© Steve Trudell

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Danny’s DNA Discoveries – Galerina s.l.
by Danny Miller

Click here for my Pictorial Key to Galerina

Introduction

Galerina has kind of been a catch-all genus for many non-descript LBMs, so it may come as no surprise that they do not all appear to be related to each other. One 2005 study found they may need to be split up into at least four genera, but that only looked at one gene at a time. Smith and Singer's 1964 Galerina monograph is able to distinguish species fairly readily microscopically, but you will need a scope and that key to have a hope of identifying your Galerina, as many are obviously very similar. Much of what we know about our local species comes from an excellent 2019 master's thesis at UBC by Brandon Landry, who sequenced many local collections. Subsequent to that, Brandon and his colleagues published a 2020 paper confirming which Galerina were actually deadly that at the same time did a much-needed multi-gene study to confirm how the genus should be split up, adding a fifth section to the list.

The caps are usually dry but occasionally viscid, usually hygrophanous, either having a veil or not maybe leaving a ring on the stem and usually adnexed gills in age but sometimes starting out decurrent like Tubaria. They are found on moss, logs, or mossy logs. Like Pholiotina (Conocybe), those with a ring may be DEADLY POISONOUS with Amatoxins. Beware all LBMs with a ring, and those without because they may have lost their ring. Most Galerina have warty spores but at least one of the groups has smooth spores.

Galerina will probably be split into at least 5 genera, which are nicknamed below, showing which sections in Smith usually (but not always) go in which clade:

Galerina - (section Galerina). These are considered the real ones only because the first one ever found was one of these. They are fragile and mycenoid, grow in moss, do not have veils and have distinctly warty spores usually 2 per basidium. They are most closely related to Naucoriopsis and have the same large, ventricose-fusoid gill cystidia, also found on the stem (caulocystidia) and sometimes on the cap (pileocystidia).

Mycenopsis - (section Mycenopsis subsection Mycenopsidae and section Calyptrospora and perhaps at least one species in section Inoderma with a fibrillose cap). They are also fragile and growing in moss. Many smooth or almost smooth spored species without tibiiform cystidia will be here (the only cystidia are cheilocystidia on the gill edges). They may have a faint veil that is hard to detect. One monkey wrench: Gymnopilus lives inside this genus, making it paraphyletic instead of monophyletic, so some will argue that Mycenopsis will have be split into a bunch of new genera to accommodate. I hope not. This is the same situation as Leucocoprinus living inside of Leucoagaricus.

Tubariopsis - (subgenus Tubariopsis and section Mycenopsis subsection Tibiicystidiae Stirps Tibiicystis). They are fragile, growing in moss and veil-less. These species have tibiiform cystidia everywhere but on the gill faces (no pleurocystidia) but must also have either no clamps (all others do) or grow specifically on Sphagnum moss.

Naucoriopsis - (section Naucoriopsis which contains some deadly species and other sections). Section Naucoriopsis species are somewhat fleshy compared to the above groups, although still LBMs, with a ring or ring zone on the stem and usually an incurved margin on the cap. The spores are usually distinctly ornamented (warty) and there are large ventricose-fusoid cheilocystidia and pleurocystidia (but no caulocystidia nor pileocystidia).

Other sections clade near enough that they could be considered part of Naucoriopsis instead of new genera of their own, although then the definition of the genus would have to be expanded. At least some species in the following sections clade nearby: section Mycenopsis subsection Tibiicystidiae stirps Triscopa (tibiiform cystidia, not in Sphagnum and distinctly warty spores), Galerina jaapii (keys out to Mycenopsis), section Inocyboides (metuloid pleurocystida), section Physocystis (round pleurocystidia) and section Porospora (with a germ pore).

Sideroides - (section Mycenopsis subsection Tibiicystidiae stirps Sideroides). These grow in non-Sphagnum moss, have tibiiform cystidia and smooth or almost smooth spores. Some can be fleshy with veil zones like Naucoriopsis.

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

Summary of Interesting Results

Here are some of the newest, most interesting results of the study:

  • TBD

Galerina

Galerina cf atkinsoniana MI - EU and WNA DNA agree, but we need type area DNA from ENA to make sure we have the same thing they do. Many sequences labeled with this name are a different species, but Landry thinks he figured out which sequences are correct, and I agree.

Galerina cf vittiformis complex EU - Landry has chosen the type sequence of G. vittiformis forma bispora WA as representing the original EU species, out of many possible genetic species that come out when EU collections of this are sequenced. This was just a guess. The PNW has this species, but it may be a complex as some sequences differ from others by as much as 4 bp.

Galerina aff. vittiformis sp. 3 - We also have an even more abundant sister species he called "aff vittiformis sp. 3" (sequenced a couple dozen times). Also, see the next species.

Galerina cf perplexa MI - a couple of collections in the G. vittiformis group keyed out to this, one from BC and one from the EU, but that's not nearly enough proof that they are correctly named. Also, the species is from ENA and no sequences are from there. If this isn't G. perplexa, we have to figure out what reports of it do represent. Landry calls this "aff vittiformis sp. 5".

Galerina cf minima NY - two Greenland sequences could possibly represent this, and Landry seems to agree, but that's not proof. Most sequences of collections turned out to be G. vittiformis instead. Smith reported it from the PNW, but we have no local sequences yet.

Galerina atkinsoniana © A and O Ceska,     Galerina cf vittiformis complex © Bitty Roy,     G. aff. vittiformis sp. 3 © Daniel Winkler

 

The following are described from the PNW, so are here by definition, without any genetic data yet (some may be duplicates, which often happens in Smith monographs):

Galerina diabolissima ID -
Galerina fontinalis ID -
Galerina funariae WA -
Galerina latispora WA -
Galerina mollis var. latifolia WA -
Galerina mollis var. mollis WA -
Galerina nordmaniana ID -
Galerina oreina WA -
Galerina pubescentipes ID -
Galerina rainierensis WA -
Galerina umbrinipes OR -

 

 

Mycenopsis

Galerina cf pumila EU (=G. vexans CA) - this is confusing, it's a Persoon species from 1801 yet Index Fungorum says the type area is Washington State, which didn't exist then. I'm assuming it's an EU species. Many UNITE sequences agree on what the DNA is. A Greenland sequence that Landry uses disagrees, but my money is on the dozen UNITE EU sequences being right. The type sequence of Galerina vexans is the same as my concept. Victoria BC DNA matching all the EU sequences has been found at least 4 times.

Galerina pumila var. subalpina WA - an original Smith collection from OR sequences to be quite different, so this may need to be promoted to species. It's been found recently in OR and 5 times in Victoria.

Galerina mniophila EU - a dozen UNITE sequences agree and match a couple of BC sequences and an ID sequence. At >1% different from G. pumila var. subalpina, Landry considers them to possibly be synonymous, but I see clear clades, so I consider them separate.

Galerina hypnorum EU - not known from the PNW before, but the DNA has shown up in south BC, matching UNITE EU sequences.

Galerina fallax WA - a half dozen local sequences seem to settle what the DNA looks like. Interestingly, the DNA has been reported from Antarctica, where very little DNA of any kind is found. This is one of those section Calyptrospora species. One collection called Galerina subfiliformis WA turned out to be this, but that doesn't mean they're the same.

Galerina paludosa EU - we have EU DNA that seems to agree, but nothing local to confirm its existence here.

Galerina subcerina ME (=G. calyptrata UK) - we have what may be the type sequence of G. subcerina, and a lot of UK sequences of G. calyptrata, and they are the same, so the latter may be a newer synonym. We have 5 Victoria BC sequences that match. It was never recorded by Smith from the PNW, but here it is. It is another section Calyptrospora species.

Galerina cf luteolosperma MI - I have no idea why Landry calls these Victoria BC sequences by this name, there are no ENA type area sequences to compare to, but for now let's say something matching the description has been found in the PNW. Smith never reported it from here. We need eastern sequences to verify.

Galerina fibrillosa WA - we have the type sequence, and this section Inoderma species is indeed in Mycenopsis.

Galerina pumila var. subalpina © Bitty Roy

 

The following, many described from the PNW and therefore here by definition, do not have any genetic data yet (some may be duplicates). Smith's section is given.

Galerina alluviana WA - Mycenopsis
Galerina lacustris WA - Mycenopsis
Galerina obscurata MI - Mycenopsis
Galerina rugisperma WA - Mycenopsis
Galerina subbadipes EU - Mycenopsis
Galerina subfiliformis WA - Mycenopsis (one purported collection sequenced to be G. fallax)
Galerina tatooshiensis WA - Mycenopsis
Galerina tundrae WA - Mycenopsis

Galerina acicola OR - Calyptrospora
Galerina anelligera ID - Calyptrospora
Galerina cerina USA - Calyptrospora (5 varieties said to occur in the PNW)
Galerina cortinarioides WA - Calyptrospora
Galerina evelata NH var. fulvipes WA - Calyptrospora
Galerina odora WA - Calyptrospora
Galerina payettensis ID - Calyptrospora

Galerina insignis WA - Inoderma
Galerina vaccinii WA - Inoderma

 

 

Tubariopsis

Galerina semilanceata WA (=G. dimorphocystis MI, =G. heterocystis Jamaica) - most sequences of all three are almost identical, so they are probably synonymous, with the oldest name being G. semilanceata. Landry also synonymizes G. clavatus EU, but EU sequences are quite distinct, so I believe that EU species is distinct.

Galerina nigripes OR - we may have the type sequence, and we do have several recent BC sequences.

Galerina cf tibiicystis MD - Smith placed this in a different section, but it clades inside Tubariopsis. We have a few EU sequences, but no ENA type area sequences, nor local sequences. We'll need both of those to figure out if reports of this species being here are true.

Galerina semilanceata © Steve Ness


Reported from the PNW, but no genetic data yet.

Galerina brunneomarginata CA (as G. brunneimarginata) -

 

 

Naucoriopsis

First, the deadly species containing alpha- and beta-amanatin, like the deadly Amanita. They clade tightly together genetically.

Galerina marginata EU complex (=G. venenata OR, =G. cinnamomea OR, =G. unicolor EU) - 163 UNITE sequences of G. marginata, including almost every EU collection ever made, sequence to the same thing, so I'm very sure we know what this is. For some reason Landry chose a different concept, of a few collections found in Asia and the US (with only a couple of soil samples showing the DNA in Europe), but I disagree. The type sequences of G. venenata and G. cinnamomea are practically identical to all the EU sequences of G. marginata, so they appear to be synonyms. G. venenata and G. unicolor had already been declared synonyms. So had G. oregonensis and G. autumnalis, but they appear to be distinct genetically. The DNA of the dozens of sequences we have are not identical. Some differ by as much as 5 bp from each other, but there are no noted ecological or morphological differences, and the differences do not work out to represent the different names that have been used for the species, so I believe it to be all one slightly variable species.

Galerina autumnalis NY - is said to be distinctly viscid. East coast collections do sequence >3% different from the above, enough that this is probably a distinct species, even though it was synonymized. It was reported from the PNW, but we have no local sequences of it yet to prove it. We need sequences of very viscid local collections to prove that is not a differentiating feature of the species and to show that G. autumnalis is not likely present here.

Galerina castaneipes OR (=G. oregonensis OR?) - the type sequence and a couple dozen recent local sequences are >2% different from the above, enough that it is probably a distinct species. It was separated out as a unique species on the basis that it lacked a veil. However, this photo and other photos show it at least has a ring zone, so I'm not sure how to identify it. If it does have a veil, how did Smith correctly note that it is a unique genetic species? Is the veil more ephemeral? This needs study. Galerina oregonensis, with a weak veil, had some collections that match this genetically, so perhaps it is a newer synonym, and this species has a weaker veil than G. marginata, which was just overlooked in the type collection of G. castaneipes.

Galerina aff. marginata sp. 4 - many collections labeled G. oregonensis match the type sequence of G. castaneipes, as noted above, so it may be a newer synonym. One collection has a unique sequence, differing by 13bp and 2 indels, that Landry calls G. aff. marginata sp. 4. It probably is not G. oregonensis, but we do need to figure out what that is as well as come up with a name for this sequence if in fact this unique sequence is correct. We'll need to see if we ever see this sequence again, I hate to make any conclusions based on a single sequence.

Galerina marginata complex © A and O Ceska,     Galerina castaneipes © Danny Miller

 

Deadly?

Galerina badipes EU (=G. cedretorum EU) - usually less fleshy than the above, but not quite mycenoid. It is the closest species to the deadly clade, this species contains gamma-amanatin, but not alpha- nor beta-, at least not in detectable amounts, but that could still make it deadly. No other Galerina in Naucoriopsis nor elsewhere were found to be deadly. The US type sequences of two American varieties, G. cedretorum var. bispora and G. cedretorum var. microspora, have the same sequence as most EU G. badipes and about a dozen recent collections of local G. badipes so all varieties of both species are probably the same genetic species. As above with G. marginata and G. castaneipes, G. cedretorum is described as a veil-less G. badipes, but it appears that there is a variable amount of veil material in the species.

Galerina badipes © Danny Miller

 

Galerina pseudocamerina MI var. fulvovelosa WA - keys out to section Sideroides. This was not properly described, so I don't know if a new name will be chosen. We only have sequences from the EU, not from ENA. Our one WA sequence is about 5 bp different. If it is correctly identified, we do have a local name to use, even if it is decided to be distinct from the type variety.

Galerina jaapii EU - keys out to Mycenopsis. We do have EU sequences to compare to, but no local sequences to confirm the reports of it here.

Galerina aff nana EU - keys out to section Inocyboides. Our local sequences are a few % different from the one EU sequence we have, so ours may be a distinct sister species. We need more reliable EU sequences to tell. Galerina BTSP #24 is probably a local sequence from a lost collection that seems to be nearest G. nana, and that DNA should be looked for again.

Galerina pruinatipes WA - keys out to section Physocystis. we have a few sequences of this, including one from WA, so I think we know what this is.

Galerina aff triscopa EU - keys out to section Mycenopsis subsection Tibiicystidiae stirps Triscopa. EU sequences are about 4% different than the one BC sequence we have, so ours may be a sister species. We do have a local form that might also represent the genetics of local type form collections: G. triscopa forma longicystis from Idaho.

Galerina pseudocamerina © Daniel Winkler,     G. aff. nana © A and O Ceska

 

The following, many described from the PNW and therefore here by definition, do not have any genetic data yet (some may be duplicates). Smith's section is given.

Galerina rudericola WA - Naucoriodes
Galerina subglabripes WA - Naucoriodes
Galerina subochracea OR - Naucoriodes
Galerina vialis WA - Naucoriodes

Galerina camarinoides OR (as camerinoides) - Mycenopsis subsection Tibiicystidiae stirps Triscopa
Galerina cascadensis WA - Mycenopsis subsection Tibiicystidiae stirps Triscopa
Galerina laticeps OR - Mycenopsis subsection Tibiicystidiae stirps Triscopa
Galerina pistillicystis NY - Mycenopsis subsection Tibiicystidiae stirps Triscopa
Galerina pteridicola WA - Mycenopsis subsection Tibiicystidiae stirps Triscopa
Galerina pulchra WA - Mycenopsis subsection Tibiicystidiae stirps Triscopa

Galerina olympiana WA - Physocystis

Galerina stagnina EU - Porospora
Galerina subdecurrens WA - Porospora
Galerina subtruncata WA - Porospora

 

 

Sideroides

Galerina cameria EU (=G.pseudobadipes EU) - we have EU sequences and they are different than the many sequences from NA that have been identified as this. The same DNA as found in the EU has not been found here yet to verify the reports of it.

Galerina cf mammilata WA - we have a purported WA sequence and a dozen BC sequences that might be this.

Galerina stylifera var. velosa ID - We have the type sequence, and it is distinct from the type sequence of G. stylifera var. caespitosa MI. The DNA has also been found once in WA and 8 times in BC, but usually mistakenly called G. sideroides.

Galerina stylifera NY - we don't have any type area sequences. EU and BC sequences are quite different than G. stylifera var. velosa, leading Landry to postulate that if most collections that people thought were G. sideroides turned out to be a variety of G. stylifera, then perhaps everybody has it backwards and the sequences that everybody is calling G. stylifera are actually G. sideroides or close to it. He thinks this type variety might not be distinct from G. stylifera var. velosa. We need reliable sequences from ENA.

Galerina stylifera var. badia ID - also, what is this variety? We don't have any DNA yet.

Galerina aff stylifera - one sequence from OR and three from BC are sister to G. stylifera var. velosa and EU sequences called G. sideroides (next). Landry is calling it G. aff. stylifera.

Galerina sideroides EU - reported from the PNW but all the EU sequences are very close to G. stylifera var. velosa. Unless everybody is identifying them backwards. This DNA has not yet been found in the PNW, although the species is reported from here.

Galerina cf stylifera (aff sideroides) - we have a dozen and a half sequences that were identified as G. stylifera but which Landry is calling G. aff sideroides. One collection called Galerina pallidispora WA sequenced to be this as well.

The bottom line is we need reliable sequences of both G. stylifera and G. sideroides to proceed. This section is currently very confusing.

Galerina pseudocamerina MI, an illegal name, has collections that sequence in Naucoriopsis even though Smith keys it out here.

Galerina cf mammilata © A and O Ceska,     G. stylifera var. velosa © Danny Miller,     G. cf stylifera (aff. sideroides) © Danny Miller

 

The following, many described from the PNW and therefore here by definition, do not have any genetic data yet (some may be duplicates).

Galerina agloea WA -
Galerina borealis ID -
Galerina castanescens ID -
Galerina fuscobrunnea OR -
Galerina occidentalis WA -
Galerina pallidispora WA - our only sequence of this turned out to be the same as G. cf stylifera (aff. sideroides).
Galerina pseudostylifera ID -
Galerina subbadia MI -

 

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