Cyphellaceae s.l.

© Kent Brothers

Danny’s DNA Discoveries – Cyphellaceae s.l. of the PNW (Marasmiineae)
by Danny Miller

Click here for my Pictorial Key to the pleurotoid and mycenoid statures, where many of these species are found.

Introduction

The Cyphellaceae s.l. include some more primitive shaped crust and cup mushrooms growing on wood (usually cups are ascos, not basidios), as well as many gilled mushrooms of pleurotoid (eccentric stems on wood) and mycenoid (conical on the ground or wood) stature (as well as a few collybioids).

Many other genera in many other families meet this family's identification criteria as well. Unfortunately, there's not much rhyme nor reason to identifying the vast multitude of miscellaneous white spored mushrooms to family, as many mushrooms in different families and even sub-orders lack distinctive traits, so they have to be learned individually.

Although an ITS tree might show these genera intermingling with the Porotheleaceae, an LSU tree in a 2022 study shows the two families as separate. They are at least close sister families to each other, and I don't think we've heard the final word on if they should be kept separate or combined into one big family.

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:

tbd

Crust and Cups (Chondrostereum, Calyptella, Cyphella, Henningsomyces, Rectipilus) - click to expand
Crust - Chondrostereum has a reflexed cap and a purple underside.
Cup - Calyptella - tiny white cups (several mm across) with a short stem on wood. 'Hemimycena 3' is closely related.
            Cyphella - tiny white stemless cups on wood, perhaps with a hairy margin.
            Henningsomyces is a cluster of minute (<1 mm across) white tubes growing on wood. A few Henningsomyces sequences show up in a different family, so it's possible that this genus needs to be split, but it looks like our species are true Henningsomyces and will not be affected. See also Rectipilus, next.
            Rectipilus are similar, little known minute white tube/cups with hairs that are easier to see. This is another controversial genus. Sequences that may be of the type species of Rectipilus and sequences that may be our local species, Rectipilus idahoensis, mix in with Henningsomyces sequences, so Rectipilus may not deserve its own genus and those species may be moved to Henningsomyces. Other species of Rectipilus, though, may need moving to a new genus, like other species of Henningsomyces. This still needs to be sorted out with type sequences. See this paper.
Most cups are Ascomycota. The only other Basidiomycota cups are found in the Niaceae.
Species mentioned: Chondrostereum purpureum. Henningsomyces candidus, puber. Calyptella capula. Rectipilus idahoensis. Cyphella marginata.

Chondrostereum purpureum EU - a crust with a reflexed cap that is purple underneath when fresh. One collection exuded red droplets, which has not been noticed before. Keep an eye out for that. WA and OR sequences match hundreds of EU and worldwide sequences quite well.
Cyphella cf marginata Australia - minute yellow cups <1 mm across with brown hairs around the rim on fruit tree twigs. This was described in Maireina, in the Niaceae, but is generally accepted in Cyphella. No DNA from anywhere.
Calyptella cf capula EU - tiny white cups (several mm across) with a short stem on wood. There are 2 competing sequences quite distant from each other for what this species is. We have no local sequences to compare yet.
Henningsomyces cf candidus EU - a cluster of minute white tubes (<1 mm across). Three EU sequences and a sequence from eastern NA and South America are 5 different genetic species so we don't know what the real species is or what to call ours. We don't have any local sequences yet.
Henningomyces cf puber EU - similar, with very tiny hairs all over that you probably need a dissecting scope to see. Two EU sequences are different genetic species so once again we don't know what this species really is nor what the few BC reports of it really are. We don't have any local sequences.
Rectipilus idahoensis ID - similar, little known minute white tube/cups with hairs. We have a French sequence purporting to be this, but who knows it that's true. It might be inside the correct genus. We need local collections.

unsequenced Chondrostereum purpureum © Kit Scates Barnhart and Danny Miller,     unsequenced Calyptella cf. capula © A and O Ceska,     unsequenced Henningsomyces cf. candidus © A and O Ceska

Pleurotoid (Cheimonophyllum and Scytinotus) - click to expand
These are the pleurotoid species in the family, on hardwoods with a short, stubby lateral stem.
Cheimonophyllum is a tiny, dry, chalky white oyster with hyphoid cheilocystidia.
Scytinotus (formerly in Pleurotopsis and Panellus) are small, sometimes pinkish oysters that may be viscid, with amyloid spores, not readily told apart from Panellus.
Species mentioned: Cheimonophyllum candidissimum, haedinum. Scytinotus longinquus, ringens. Pleurotopsis longinqua. Panellus ringens.

Cheimonophyllum candidissimum EU - a tiny, dry, chalky white oyster on hardwoods usually with a short, stubby lateral stem. Our AK and WA sequences match 100% with a few EU sequences, except one EU sequence has a very long indel. A recent paper says that this is a newer synonym of Cheimonophyllum haedinum from Cuba, but they did not obtain any sequences from Cuba, just one from Europe to justify that, so I await better evidence.
Scytinotus longinquus Cape Horn (='Pleurotopsis' longinqua, ='Panellus' longinquus) - viscid, white when young to dark pinkish tan when old (spotting pinkish brown in the interim) with a definite eccentric to lateral stem on hardwoods. Our BC, WA and OR var. pacifica sequences have the exact same ITS sequence as Chilean var. longinquus so it may or may not deserve varietal status. 'Mitellus' mitis is not viscid and the cap and gill edges peel.
Scytinotus ringens EU (='Panellus' ringens) - dry, rose-grey cap, almost stemless on hardwoods. We have EU sequences and an ENA sequence that match fairly well, with some ambiguities but no local sequences to confirm that our couple of reports of it are true.

Cheimonophyllum candidissimum © Ed Barge, Scytinotus longinquus (unsequenced young and sequenced old) © Fred Rhoades and Danny Miller

Atheniella - click to expand
Atheniella and the very difficult to differentiate Phloeomana, below, were both separated from Mycena on the basis of having inamyloid spores (and simple cystidia). Atheniella have long been thought of as colourful, but the all white former 'Hemimycena' delectabilis is here now too, with well developed cheilocystidia and no pigment, so we now know they don't all have simple cystidia nor are they all colourful. Conversely, the colourful inamyloid 'Mycena' acicula seems like it should belong here, but it is closer to the currently all-white genus Hemimycena. Hemimycena and its segregate genera are going to be difficult to differentiate as well.
Species mentioned: Atheniella adonis, amabilissima, fusipes, flavoalba, aurantiidisca. Hemimycena delectabilis.

Atheniella 'adonis PNW01' (=Mycena fusipes WA?) - scarlet cap that does not fade to yellow (but may fade to white). White to pale pink stem. There are at least 2 competing concepts for A. adonis in the EU. Our one WA sequence averages 6 bp and 5 indels different than one of them, so I'm going to assume that even if that ends up being the real one, ours may need a new name or varietal status. Back east they have the name A. amabilissima, thought to be a synonym but probably not. There are 3 genetic species passing as A. adonis/amabilissima back east, but none of them match our sequences, so we cannot use that name. We do have a locally described species, Mycena fusipes, also though to be a synonym of A. adonis, but as it looks like our species is unique perhaps the name Atheniella fusipes should be used for it. We need a type sequence to prove it, though. It is also possible that 'Mycena' fusipes could refer to Atheniella 'flavoalba PNW02' or Atheniella PNW03.
Atheniella 'flavoalba PNW02' - pink cap that fades to yellow and then to white. White stem. There are several competing concepts for A. flavoalba in the EU, and ours is quite distinct from all of them and definitely needs its own name. In the EU, pink colours are rarely found on fresh fruitbodies, but our species seems to often start out pink. Perhaps there are more species here. We should sequence collections that start out yellow and never have any pink and see if they are the same thing.
Atheniella PNW03 - found on sedges in a fen. This delicate pale pink capped Atheniella with a white stem has fewer gills than the above two species. It can fade entirely to white and then be mistaken for Hemimycena.

Atheniella 'adonis PNW01' © NAMA and the Field Museum of Natural History, A. 'flavoalba PNW02' © iNaturalist user lcanedo, A. PNW03 © Connor Dooley (2 images)

Atheniella aurantiidisca OR - tri-toned orange cap (dark orange disc, orange-yellow in the middle and pale yellow rim). Yellowish stem. The cap might be mostly bright yellow, without much orange, but no other species is bright yellow.
Atheniella cf delectabilis NY (='Hemimycena' delectabilis) - pure white, decurrent gills, prominent bleach odor. Larger than many other 'Hemimycena' which are usually <1 cm across. A bleach odor was detected in Hemimycena aff. lactea below, complicating how to recognize this species. We don't have NY sequences where it was described. West coast WA and CA sequences are 2% different than a couple of EU sequences, so we need east coast sequences to confirm that we have the actual species here.

Atheniella aurantiidisca © Kent Brothers, Atheniella delectabilis © Fred Rhoades

Hemimycena s.l. - click to expand
Very small mycenoids (after all, the name does mean "half of a Mycena") with inamyloid spores, supposedly all white. These do not include some all white mushrooms that are real Mycenas, which have either a basal disc at the base of the stem or a thorny/hairy/sugary appearance (and amyloid spores). This genus needs to be split and all of the genera will be difficult to distinguish from the other genera that were split from Mycena for having inamyloid spores, including Atheniella and Phloeomana.
'Mycena' acicula, one of the smallest colourful Mycenas with inamyloid spores, belongs near the true Hemimycena and is treated here, although it is not yet certain if it will get its own genus or end up in Hemimycena.
'Hemimycena' delectabilis was moved to Atheniella.
Other 'Hemimycena' need moving to either new genera and/or perhaps an expanded Phloeomana and/or Calyptella. Many others are still unknown.
Mycenella needs moving into Hemimycena s.s. to avoid splitting Hemimycena. It is covered below.
Species mentioned: Hemimycena albicolor, albidula, cyphelloides, delectabilis, gracilis (Helotium immaculata, Omphalia papillata), hirsuta, lactea (delicatella), leucophaea, mauretanica, nebulophila, pseudocrispula, subimmaculata (albissima, macmurphyi), substellata, tortuosa. Atheniella delectabilis.

Hemimycena1 - these belong in Hemimycena, although it either needs to be split, or Mycenella and Lactocollybia sect. Albidae need to move to Hemimycena (more likely).
Hemimycena lactea EU - on debris, a local collection from Vancouver Island BC matches a bunch of EU sequences.
Hemimycena 'lactea PNW04' - one OR and one WA sequence are 2% different, the OR collection with a bleach odor like Atheniella delectabilis so this needs further study to distinguish it from that species. This could potentially be one of the unknown species below. Spores 6.5-7 x 2.5-4u. One short ITS2 sequence is 2 bp different than the other two full ITS sequences of this species.
Hemimycena 'lactea PNW07' - one lone WA sequence is 6 bp or so different from every PNW04 sequence (even the short one), so I'm giving it its own code for now. It's possible that PNW04 and PNW07 are just H. lactea, if that species has variable ITS.

Hemimycena 'lactea PNW04' © Michael Beug and iNaturalist user cinthomas,     H. 'lactea PNW07' © Yi-Min Wang

These Hemimycena species would need a new genus if Hemimcyena was split. Otherwise, 'Mycena' acicula group and Mycenella belong in Hemimycena too.
Hemimycena tortuosa UK - on wood and bark, with unique tortuously spiraled cystidia. A couple of WA sequences match a couple of UK sequences.
Hemimycena CA01 - It is not yet known which of the species reported from here (listed below) this might be, but it is interesting in that it isn't always white, it can be pale orange.

Hemimycena tortuosa macro and micro © Fred Rhoades,     H. CA01 © Yi-Min Wang and Christian Schwarz

'Mycena' acicula PNW01/PNW021 - on debris, two sister species 3% apart in ITS occur here. We have no EU sequences to tell which species (if either) is the real thing. PNW01 was found twice in WA, and has a microscopic workup. PNW02 was found in OR and CA. We need a microscopic workup to compare to PNW01. They need new names in Hemimycena, unless Hemimycena is to be split.

'Mycena' 'acicula PNW01' © Fred Rhoades,     'Mycena' 'acicula PNW02' © Jonathan Frank

Hemimycena2 (Phloeomana?) - the following probably all belong in an expanded concept of Phloeomana along with 'Mycena' oregonensis, but it's also possible somebody will want to give them both their own genera.
'Hemimycena2' PNW06 - found in WA and CA, it has few gills like species of Hemimycena3.
'Hemimycena' albicolor OR - we have the type sequence, but no recent collections nor photos.
'Hemimycena' cf. gracilis EU - one CO sequence of a collection resembling H. gracilis may represent this EU species, but as it's from nowhere near the type area and as they are hard to identify, this doesn't yet prove anything. It is still only a theory that H. gracilis may belong here.
uncultured 'Hemimycena' OR - one environmental sequence is so far unique. It may be one of the unknown species below, assuming it actually fruits here.

'Hemimycena2' PNW06 © Yi-Min Wang

Hemimycena3 - these species are sister to Calyptella and may have fewer and/or more rudimentary gills than the other genera (although 'Hemimycena2' PNW06 has few gills too), in other words a kind of missing link between other Hemimycena s.l. with well developed gills and Calyptella without gills and with inverted stem (coming out of the top of the cap). It could belong in an expanded definition of Calyptella, but then that genus would include both gilled mushrooms and cups, so it is more likely that Hemimycena3 will get its own genus name.
'Hemimycena3' cf. crispula EU - one CA collection was a micro match to H. crispula (which is thought to be a synonym of Hemimycena hirsuta, a species with hairs all over the cap and stem, and no gills whatsoever - it's just empty under the cap. That does not mean it was H. crispula/hirsuta, but it could mean that species belongs in this genus. The lack of gills certainly supports this theory.
'Hemimycena3' PNW02 - with few gills (but well defined). Hemimycena pseudocrispula EU matches this species very well, and could be this. Sequenced from WA and OR.
'Hemimycena3' PNW03 - with just a few rudimentary gills (not fully formed). Our one WA collection is 2% different in ITS from both of the above.

'Hemimycena3' PNW02 © Connor Dooley and Jacob Kalichman,     'Hemimycena3' PNW03 © Yi-Min Wang

Hemimycena4 (Phloeomana?) - this also probalby belongs in an expanded concept of Phloeomana, or perhaps by itself. We have one ITS sequence that appears to be in a distinct genus.
'Hemimycena4' PNW05 - found once in WA, a dainty long stemmed species with adnate to decurrent distant gills.

Hemimycena4 PNW05 © Yi-Min Wang

Unknown - we need both type area and local sequences of all of these to confirm reports that the following occur in the PNW, and to figure out what genus they actually belong in. Hopefully they all fit in one of the above and don't require even more new genera.
Hemimycena albidula NY

Hemimycena cyphelloides EU

Hemimycena gracilis EU (=H. immaculata NY?, =Mycena papillata NY?) - these synonymies need confirmation.

Hemimycena hirsuta EU - mentioned in Hemimycena3.

Hemimycena lactea EU (=H. delicatella NY) - these synonymies need confirmation.

Hemimycena leucophaea WA - described from here and occurs here by definition. But what is it really?

Hemimycena mauretanica EU

Hemimycena nebulophila BC - described from here and occurs here by definition. But what is it really?

Hemimycena pseudocrispula EU - probably 'Hemimycena3' PNW02.

Hemimycena subimmaculata WA (=Mycena albissima CA?, =Mycena macmurphyi CA?) - also here for sure, and these synonymies need confirmation.

Hemimycena substellata EU

Phloeomana, Mycena oregonensis and Mycenella - click to expand
Phloeomana was also separated from Mycena on the basis of inamyloid spores and simple cystidia, just like Atheniella, but are genetically different. (The various Hemimycena genera are difficult to distinguish as well). Our species is recognized by a pale grey pleated cap, a few decurrent gills and hairs at the base of a stem that may have a hint of yellow.
Hemimycena2, Hemimycena4 are near Phloeomana, and may be included in an expanded definition of Phloeomana.
'Mycena' oregonensis is also near Phloeomana, and may be included in an expanded definition of Phloeomana. It is entirely orange with orange marginate gills.
'Mycenella' has warty inamyloid spores (like Gamundia), and is hoary/pruinose over the entire fruitbody. Our species has a rooting stem and red staining gills. It is inside Hemimycena, so either that needs to be split (probably not) or Mycenella will be moved into an expanded definition of Hemimycena.
Species mentioned: Phloeomana speirea, alba, hiemalis. Mycena oregonensis. Mycenella nodulosa.

Phloeomana spierea EU - a very pleated capped 'Mycena' with decurrent gills often found on bark. we have WA, OR and CA sequences that match dozens of EU sequences and other sequences worldwide.
Phloeomana alba EU - not known from the PNW until found by DNA. One OR sequence (and one close CO sequence), matching a couple of type area EU sequences. The older P. hiemalis, according to one lone EU sequence, may be similar or the same, so that should be investigated.
Phloeomana minutula NY - not known from the PNW until a BC collection was sequenced. It is a small, white species likely to be mistaken for Hemimycena.
'Mycena' oregonensis OR - a pretty, entirely orange 'Mycena' with orange marginate gill edges. This may belong in an expanded Phloeomana.

Phloeomana spierea © Bitty Roy, Phloeomana alba © Heather Dawson, Phloeomana minutula © Stephen Russell, 'Mycena' oregonensis © Steve Trudell

'Mycenella' nodulsa WA - the round warty inamyloid spores and entirely hoary/pruinose fruitbody probably means this was placed in the correct genus but we have no local sequences to prove it. We need some. Our species also has a rooting stem and red staining gills. As stated above, Mycenella may be moved into Hemimycena.
'Mycenella' margaritispora EU - differs microscopically. One UK sequence may be this, but we have no local sequences yet.

Mycopan - click to expand
Hard to separate from Mycena, also possessing amyloid spores. Hydropus, which was split from Mycena for often possessing a different cap cuticle, was split again when some species turned out to belong in this family instead of with the other Hydropus. 'Hydropus' pseudotenax (similar to Mycopan scabripes) and 'Mycena' quinialtensis (a dark, viscid 'Mycena') belong here. Note that Leucoinocybe is also hard to distinguish from Hydropus and Mycopan.

Species mentioned: Mycopan scabripes. Hydropus pseudotenax. Mycena quinialtensis.

Mycopan scabripes NY - olive-brown conical cap, perhaps not as fragile as a Mycena, stem pruinose to fibrillose at first. We have a newly designated epitype sequence (the holotype was too old), and one matching sequence from OR (in albino form).
'Hydropus' pseudotenax OR (='Mycena' pseudotenax) - similar tough mycenoid with an olive-yellow-grey cap, smaller spores and yellow mycelium at the base of the stem. We have the holotype sequence and one modern WA collection. It needs to moved to Mycopan.
'Mycena' quinialtensis WA - a sometimes very dark, entirely viscid 'Mycena', but difficult to ID when paler capped and much resembling other viscid Mycenas like Mycena vulgaris (which has more decurrent gills). It needs to be moved to Mycopan. It's surprising that this ends up here, although it has been compared to Mycena tenax, which has been compared to 'Mycena' pseudotenax which has been compared to Mycopan scabripes.
Mycopan IN01 - seemingly with a colouration that is dark grey like M. quinialtensis, and also viscid on its cap and stem like M. quinialtensis. It was found once in OR with matching DNA from California that they thought was M. quinialtensis, due to the similarities and not knowing about this lookalike. Then it was found in WA.

Mycopan scabripes © Matthew Koons (normal) and Connor Dooley (albino form), 'Hydropus' pseudotenax © Matthew Koons, 'Mycena' quinialtensis © Star Li

Mycopan IN01 © Connor Dooley and Matthew Koons

Baeospora - click to expand
Small crowded gilled collybioids. One small species grows on conifer cones like Strobilurus but has a uniform pink stem (and more crowded gills). The second not-so-small species is a lilac-tan mushroom on wood.
Species mentioned: Baeospora myosura, myriadophylla.

Baeospora myosura EU (=Baeospora occidentalis ID?) - a similar small mushroom (< 2cm) to the more common Strobilurus on conifer cones but with a uniform pinkish stem and crowded gills. WA sequences match dozens of EU sequences quite well, as well as ENA sequences.
In 2012, a local species from Idaho was described, B. occidentalis, which they thought was different, as it was a snowmelt species with a darker brown cap that didn't grow directly on cones. They did not sequence it, but now we have, and it looks like you can't tell it apart from B. myosura by ITS DNA. Does this mean they are the same or are they actually distinct and other genes will differentiate them? We don't know yet.
Baeospora myriadophylla NY - not quite as small, (~2.5 cm) tan coloured mushroom with distinctive lilac gills when young and a lilac tinted cap when even younger. On wood. It becomes hard to recognize once all the lilac has faded, but still has crowded gills. One WI sequence from back east matches a WA sequence and some EU sequences fairly closely.

Baeospora myosura © Noah Siegel, B. myriadophylla© Shannon Adams

Pleurella - click to expand
Pointy cap and rooting stem like the brown spored Phaeocollybia. Rhodocollybia and Paraxerula are also white spored and have long rooting stems but perhaps not as pointy a cap.
Species mentioned: Caulorhiza umbonata, hygrophoroides. Pleurella microspora.

Pleurella 'microspora PNW01' - Pointy, hygrophanous, orange-brown cap. Crowded adnexed gills. Long, twisted striate stem, rooting stem. Caulorhiza umbonata is a similar California redwood species not yet found here.
Pleurella microspora is only known from Italy, where it does not have a rooting stem. Our ITS sequence differs from the type sequence of P. microspora by 3 bp and 3 indels. Our LSU sequence differs from the type by 2 bp and 1 ambiguous location. So given that, and the fact that the rooting stems do not match the species description, we should consider that our species with a rooting stem needs its own name.

Pleurella 'microspora PNW01' © Andrew Parker

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