Danny’s DNA Discoveries – Ramaria of the PNW
Click here for my Pictorial Key to the Ramaria
Ramaria are corals with striate, spiny, or warty spores. They may be quite colourful or blah, and found either on the ground or on wood. Clavulina are blah corals on the ground with smooth spores. Lentaria are blah corals on wood with smooth spores. Other coral fungi are found in various other places in the fungal tree, it is a somewhat common shape.
Phaeoclavulina - with spiny spores (sharper than the warts found in the various subgenera of Ramaria. They have rhizomorphs and are found on the ground and the colours are usually yellowish blah. They seem to form a well supported clade together, but the true Lentaria (Lentaria A) with smooth spores are found near or inside this genus and a multi-gene study needs to sort out that relationship.
Ramaria subgenus Ramaria - with striate spores, usually colourful on the branch tips only and with amyloid flesh. Most are of the cauliflower shape with a single large stem and only short abortive branches growing out of it. Found on the ground. They form a well supported clade together.
Ramaria subgenus Lentoramaria - usually with warty spores (duller and rounder than the sharp spines of Phaeoclavulina) usually blah to yellowish-blaw coloured (occasionally pinkish), with rhizomorphs, and found either on the ground or on wood.
Ramaria subgenus Laeticolora - also usually with warty spores, but usually brightly coloured both on the branches and their tips. The flesh is usually not amyloid. Many are not strongly cauliflower shaped, but have some other coral form. Found on the ground, usually without rhizomorphs. This large subgenus has been divided into two based on whether a species has clamp connections or not, but that has little or no taxonomic value. It can, however, greatly aid in identification.
Unfortunately, Ramaria subgenus Lentoramaria may have to be split into 4 cryptic genera, unless Clavariadelphus (clubs, not corals), our Lentaria B species, and associated crust genera are all lumped together as Clavariadelphus, but then that genus would contain both clubs and corals. A multi-gene study is needed to sort this out.
Even more unfortunately, Ramaria subgenus Ramaria and Ramaria subgenus Laeticolor are in one big clade with Gomphus, Turbinellus, and Gautieria, etc. This means that either false chanterelles and corals will all be in one large genus Gomphus (the oldest name), or Ramaria subgenus Laeticolor may have to be split into a bunch of cryptic genera (my first guess is 9 genera)! Neither is an ideal solution. It is fascinating to have learned that false chanterelles and corals evolved back and forth so many times. It's easier to see how gilled and pored mushrooms evolved back and forth, you can picture pores as gills that are pinched together every once in a while to form the pores. But I cannot fathom why the two very different shapes of Gomphus/Turbinellus and Ramaria evolved back and forth so many times. See this tree.
DO NOT BOTHER to try and identify a Ramaria if it is not young and fresh. Every species soon becomes the same dull yellow-orange-brown. Things you need to know to identify a Ramaria:
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.
Phaeoclavulina - click to expand
Species mentioned: Phaeoclavulina eumorpha, corrugata, myceliosa, curta, abietina, argentea, glaucoaromatica, mutabilis, incognita.
Phaeoclavulina 'abietina PNW01' - a small (<5 cm tall) blah coral that stains green. Ramaria apiculata and Ramaria tsugina, below, may have greenish branches but they are found on wood. Phaeoclavulina abietina is a complex of species, and the one that occurs here is not the same as the real one that occurs in the EU (although the real one occurs in Northern Alberta). Holzapfel calls ours "complex B-IV". Sequenced once from OR so far.
Phaeoclavulina PNW02 - also stains green. It should be investigated if this is Phaeoclavulina glaucoaromatica TN or Phaeoclavulina mutabilis EU. Sequenced a few times from WA and OR.
Phaeoclavulina PNW03 - also stains green. One WA collection with unique DNA (the only thing similar is a sequence from Montana 7 bp different). It seemed especially yellow, and not as blah as expected. It had white rhizomorphs and some green staining where cut or handled.
Phaeoclavulina glaucoaromatica TN - a larger green staining species (5-15cm tall) with a white stem base that smells of maple syrup when dried, with large spores (~10u long). It has been reported from ID. No DNA yet from anywhere.
Phaeoclavulina mutabilis EU - 5-15 cm tall, staining green only on the stem. Spores average 6.5u long. It has been reported throughout the PNW. No DNA yet from anywhere.
Phaeoclavulina eumorpha EU - a blah coral with yellow rhizomorphs, spores averaging ~7.5u long. We have EU type area DNA and local DNA from BC and WA to prove it's here.
Phaeoclavulina curta EU - a small (<5 cm tall) blah species with the smallest spores (<5u long) that have the smallest spines (<0.3u). We have some EU sequences that have a good chance of being this species, one one BC sequence is 2 bp and an indel or two different.
Phaeoclavulina myceliosa CA - much like R. curta, with similar spores, but this coral has branches that are a little more open. We don't have any western DNA, and we don't really know if this is distinct from P. curta or not. Some say it is, but we have a few EU sequences purporting to be this, and those sequences are distinct from R. curta. We need collections.
Phaeoclavulina argentea EU - another blah coral with a stem that bruises brown and spores averaging ~6.5u long with spines <=1.2u long. It was reported from ID. We have ENA DNA purporting to be this, but no EU type area nor local DNA.
Phaeoclavulina incognita TN - very similar to P. argentea and some synonymize them. It is said the tips are less likely to turn white in this species. The spores are said to average ~7.5u long with spines <=2u long. We have no DNA yet from anywhere yet, to show if this is indeed a distinct species.
Phaeoclavulina PNW02 © iNaturalist user ellenkr, P. PNW03 © Steve Ness (2 images), P. eumorpha © Daniel Winkler
Ramaria subgenus Lentoramaria - click to expand
These species appear in four different clades in the tree, so if the genus needs to be split, I have numbered the clades from 1 to 4.
Species mentioned: Ramaria (1) gracilis, testaceoflava, apiculata, rainierensis, tsugina, (2) stricta, concolor (3) rubella (4) suecica.
Ramaria gracilis EU - found on the ground, with the smallest spores in the whole subgenus, averaging ~6u long. Easily confused with various Phaeoclavulina. We have a bunch of EU type area sequences, and a couple of OR sequences match them.
Ramaria rainierensis WA - similar, also found on the ground, nondescript and very much like various Phaeoclavulina and R. suecica, but with dimitic rhizomorphs and with oddly ornamented clamps. We have the type sequence.
Ramaria testaceoflava EU var. brunnea NM - a rather erect coral found on the ground with somewhat chocolate branches with yellowish tips, staining brown where handled. DNA recently showed this is a Lentoramaria and not a Laeticolora (it always was rather blah for a Laeticolora and it does have rhizomorphs). EU type area sequences of the type variety that does not stain brown are about 3 bp different than our local variety sequences, so I am considering ours a distinct variety.
unsequenced Ramaria gracilis © C D Marr, Ramaria testaceoflava var. brunnea © Mary McCallum
Ramaria 'apiculata PNW45' - found on wood, upper branches and tips greenish like some green terrestrial Phaeoclavulina. Our one local sequence purpurting to be the EU species R. apiculata does not match a couple of EU sequences and may need a new name. We should also check if the real thing occurs here too.
Ramaria tsugina WA - also on wood and may have green tones, but with dimitic rhizomorphs and slightly smaller spores averaging ~9u long. We have the type sequence and matching recent sequences.
Ramaria tsugina © Shannon Beers and Steve Ness
Lentoramaria2 - erect on wood
Ramaria stricta EU - an tall, erect coral on wood with yellow tones in the branches, dimitic hyphae, and spores averaging ~8u long. We have a bunch of EU type area sequences, and matching local sequences.
Ramaria stricta © NAMA and the Field Museum of Natural History, Ramaria PNW14 © Danny Miller
Lentoramaria3 - pink tones
Ramaria rubella EU - with pinkish branches when fresh, a greyish-brown stem, and rhizomorphs that turn pink in KOH. We have a bunch of EU type area sequences, and matching local sequences.
Ramaria rubella forma blanda CA - also with pinkish branches, but with a whiter stem and rhizomorphs that turn yellow in KOH. It is reported from BC and WA. It has the same ITS sequence as the type variety.
Ramaria rubella © Steve Ness
Ramaria suecica (=Ramaria circinans var. anceps WA) - found on the ground, nondescript and very much like various Phaeoclavulina but with different spore ornamentation. It is also like R. rainierensis, but with monomitic rhizomorphs and with regular, unornamented clamps, and like R. gracilis but with bigger spores. R. suecica has a slight variation in ITS, maybe 1%, between the EU type area and our area. We likely have the real thing, but just in case we don't, we do already have a local name for our species, Ramaria circinans var. anceps WA. The type variety of R. circinans NY is distinct in LSU.
unsequenced Ramaria suecica © Ron Exeter
Ramaria subgenus Ramaria - click to expand
Species mentioned: Ramaria botrytis, subviolacea, rubrievanescens, rubripermanens, aurantiiramosa.
Ramaria 'subviolacea PNW01' complex - violet tinged tips only when young, later mostly off-white. Other purple Ramaria are not as cauliflower-like, have purple not just on the tips, and lack amyloid flesh. This is our best guess as to what R. subviolacea ID is, with spores averaging 14u long. If it's not in this complex, we need to figure out what it is. Either this is a species with variation in ITS of 2%, or there are 3 species in this complex.
Ramaria rubrievanescens WA - with pink branch tips only when young and fresh (fading to whitish like the branches themselves). The stem discolours brown and the spores average 12u long. Only LSU has been sequenced of the type. We need ITS sequences.
Ramaria 'rubrievanescens PNW09' - 3.5% different in LSU from the type sequence, we have ITS of this species. I don't know how to tell it apart from R. rubrievanescens.
Ramaria PNW08 - mostly off white when mature, perhaps the branch tips are a little peachy when young.
Ramaria rubripermanens WA - with reddish (or at least dark) branch tips even when somewhat mature, sometimes found in the spring. The spores average 12u long.
Ramaria 'botrytis PNW02' - also with reddish (or at least dark) branch tips even when somewhat mature, but with larger spores averaging 14u long. Our species is distinct from the EU species and probably needs a new name.
Ramaria aurantiiramosa WA - similar to R. botrytis with similarly sized spores, but with orange branch tips when young. It was formerly a variety of R. botrytis, but it is not closely related enough.
Ramaria PNW04 - a spring yellow cauliflower-like coral easily mistaken for R. rasilispora and R. magnipes in subgenus Laeticolora, but this species will likely have amyloid flesh and the branches themselves should be paler (mostly the tips show the colour).
Ramaria PNW07 - no information yet. Sequenced twice from OR.
Ramaria PNW10 - no information yet, except that the branches might be rather stubby. Sequenced once from BC and once from OR.
Ramaria 'subviolacea PNW01' © Noah Siegel (fresh) and Michael Beug (not as fresh)
Ramaria PNW08 © Mary McCallum, Ramaria PNW04 © Mary McCallum
Ramaria 'botrytis PNW02' © Jonathan Frank and Steve Ness, Ramaria rubripermanens © Jenny Lippert, unsequenced Ramaria aurantiiramosa © C D Marr
Ramaria subgenus Laeticolora - click to expand
Species mentioned: Ramaria botrytoides, aureoprimulina n.p., gelatiniaurantia, leptoformosa, rubricarnata, sandaracina, spinulosa, marrii, fumosiavellanea, acrisiccescens, rubribrunnescens, rasilisporoides, celerivirescens, claviramulata, rubiginosa, vinosimaculans, velocimutans, synaptopoda, gelatinosa var. oregonensis, stuntzii, maculatipes, cyaneigranosa, magnipes, rasilispora, coulterae, verlotensis, araiospora, longispora, fennica, violaceibrunnea, formosa, flavobrunnescens, cartilaginea, caulifloriformis, conjunctipes, largentii, aurantiisiccescens, amyloidea, flavigelatinosa, thiersii, cystidiophora, armeniaca, distinctissima var. americana, hilaris var. olympiana, purpurissima, raveneliana.
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