pine species with a dark vinaceous blackish-brown cap that is usually less indented than the cap of S. imbricatus (which also paler and lacks violaceous tones). One south Vancouver Island BC sequence is 3 bp different than many EU sequences, so that probably represents the real species.
Sarcodon 'imbricatus PNW07' - one northern BC sequence is 6 bp different than many EU sequences. No sign of it further south yet.
Sarcodon 'imbricatus PNW03' - one scaly capped collection from WA has quite a unique sequence. It was thought to be S. imbricatus.
Sarcodon 'imbricatus PNW04' - one scaly capped collection from southern OR also has quite a unique sequence.
Sarcodon imbricatus EU is reported as common but we need more collections to see which species are actually here, and how to tell them apart, as the sequence of S. imbricatus itself has not yet been found here.
Sarcodon squamosus © SVIMS, unsequenced Sarcodon scabrosus © Noah Siegel
Sarcodon 'imbricatus PNW03' © NAMA and the Field Museum of Natural History (2 images), S. 'imbricatus PNW04' © Jonathan Frank (2 images)
Smooth cap - (kept in Sarcodon due to genetics, but not easily distinguished from Hydnellum except possibly by spore length, and not yet distinguishable from each other)
Sarcodon 'leucopus PNW01' - Sarcodon leucopus is grayish brown, discolouring purplish. Alaska and northern BC have collections 6 bp different in ITS. We need PNW collections to see how they fit in, as it has been reported from here. The spores of the real S. leucopus are reported small for the genus, at a length of 6.5 - 7.5u.
Sarcodon PNW02 - our single WA collection is greyish tan, with irregular darker orange-brown stains on the cap. It sequences close to the scaly capped species S. imbricatus and S. squamosus, but PNW02 seems to have a smooth cap.
Sarcodon 'scabripes PNW05' - another greyish tan-brown species, with white flesh. Our one OR collection has a sequence 10 bp from one NC sequence of this NY species, so it appears we might have a sister species here The OR sequence has also been found in Mexico. S. scabripes has never been reported from the PNW.
Sarcodon PNW06 - with a smooth, bright purplish cap, less dark and less sordid than purplish Hydnellums, and with brown spores unlike the similar Bankera 'violascens PNW01'. Found and sequenced once from WA.
Sarcodon PNW02 © Daniel Winkler, S. 'scabripes PNW05' © Jonathan Frank (2 images)
Sarcodon PNW06 © Steve Ness (2 images)
Hydnellum - vaguely cap and stem shaped - Hydnellum usually have spore lengths in the 4.5 - 7u range. Note that both this form and the top shaped form are mixed together throughout the tree. The two forms seem to have evolved back and forth regularly.
Dark vinaceous-grey species
'Sarcodon' rimosus OR - dark vinaceous grey-brown cap, but paler spines and flesh. This still needs to move to Hydnellum. It's a local species the study probably never heard of, so they neglected to move it.
Hydnellum fuscoindicum WA - very dark purple throughout the flesh and even in the spines. The taste should not be bitter.
Hydnellum 'fuligineoviolaceum PNW04 - one OR collection sequenced about 3% different in ITS to this very similar dark vinaceous black species. Supposedly this species will taste much more bitter than H. fuscoindicum. This species had never been reported from the PNW before. Perhaps it was mistaken for H. fuscoindicum, or perhaps it is 'Sarcodon' subincarnatus, below. An uncultured soil sample sequenced to a unique species in this group as well, but I don't know if it has ever fruited here.
'Sarcodon' subincarnatus OR - said to be vinaceous brown, never photographed. Based on spore size, we suspect this native species needs to be moved to Hydnellum, but we have no sequences yet and need collections. It should be investigated if H. 'fuligineoviolaceum PNW04' is this species.
'Sarcodon' rimosus © Andrew Parker and Bruce Newhouse, Hydnellum fuscoindicum © Noah Siegel
Stem base and flesh sordid green, and/or instrinsically scaly cap
Hydnellum scabrosum EU - intrinsically scaly cap, not just cracking into scales. The other intrinsically scaly capped species are true Sarcodons, above. Stem base and base flesh dark greenish.
Hydnellum glaucopus EU - much like Hydnellum scabrosum, usually (but not always) with a green stem base, but without the intrinsically scaly cap of that species. However, the cap does crack and break up into scales. This species was never suspected to be in the PNW, it was probably assumed to be H. scabrosum, but now we know both species occur here. One Victoria BC collection sequenced within 3-4 bp of a couple dozen EU sequences of H. glaucopus.
Hydnellum underwoodii CT - paler brown cap, also distinctly scaly like H. scabrosum, and said to be sometimes greenish at the stem base (otherwise hard to ID). We need local collections to see if this really is here, or if this is a case of confusion with H. scabrosum or H. illudens. We have ENA DNA to compare with, but that sequence has never been found out west yet.
Hydnellum illudens EU - a coarsely scaly species with a yellow-orange-brown cap and a farinaceous taste (most others are bitter). Stem base not reported as green. Our few local sequences sometimes come back dirty, but cleaned sequences are clearly matching H. illudens. It is another species that had never been suspected to be here before. It is described much like H. underwoodii so I wonder if reports are that were really this species.
Hydnellum scabrosum © iNaturalist user stevef (2 images), unsequenced H. scabrosum © Michael Beug
Hydnellum illudens © Regina Johnson
Other Orange-brown capped species
Hydnellum stereosarcinon NS - with an orange-brown somewhat zoned cap. Note that is was placed in Sarcodon because it can have a more regular cap and stem shape, but based on our local photos, it may also appear somewhat top shaped. Our sequences match a Quebec sequence of this Nova Scotia species within 3 bp in ITS, and the micro measurements match too.
Hydnellum 'versipelle PNW08' - probably looks a lot like H. stereosarcinon, because collections of the latter have been mistaken for the former, with a pinkish-orange cap that is not zoned. WNA and ENA sequences differ from EU sequences by 5 bp and 1 indel, so it should be investigated if ours is a distinct species. We have one Victoria BC sequence locally, but no photo. Hydnum crassum NS is currently a synonym, but it's possible they are not and that could be the name of our fungus.
Hydnellum stereosarcinon © Michael Beug and Richard Bishop
'Sarcodon' indurescens WA - dark orange-brown. KOH turned dark blue-green-black, but it may do that in a number of other species too. Three local collections have shown us what the sequence of this PNW native is. As a native it was not included in the Sarcodon study and has yet to be formally moved to Hydnellum.
'Sarcodon' indurescens © Kendra Dedinsky
Hydnellum lundellii EU - another species not known from the PNW until one collection from ID was sequenced. Our collection was dark brown but in the EU it is reported as orange-brown, making it resemble the above species. I don't know much about how to recognize it yet. There are 2 bp differences and 2 indels in ITS compared to a half dozen or so EU sequences, one of them a long indel.
Hydnellum lundellii © Joe Matanzas
'Sarcodon' calvatus OR / 'Sarcodon' calvatus var. odoratus ID - tan to cinnamon brown cap, especially large, caps up to 25-35cm across (most other species max out at 10-20cm). So far collections thought to be this local species have turned out to be something else. We need a type sequence or other reliable collections to find out what the sequence of this species is and get a verified photo. The var. odoratus is spicy/fragrant similar to H. suaveolens. Based on spore size, we suspect it needs to be moved to Hydnellum.
Hydnellum - top shaped - note that both this form and the top shaped form are mixed together throughout the tree. The two forms seem to have evolved back and forth regularly.
Hydnellum 'caeruleum PNW07' - this EU species is blue only on the young cap and perhaps a bit in the young teeth, with rusty orange stem flesh. Our 2 local sequences are at least 3 bp and 1 indel (maybe a few more indels) from many EU sequences, but some EU sequences are quite different, so I can't be sure our species is the real thing. Hydnellum cyaneotinctum ME is an ENA species that is currently a synonym, but if our west coast species turns out to be distinct from H. caeruleum, we should investigate if we can use the Maine name for it, or if ours needs a new name.
One WA sequence (from an old collection without any blue) differs by a few characters in ITS2 only, and has a string of differences near the beginning of ITS1 that might not be real. It may just be an aberrant sequence, but I am looking into whether or not there could be another species in this group.
Hydnellum 'suaveolens PNW01' - blue mostly on the stem, with zoned blue flesh, otherwise whitish darkening in age. Strong licorice odor. Our local species is 3% different in ITS from EU type area sequences, so perhaps our species is distinct. Hydnellum rickeri ME is currently a synonym, but it's possible it is not and that could be the correct name for our species.
Hydnellum cyanopodium CA - blue tones throughout the mushroom, including the flesh, with the margin of the cap beaded with red drops when fresh. Strongly aromatic, but not like licorice. One WA and one OR collection probably represent this CA species.
Hydnellum regium OR - many rosettes of purple black caps from a single stem. Two OR sequences of this OR native give us the sequence of this species.
Hydnellum nigellum NS - a small, thin, black species that resembles Phellodon except for having brown spores. We need PNW collections to see if the report that it occurs here is true.
Hydnellum 'caeruleum PNW07' © Lauren Ré, H. 'suaveolens PNW01' © Autumn Anglin and Mary McCallum
Hydnellum cyanopodium © Mary McCallum (2 images), H. regium © Bruce Newhouse
Hydnellum aurantiacum group - orange species
Hydnellum aurantiacum EU - orange cap, stem, and flesh. Whitish teeth and cap rim. Sequenced from WA and OR.
Hydnellum 'aurantiacum PNW09' - our one photograph of this sister species has brighter, paler orange cap, and a unique sequence. Perhaps reports of H. aurantile, described with a brighter orange cap than H. aurantiacum, teeth that are more orange when young, and smaller spores, is actually this species, but these teeth are white. So far H. auratile DNA has not been found here, but see PNW08 below. PNW09 is found so far in BC and WA.
Hydnellum 'aurantiacum PNW10' - another sister species to H. aurantiacum, found in BC. We don't even have a photo yet.
Hydnellum PNW07 - a thin fleshed orange mushroom, perhaps this is H. conigenum, below. If not, it probably needs a name. Found in BC.
Hydnellum 'auratile PNW08' - this collection with orange teeth (but caps not brighter orange) sequenced close to H. auratile, described as having orange teeth and a smaller, brighter orange cap than H. aurantiacum. Hydnellum auratile is reported from WA, we need collections of it to see what it actually is.
Hydnellum conigenum ID - a smaller, more thinner fleshed mushroom. No DNA data yet on what this is. Perhaps it is H. PNW07 above. It is assumed to be in this clade.
Hydnellum complectipes WA / Hydnellum complectipes var. disjunctipes WA - rosettes of many caps on a single stem, or each with their own stem in var. disjunctipes. No DNA yet, we need collections. It is assumed to be in this clade.
Hydnellum complicatum NY - with several caps on a single stem, but the one report of this was made a long time ago. We need local collections to see if this is really here. We do have ENA DNA to compare to.
Hydnellum cumulatum NS - not in the same clade as H. aurantiacum and the others. It also consists of many caps on a single stem, but perhaps not as distinctly orange as species in the H. aurantiacum group. EU and ENA sequences agree, but we have no WNA sequences to confirm Hall's report of this species here. Perhaps the similar H. complectipes has been mistaken for this species, or perhaps they're the same.
Hydnellum aurantiacum © NAMA and the Field Museum of Natural History, & Mary McCallum, H. 'aurantiacum PNW09' © NAMA and the Field Museum of Natural History
Hydnellum PNW07 © SVIMS, H. 'auratile PNW08' © Noah Siegel
Hydnellum scrobiculatum group - brown to red-brown, rough pitted cap (sometimes concentrically zoned), dark red-brown flesh with zones of different colours, sometimes bleeding red, but not dramatically like H. peckii.
Hydnellum scrobiculatum EU - cap not concentrically zoned. We don't know for sure what this sequence looks like, but so far our best guess doesn't match any species found yet in the PNW. It is regularly reported from the PNW but we need sequenced collections.
Hydnellum 'scrobiculatum PNW02' - one uncultured soil sample from OR has a unique sequence that may or may not represent a species that can fruit here.
Hydnellum 'scrobiculatum PNW03' - one OR collection is a unique species in this group. It was quite small, resembling a Phellodon (perhaps it was young?) and the caps seemed slightly zoned.
Hydnellum subsuccosum WA+OR - with a concentrically zoned cap. We have ENA and EU sequences purporting to be this, but no local collections sequenced. We need some.
Hydnellum concrescens EU - also with a concentrically zoned cap, is often reported in the PNW. This is a complex of 4 or more species in the EU, so there's no reason to believe this species is actually here. We do, however, need to document which other species with concentrically zoned caps are here besides H. subsuccossum. Hydnellum parvum AL is a synonym of H. concrescens right now, but it should be investigated if they really are synonyms, and if not, if that is one of our species.
Hydnellum 'scrobiculatum PNW03' © Joseph D. Cohen
Hydnellum subzonatum NS - with zoned, chestnut brown caps, that could be mistaken for zoned species in the H. scrobiculatum group, but the flesh of this species is not zoned. We don't have any sequences from anywhere yet, but it has been reported here once.
unsequenced Hydnellum subzonatum © Noah Siegel
Hydnellum 'peckii PNW05' NY - "strawberries and cream". Species in this group are easily recognizable due to their incredibly hot taste and the presence of many red drops on the cap when young. Caps somewhat reddish-brown. There is some confusion about what this species is. One NC sequence provided by Baird matches our sequences, so we may be right that this species occurs here. But other studies by Baird called sequences of a different lookalike by the name H. peckii, so until I see a type sequence, I'm not sure what this species should be called.
Hydnellum 'diabolus PNW06' ME - some local collections have sequences that instead match a Nova Scotia sequence labeled H. diabolus, currently thought to be a synonym. There are definitely a few species in this complex, and we have two of them, but without type sequences I'm not sure what to call them, so for now they both get codes as I'm not sure I'm applying the names correctly. Interestingly, these 2 are not sister species, so hopefully differences between them will be found in the future, and their real names revealed.
Hydnellum ferrugineum EU - said to look the same (many red droplets) but taste mild. There are 2 soil sample sequences from southern BC >3% different in ITS than EU sequences of this species. I don't know if it fruits here, but be on the lookout for something in this group that tastes mild. Unfortunately, to find out, you'll have to endure a lot of discomfort trying a lot of very, very spicy collections. This species could easily be mistaken for something in the H. scrobiculatum group.
Hydnellum 'peckii PNW05' © NAMA and the Field Museum of Natural History, H. 'diabolus PNW06' © Autumn Anglin
Hydnellum mirabile EU - with a white hairy cap that turns yellow and then dark brown, and exuding a brown juice when squeezed. One OR and once northern BC collection match many EU sequences well.
Hydnellum cf geogenium EU - bright yellow caps, we need local sequences to see if this is really what is found here. The real EU species is also found on the east coast, but collections in the Pacific Northwest are more spiky tubular and less fan-shaped than those in eastern North America, so we need to verify what our species really is.
Hydnellum mirabile © Leah Bendlin, unsequenced H. cf. geogenium © Noah Siegel
Phellodon (Bankera) - click to expand
Phellodon - small to medium sized, white spored mushrooms (so expect the spines to stay paler in age then in Hydnellum/Sarcodon), somewhat top shaped and thin fleshed, like diminutive Hydnellums, that dry to have a strong curry odor.
Bankera (now inside Phellodon) - large and vaguely cap and stem shaped like Sarcodon (with the same white spores and strong curry odor, especially when dried, as Phellodon). Bankera is paraphyletic inside Phellodon and they have been moved to Phellodon for those who don't accept such situations (like how Leucocoprinus is inside Leucoagaricus). This is another example showing how the top shape and the cap and stem shape evolved regularly back and forth inside this family.
Small top shape
Phellodon atratus CA - bluish black cap, spines, stem and flesh. This west coast native has been sequenced from OR and BC.
Phellodon melaleucus EU - blackish brown zoned cap (perhaps with purple tones) with a white margin, thin, dark stem and flesh but whitish spines when young. Also thinner than P. atratus. Some sequences (both in the EU and BC) seem to be 4 bp different, but close examination shows there might only be 2 significant differences so I think it's all one species. In Europe they think of it as a species complex, so it's possible ours might not be the real thing, but so far, I think it is.
Phellodon 'niger PNW03' EU - similar to P. melaleucus, but somewhat blue-black with thin white margin, a tomentose stem base and duplex flesh (if you cut the bottom of the stem off, the inner flesh is firmer and darker than the outer flesh). One soil sample sequence found one concept of Phellodon niger in BC, but I don't yet know if it can fruit here. It should be looked for. There are at least 4 concepts in the EU of this species, so it seems to be a complex. Our matches the most common one, but that's no guarantee that it's the right one, so I'm giving it a code.
Phellodon tomentosus EU - brownish concentrically zoned cap with a white margin, much like P. melaleucus, but often with a paler brown cap and thicker stem. Most EU collections agree on what the sequence is, but not all, so I'd like a type sequence to verify that we have the right concept of this species. We have a WA sequence.
Phellodon 'tomentosus PNW01' - one OR sequence is 6 bp different, so this sequence should be looked for again.
Phellodon PNW02 - much like P. tomentosus, with a brown concentrically zoned cap with a white margin, but one collection implies the spines are dark instead of white. Our one Victoria BC sequence is a unique species not sister to P. tomentosus, but perhaps to Bankera.
Phellodon atratus © Ann Goddard, Phellodon melaleucus © Jared LeBoldus, Stephanie Langlois, and Reva Shenwai
Phellodon tomentosus and Phellodon 'tomentosus PNW01' © NAMA and the Field Museum of Natural History, Phellodon PNW02 © Marty Kranabetter
Large cap and stem shape - often confused with Sarcodon/Hydnellum unless the spore print is checked.
Phellodon fuligineoalbus (Bankera fuligineoalba) EU - starts out pale but turns orange-brown and usually has debris incorporated into the cap.
Phellodon (Bankera) 'violascens PNW01' - pale violet grey cap, grey spines. Our three west coast sequences are consistently 4 bp different than one EU concept of this EU species and 6 bp different than another. Other insignificant changes in ITS make it look like there are many more differences. I asumme this is geographical isolation, but if our species is distinct, Phellodon carnosus NY is currently a synonym and could possibly be a name for it.
Bankera fuligineoalba © OMS (2 images), Bankera 'violascens PNW01' © Andrew Parker
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