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Danny’s DNA Discoveries – Hygrophorineae (Waxy Caps) of the PNW
by Danny Miller

Click here for my Pictorial Key to the Hygrophoraceae

Summary of Interesting Results - click to expand

Here are some of the newest, most interesting results of the study:

Summary of Results from South Sound Mushroom Club Collections - click to expand

Steve Ness, Lauren Ré, Sadie Hickey and Eric Chandler are leading a study of waxy caps with the South Sound Mycological Society and lots of the information we know about this family is coming from the collections they have been making and sequencing.


Cuphophyllus - click to expand

Formerly Camarophyllus, this genus is defined by interwoven gill hyphae, but without a microscope, you may recognize them by dry stems, pastel colours and decurrent gills that are widely spaced apart. Some caps are <5cm across, but others can grow up to 10cm. Other genera in the family may also have these characters, so it's best to browse this section first before moving on to find your mushroom elsewhere. Note that this genus of waxy caps is separate from the other two clades.

Species mentioned: Cuphophyllus borealis, virgineus, niveus, russocoriaceus, lacmus, subviolaceus, rainierensis, cinereus, cinerellus, colemannianus, recurvatus, subradiatus, pratensis. Hygrophorus burgdorfensis, lawrencei, nordmanensis, graveolens, subaustraligus. Camarophyllus cremicolor.

Hygrophorus - click to expand

All of these species have divergent gill hyphae. Most have a viscid cap unless noted. A viscid stem will be noted. The stems are often less colourful than the caps, but if they have a colourful stem they will usually be larger than a Hygrocybe s.l. Some have a partial veil (that never happens in the other two groups). Note that this genus is a separate clade from Cuphophyllus and all the others.

Species mentioned: Hygrophorus caeruleus, subalpinus, albiflavus, sordidus, penarius, siccipes, hypothejus, boyeri, purpurascens, velatus, olivaceoalbus, olivaceoalbus var. gracilis, fuscoalboides, whitei, fuscoalbus,  inocybiformis, agathosmus, agathosmoides, albofloccosus, odoratus, bakerensis, variicolor, secretanii, monticola, vinicolor, erubescens, erubescens var. gracilis, neoerubescens, persicolor, capreolarius, amarus, russula,  parvirussula, piceae, eburneus, cossus, chrysaspis, glutinosus, gliocyclus, flavodiscus, melizeus forma minor, sordidus, penarius, chrysodon, ellenae, pusillus, subpungens, goetzei, goetzii, saxatilis, karstenii, karstenii forma minor, pudorinus, fragrans, speciosus, speciosus var. kauffmanii, pacificus, vernalis, discoideus, occidentalis, tephroleucus, pustulatus, pustulatoides, morrisii, marzuolus, camarophyllus, calophyllus, megasporus, avellaneifolius. Clitocybe odora.

Hygrocybe s.l.

Hygrocybe s.l. have parallel gill hyphae, and both the smaller and the larger species have bright colours on the stems. They never have a partial veil. It has been decided to split this genus up into a number of other genera (including Chromosera, Gliophorus, Gloioxanthomyces and Humidicutis as well as Hygrocybe), although they are all closely related so you could argue that the split didn't need to happen. This becomes especially true because the individual genera are hard to characterize and recognize, which is less than ideal for a genus. This is the third and largest clade of waxy caps.

Note that Chrysomphalina is in this clade, but as they are semi-waxy and not typically thought of as waxy caps (and grow on wood, which is unusual for mushrooms on this page except for Chromosera) I treat them on the non-waxy Hygrophorineae page.

Hygrocybe s.l. - Chromosera - click to expand

Chromosera are viscid all over with decurrent gills. To distinguish between similar Gliophorus, Gloioxanthomyces and Hygrocybe you are looking for gills at least as brightly coloured as the stem and cap, only yellow or lilac colouration, and a stem that is not long and thin. One species grows on wood (the only species on this page that does).

Species mentioned: Chromosera cyanophylla, citrinopallida. Mycena lilacifolia.

Hygrocybe s.l. - Gloioxanthomyces - click to expand

We have one rare species in this genus here, and it is viscid and bright yellow everywhere, including in the gills, with an indented cap, decurrent gills and a very long, thin stem.

Species mentioned: Gloioxanthomyces nitidus

Hygrocybe s.l. - Humidicutis - click to expand

A rare, entirely orange (with olive colours when fresh) species with a dry cap. The gills are especially bright orange and stay orange, at least on the margins, long after the colour has faded elsewhere.

Species mentioned: Humidicutis marginata var. marginata and var. olivacea

Hygrocybe s.l. - Gliophorus s.l. - click to expand

Gliophorus species are slimy-viscid all over, sometimes with decurrent gills. Similar mushrooms are found in Chromosera, Gloioxanthomyces and Hygrocybe, but those are usually not quite as slimy-viscid. Usually. Gliophorus are never red like Hygrocybe can be (they probably lack the ability to produce betalain pigments). A famous green mushroom is found here. Gliophorus does not appear to be monophyletic, so perhaps some of the three groups will be split into additional genera.

Species mentioned: Gliophorus psittacinus, laetus, sciophanus, irrigatus, unguinosus. Hygrophorus subaromaticus. Hygrocybe hondurensis, flavifolia.

Hygrocybe s.s. - click to expand

The rest of the species are still properly in Hygrocybe itself.

Species mentioned: Hygrocybe virescens, miniata, constans, squamulosa, substrangulata, phaeococcinea, reidii, cantharellus, turunda, turunda var. sphagnophila, coccineocrenata, singeri, conica, californica, persistens, cuspidata, acutoconica, acutoconica var. microspora, flavescens, chlorophana, ceracea, subceracea, coccinea, marchii, aurantiosplendens, laetissima, punicea, mucronella, reae, parvula, minutula, subminiata, fenestrata, glutinipes, constrictospora, americana, luteo-omphaloides, atro-olivacea. Gliophorus minutulus, fenestratus. Omphalina occidentalis. Hygrophorus albicarneus.

One study showed that the DNA of the following Hygrocybe species were found inside seeds in either Washington or Oregon, far away from their normal range, and never known to have fruited (so the mushrooms may never be found here).

H. chloochlora, H. nigrescens, H. occidentalis, H. papillata, H. caespitosa, H. glutinipes, H. noninquinans

Summary of Future Studies Needed

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