© Steve Trudell
Danny’s DNA Discoveries – Galerina s.l.
by Danny Miller
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Danny’s DNA Discoveries – Galerina s.l.
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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.
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).
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. |
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Summary of Interesting Results
Here are some of the newest, most interesting results of the study:
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Galerina This mycenoid moss-dwelling group of species are small, with caps usually <1cm across, with stems that darken from the base up, two features that differentiate it from the Galerina semilanceata group. Galerina 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 vittiformis forma bispora WA - 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. That was a guess. I do know that this local forma bispora is found in the PNW as it was described from here and we have recent matching sequences, but I don't know what Galerina vittiformis itself is or if it occurs here. This means that our form name might need promoting to species level. Some sequences differ from others by as much as 4 bp, so we may have more than one form or variety here. Galerina 'vittiformis PNW10' - We also have an even more abundant sister species he called "aff vittiformis sp. 3" in his 2020 paper (a different number than he used in 2019). This has been sequenced a couple dozen times. Also, see the next species. Spores ~10-11 x 5.5-6.5u and ~8.5-11 x 5-6u in another collection. Galerina 'vittiformis PNW11' - a deeper brown in the one collection we have photographed. This is "aff vittiformis sp. 2" in his 2020 toxins paper, where the clades are named differently than in his original 2019 paper. Galerina 'perplexa PNW13' - 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 MI and no sequences are from ENA. 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 - a Greenland sequence 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 vittiformis f. bispora © Bitty Roy, G. 'vittiformis PNW10' © Daniel Winkler, G. 'vittiformis PNW11' © Connor Dooley
Galerina 'perplexa PNW13' © Yi-Min Wang (2 images)
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 - 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, an ID sequence and a WA 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 - one OR collection matches a number of EU sequences very well. Galerina subcerina ME (=G. calyptrata UK) - is a rather bright orange-brown mycenoid moss species like G. vittiformis and G. semilanceata best differentiated microscopically. 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 and one WA collection that match. It was never recorded by Smith from the PNW, but here it is. It is another section Calyptrospora species. Galerina 'luteolosperma PWN14' - I have no idea why Landry calls these Victoria BC sequences by this name, there are no MI 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 lubrica CO - we have the ITS1 type sequence and matching sequences that provide ITS2. It had been reported from BC, but now we have a sequenced collection from WA to prove it is here. Galerina PNW12 - close to G. lubrica. One WA collection. Galerina allospora MI - one OR collection has a sequence matching EU and Chinese sequences, but we don't have any MI type area sequences yet to prove that it's the same species worldwide. For now, I'm assuming it is. Smith had trouble placing this in the correct section of Mycenopsis. He finally chose Sideroides (which are going to be split into genus Sideroides) but it actually belongs here with the core of Mycenopsis, where he also keyed it out. It was found in sphagnum, where Sideroides are not usually found. Galerina PNW15 - one WA collection Galerina PNW18 - one of the little Mycena-like species in moss with few gills. It is known from only one WA collection.
Galerina pumila var. subalpina © Bitty Roy, G. fallax © Yi-Min Wang (2 images), G. paludosa © Connor Dooley, G. subcerina © Yi-Min Wang Galerina lubrica © Yi-Min Wang (2 images), G. PNW12 © Yi-Min Wang (2 images), G. allospora © Connor Dooley
Galerina PNW15 © Yi-Min Wang, G. PNW18 © Anne Polyakov
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 acicola OR - Calyptrospora Galerina insignis WA - Inoderma
Tubariopsis Galerina semilanceata WA (=G. dimorphocystis MI?) - this mycenoid moss species has caps that will often reach >1cm across and stems that don't darken, which contrasts it with the similar Galerina vittiformis group. I suspect that the east coast species G. dimorphocystis is the same, although it is described without a veil and G. semilanceata is described with a very fleeting veil. We need sequences of very young collections without a veil and/or east coast type area sequences of G. dimorphocystis to prove the synonymy. Brandon declared them probably synonymous, but all of his collections of G. dimorphocystis were from the PNW, not back east, and they do not appear to have been identified on the basis of a lack of a veil. The older of the two names is G. semilanceata, although Brandon mistakenly thought that G. dimorphocystis was older because G. semilanceata went by Galera until 1964. The spores are 8-10x5-6u. G. heterocystis Jamaica (=G. clavata EU?) - the spores are much longer (11-17x6.5-8.5u) than in G. semilanceata, and at least here in the PNW, this species has few complete gills, and oddly short tiers of sub gills. Landry synonymizes G. heterocystis with G. dimorphocystis, but that is incorrect as he had no type area collections, only PNW collections, which were misidentified. They were called G. heterocystis on the basis of apparently lacking pileocystidia, ignoring the fact that the spores were way too small for G. heterocystis. Many people think that G. clavata is the same thing as G. heterocystis, but we can't prove it yet without Jamaican type area sequences of G. heterocystis. I can tell you that our sequences do match EU sequences of G. clavata, and right now I am assuming that the older G. heterocystis is in fact a synonym because G. clavata is not restricted to the EU, so it could conceivably be in Jamaica. If not, we should call our collections G. clavata. Galerina nigripes OR - the often blackening stem base is far from unique in this species. We may have the type sequence, and we do have several recent PNW sequences. Galerina tibiicystis MD - Smith placed this in a different section, but it clades inside Tubariopsis. We have a few EU sequences, and one local OR sequence that matches them, but no ENA type area sequences to prove that's what we have. Galerina hybrida EU - found 2 years in a row in the same montane region in OR in sphagnum moss, one year with literally thousands of them growing at once. Smith did not find this species here, but our sequences match several EU type area sequences except for 2-3 bp and 1 chunk of indel. My ITS trees place it here in Tubariopsis near G. tibiicystis. Galerina PNW19 - two OR collections.
Galerina semilanceata © Steve Ness, G. heterocystis © Yi-Min Wang (2 images), G. nigripes © Yi-Min Wang, G. tibiicystis © Connor Dooley
Galerina hybrida © Connor Dooley (2 images), G. PNW19 © Jonathan Frank
Galerina brunneomarginata CA (as G. brunneimarginata) -
Naucoriopsis First, the deadly species containing alpha- and beta-amanitin, like the deadly Amanita. They clade tightly together genetically. Galerina marginata EU (=G. venenata OR, =G. cinnamomea OR, =G. unicolor EU) - usually a distinct ring or ring zone, usually on conifers. 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?) - differs from G. marginata by supposedly a weaker ring zone and growth usually on hardwoods. 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 it will be difficult to separate macroscopically from G. marginata. 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. G. castaneipes may have slightly smaller spores than G. marginata, but perhaps not small enough to reliably differentiate them that way. Galerina 'marginata PNW07' - one collection labeled G. marginata has a unique sequence, differing by 13bp and 2 indels, that Landry calls G. aff. marginata sp. 4. It is probably not G. oregonensis, but we do need to see if we ever see this sequence again and if so, figure out what it is.
Galerina marginata © A and O Ceska, Galerina castaneipes © Danny Miller and Yi-Min Wang
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-amanitin, 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 CA02 - this is close enough to G. marginata and G. badipes that it should be tested to see if it is deadly poisonous. It could have alpha- or gamma-amanitin. It is known north of CA from one WA collection. Galerina PNW02 - from moss covered swamp muck, gills somewhat widely spaced, stem somewhat clavate. This too is close to G. marginata and G. badipes and should be tested for amatoxins.
Galerina badipes © Danny Miller and Yi-Min Wang, G. CA02 © Yi-Min Wang (2 images), G. PNW02 © Buck McAdoo
Galerina pseudocamerina MI (var. fulvovelosa WA) - keys out to section Sideroides, but sequences here in Naucoriopsis, if we can believe the sequences. We only have sequences from the EU, not from ENA. One WA sequence is about 5 bp different from the EU sequences, but another WA sequence has some ambiguous locations and may only be 3bp different. If it is correctly identified, the fact that WNA and EU sequences are close to each other tells me perhaps ENA sequences will be similar too, and that we do have this species here. If ours is different enough to deserve its own variety, we do have a local variety name to use, var. fulvovelosa. G. pseudocamerina was not properly described, but nobody seems to care as the name is still being used. 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 'nana PNW03' EU - keys out to section Inocyboides. Our local sequences are a few % different from the one EU sequence we have, so ours seems to be a distinct sister species. We need more reliable EU sequences to tell. Galerina 'nana PNW04' - one sequence is unique, from a lost collection without a photo. It 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 'pruinatipes PNW05' WA - found once, a sister species >4% different in ITS from G. pruinatipes. Galerina 'triscopa PWN06' EU - keys out to section Mycenopsis subsection Tibiicystidiae stirps Triscopa. EU sequences at first seemed to be 4% different than the one BC sequence we have, implying ours may be a sister species. And there were 2 clades of local sequences about 2% apart. But then we found WA collections with many ambiguous locations that were a hybrid of our 2 clades, and showing that except for the known ambiguous locations, it was <2% different than both the EU and BC sequence. We do have a local form described but we don't know its genetics: G. triscopa forma longicystis from Idaho. However both that and the type variety are reported from the PNW, and it seems our varieties can hybridize.
Galerina pseudocamerina © Daniel Winkler, G. 'nana PNW03' © A and O Ceska, G. 'pruinatipes PNW05' © Jacob Kalichman, G. 'triscopa PNW06' © Yi-Min Wang
Galerina 'Naucoriopsis PNW01' - we haven't investigated what species this sequence matches up with. By the appearance of the roughened gill edges, it might have gigantic cystidia. This and the next species could be one of the little understood species listed below.
Galerina 'Naucoriopsis PNW01' © Yi-Min Wang (2 images)
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 - Naucoriopsis Galerina camarinoides OR (as camerinoides) - Mycenopsis subsection Tibiicystidiae stirps Triscopa Galerina olympiana WA - Physocystis
Galerina stagnina EU - Porospora (this is actually Phaeogalera
stagnina).
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 mammillata WA - Brandon Landry provided purported sequences of this local species, and I agree all collections with matching ITS match the description well. 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 the type sequence, but collections with this name are turning out to be var. velosa, so it's possible this doesn't deserve distinct varietal status from an ITS perspective either. Galerina 'stylifera PNW08' - 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 'sideroides PNW09' - 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. Galerina 'sideroides PNW16' - one WA collection is a sister species to PNW08 and PNW09. Galerina 'sideroides PNW17' - another WA collection has a unique sequence. 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 - has collections that sequence in Naucoriopsis even though Smith keys it out here, so it is discussed above under Naucoriopsis.
Galerina mammillata © A and O Ceska, G. stylifera var. velosa © Danny Miller, G. 'stylifera PNW08' © Yi-Min Wang, G. 'sideroides PNW09' © Danny Miller, G. 'sideroides PNW16' © Yi-Min Wang
Galerina 'sideroides PNW17' © Cady Adair (2 images)
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 - |
