Tubaria, Flammulaster and Phaeomarasmius

© Christian Schwarz

Danny’s DNA Discoveries – Tubaria, Flammulaster and Phaeomarasmius of the PNW
by Danny Miller

Click here for my Pictorial Key to Tubaria and Flammulaster and Phaeomarasmius

Introduction

These three related genera definitely qualify as LBMs. They are usually little (<2.5 cm across), brown spored and usually brown overall, except for some striking red Tubarias.

Tubaria - hopefully recognized by slightly decurrent gills, and by being the most hygrophanous mushrooms I've ever seen, with a striking colour change from dark when wet to very pale when dry (check out the photo at the top left for the extreme range of colours). Otherwise they can be hard to recognize without learning them all, especially when they get >2.5 cm across and aren't as little as you expect an LBM to be. Some are abundantly common. A couple of bright red Tubarias are the most easily recognized.

Flammulaster/Phaeomarasmius - stand out from most other LBMs by usually being granular or floccose scaly over the entire fruitbody. They are rare enough that we had not noticed any reports of Flammulaster from the PNW until I found one at a Key Council meeting in 2012, and we started looking into it further. The ones found on wood are probably most easily confused with the scaly Pholiota, especially the small, pale capped Pholiota scamba, but most others are usually larger and those all have viscid caps (everything in this family is dry capped).

However, there does not appear to be a real genetic distinction between Flammulaster and Phaeomarasmius and in order to avoid making a bunch of new, tiny, obscure genera, it may be best to think of both of them as Phaeomarasmius, the older genus. The famous Moncalvo paper "117 Clades of Agarics" only had one Flammulaster and one Phaeomarasmius sequence, so they couldn't tell that those two genera weren't distinct. More recent papers like Matheny's "Taxonomy of Displaced Species of Tubaria" had more sequences and showed that the Flammulaster and Phaeomarasmius sequences intermingle, and we're not sure how many total clades there will be. Indeed, although they clearly separate from Tubaria, it may be necessary in the future to consider all three of them as Tubaria to avoid needing new genera to avoid making a combined Phaeomarasmius paraphyletic - we'll have to see. But for now, let's consider them as two genetic groups - Tubaria without true scales, and Phaeomarasmius/Flammulaster with scales all over (granular or squarrose).

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:

Tubaria furfuracea and hiemalis may be distinct but can hybridize easily, so maybe not.

Our local species of Tubaria conspersa and Tubaria confragosa may need new names.

Phaeomarasmius and Flammulaster are not distinct genera, and they may need to all be combined into Phaeomarasmius or split into several additional genera.

We have more than a half dozen undescribed Phaeomarasmius/Flammulaster in the PNW.

Phaeomarasmius/Flammulaster - click to expand
Granular or floccose scales all over, with dry caps. Although genetically it may be best to think of them all as Phaeomarasmius for now, I will talk about them using the name they were first given. Traditionally, Phaeomarasmius was separated by having tougher fruitbodies that don't decay as easily as Flammulaster, and by a bark habitat and lack of a germ pore on the spores. Also, Flammulaster needn't be found directly on wood. But none of these differences may be genetically relevant.
Species mentioned: Phaeomarasmius erinaceus, erinaceellus, proximans. Flammulaster carpophilus, rhombosporus, muricatus, granulosus, limulatus.

Flammulaster cf carpophilus group - we have a lot of sequences from the EU of this EU mushroom, and it doesn't appear to occur here, but we do have 3 genetic species that seem somewhat similar and are somewhat closely related to it. One of them was studied microscopically and was a good microscopic match too (sp. 5). Possible defining macroscopic characters are that the colour is paler than some other species, and the scales are granular. One of these species could be Flammulaster rhombosporus NY, an east coast mushroom with no sequences yet that is currently regarded as a synonym but is probably a distinct species and more likely to be what is found here than F. carpophilus.

Flammulaster sp. 5 © A & O Ceska, F. sp. NAMA2014 © NAMA and the Field Museum of Natural History, F. sp. BTSP © Daniel Winkler

Flammulaster cf muricatus group - these might clade close to the EU species F. muricatus, if one EU sequence is to be believed, but that should be taken with a grain of salt. What we do know is that sp. 2 was a good microscopic match to F. muricatus, and sp. 3 is a sister species that was a good microscopic match to Flammulaster granulosus. However, if a couple of EU sequences are to be believed, both are closer genetically to F. muricatus than they are to F. granulosus. When we first started finding Flammulaster species in the PNW, we called them all F. granulosus, as that was the only species we were familiar with, but there's no evidence yet that it actually exists in the PNW.

Flammulaster sp. 2 © A & O Ceska, F. sp. 3 © A & O Ceska

Phaeomarasmius cf erinaceus EU - this seems to be by far the darkest, most densely scaly species we have. we don't have any local sequences yet to see if our species is really this EU species, or the similar east coast Phaeomarasmius erinaceellus NY (or neither). We need sequences. It has been found in CA and OR infrequently and reportedly from WA and BC as well.

Phaeomarasmius cf erinaceus © Leah Bendlin

Phaeomarasmius proximans MI - our sequences match more than a dozen ENA sequences of this, so it seems very likely we do have this species here. It was unknown from the PNW until two recent WA collections. It is found on wood. The first collection had a cap covered in tiny, dark scales with a yellow-brown ground colour, but wasn't as prominently scaly on the stem. The second collection had more red-brown scales on the cap , a somewhat scaly stem and a stem bulb.

Phaeomarasmius proximans © Lauren Ré and Yi-Min Wang

Flammulaster sp. 1 (aff F. limulatus/P. proximans) - this species is 5% different from Phaeomarasmius proximans MI and 3% different from Flammulaster limulatus EU, and microscopically matched Flammulaster limulatus var. litus pretty well. It was sequenced once from OR and once from WA. At least from the one photo we have, it is a brighter orange brown than the others.

Flammulaster/Phaeomarasmius sp. 1 © A & O Ceska

Tubaria - click to expand
No granular nor floccose scales all over. Often decurrent gills. Dry caps that are often extremely hygrophanous.
Species mentioned: Tubaria furfuracea, hiemalis, tenuis, abramsii, conspersa, confragosa, punicea, vinicolor.

Tubaria furfuracea EU/hiemalis EU (tenuis CA?) - at first glance, there do appear to be two species, differing by a dozen bp in ITS, and judging by the names usually attached to the EU sequences, one is T. furfuracea and the other is probably T. hiemalis. Some have synonymized them, but sequencing in the EU appears to support that there are two species. Over a dozen sequences of each from the PNW have been found, showing that both are abundant here. However, telling them apart is difficult. T. furfuracea is supposedly a summer species found on soil with a strong white partial veil. T. hiemalis is supposedly a winter species found on wood with a weak ochre partial veil. But our sequenced collections don't seem to always bear that out, so more study will have to be done to determine the real way to tell them apart.

Another interesting observation is that there are very different looking kinds of collections - some almost brick red with lots of veil tissue hanging off the cap margin, and others pale tan/buff with seemingly thinner flesh and striate cap margin. Both will sequence the same. Now, Tubaria furfuracea is probably the most hygrophanous species I have ever seen, so you can expect drastic colour changes as it dries, but the pale, thin collections always seem to be fresh when found and I can't say for sure that there aren't distinct colour forms. Compare photos below.

One complication is that some think that the older California name Tubaria tenuis is a synonym of the newer, but more popular name Tubaria hiemalis. If so, that's the name that should be used for it. However, with all the confusion around these species, that's just a guess for now, as we have no sequences of it. If they are all the same species, T. furfuracea is still the oldest name of the three.

Unfortunately the story is further complicated by the fact that these species appear to hybridize commonly. I have three sequences with 4, 8 and 12 ambiguous locations respectively that sequence in between the two species, the latter sequence having ambiguous nucleotides in virtually all 12 of the places that the two species differ, making it a complete hybrid of equal parts both species. This was an important find. If they can mate with each other that easily to produce hybrids, and if the hybrids are shown to be able to reproduce, and if the ecological and morphological differences that supposedly exist are shown to not be cut and dried, then it may not be a bad idea to think of it as one variable species, using the oldest name, Tubaria furfuracea.

Tubaria hiemalis (thin, pale version) © Daniel Winkler, Tubaria furfuracea (dark version with veil remnants) © Joe Miller

Tubaria cf abramsii CA - this one collection from OR still had a sequence 6 bp different from both previous species, even after all 20 ambiguous locations were accounted for, so it may be a distinct species. If so, one species it somewhat resembles is the California species T. abramsii, but we have no sequences of that to be able to tell for sure.

Tubaria cf abramsii © Michael Lainoff

Tubaria aff. conspersa - our local species is >4% different in ITS than the EU species Tubaria conspersa, so it likely needs a new name. We have 4 sequences from WA and OR, and one of them is 5 bp different than the others (which are within 2 bp of each other) so it should be verified that there is only one species here. It is recognized by its even smaller size and fibrils all over the cap and stem, not true scales like Phaeomarasmius/Flammulaster, just fibrils. The cap margin will be somewhat ragged with veil material.

Tubaria aff. conspersa © Buck McAdoo

Tubaria aff. confragosa - similarly, our local species is about 3.5% different in ITS than the EU species Tubaria confragosa, and may need its own name. Smith described Pholiota canescens as being very similar and that species may well actually be a Tubaria, perhaps this Tubaria, but unfortunately the name Tubaria canescens is already taken for an east coast species, so we cannot use that epithet for our local species no matter what. We have 3 WA sequences. Found on wood, it usually has a veil that leaves a ring on the stem.

probable Tubaria aff. confragosa © Buck McAdoo

Tubaria punicea OR - this bright burgundy mushroom is found on Madrone. We have sequenced it from CA and BC.

Tubaria punicea © Noah Siegel

Tubaria vinicolor CA - this lookalike is not always quite as bright red, especially in the stem, and is found on other kinds of wood. We have sequences from CA and WA.

Tubaria vinicolor © Christian Schwarz

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