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Clavulinopsis subgenus Clavulinopsis

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This subgenus includes the type species, Clavulinopsis sulcataand is characterized by minutely apiculate, thin-walled spores that are invariably globose to subglobose in the well known species (Figure 1) though some poorly known species are reported to have spores tending toward the ellipsoid spectrum). This is not a particularly useful character for field identification unless you have very good eyesight so don't worry, there are some other clues that can be pieced together to arrive here with out laborious microscopy. Fruiting bodies are typically large and "inflating" with brightly colored hymenia in the red-orange-pink-yellow series though there are pale to white representatives (e.g., Cu. bicolor). These pigments appear to be chemically different from those found in other subgenera of Clavulinopsis (carotenoids) and fade drastically in herbarium specimens which makes screening preserved collections easy (Figure 2). This can sometimes be seen in the field in somewhat dry conditions and helps differentiate yellow representatives from the macroscopically similar Clavulinopsis species in subgenera (e.g., fusiformis groups, laeticolor group, and a (sub)simple corniculata-related species found in Western North America).

 A

IMG_3957_edited.jpg

B

Figure 1. Comparison of the apiculus between subgenus Clavulinopsis and subgenus Acularia. A. Small, "papillate" apiculus diagnostic of subgenus Clavulinopsis (Cu. aurantiocinnabarina complex: JMB081223-4/iNat 178452312. B. Large, "conical" apiculus (a somewhat exagerated example) typical of subgenera Acularia and Donkella (Clavulinopsis fusiformis complex: Buck Raeder 05202309/iNat 162717778). Note: The spores of subgenus Paraclavaria are typically intermediate in size and not pictured here.

 A

B

JMB081223-18_edited.jpg

C

D

Figure 2. Comparison of the color of fresh and dried fruiting bodies showing the change in color intensity. A. Clavulinopsis aurantiocinnabarina group fresh (JMB081223-4/iNat 178452312), B. Clavulinopsis aurantiocinnabarina group dried (JMB081223-4/iNat 178452312), C. Clavulinopsis laeticolor group fresh (JMB081223-18/iNat 178452361), D. Clavulinopsis laeticolor group dried (JMB081223-18/iNat 178452361​)

This assemblage was noted as homogenous and uniquely difficult to parse in Clavulinopsis due to the near invariability in microscopic features but the strikingly wide range of color variation lead to tension between splitting every color-morphology or lumping them all under one "mega-species" of sorts with mycological heavy-weights like Petersen and Corner never quite arriving at a satisfactory conclusion. The question remains whether discrete categories can be distinguishing based on color, but this will have to be done very carefully as the color begins to fade very quickly in dry conditions and after picking if not transported with humidity.

Interestingly, this subgenus appears to be circumglobal EXCEPT for Western North America and Europe where no reports are currently known. The subgenus appears to be exceptionally prolific in Eastern Asia through Australasia and in Eastern North America. It is present but less well known in South/Central Asia, Sub-Saharan Africa, Central America, and South America.

The two most commonly used names are: Clavulinopsis aurantiocinnabarina (represented in the phylogeny by about 6 different species level clades) and Clavulinopsis sulcata (with sequences in 4 different locations in the phylogeny). Otherwise, there are many names published from East Asia through Australasia that need clarification.

Phylogeny Reconstructed using the ITS region

appalachiensis/subaustralis?

sp.

sp.

Clavulinopsis appalachiensis/subaustralis

This species is distinguished from other Clavulinopsis species by its softer colors in the peachy-salmony range with a somewhat more intensely colored sterile portion in the orange range though also sometimes lemon colored (maybe I am hungry but this sure is a food-centric range of colors here: Easy Citrus Salmon, Peach Glazed Salmon). It also grows in single or in small clusters, never(?) cespitose as often seen in the Cu. aurantiocinnabarina complex. 

The name Cu. appalachiensis (Coker) Corner is being used for this organism and I tentatively agree but think that a detail comparison with Cu. subaustralis R.H. Petersen should be presented:

Souces: Coker 1923, Petersen 1968 and 1971

What I think becomes apparent is that the species typically collected and referred to as Cu. appalachiensis matches the coloration described for Cu. subaustralis better. The questions then follow: How consistent are colors in this group? Is the color description for Cu. appalachiensis correct? How consistent a character is the inconstancy of clamp connections at the contextual septa?

Colors do vary based on moisture alone and some collections tend toward the yellow end of them spectrum. In fact, the tendency of species in the subgenus to fade to much paler colors when dried could be responsible for the very pale yellow colors described by Coker in the protologue of Cu. appalachiensis. Petersen notes that dried specimens of Cu. subaustralis become "cream buff or cream color" in the protologue of his species.

What to do about the character of clamp connection constancy. It may not be that constant after all. Completely clampless versions of otherwise clamped species of Clavulinopsis are known (Petersen 1978) though a partially clamped state has not been reported to my knowledge, until possibly now. A more thorough study of specimens, particularly those putatively with prevalent clamp connections (identified as Cu. subaustralis), may help elucidate this situation.

With all this in mind, I think it is reasonable to consider these two species conspecific which would give Cu. appalachiensis priority. Specimens with colors typical of Cu. subaustralis but with the clamp inconstancy, otherwise a unique character(?) of Cu. appalachiensis, seem to support this, especially given the possibility of the miss-application of the dried color of the latter species to that of its fresh state. I cannot, however, rule out that there may be a separate, related species, sharing clamp inconstancy which does indeed have a very pale yellow coloration in its fresh condition, lacking pink colorsbetter matching Coker's species and leaving this species more likely with the name Cu. subaustralis.

A collection from Florida may be conspecific. It has >98.5/<90.0% similarity to the other ITS regions.

Distribution: Originally described from North Carolina and Pennsylvania, it is now known from Tennessee, New York, and possibly Florida. Apparently uncommon to rare.

Verified Collections

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Clavulinopsis aurantiocinnabarina group

The species "Clavaria aurantio-cinnabarina" was somewhere around the 8th species of clavarioid fungus described from the United States by Lewis David de Schweinitz (1832), the first  American mycologists. He described a large, fasciculate, flexuous, compressed (up to 2.5 cm), fusiform but obtuse, orange to cinnabar colored, unbranched clavarioid with a white pruinose to subbyssoid base on bare soil near Rhododendron on the campus of Lehigh University in Bethlehem, Pennsylvania. This group of species exhibits some of the most beautiful and striking colors in the fungal world, rivaled by only some species of Cantharellus and Hygrophoraceae in my opinion.

While this group is strikingly recognizable, the differences between the species are less so. Hopefully with careful study differences can be found. If they exist, they are not readily recognizable. Please, let me know if know how to tell them apart! Would the real Cu. aurantiocinnabarina please stand up?

Distribution: Species in this group are found through-out the Eastern United States east of the great plains, extending up into Eastern and Maritime Canada. Quite common. The name has occasionally been applied to Asian organisms.

Observations and notes for each member so far known are listed below

Clavulinopsis aurantiocinnabarina NN01

The two collections with photographs of a very orange species, though both are a bit dry. They both seem to have a somewhat noticeable, orange sterile portion.

 

Distribution: This species appears widely distributed across the Eastern United States from Florida up to New York.

Verified Occurrences

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
aurantiocinnabarina NN01
FLAS-F-69299
FLAS-F-69299
USA-FL
OQ725183.1
Possible
Clavulinopsis
aurantiocinnabarina NN01
AO20
-
USA-IN
ON650113.1
Certain
Clavulinopsis
aurantiocinnabarina NN01
23983
-
USA-NY
ON416905.1
Certain

Clavulinopsis aurantiocinnabarina NN02

This group is based on four collections of mine from the Southern Appalachians. My brief notes indicate that these were "orange", and I don't have much more until I can find the full descriptions.

Distribution: The four collections of mine are all from Tennessee but there is no reason at this point to assume it is an endemic. Sequencing of a lot more "aurantiocinnabarina" need to be done before these things can be sorted out.

Verified Occurrences

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
aurantiocinnabarina NN02
JMB08071204
TENN-F-067402
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina NN02
JMB08171004
TENN
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina NN02
JMB08240901
TENN
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina NN02
JMB08061204
TENN-F-067586
USA-TN
Need
Certain

Clavulinopsis aurantiocinnabarina NN03

This group is composed of two collections from the Southern Appalachians. My collection had pigments primarily residing in the contextual elements while the hymenium was pale. This seems to be Cu. miniata sensu Petersen 1968 and I am curious how consistent pigment distribution is.

Distribution: Only known from Tennessee but unlikely to be restricted to there.

Verified Occurrences

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
aurantiocinnabarina NN03
JMB08101205
TENN-F-067667
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina NN03
SAT0921319
TENN-F-064103
USA-TN
Need
Certain

Clavulinopsis aurantiocinnabarina NN04

This is yet another group producing orange fruiting bodies.

Distribution: Appalachian Tennessee to Pennsylvania so far.

Verified Occurrences

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
aurantiocinnabarina NN04
JMB08031201
TENN-F-067378
USA
Need
Certain
Clavulinopsis
aurantiocinnabarina NN04
iNat 133671306
-
USA-PA
Need
Certain
Clavulinopsis
aurantiocinnabarina NN04
SAT0921301
TENN-F-064102
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina NN04
PBM3010
TENN
USA-WV
Need
Certain

Clavulinopsis aurantiocinnabarina/sp OH-01

Ok, this is probably pretty clearly not aurantiocinnabarina since it looks like the yellow coloration is consistent. I have called it "aurantiocinnabarina Lemon" in my head for a while but with only two specimens examined at that point, I was unsure of the consistency of this coloration. Now it looks to be a potential trait with 4 specimens that are all yellow. It is tempting to call this Clavulinopsis aurantiocinnabarina f. amoena R.H. Petersen (1968) given its coloration, but it is unclear to me if Petersen meant the form for a truly yellow species as the name implies, or a somewhat paler or faded orange species as the description and picture show. Either way, the true Cu. amoena does not appear to be in this subgenus, though the species typically ascribed to that name cetainly does (e.g., Corner 1950).

In the field, this species is probably hiding under the name Cu. fusiformis of subgenus Acularia. It cold be distinguished dried at a glance (pale as opposed to maintaining the yellow coloration) which can sometimes be noticed in the field when conditions are dry. Additionally it has mildly sweet taste (as opposed to bitter/farinaceous), likely doesn't turn greenish in iron salts nor orange in KOH, and is clearly different based on the spore apiculus.

Distribution: Tennessee up through the midwest (Michigan, Ohio) extending at least into Quebec. Likely more widely distributed and hiding under the name Cu. fusiformis.

Verified Occurrences

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
aurantiocinnabarina/sp OH-01
HRL2567
-
CAN-QUE
ON943319.1
Certain
Clavulinopsis
aurantiocinnabarina/sp OH-01
STZ11451
WTU
USA-MI
Need
Certain
Clavulinopsis
aurantiocinnabarina/sp OH-01
66236
PUL F27206
USA-OH
Need
Certain
Clavulinopsis
aurantiocinnabarina/sp OH-01
JMB08101209
TENN-F-067692
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina/sp OH-01
JMB08171006
TENN
USA-TN
Need
Certain
Clavulinopsis
aurantiocinnabarina/sp OH-01
JMB08061202
TENN-F-067599
USA-TN
Need
Certain

Clavulinopsis aurantiocinnabarina IN03

This one is also orange but maybe with a salmon tint? Would need a lot more data to consider this diagnostic.

Verified Occurrences

Click on the collections below to see link to verified observations (if applicable on iNaturalist or Mushroom Observer).

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
aurantiocinnabarina IN03
2273
-
USA-IN
Need
Certain
Clavulinopsis
aurantiocinnabarina IN03
BPL253
TENN-F-067381
USA-TN
Need
Certain

Clavulinopsis bicolor

This is a pleasant little species with white to off-white hymenial coloration and a distinct yellow brown sterile portion recent described from China based on a single specimen (Yan et al., 2023).

Verified Occurrences

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
bicolor
HOLOTYPE
MHHNU10381
MHHNU10381
China-Hainan
OQ703780.1
TYPE

Clavulinopsis incarnata

This beautiful and recently described species is characterized by its sparse (non-fasciculate) growth and soft pink fruiting bodies (Yan et al., 2023). There are several older names for pinkish Clavulinopsis for subtropical to tropical Asian species that need to be clarified before this name is completely safe in my mind.

Verified Occurrences

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
incarnata
PARATYPE
MHHNU9314
MHHNU9314
China-Hunan
OQ703785.1
PARATYPE
Clavulinopsis
incarnata
PARATYPE
MHHNU11331
MHHNU11331
China-Yunnan
OQ703788.1
PARATYPE
Clavulinopsis
incarnata
PARATYPE
MHHNU9813
MHHNU9813
China-Yunnan
OQ703786.1
PARATYPE
Clavulinopsis
incarnata
HOLOTYPE
MHHNU11330
MHHNU11330
China-Yunnan
OQ703787.1
HOLOTYPE

Clavulinopsis novo-zealandica

This species is described as fleshy and inflated like Cu. sulcata. It has a pale yellow to cream fertile portion with a brighter yellow sterile portion.

Distribution: Known only from New Zealand.

Verified Collections

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
novozealandica
HOLOTYPE
TENN043575
TENN
NZ
Need
HOLOTYPE

Clavulinopsis sulcata/miyabeana/etc.

It looks like the issue we have in eastern North America with a high amount of diversity passing under 1 or 2 names is not unique. Most of these are probably in the pink-orange color series given the names of the species used. Somewhere in here is probably the type species, Cu. sulcata but I couldn't hazard a guess which. More work in East Asia/Australasia will be needed to sort this pile of species out. There are A LOT of names available for white-yellow-pink-orange-red species this region (see below) and the best of luck to the brave souls that take on the project of sorting all of these abandoned, poorly known, or otherwsie confused taxa!

Collections in this group

Genus
species
Type
Collection
Herbarium
Location
ITS Genbank
Certainty
Clavulinopsis
sp.
WML-48
WML-48
-
OK584470.1
Clavulinopsis
sp.
KN23
KN23
-
OL638253.1
Clavulinopsis
sp.
WML2021-8-25-2
WML2021-8-25-2
-
OL998888.1
Clavulinopsis
sp.
KFRI MH1519
KFRI MH1519
-
OR224623.1
Clavulinopsis
sp.
LD2021-6-9
LD2021-6-9
-
OK643821.1
Clavulinopsis
sp.
MEL 2382732
MEL 2382732
AUS-NT
KP012914.1
Clavulinopsis
sp.
MCCNNU 00948
MCCNNU 00948
China
MT587808.1
Clavulinopsis
sp.
MCCNNU 00952
MCCNNU 00952
China
MT587809.1
Clavulinopsis
miyabeana
ZP-2118
ZP-2118
China-Hunan?
MK427059.1
Clavulinopsis
sulcata
ZP-2119
MHHNU 8619
China-Hunan?
MK427060.1
Clavulinopsis
sp.
ZP-2400
MHHNU 8900
China-Hunan?
MK427063.1
Clavulinopsis
sulcata
HFJAU-TD330
HFJAU-TD330
China-Jiangxi
MN622711.1
Clavulinopsis
sulcata
26
26
China?
OL616149.1
Clavulinopsis
miyabeana
198-280
198-280
Japan
AB509666.1
Clavulinopsis
miyabeana
198-506
198-506
Japan
AB509804.1
Clavulinopsis
miyabeana
197-490
197-490
Japan
AB509796.1
Clavulinopsis
sulcata
PDD 107085
PDD 107085
NZ
Need
Clavulinopsis
miyabeana
BMS12BM
BMS12BM
Philippines
OR099734.1
Clavulinopsis
sp.
BMS13AM
BMS13AM
Philippines
OR099733.1
Clavulinopsis
miyabeana
BMS12F
BMS12F
Philippines
OR099726.1
Clavulinopsis
sulcata
MG10
MG10
Philippines
OK446763.1
Clavulinopsis
sp.
BMS13F
BMS13F
Philippines
OR099728.1

Non-exhaustive list of other species possibly or likely in this subgenus

Clavulinopsis amoena sensu Corner et al. (= Cu. spiralis?) - Yellow, widespread paleotropical

Clavulinopsis arenicola - pale buff, Singapore

Clavaria brevipes - white, Singapore

Clavaria caepicolorosa - red, Australia

Clavaria cardinalis - red, Australia

?Clavulinopsis inflatissima - yellow?, Bolivia

Clavaria laeta - red, Sri Lanka

Clavulinopsis liguloides - yellow-brown, Indonesia

?Clavaria militina - red, India

Clavulinopsis miniata var. rosacea, pinkish flesch colored, Malaysia? Madagascar?

Clavulinopsis miniata var. sanguinea, red, Malaysia

Clavaria miyabeana - red, Japan

Clavaria pallidorosea - orange-pink, Australia

Clavaria phoenicea - pinkish purple? Orange-red?, Indonesia

Clavaria phoenicea var. ealensis - pinkish purple, Democratic Republic of the Congo

Clavaria rosacea - pinkish flesch colored, Madagascar

Clavaria sanguinioacuta - red, Malaysia

Clavaria schimadai - orange, Japan

Claulinopsis spiralis - white, pale yellow, Indonesia

?Clavaria subargillacea - yellow, Japan

?Clavaria subaurantiaca - orange, Indonesia

Literature Cited:

Coker, W. (1923). The Clavarias of the United States and Canada. University fo North Carolina Press.

Corner, E. (1950). A Monograph of Clavaria and Allied Genera. Oxford University Press.

Petersen, R. (1971). Notes on Clavarioid fungi—IX. Addendum to Clavulinopsis in North America. Persoonia, 6(2), 219–229.

Petersen, R. (1978). Notes on clavarioid fungi. XVI. Clampless taxa in Clavulinopsis. Mycotaxon, 6(9), 503–507.

Petersen, R. H. (1988). The Clavarioid Fungi of New Zealand. Science Information Pub. Centre, Department of Scientific and Industrial Research.

Schweinitz, L. (1832). Synopsis Fungorum in America Boreali media degentium. Secundum Observationes. Transactions of the American Philosophical Society, 4, 141–316.

Yan, J., Wen, J., Li, G.-W., Wu, S.-W., & Zhang, P. (2023). Taxonomy and Phylogenetic Relationships of Clavulinopsis (Clavariaceae, Agaricales): Description of Six New Species and One Newly Recorded Species from China. Journal of Fungi, 9(6), 656. https://doi.org/10.3390/jof9060656

Additional Notes

Tree in Newick Format

Version 2308.22 (fixed typos and tables).

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