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Danny’s DNA Discoveries – Ramaria of the PNW
Click here for my Pictorial Key to the Ramaria |
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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 (formerly subgenus Echinoramaria) - with spiny spores (sharper than the warts found in the various subgenera of Ramaria. They have rhizomorphs and are found on the ground litter and the colours are usually yellowish blah (sometimes greenish). They seem to form a well supported clade together in a multi-gene tree. I am assuming they are not mycorrhizal, as they are relatively basal in this order, but some wonder if they have a dual ecology. This has not been answered yet. Lentoramaria (+Rubellaria, Graciramaricus, Acutiramaria, Brachyramaria) (formerly subgenus Lentoramaria) - usually with warty spores (duller and rounder than the sharp spines of Phaeoclavulina) usually blah to yellowish-blaw coloured (occasionally pinkish or greenish), with rhizomorphs, and found either on the ground or on wood. This section had to be split into multiple genera. 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, and most are assumed to be mycorrhizal. Very few mycorrhizal Ramaria species are found both in the EU and in NA (see the discussion under Laeticolora). Gautieria and Destuntzia truffles are part of Ramaria subgenus Ramaria. 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. Most are assumed to be mycorrhizal. Very few mycorrhizal Ramaria species are found both in the EU and in NA as they are highly tree specific. The local Ramaria studies guessed that perhaps 2 out of 75 EU species were found here, but it turned out none of them are. It took a lot of genes to figure out that they are (mostly) all together in the tree next to subgenus Ramaria so thankfully they can keep the name Ramaria. Gomphus - a few purplish Laeticolora turn out to belong in Gomphus with the purple chanterelle. 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 and this study that formalized the names. 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, coniferarum, argentea, glaucoaromatica, mutabilis, incognita.
Might be green Phaeoclavulina sp. 'abietina-PNW01' - a small (<5 cm tall) blah coral that usually has greenish tones, either inherently or from staining. 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 sp. 'PNW02' - also has green. It should be investigated if one of our unnamed species is Phaeoclavulina glaucoaromatica TN or Phaeoclavulina mutabilis EU. Sequenced a few times from WA and OR. Phaeoclavulina argentea EU (formerly PNW03) - can have green when cut or handled, has white rhizomorphs, and is sometimes quite yellowish, but is described as another blah coral with a stem that bruises brown and spores averaging ~6.5u long with spines <=1.2u long. We have ITS2 of the type sequence, and matching local sequences. Phaeoclavulina glaucoaromatica TN - a larger greenish 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 coniferarum EU - not known from the PNW until sequenced once from ID and once from WA. The first was quite yellowish, the second was quite green. The 2 local sequences match each other, but are 4 bp and 3 ambiguous locations from the type sequence. Phaeoclavulina mutabilis EU - 5-15 cm tall, with green only on the stem. Spores average 6.5u long. It has been reported throughout the PNW. No DNA yet from anywhere.
Never noted to be green 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 show it's here. Phaeoclavulina curta (='curta-CA01', =CA08) - a small (<5 cm tall) blah species with the smallest spores (<5u long) that have the smallest spines (<0.3u). We have some EU type area sequences that have a chance of being this species, one one BC sequence is 2 bp and an indel or two different. A photographed OR collection is also very close. Phaeoclavulina minutispora EU - one BC and one CA collection match the type sequence in ITS2 but are somewhat different in ITS1. Phaeoclavulina corrugata EU - a couple of BC collections were discovered through DNA. 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 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 sp. 'PNW02' © iNaturalist user ellenkr, P. sp. 'PNW03' © Steve Ness (2 images), P. coniferarum © Ed Barge Phaeoclavulina eumorpha © Daniel Winkler, Phaeoclavulina curta © Dominic Kurdziolek
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Lentoramaria (Rubellaria, Graciramaricus,Acutiramaria,Brachyramaria) - click to expand
Species in former subgenus Lentoramaria were split into several genera. Species mentioned: Ramaria stricta, concolor, apiculata, tsugina var. prasina, rubella, suecica, gracilis, testaceoflava, rainierensis.
Lentoramaria - erect on wood Ramaria stricta EU (=Ramaria stricta var. concolor ID, =Ramaria concolor ID) - an tall, erect coral on wood with yellow tones in the branches, dimitic hyphae, and spores averaging ~8u long. Both type sequences match except for a couple bp, and some sequences fall in between or have ambiguities. Perhaps R. concolor, lacking yellow tones, should just be considered a colour form and not a true variety. Ramaria stricta var. concolor forma marrii TN - less erect, but branches more curved and open. We have no ITS sequences, but I suspect this has no meaningful genetic difference either. It has been reported from ID and WA. Ramaria sp. 'stricta-IN01' - more common back east, but a few collections in the PNW are this sister species. Ramaria sp. 'PNW14' - no information yet. ~70% support for being in Lentoramaria s.s. It is known from a couple of WA and a couple of OR sequences. Ramaria stricta © NAMA and the Field Museum of Natural History, R. stricta var. concolor © Steve Ness, R. sp. 'stricta-IN01' © Buck McAdoo, Ramaria sp. 'PNW14' © Danny Miller
Acutiramaria - Lignicolous, possible green tones. Ramaria sp. '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 matches an epitype sequence in ITS1, but not in ITS2.
Ramaria tsugina NY var. prasina 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. I have no data on the type variety, nor if it is distinct, nor if it occurs here too. Ramaria sp. 'WA03' - one WA collection with some discolouration that could be interpreted as greenish. Ramaria sp. 'IN11' - BC collections showing peachy, orange, or yellow colours, but no green has been noticed so far. Ramaria sp. 'WA05' - found both white and bright yellow. Ramaria tsugina var. prasina © Shannon Beers and Steve Ness
R. testaceoflava is sister to Acutiramaria, but is not lignicolous. Nor does it have the small size or sharp tips of other Acutiramaria. However, I am assuming for now that it will be considered part of it. 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. Ramaria testaceoflava var. brunnea © Mary McCallum
Rubellaria - 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
Brachyramaria - Terrestrial Ramaria suecica (=Ramaria circinans var. anceps WA, =PNW54) - 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
Graciramaricus - Terrestrial 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 and ID sequences match them. Ramaria rainierensis WA - 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. You need quite a few genes unsequenced Ramaria gracilis © C D Marr, R. gracilis © Ed Barge |
Ramaria subgenus Ramaria - click to expand
Gautieria and Destuntzia truffles are in this section and need to be moved to Ramaria. Species mentioned: Ramaria botrytis, subviolacea, rubrievanescens, rubripermanens, aurantiiramosa.
Ramaria sp. '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 and photographs. Ramaria sp. 'CA05' - seems similar to R. rubrievanescens. We need LSU of this or ITS of R. rubrievanescens to verify they are distinct. It has been found in OR and CA, but not WA. Ramaria sp. '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 sp. 'PNW08' - mostly off white when mature, perhaps the branch tips are a little peachy when young like R. aurantiiramosa. 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 sp. '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. This is one of only 2 mycorrhizal EU species thought to be found in the PNW (the other was R. formosa) but neither of them actually do. There is very little overlap between EU and NA species of the mycorrhizal subgenera Ramaria and Laeticolora. Ramaria aurantiiramosa WA - similar to R. botrytis with similarly sized spores, but with orange branch tips when young like R. PNW08. It was formerly a variety of R. botrytis, but it is not closely related enough. Ramaria sp. 'PNW04' - a spring yellowish (or reddish!) 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). Also, the colour in the tips might fade like others in this subgenus. Most notably, it can also be red, explaining why it has gone undetected until the DNA age. Only when it is yellow will it stand out. Ramaria sp. 'PNW07' - no information yet. Sequenced twice from OR. Ramaria sp. 'PNW10' - no information yet, except that the branches might be rather stubby. Sequenced once from BC and once from OR. Ramaria sp. 'subviolacea-PNW01' © Noah Siegel (fresh) and Michael Beug (not as fresh) Ramaria sp. 'rubrievanescens-PNW09' © Ed Barge, R. sp. 'PNW08' © Mary McCallum, R. sp. 'PNW04' © Mary McCallum Ramaria sp. 'botrytis-PNW02' © Jonathan Frank and Steve Ness, Ramaria rubripermanens © Jenny Lippert, Ramaria aurantiiramosa © Mary McCallum
Ramaria sp. 'PNW51' - a red tipped species like R. botrytis, but this sequence is out on a long branch by itself, almost like it deserves its own genus/subgenus. I have doubts that it is subgenus Ramaria, but I can't rule that out, and it keys out here. We need to test the flesh to see if it's amyloid. Ramaria sp. 'PNW44' - in the same distant clade as PNW51. There is no photo, but it was mistaken for R. rubrievanescens, so it appears to also resemble a subgenus Ramaria species. Ramaria sp. 'PNW51' © Kathleen Dobson |
Gomphus - click to expand
A few purplish Laeticolora turned out to belong in Gomphus with the purple chanterelle. Species mentioned: Ramaria violaceibrunnea, fennica.
Ramaria violaceibrunnea WA (=Ramaria fennica EU var. violaceibrunnea WA) - duller smoky-purple upper branches and tips (whiter base), not cauliflower-like. Clamps. R. violaceibrunnea was formerly a variety of Ramaria fennica EU, and is probably closely enough related that it could still be considered a variety of it. We have the type sequence and several recent local matching sequences. Ramaria sp. 'ID02' - was an dull orange collection, but the stipe had fresh purple colours. Ramaria violaceibrunnea © Mary McCallum |
Ramaria subgenus Laeticolora - click to expand
Species mentioned: Ramaria botrytoides, aureoprimulina n.p., gelatiniaurantia, leptoformosa, rubricarnata, sandaracina, spinulosa, marrii, fumosiavellanea, acrisiccescens, rubribrunnescens, foetida, 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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Introduction - click to expand