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Danny’s DNA Discoveries – Hebeloma of the PNW
by Danny Miller

Click here for my Pictorial Key to Hebeloma


Hebeloma - viscid capped, boring brown, brown spored mushrooms found on the ground, never with a ring but some with a cortina (cobweb veil) like Cortinarius. They are usually fragrant, like radish or sweet smelling (famously one is chocolaty). The ones that don't have the odor are easily confused with the less common odorless Meottomyces and Phaeonematoloma.

Dr. Henry Beker is leading a team of people creating an amazing project documenting worldwide Hebeloma. Single handedly, they are transforming Hebeloma from one of the least understood genera of LBMs on the planet to one of the best documented genera. If Henry and his team can do this for Hebeloma, there's no reason why almost any other genus can't be similarly documented. They have already released "Fungi Europaei Volume 14 - Hebeloma" for Europe and are working on North American monographs now. Soon, they will have a public website at www.hebeloma.org where you can access all of their data and see the amazing tools they have created. Here are just some of the amazing things they can do with the data:

  • computer generated descriptions of each mushroom that read like prose as if a human wrote them, created automatically from looking at the characters of every collection ever made of that species. The description is re-created every time a new collection is made to keep it updated. It can create descriptions like "usually between 2 and 4 cm across, but rarely bigger than four. Often found with spruce but rarely with hardwoods".
  • You can graph anything you can imagine, such as latitude of a find vs. altitude, to see logical correlations such as how the further south you go, the same mushroom will be found higher up. Playing around with the plotter has discovered some fascinating facts such as the hotter the temperature where a certain Hebeloma was collected, the smaller the spores and the more gills it has.
  • computer assisted identification using a synoptic key (similar to MycoMatch but specific to Hebeloma and focusing on only the few features they calculated that are necessary to know to best identify your Hebeloma).
  • it can help figure out the actual tree associations by looking at all the trees near each collection of a species and noticing what the overlaps are.
  • Excellent search tools to look at things like where each species grows and how common it is based on over 10,000 worldwide collections so far.
  • The computer can figure out the actual differences between the species and create a new, updated binary key automatically every time a new collection is made with more data.

This is a very difficult genus of species mostly separated microscopically. In fact, Henry's data analysis has shown that the only useful macroscopic character for telling them apart may be the number of complete gills. For now, I am allowed to say that "unpublished research indicates the presence of the following species in the PNW".

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.

Section Hebeloma subsection Hebeloma - veiled (cortinate) - radish odor, dark disc, somewhat stout?, almost smooth, almost non-dextrinoid spores

Along with the cobweb veil and typical radish odor, the caps are somewhat darkly pigmented, at least on the disc, although often paler on the rim (two toned). We usually identify any mushroom in this section and the next (veiled with a dark disc) as Hebeloma mesophaeum. It turns out that a number of species share almost identical ITS and so they cannot be distinguished by DNA sequencing of that region. Yet they do have clear microscopic or ecological differences and Dr. Beker is keeping them as separate species. If you don't have all that data and don't want to wade through the keys and descriptions of all of them, you would be forgiven for thinking of all of the following species simply as subsection Hebeloma. Just be sure you don't have something in subsection Amygdalina, described next, microscopically different and perhaps with a slightly more slender stem. Subsection Hebeloma are common as a group.

H, mesophaeum EU (=H. lateritium WA, =H. vinaceogriseum ID) - known from every state and province in the PNW BCx2, WAx6, ORx1, IDx1

H. excedens NY (=H. sterlingii ID) - only known recently from 2 south BC collections (photo Oluna MO#258069)

ITS nor any known gene can distinguish the former two yet

H. angelesiense WA (=H. pseudostrophosum WA) - only known recently from 1 WA (Olympics, the type area, me) and 1 OR collection (Ron P)

H. velatum NY (=dunense EU) - only known from 1 WA collection (Noah 4268 photo)

H. alpinicola ID (=H. nigromaculatum OR, =H. perigoense CO, = H. strophosum var. occidentale, = H. pseudofastibule var. distans) - known recently from 1 Whistler BC and a 7-9 WA collections (Buck 492, RHM-19-26, Shannon 13972?, Noah 1238, 3423, 4505, me @ Barlow Pass photo)

H. marginatulum EU (=H. ollaliense OR) - only known recently (post website) from 1 BC and 1 WA collection, no photo or info yet

H. harperi CA - Oluna Victoria x6 (including F29522) plus JLF OR iNat79807601

ITS DNA will not well distinguish any of the above species for me, but Henry's tree seems to (barely)


H. colvinii NY - awaiting type sequence MN017797. we only have old, almost 100 year old rumours of this species being here, until I heard Noah found it in WA (photo but no DNA) but now I can't find where I heard that and it's not reported on the website, so take that with a grain of salt. The only species easily separable by ITS.


Hebeloma section Hebeloma subsection Hebeloma © Noah Siegel


Section Hebeloma subsection Amygdalina - similarly veiled with dark discs (perhaps more slender?, with distinctly but finely verrucose distinctly dextrinoid spores and perhaps more amygdaline shaped spores)

Species in this section differ microscopically from subsection Hebeloma and may appear more slender (but don't count on it). Except for some collections of H. praeolidium which sometimes has a sweetish odor instead of the usual radish odor, these have probably all been mistakenly called H. mesophaeum in the past. They are much more uncommon here than those in subsection Hebeloma.

H. paludicola NY (=H. hygrophilum EU, =H. oregonense var. oregonense OR) -  the cap can go through a wide range of colours, no wonder if was redescribed a couple times. We have 2 recent collections from OR  and 1 from WA.

H. palustre NY (=H. clavulipes EU, =H. lutescentipes OR, =H. idahoense ID, =H. occidentale OR, =H. stanleyense ID) - only recently found once in WA and once in OR - no DNA of the type, but microscopic and ecological examination showed H. clavulipes etc. are most probably younger synonyms.

H. praeolidum WA - may have a sweetish or radish odor - many recent collections have been from BC, WA and OR. Buck 148 #5, Buck 830 photo

H. fuscatum EU - one recent WA collection (JFA9881, no photo) - now, even Henry can't distinguish from H. palustre ITS.

ITS DNA will not well distinguish any of the above species, nor H. monticola from boreal BC and AK. Henry can sometimes do it (except for H. palustre/fuscatum), but not me.

H. paludicola © Noah Siegel (2 images)


Hebeloma palustre © Noah Siegel and Buck McAdoo (2 images),     H. praeolidum © Buck McAdoo



Section Scabrispora - scant veil (cortinate), paler cinnamon caps - (range of scents, some are bitter?)

Hebeloma viscidissimum OR (=amarellum CA, =H. pinetorum OR) - these species have scanter veils and paler caps than the other veiled sections above usually with a cinnamon tinge. This species usually has a bitter taste (originally described as Gymnopilus) and lacks much of an odor, but H. pinetorum's type was fragrant. The stems seem to get very dark from the bottom up. 2 recent collections from BC and a few from WA.

H. immutabile WA - no odor or taste, many recent Haida Gwaii BC collections, 2 recent Vanc BC (KJ146710) and WA (NS4592) collections for PNW.

Hebeloma parcivelum OR - this species is said to lack much of an odor or taste, perhaps weakly bitter. 1 recent NBC and WA collection.

Hebeloma olympianum WA (=H. aurantiellum OR, =H. subhepaticum WA) - weak odor and taste, recently collected once in Haida Gwaii BC and once in WA

Hebeloma viscidissimum © Noah Siegel,     H. immutabile © Priyashni Goundar,     H. parcivelum © Noah Siegel,     H. olympianum © Noah Siegel



Section Denudata - pale caps (poison pie), no veil, spores not dextrinoid, stems not usually bulbous

These are some of the species commonly referred to as "poison pie", along with species in section Velutipes. You should be able to figure out which of the two sections your poison pie is in, but getting to species is currently very difficult, even with DNA, as some species share DNA with each other in each section.

H. crustuliniforme EU - recently BCx1, WAx1, ORx1

Hebeloma aanenii EU - 1 recent WA collection (NS4769) no photo yet

H. eburneum Morocco - 2 near AK, 1 recent in OR, 3 recent in WA (Buck 433, RHM-19-27) - ITS DNA will not well distinguish these three species.


H. vaccinum EU - known recently from 2 OR collections, 1 not in DB -

H. cavipes EU - many recent BC+WA collections. - ITS DNA will not well distinguish these two species.


H. ingratum EU - white or tan with a darker brown disc. recent NBC + 2 WA - same ITS as H. album

H. album NY (=H. fragilipes Fr)uniformly white or tan without the darker disc. 1 from WA (need photo of RHM-18-22) +ORx3, also NBC - the type of synonym H. fragilipes is available, no DNA from H. album type but close examination synonymized them - same ITS as H. ingratum


H. helodes EU - 1 recent Whistler BC and 1 recent WA

H. sordidulum NY (=H. arenosum WI) - we have the NY type sequence. Henry confirmed it twice from Vanc Isl. (collections from the 60s and 70s). We have sequences that match very well from CA and WA.

H. limbatum EU - we have the Italian type sequence and local collections NS4626 and 4628 from OR and CA oak.

H. hiemale EU - ITS is the same for EU type area collections and our local collections, but some gene regions do differ. However, with no ecological nor morphological differences noted, ours is assumed to be the real species. Found commonly in BC, WA, OR.

Hebeloma cavipes © A & O Ceska,     H. album © Richard Morrison,     H. sordidulum © Yi-Min Wang,     H. hiemale © Buck McAdoo



Section Velutipes - usually pale caps (poison pie), no veil, spore dextrinoid, stems often bulbous

This is the other section containing "poison pie" like species, also difficult to differentiate beyond section. Besides having dextrinoid spores, the species here are more likely to have a stem bulb.

H. velutipes EU (=H. latisporum ID, =H. salmonense ID) - found everywhere

H. incarnatulum MI - recent 3 BC + 5 WA

H. subconcolor EU - a darker capped species under high elevation willow, only 20-32 complete gills. Only found twice in the PNW in WA.

H. leucosarx UK - recent 1 boreal BC + 1 WA, need ITS (even though it's not unique)

H. albidulum NY - awaiting type sequence MN017799. Old rumour from BC, Henry hasn't found it here (east coast only). This ITS is unique so we will recognize it if we find it, but we probably won't.

ITS DNA will not well distinguish most species in this section. H. velutipes may be the most common species of Hebeloma in the world. Our local collections have many ambiguous locations in their sequences (up to 20!)

H. neurophyllum NY - known in the PNW from a single recent WA collection. This species is in fact distinguishable by ITS.

Hebeloma velutipes © Buck McAdoo,     H. subconcolor © Yi-Min Wang (2 images),     H. neurophyllum © Yi-Min Wang (2 images)



Section Sinapizantia - very large, scaly stem, no veil

Hebeloma albomarginatum TN (=H. megacarpum MI) - this species is easy to recognize with its large 15cm cap and scaly stem up to 3cm thick. It also has a strong radish odor. We used to call it Hebeloma sinapizans until genetics figured out the correct name for our species. A few collections from BC, WA and OR.

Hebeloma albomarginatum © Buck McAdoo



Section Sacchariolenta - sweet odor, no veil

Hebeloma sacchariolens EU - this sweet smelling  (but not usually like chocolate) species, unlike H. praeolidium above, has no veil, and always smells sweet. One sequence differs by 4 bp from the others, but shares morphology and ecology and is likely the same species. Several recent collections from BC, WA, OR.

Hebeloma ischnostylum EU - a similar species that can be a little smaller, and a little paler, perhaps with pinkish or yellowish tones, perhaps more likely to be found in urban areas (but H. sacchariolens can be found in urban areas too). We have sequences from near Victoria BC, WA and OR.

Hebeloma sacchariolens © Michael Beug,     H. ischnostylum © Noah Siegel and Yi-Min Wang



Section Theobromina - occasional odors of stale chocolate (pungent) give this section its name

Hebeloma kelloggense ID - finds of this species may have been going by the name Hebeloma theobrominum EU but that species does not seem to occur here, and nobody noticed we had our own species already described from Idaho which has also been found recently 4 times near Victoria, BC. Photo MO#188815. If these species do have a veil, it is hard to detect.

Hebeloma kelloggense © A & O Ceska



Section Naviculospora - best recognized microscopically?

Hebeloma avellaneum WA - this section as a whole is defined microscopically (like all sections) but this species might be recognized by an orange tinged pale cap that is somewhat frosted with canescense. 5 recent WA collections.

Hebeloma pungens OR - also best recognized microscopically, this species may have a pungent radish-like odor and a scaly stem. Only known from the type sequence, plus a CA sequence. No photo.

Hebeloma avellaneum © Noah Siegel (2 images)


Section Syrjense - "corpse finders"

Species in this section are famous for being able to fruit above human remains, but otherwise need to be identified microscopically. Hebeloma syrjense EU DNA was found on root tips of trees in the interior of BC, so it's possible it could fruit here too. It fruited in AK.

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