Bolbitiaceae

© Kit Scates Barnhart and Steve Trudell

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

Click here for my Pictorial Key to Bolbitius, Conocybe and Pholiotina

Introduction

This family is made up of "Little Brown Mushrooms" with a cellular cap cuticle (fragile little things where the cap can break in any direction and tends to glisten and wrinkle, as explained in more detail in the Introduction on my Psathyrella page). The spore print can often be a light, bright cinnamon brown. They are often conehead shaped, in fact, Conocybe literally means "cone head". While Conocybe and Pholiotina s.l. are in fact little brown mushrooms with dry caps, Bolbitius are viscid, pretty little yellow or lilac mushrooms (rarely other colours too). They are all mostly urban mushrooms, growing in grass, gardens, straw, wood chips, dung and maybe rotting wood. Conocybe itself can be told by the presence of capitate cheilocystidia. They look like bowling pins. Pholiotina s.s. can be told by the presence of a ring, or usually at least a partial veil.

There only used to be two genera in this family known from the PNW, Bolbitius and Conocybe, with all Conocybes thought to be deadly poisonous, until genetic evidence showed that Conocybe needs to be split into four genera: Conocybe, Pholiotina and two as yet unnamed genera. Pholiotina contains species with deadly amatoxins, the principle toxins in the death cap, Amanita phalloides. Some Conocybes, like C. alpala, contain a less concerning phallotoxin (which they say is not absorbed through the stomach). The unnamed genera are not really studied yet so we don't know their edibility, and Bolbitius seems to be non-toxic.

A good paper on how this family is arranged, by Toth, can be found here.

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:

Conocybe and Pholiotina s.l. have been little studied here, and the species we're finding through sequencing do not always match the species reported from here.

Conocybe tenera, a widely misapplied name, has not been found here, but our most common species could be the similar Conocybe semiglobata.

A similar common species, differing microscopically with a paler stem needs a new genus name, for now called Pholiotina III.

Our subtly bluing, slightly hallucinogenic species are in need of a new genus name, for now called Pholiotina II.

Bolbitius - click to expand
Viscid, yellow or sometimes lilac LBMs, usually more colourful than the other genera in the family, which are dry capped and will at most be white or golden brown. Any of the species can wrinkle or become reticulated. Rarely, red, orange or green colours are found.
Species mentioned: Bolbitius titubans, vitellinus, titubans var. olivaceus, variicolor, reticulatus, aleuriatus

Bolbitius titubans var. titubans EU (Bolbitius vitellinus) - yellow viscid caps, fading to very pale tan. This EU species is found here.
Bolbitius titubans var. olivaceus EU (Bolbitius variicolor) - also yellow caps, fading to very pale tan, but supposedly with a hint of olive in some fresh caps, that are more likely to be a little more sturdy with some reticulation. Some collections with those differences from CA and the EU do sequence to be a distinct sister variety, described from the EU, a bit less than 2% different in ITS DNA. Unfortunately, not all specimens seem to show any of these traits, as shown in the following pictures, in which the proper variety could not be distinguished by the photos. We do not have local PNW DNA of this variety, but it has been reported once from BC so we need to confirm this.
Bolbitius 'titubans PNW02' - one BC sequence and some Russian and Chinese differ a lot in one section of ITS1 and may be a unique species.

Bolbitius titubans var. titubans © Julie Jones, two collections of var. olivaceus from CA © Joanne Schwartz, B. 'titubans PNW02' © Anne Polyakov

Bolbitius aff reticulatus ("aleureatus sensu CA") - lilac viscid caps. Bolbitius aleureatus is a newer synonym of the EU species B. reticulatus. CA sequences are distinct from EU sequences by more than 5% in ITS, so our species is likely an unnamed sister species. It has been reported from BC and WA, but we don't have local DNA yet.

Bolbitius aff reticulatus © Christian Schwarz

Bolbitius PNW01 - first found in CA and now in Portland, OR and ID, with a green to olive-brown cap and green to orange stem. The gills seem fairly widely spaced for the genus. The closest known species are other unusually colourful Bolbitius.
Bolbitius CA01 - another rare species has been found in CA, with a beautiful red-orange cap and yellow stem. Not yet known from the PNW.

Bolbitius PNW01 © Christian Schwarz (2 images), Leah Bendlin (2 images), and Ed Barge (2 images)

unsequenced B. PNW01 © Christian Schwarz, Bolbitius CA01 © Christian Schwarz

Conocybe - click to expand
Little brown dry capped coneheads without a ring (nor any kind of partial veil). Some Pholiotina s.l. may not have a ring, so microscopically, Conocybe can be told by capitate cystidia (sterile cells on the gills with prominent round heads). Species that turn blue are probably not Conocybe, but ringless Pholiotina II. At least some Conocybes contain phallotoxins, not as concerning as amatoxins, but still to be avoided.
Species mentioned: Conocybe alpala, lactea, aurea, tenera, coniferarum, rickenii, fuscimarginata, pubescens, semiglobata, fibrillosipes, brunnea, michiganensis, hausknechtii, pallidospora

"Bowling pin" capitate cheilocystidia of Conocybe © A and O Ceska

"Colourful" Conocybes (take that with a grain of salt)

Conocybe apala EU (=C. lactea) - This whitish (at least in age) Conocybe is found in grass. Reliable ITS EU sequences match a couple of Oregon sequences quite well, one within 1 bp and 1 indel, so we do appear to have this EU species here.

Conocybe 'apala PNW12' - one WA sequence is 4 bp and 2 chunks of indels different than any other known sequence from anywhere. It seems to be a high quality read, so I am keeping my eye out to see if we see this sequence again. It had a very tall, slender cap.

Conocybe aurea EU - This has a golden brown cap and somewhat of a paler or brighter stem (compared to the many boring brown Conocybes). This EU species is found in ENA and we now have a WA sequence to confirm its presence here too.

Conocybe apala © Buck McAdoo,     C. aurea © Yi-Min Wang

Scaly-stemmed

Pholiotina PNW06 "scaly stem, capitate cheilos" - this species has a distinctive scaly stem. It has been found twice in WA. The extremely capitate cystidia and lack of a ring make it key out to Conocybe, but the DNA sequence is clearly inside Pholiotina. I don't know what to make of this.

Pholiotina PNW06 "scaly stem, capitate cheilos" © Connor Dooley

LBMs - boring brown ones

Conocybe cf tenera - This is what almost every Conocybe in North America (never mind the PNW) is assumed to be, for some reason, but there's no evidence yet that it exists here in the PNW. It seems to be a complex of species in Europe, with DNA varying by 5% or so in the group.

Conocybe semiglobata complex (=Conocybe subpubescens?) - our most common species, it has a brownish stem. The similar Pholiotina III sp., below, seems to have a paler stem and may be our next most common species. There are four clades where sequences cluster, all within 1.5% of each other, so unless morphological and/or ecological differences are found between the clades, this may all be one species. But just in casefour possible species. Clade 4 is where most EU sequences are of both the type variety and var. campanulatus. Most local sequences are also in this clade. Many EU sequences of the older species Conocybe subpubescens fall in clade 4 as well, so it's possible that C. semiglobata is just a newer synonym, but we'll need type sequences to prove it. Clade 4 spores measured larger than clade 3 spores. Clade 1 is 4 bp different found in WA, clade 2 is 10 bp different found in OR, and clade 3 is about 6 bp different and found in OR and WA. Clade 3 spores were smaller than clade 4 spores. We had only have 1 sequence so far each of clades 1 and 2, and 2 sequences of clade 3. We need an official determination of what this species is to be called and if it needs to be split into varieties or possibly even new species.

Conocybe cf semiglobata © Alan Rockefeller and Jonathan Frank

Conocybe 'coniferarum PNW03' - a sequence 5 bp from the type sequence of this Russian species was a recent discovery in WA. Notably, it had a small bulb on its stem.

Conocybe velutinomarginata EU - with a unique mottled cap and spores 16x9u. A couple of PNW sequences match a sequence seemingly provided by the author of this species. This sequence has been given several species names before, so it's possible there's an older synonym.

Conocybe 'coniferarum PNW03' (note the bulb) © Danny Miller,     C. velutinomarginata (mottled cap) © Yi-Min Wang

Conocybe PNW09 - one WA sequence is almost 10% different in ITS from any other known species.

Conocybe PNW01/PNW02 - these two species aren't close enough to any other known species for me to give them any kind of a name, but they are clearly inside Conocybe and clearly related to each other. They may be one of the named local species on this page about which nothing much is known. The first one from OR was from an old burn site (I don't know if that's relevant) otherwise they were found on the forest floor.

Conocybe pubescens EU - one WA sequence is a match to reliable EU sequences of this EU species, so we appear to have this species here.

Conocybe 'hausknechtii PNW04' - one WA sequence is 3-4 bp and 1 indel different from the EU type sequence, so it could be that. It's even closer to one of three genetic species going by the name C. pallidospora, but that sequence is not likely labeled correctly.

Conocybe cf fuscimarginata EU - one southern BC ITS sequence differs in a handful of ambiguous places from some EU sequences of C. fuscimarginata, but this appears to be a complex of at least 3 species in the EU, so ours does not necessarily match the "real thing", but it is in that complex. No known photo.

Conocybe 'fragilis PNW10' - a very rich red-brown at first. The stem isn't always as opaque and richly coloured as the cap as it is in these photos, but it does seem to be an especially richly coloured species. If a TN and SC sequence are to believed, we have found C. fragilis in the PNW. It is a KS species. We don't have a type, but some people seem to believe that's what the ENA collections are. For now I'm giving it a code number.

Conocbye PNW11 - seems to be somewhat olive brown. It is close to some collections labeled C. cylindracea and C. siennophylla, but I don't know if they're labeled correctly.

Conocybe PNW13 - with a somewhat olive toned wrinkled cap (strongly so when fresh) and a cinnamon stem. We have one WA sequence, different from anything else in the PNW by >1% in ITS.

We have not worked out any clear, differentiating features of these species yet.

Conocybe PNW09 © Yi-Min Wang,     C. PNW01 © Bitty Roy,     C. PNW02 © Yi-Min Wang,     C. pubescens © iNaturalist user kellim,     C. 'hausknechtii PNW04' © Richard Morrison

Conocybe 'fragilis PNW10' © Aaron Hilliard,     C. PNW11 © Yi-Min Wang (2 images),     C. PNW13 © Damon Tighe and Yi-Min Wang (2 images)

Conocybe 'michiganensis PNW05' - described from MI as a Psathyrella because it seems kind of halfway between Psathyrella and Conocybe (it has somewhat dark spores like Psathyrella, but does have the cystidia of a Conocybe. It sequences inside Conocybe. In OR we have found perhaps a sister species differing by 2%, and with a very pleated cap, which the Michigan species is not supposed to have.

Conocybe 'michiganensis PNW05' © Jonathan Frank

Conocybe PNW07 - this seems to have a somewhat fibrillose cap (not entirely smooth) with a darker disc. It also has a somewhat two-toned stem. One collection had a greyish cap. It's closest known relative is Conocybe 'michiganensis PNW05'.

Conocybe PNW07 © Yi-Min Wang

Also reported:

Conocybe fibrillosipes WA - described from WA, but we have no genetics yet. It has capitate cystidia, but a sheathing veil, which Conocybe does not typically have, so it is just a guess that this species actually belongs in Conocybe. We have no purported sequences.

Conocybe brunnea EU - this EU species was reported by Smith as "rarely in WA" but the spore sizes he gave don't match that species, so I don't know what he found.

Pholiotina s.l. - click to expand
Pholiotina s.s. are usually ringed little brown mushrooms (usually at least with a partial veil). Two new genera that still need to be named are not ringed nor do they have partial veils, but will not have bowling pin shaped capitate cheilocystidia like Conocybe (but they may be somewhat rounded). Ringless, white stemmed species that may turn blue at the base are likely in Pholiotina II. Rings are rare on small brown spored mushrooms, and for some reason usually indicate "deadly amatoxin poisons", as in the case of Pholiotina as well as the Galerina marginata group.
Species mentioned: Pholiotina filaris, rugosa, gracilenta, utricystidiata, flexipes, pinguis, fimicola, stercoraria. Pholiotina II cyanopus, smithii, aberrans, coprophila,

The true Pholiotina s.s. is what Toth called Pholiotina III. I will call the new genera Pholiotina II (Toth Pholiotina II) and Pholiotina III (Toth Pholiotina I). (I wish Toth had called the true Pholiotina "Pholiotina I". That would have been less confusing). There is no good way yet to tell Pholiotina II and Pholiotina III apart (nor from the few Pholiotina s.s. species without a veil). Genetic work will have to be done to see if either of the 2 new genera could possibly be called Pseudoconocybe, Galerella or Tubariella, or if they will need entirely new names.

Pholiotina PNW06 "scaly stem, capitate cheilos" - a distinctive, mysterious species with a scaly stem, no ring, and with extremely capitate cheilos like Conocybe has a sequence clearly inside Pholiotina, and is also described above.
Pholiotina PNW07 - known from a single WA sequence, another odd Pholiotina with slightly capitate cystidia and no ring, it might be mistaken for Conocybe, Pholiotina II, or Pholiotina III.
Pholiotina PNW08 - one WA sequence, and one OR sequence (this collection with an appendiculate margin).
These odd Pholiotina are close enough to sequences used in Toth's muti-gene study that he placed firmly inside Pholiotina s.s. that I am sure this is where they belong.

Pholiotina PNW06 "scaly stem, capitate cheilos" © Connor Dooley (2 images),     P. PNW07 © Yi-Min Wang (2 images),     P. PNW08 © Kathryn True

Pholiotina 'rugosa PNW01' - as with Conocybe tenera for the non-ringed collections, almost every ringed collection in North America has been called Pholiotina filaris, but that is an EU species not found here so far. This species is probably P. rugosa, a year-round urban or wild species that may or may not be reticulated. One CA sequence, iNat#15087069, has 3 ambiguous locations that show that sequences of this species from CA and WA almost exactly match east coast sequences from a park in PA, back east, the part of the country where P. rugosa is described. I have not found DNA of other species back east yet for there to be another possibility. This species also closely matches the newer New Zealand species Pholiotina gracilenta which may be a newer synonym. For lookalikes, keep reading.
Pholiotina 'rugosa PNW02' - a species that differs slightly but clearly by at least 5 bp as well as the 3 ambiguous locations. It too grows in urban and wild places and may or may not have a reticulated cap, but anecdotally seems to be more likely to have a reticulated cap. As of now it has been found in all four seasons just like PNW01. Three WA sequences and 2 CA sequences are all about 5 bp different from each other so there seems to be some ITS variation in this species, but all of them cluster together away from PNW01 and seem to be distinct genetically. However, it will have to be decided if there are enough morphological or ecological differences to justify this being treated as a distinct species.
Pholiotina PNW04 - a spring wilderness species, but perhaps not as likely to have cap reticulation but we need more collections to really know. It may be capable of creating quite an impressive sized ring. We have three recent WA collections. This and 'rugosa PNW02' may be one of the little known species mentioned below.
Pholiotina 'pinguis PNW05' WA - a ringed species described from WA found on rotting hardwood logs in the spring. We have one spring collection that theoretically might represent P. pinguis found under alder. If it isn't P. pingus, then we still need DNA and photos of the real P. pinguis. It is microscopically similar to P. 'rugosa PNW01', and I don't know how to distinguish it from the other spring wilderness species in this group described above.

Pholiotina 'rugosa PNW01' © Danny Miller,     P. 'rugosa PNW02' © Yi-Min Wang,     P. PNW04 © Yi-Min Wang,     P. 'pinguis PNW05' © Sharon Squazzo

Pholiotina PNW03 - this species leaves its veil material on the cap margin instead of making a ring. It has a somewhat viscid cap. No species reported from the PNW lacks capitate cystidia and has an appendiculate margin and a tacky cap, so I can't find any existing named species that this might be. A full microscopic workup is available for one observation. It has been found in WA and OR (shown).

Pholiotina PNW03 © iNaturalist user fungi_fanatic

Pholiotina aff utricystidiata - One Vancouver BC sequence growing indoors in potted plant soil is 4 bp different than a reliable Toth sequence of the EU species P. utricystidiata, plus one location that is ambiguous. But it hasn't been found in the wild here yet.
Pholiotina flexipes WA - ringed species described from WA found on avalanche debris. No photos nor DNA yet.
Pholiotina fimicola WA - ringed species described from WA on dung in the spring, but we have no photos nor DNA yet.
Pholiotina stercoraria WA - ringed species described from WA on dung in the spring and fall, with larger spores. No photos nor DNA yet.

Pholiotina II - unless white stemmed and subtly bluing, must be separated microscopically from Conocybe, and then difficult to separate at all from Pholiotina III.
'PholiotinaII' cf cyanopus NY - This ringless species may subtly turn blue at the base of the stem (which is notably white) due to the presence of hallucinogenic Psilocybin. Usually in grass, sometimes in moss. Toth has a sequence from the EU that might be this, but we need ENA sequences to know for sure what this species is (the type is from NY). It was reported from BC + WA by Stamets, but we have no local DNA either.
'PholiotinaII' smithii MI - A very similar ringless, subtly bluing, white stemmed species more usually found in mossy bogs than grass. We have east coast sequences and matching PNW sequences from WA, OR, and ID.
'PholiotinaII' coprophila EU - EU species found on dung that Toth has provided a sequence for, but his sequence is odd compared to most other EU sequences. A Smith collection from ID and a modern BC sequence from the interior of BC match most EU sequences.
'PholiotinaII' aberrans EU - EU species found on rotting conifer logs that Toth has provided a sequence for, reported by Smith from ID, but we need a local sequence to confirm that's actually what he found.

'PholiotinaII' smithii © Ed Barge,     'PholiotinaII' coprophila © Anne Polyakov

Pholiotina III - must be separated microscopically from Conocybe and difficult to separate from Pholiotina II at all.
'PholiotinaIII' PNW01 - We have found our first as yet unnamed species in this genus three times in WA and once in OR. An additional OR sequence is somewhat different but appears dirty and may not represent a different species. It appears to have a white pruinose stem, paler than the common Conocybe cf semiglobata, above.
'PholiotinaIII' PNW02 - appears to have a paler cap and thinner stipe. One sequenced collection from OR.

'PholiotinaIII' PNW01 © Danny Miller, Daniel Winkler and Stephen Russell,     'PholiotinaIII' PNW02 © Connor Dooley

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