Danny’s DNA Discoveries – Lycoperdaceae of the PNW
Puffballs are roughly spherical gastroid mushrooms that typically fruit on the ground and above ground, unlike most truffles and false truffles. They often (but not always) have a thin, soft exterior and the uniform texture of a white marshmallow inside when fresh. The spongy white interior discolours to olive-brownish (or other shade of brown) then eventually turns to a dark powder and can escape the puffball, sometimes puffing out through a single small pore at the top. If you step on an old one, you'll create a cloud of smoke (see photo above). There may be a sterile base area, a part of the interior that doesn't turn to powder like the rest of it, but otherwise the interior is uniform - no columella (aborted stem) inside, and no outline of a gilled mushroom (like an Amanita egg that might be deadly poisonous). Some people eat puffballs in their fresh, white marshmallow stage, but their quality varies dramatically from species to species and they are easily mistaken for poisonous earthballs, false truffles and Amanita eggs.
Most puffballs have two distinctive layers of what I sometimes call a skin or rind holding the puffball together, the exterior exoperidium and the interior endoperidium. When I say a puffball is thin skinned or has a thick rind, I am referring to both layers collectively. If either of the layers is thick and/or hard, or if collectively they are significantly thick or hard I will refer to it as a thick skinned puffball. Often what happens is the exterior layer eventually falls away and a pore opens up in the interior layer, letting the spores "puff" out. Sometimes both layers break away unevenly exposing the spores inside.
Those with thicker rinds, that don't open up a pore are easily confused with earthballs (with a purple-black interior that does not turn to powder). Some false truffles are also spongy with thin rinds, but they do not eventually turn to powder inside, and are more likely to be found at least partially buried.
Puffballs are in the dark spored sub order of gilled mushrooms, along with the bird's nests.
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.
Abstoma, Apioperdon, Bovista, Disciseda, Mycenastrum - click to expand
See descriptions of each genus below.
Species mentioned: Lycoperdon pyriforme.
This genus is hard to characterize and differentiate from other genera. One notable feature is the outer skin is usually covered in sand that is often trapped in the growing tissue. Part of the top of the puffball ruptures irregularly to let the powdery spores out, which is common in other genera. The species usually grow up to 6cm across.
Abstoma cf reticulatum Australia - No DNA from anywhere, but reported from ID.
Abstoma townei CA - No DNA from anywhere, but reported from ID and OR.
unsequenced Abstoma sp. © iNaturalist user yury_rebriev
Recognizable by growth on wood (unusual in Lycoperdon), but when growing on buried wood it will be harder to identify. It is a pear shaped, relatively smooth (but may have small scales and a texture) brownish puffball with rhizomorphs connecting the base to the wood and a sterile base (inside the thin part of the pear shape it remains white and does not turn colour). Eventually a pore area opens up on top, but not as small and circular as the pores of some Lycoperdon (where this species used to be placed). Odd looking collections have been found much paler than normal and with a well developed base that is much more than pear shaped (see second photo) that could be confused with Lycoperdon perlatum, but that species has prominent spines that leaves scars after they fall off.
Apioperdon pyriforme EU - Our local sequences match hundreds of EU sequences.
Apioperdon pryiforme © NAMA and the Field Museum of Natural History (typical collection) and Sharon Squazzo (unusual collection)
This genus is almost perfectly round without any sterile base. The thin outer skin has a white exterior, and when it wears away the thin inner skin is a distinctive lead/bronze/metallic colour. A pore area opens up on top of the lead coloured inner skin, but the spores don't release as quickly or puff out quite as easily as in Lycoperdon as the interior supposedly doesn't become a pure powder. Most collections are found in grass.
Bovista pila WI - a grass species that supposedly has a small cord on the bottom attaching it to the ground. It is <10 cm in diameter. The metallic inner skin is copper/bronze coloured. One OR sequence matches what I think is an east coast sequence.
Bovista plumbea EU - a smaller grass species (<5 cm in diameter) with a tuft of threads attaching to the ground instead of a single cord. The metallic inner skin is lead coloured. We have a BC sequence that matches a couple dozen EU sequences.
Bovista californica CA - a small species (~1cm in diameter) recorded from wet open alpine meadows and next to alpine lakes and streams once each in OR and WA as of this writing. No DNA yet.
Bovista leucoderma EU/Bovista dakotensis Dakotas/Bovista minor OH&NE/Bovista aestivalis EU - desert species recorded from SW Idaho and/or eastern Oregon but none of these reports have been verified. We have one sequence from an unknown place labeled B. minor, and a bunch of worldwide sequences of B. aestivalis that shows it is found in AZ (although some ambiguous characters make it look like some of the sequences don't match).
unsequenced Bovista pila and Bovista plumbea © Michael Beug
Known as the "acorn" puffball because the top half of the outer layer wears away, but usually not the bottom half, so it ends up looking like an acorn in one stage of growth. The bottom half is often embedded in the soil, partially underground. It's also relatively small like an acorn (<2 cm across). The skin is thin and a pore opens up on top with an elevated margin.
Disciseda cf candida - The spores are round and usually <5u in diameter. I don't know if the type area is the EU or ENA, but purported sequences from various locations don't agree with each other so we need local collections plus reliable sequences of the real thing to compare to.
Disciseda cf subterrana Dakotas - has bigger spores, round with diameter usually >5u. These two species have been reported in the PNW, but we really don't know which species occur here, so we need collections. We have one short ITS2 sequence from CO purporting to be D. subterrana.
unsequenced Disciseda sp. © iNaturalist user veritisia
About 5-20 cm across with a smoothish, white, thick, felty exterior that develops areolate scales and breaks apart somewhat in rays like an earthstar. Calvatia and Lycoperdon species are not felty and don't break apart in rays.
Mycenastrum corium EU - Worldwide sequences seem to all match EU sequences (including AZ) so we probably have this species here, but we don't have any local collections sequenced yet.
Mycenastrum corium © Jonathan Frank (2 images), unsequenced M. corium © Ayla Barten
Calvatia - click to expand
Usually white, often somewhat large (>6 cm in diameter), where the skin breaks away irregularly, not forming a pore at the top. Usually there is not a clear outer and inner layer breaking apart separately, or if so, the interior layer is not shiny (except in the distinctive species C. rubroflava).
Some species of Calvatia have ended up sequencing inside of Lycoperdon, so the actual distinction between the two genera is blurry right now and you'll need to check Lycoperdon too. It should be investigated if the Calvatia-like species that moved to Lycoperdon are the ones where you can easily separate the outer and inner skin (or that have a shiny interior skin). Also consider Mycenastrum, only distinguishable by breaking up into rays like an earthstar when mature and by being somewhat felty when young.
Species mentioned: Langermannia. Calvatia booniana, gigantea, pachyderma, cyathiformis, fragilis, rubrotincta, candida, rubroflava, rugosa, lacerata, paradoxa, owyheensis, pallida, tatrensis var. gruberi, ochrogleba, lloydii.
Some people use the genus Langermannia for species of Calvatia whose exterior can entirely wear away (as opposed to pieces only), but most people, including me, consider it a subgenus especially since I don't think it's true that those species will entirely wear away their outer layer. Those species in that subgenus include C. booniana, C. gigantea, and C. pachyderma.
Calvatia booniana OR - the western giant puffball is usually 20-60 cm across! It is white and covered in polygonal warts/scales which make up a thick exterior layer. It is found in dry or desert climates usually east of the Cascades under sagebrush, juniper or in grass. The exterior eventually wears away exposing the spores. We have an ITS sequence from CO that probably represents C. booniana, but need local collections.
Calvatia gigantea EU - an equally giant desert puffball but with a nearly smooth thinner exterior layer. Most if not all rumours of this species are probably actually C. booniana, so if you think you actually find this species in the PNW, save it. We have many EU sequences to compare to.
Calvatia pachyderma AZ - usually 7-17 cm across, this is another desert species with a nearly smooth thick and hard exterior layer that eventually cracks and breaks off in chunks. Some Lycoperdon are similar, but smaller. Mycenastrum corium is very similar but the outer layer is felty and breaks apart somewhat in rays. We have a CA and an EU sequence that probably represents this, but there has been confusion over what this species is so a type sequence would be nice to prove it is what we think it is. We need local collections to verify its presence here.
unsequenced Calvatia booniana © Andrew Parker
Calvatia cyathiformis EU - the purple puffball, a large initially smooth and whitish desert species 7-13 cm across, eventually cracking into scales and exposing a somewhat thick purple inner skin (although the description says thin). The spores turn the gleba purple in age as well. This species and C. fragilis, described next, may or may not have a significant sterile base that does not change colour like the rest of the interior. Reports that you can separate the two species by the amount of sterile base may not be true. This should be investigated. We have OR and other WNA ITS sequences that match EU sequences of C. cyathiformis.
Calvatia fragilis EU - very similar to C. cyathiformis, and once thought to be a synonym, but the DNA shows it is unique. It is usually about half the size, 4-7 cm across. We do not have any local sequences yet, but we do have reports that sound reliable. Save any small specimens.
Calvatia rubrotincta OR - said to be a Calvatia cyathiformis that had red spots on it (which could have simply been a bacterial infection) and spores that seemed more strongly dextrinoid. It is known only from the type collection, so we need that sequenced, because I don't think it really is a distinct species.
Calvatia cyathiformis © iNaturalist user insectology
Calvatia candida EU - a white species with a smooth, thin skin that sloughs off, between 5 and 10 cm across when grown and the spore mass turns orange before turning olive-brown. One EU sequence suggests this does belong in Calvatia. No local DNA to confirm the report from SW ID.
Calvatia rubroflava ENA - very similar but the exterior stains yellow to orange to red where handled, and the spore mass is orange when mature. Some think it is a synonym of C. candida, but that needs confirmation, as collections labelled with both names do not have the same sequence. We have ENA type area DNA, but no local sequences yet. It is not known if the PNW has only this species or if it also has C. candida. Some synonymize this species with the older species Calvatia rugosa Cuba, but the truth is unknown. One South African sequence purporting to be C. rugosa is a distinct species, but that may not be identified correctly.
Calvatia rubroflava from Indiana © Stephen Russell
Poorly understood species that may not be in Calvatia (many are probably in Lycoperdon).
Calvatia lacerata OR - a white species with a smooth, thin skin that breaks somewhat into rays like the thick skinned Mycenastrum. Found in a Doug fir forest. It is known only from the type collection, so we need that sequenced. Genus not confirmed.
Calvatia paradoxa CO - a 5cm or less whitish puffball with a smooth leathery skin (no report on its thickness) that also opens by radial tearing (somewhat into rays). It was reported from ID. Found in a mountain pasture. We need the type sequenced. Genus not confirmed.
Calvatia owyheensis ID - a 5cm or less white puffball with a warty upper half of the exterior and a furry lower half. Found on soil. No word on how thick the skin is. It is known only from the ID type collection, so we need that sequenced. Genus not confirmed.
Calvatia pallida WA - a 5cm or less whitish puffball with a mostly smooth and very thin skin. The mature spores have a hint of purple and the species was found at high elevation in an open wood. It is known only from the WA and ID type collections, so we need those sequenced. Genus not confirmed.
Calvatia tatrensis EU var. gruberi OR - another rather non descript whitish puffball from 5 to 10 cm across with a smooth, only somewhat thick skin. C. tatrensis is thought by some to be the same as Lycoperdon utriforme EU, so the type of our local variety needs to be sequenced to find out what it really is. We do have many reliable sequences of L. utriforme to compare to, but none from here. See its entry under Lycoperdon for more detail.
Calvatia ochrogleba OR - differs microscopically from C. tatrensis. Found in a meadow. It is only known from the type collection, so we need that sequenced. It is probably a Lycoperdon.
Calvatia lloydii CA - perhaps with a thinner skin, but also said to differ chiefly microscopically from C. tatrensis. It was originally found in ID and CA, but is otherwise unknown. Found in a conifer forest. We need the types sequenced. Genus not confirmed, but probably also a Lycoperdon.
Note that Calvatia sculpta CA, Calvatia subcretacea OR, and Calvata elata New England (=Lycoperdon excipuliforme?) have been confirmed to belong in Lycoperdon and are discussed there.
Lycoperdon - click to expand
Latin for "wolf fart". Typically thought of as smaller species <6 cm in diameter that form a pore at the top to let the spores puff out. I will describe those species first.
Some species of Calvatia and all species of Calbovista, Gastropila, Handkea and Vascellum that I know of have sequenced inside of Lycoperdon so the this genus is now hard to characterize. After the typical species, I will describe those that are larger (6-15 cm across) and do not open up a pore to release the spores. As those resemble Calvatia, check that section too. Finally, some Calvatia that haven't been sequenced yet no doubt still need to be moved to Lycoperdon. It should be investigated if the Calvatia-like species that moved to Lycoperdon are typically the ones where you can easily separate the outer and inner skin (or that have a shiny interior skin). That does not seem to always be the case, as L. vernimontanum n.p. does not have separable layers. One final note: the typical and atypical Lycoperdon do not cluster into two distinct clades, but seem to be somewhat mixed in the tree together, which somewhat surprised me.
Species mentioned: Lycoperdon curtisii, marginatum, pulcherimum, pyriforme, perlatum, dermoxanthum, floccosum, molle, nettyanum, nigrescens, foetidum, rimulatum, subincarnatum, umbrinum, subumbrinum, vernimontanum n.p., excipuliforme, utriforme, bovista. Bovistella utriforme. Calbovista subsculpta. Calvatia bovista, sculpta, subcretacea. Gastropila fumosa, hesperia. Handkea. Vascellum pratense, lloydianum.
Apioperdon pyriforme is described above and usually grows on wood or buried wood, (unusual in Lycoperdon).
These white (when fresh) species are well decorated with pyramidal warts and/or long spines whose tips often touch in one stage of growth.
Lycoperdon curtisii NC/CT - This species is supposedly round without a sterile base, with the smallest diameter (<2 cm) and has the smallest spores, 3-3.5u in diameter. It is often found in pastures. There are 2 clades of sequences with this name with sequences from ENA. One of them has identical ITS to Lycoperdon pratense, which may be more decorated when young than we realize. A third clade only has sequences from CA, far from the type area, so is likely not the real thing, but it may occur here and I don't know what it is.
Lycoperdon marginatum EU - This species has a sterile base-like stem area, and larger (<5 cm in diameter) and has spores 3.5-4u in diameter. After the spines/warts shed, a netted pattern is left on the inner skin, unlike the other two species. It is often found in sandy soil. We have many EU sequences and do know what this ITS sequence is. It has been found in ENA. So has a lookalike species with distinct DNA and no known name yet.
Lycoperdon pulcherimum PN - This species also has a sterile base-like stem area, is also <5 cm in diameter, and has the largest spores, 4-5u in diameter. It is found in forests or clearings. We have one AZ sequence purporting to be this species, but no reliable sequences.
No local DNA yet from any of these well decorated species, but they all have been reported. We need collections.
Lycoperdon pratense EU (=Vascellum pratense EU) - a white compressed sphere (flattened on top) also <5 cm across found in grass. Much of the bottom half is sterile and this can be noticed if it is cut in half. After maturity, a kind of diaphram separates the sterile part from the fertile part. The top opens somewhat irregularly into a large hole, not as small and well-formed as a pore. It shares ITS with many ENA sequences of collections labeled L. curtisii that are indeed highly decorated, but here it is mostly found without (or having lost) its spines. We have many EU type area and worldwide sequences that match some of our local collections.
'Vascellum' lloydandium WA - differs by having the diaphram visible only later in maturity than in L. pratense, plus some microscopic differences. Jarvis' master's thesis showed that it has distinct ITS (but those sequences were never published). We need collections.
Three different species of well decorated puffballs from Indiana © Stephen Russell, Lycoperdon pratense © Bitty Roy
Lycoperdon perlatum EU - Easily recognized by its gem-like spines and large sterile base (one of the largest in the genus). It is <6 cm across (but can be taller due to the base). It starts out pure white but the spines and finally the rest of the fruitbody can blacken in age. After a spine falls off, it leaves behind a scar, as seen in the photo. A couple of EU type area sequences are about 3 bp different than each other, and our own sequences vary by about that much. Some are within 1 bp of an EU sequence, so this appears to be a widespread species.
Lycoperdon perlatum © Buck McAdoo
Lycoperdon dermoxanthum EU - a small (1-2 cm in diameter), almost perfectly round puffball (unusual for Lycoperdon) in grass or waste soil that is white turning brownish in age. It seems relatively smooth but does have small warts that fall away to reveal lighter brown scars on a darker brown background. This species is rarely identified in the PNW, but DNA shows it is common, with many local sequences matching Jeppson's official EU sequences, so we must have been calling it something else or giving up on identifying it to species many times.
Lycoperdon dermoxanthum © Bitty Roy, and NAMA and the Field Museum of Natural History
Lycoperdon nigrescens EU (=Lycoperdon foetidum EU) - significant, dark spines that leave a scar when they wear away on a dark exterior, even when fresh. When young, the colours are almost black. Almost round (<5 cm across). When L. perlatum darkens in age it always has a large base. WA and BC sequences match EU sequences.
Lycoperdon umbrinum EU (=Lycoperdon subumbrinum EU?) - somewhat spiny (but the spines aren't as big as L. perlatum or L. nigrescens). The spines usually don't leave a scar when they wear away, making a mature one appear to be a smooth skinned species. The spines are blackish on a yellow-brown exterior, although the colours seem to be able to be dark or light. The powered mature spore mass is usually olive-brown.
Jeppson described L. subumbrinum because he had a bunch of sequences that were very different than his sequences of L. umbrinum (not even in the same part of the tree). But his sequences of L. umbrinum were mislabeled! They were actually 'Gastropila' fumosa. I have no idea how that mistake was made. He acknowledged that the only difference he could find was coarser ornamentation on the spores of L. subumbrinum than in L. umbrinum. Since spore ornamentations measure in tenths of a micron, to be coarser, we're talking a difference of only some hundreths of a micron! That's no basis to describe a new species. His sequences of the type collections of L. subumbrinum are indeed about 4 bp different than actual sequences of L. umbrinum, but that species seems to have some genetic variation as EU sequences of that species differ amongst themselves by the same amount. And our local sequences differ by the same amount from sequences of both species. I bet it's all one species, and that we have L. umbrinum here in the PNW, and that L. subumbrinum is not a distinct species.
Lycoperdon floccosum ENA - The exterior granules easily wear away to make it appear smooth. There is a sterile base and many rhizomorphs on the bottom. Smith considered it a variety of L. umbrinum. There is one unsubstantiated report that it might have been found here in 1980.
Lycoperdon molle EU - similar to L. umbrinum but the background colour of the exterior may not have yellow tones and the mature spore mass may be reddish brown instead of olive brown. We have EU sequences to compare to, but no local sequences yet of this supposedly common species. We need some.
Lycoperdon nigrescens © Michael Beug (unsequenced), SVIMS (sequenced), and NAMA and the Field Museum of Natural History (sequenced)
Lycoperdon umbrinum © NAMA and the Field Museum of Natural History, and Danny Miller
Lycoperdon nettyanum WA - with a subtly spiny exterior at first, an obvious sterile base and the exterior and interior layers of the skin separating easily. It is similar to Apioperdon pyriforme, reported once from WA. No DNA from anywhere.
Lycoperdon rimulatum WI - similar to L. nettyanum, but the exterior is even smoother. It was reported from OR. We have one possible sequence of this from AZ, but nothing reliable and no local DNA yet.
Lycoperdon subincarnatum NY - small and mostly round (<3 cm in diameter), with dark spines on a pinkish tan background that leave a scar after they fall off. The spore mass is purplish-brown at maturity. Said to be found on hardwood, not on the ground. There are vague reports of this being in the PNW. We have one NY type area sequence that might be this, but no local DNA yet.
Lycoperdon vernimontanum n.p. WA (='Calvatia lycoperdoides' WA nom. illeg., =Handkea lycoperdoides WA) - a round species (<5 cm in diameter) that is white turning brown, delicately decorated with the exterior somewhat cracking into warts. No sterile base. There already was a Calvatia lycoperdoides when Smith described the one from WA, so that name was never legal. I haven't seen sequences of this yet.
Lycoperdon excipuliforme EU (=Calvatia elata New England) - a brownish puffball about 5 cm across on a substantial stem, but with a relatively smooth exterior (unlike L. perlatum). The two are generally regarded as synonyms, and sequences of L. excipuliforme from the EU, ENA, Mexico and here in the PNW all match, so this appears to be a worldwide species so the New England species probably is a synonym. We had only vague rumours that it might be here, but then it was found and sequenced from Victoria, BC.
Lycoperdon utriforme EU (=Bovistella utriforme EU, =Lycoperdon bovista EU, =Calvatia bovista EU) - another whitish puffball from 5 to 10 cm across with a somewhat thick skin that is covered in irregularly shaped scales that form a mosaic and wear off in age. It sits on a somewhat squat sterile base and is found in grass and waste places. This should be looked for in the PNW as Calvatia tatrensis EU is thought by some to be a synonym, and we need to figure out what local variety Calvatia tatrensis var. gruberi OR is. We do have many reliable sequences of L. utriforme to compare to, but none from here.
Lycoperdon excipuliforme © SVIMS
Atypical Lycoperdon formerly/currently in other genera (usually because they are large (>6 cm across) and don't form a pore)
'Gastropila' fumosa OR (=Handkea fumosa OR) - a smooth, thick skinned round, whitish puffball, usually 3-8 cm in diameter, found in conifer forests. The skin may crack a bit like dried mud. When mature, this puffball smells terrible, like a pit toilet. It doesn't always open up much by itself, but the smell can attract rodents to eat it and let the spores out. It needs to be renamed to Lycoperdon fumosum. We have OR, ID, and CA sequences.
'Gastropila' hesperia CA - reported from ID in a more open place than you would find G. fumosa, with harder to find ornamentation on the spores. No word on how bad it smells. It probably needs to be renamed to Lycoperdon hesperia. No DNA yet.
Lycoperdon sculptum CA (=Calvatia sculpta CA) - The exterior warts on the very thick outer skin are crazy looking. It is a large (8-15 cm across) white puffball with a sterile base from high elevation conifer forests. The skins cracks between the warts unevenly to expose the spores. We have an OR sequence that probably represents this CA species.
'Calbovista' subsculpta CA - A large white roundish puffball (6-16 cm in diameter) with large, prominent pyramidal warts that make up a very thick outer layer. It is from high elevation conifer forests. The skin cracks between the warts to unevenly expose the spores. It needs to be renamed to Lycoperdon subsculptum. We have an OR sequence that probably represents this CA species.
'Calbovista' subsculpta var. fumosa ID - a variation with gray color, very rudimentary squamule formation, and spores >5u diameter. We need a type sequence as it is only known from the type collection. It probably needs to be moved to Lycoperdon along with the type variety.
Lycoperdon subcretaceum EU (=Calvatia subcretacea EU) - this round white puffball (2-5 cm in diameter) has pyramidal warts with black tips and a very thick skin. The skin then breaks up irregularly to expose the spores. Jeppson provides official sequences of this species, and we have a matching WA sequence.
'Gastropila' fumosa © Andrew Parker, Lycoperdon subcretaceum © Yi-Min Wang
The "Poorly Understood" Calvatia section contains more species that likely need to be moved to Lycoperdon or that may be duplicates of other other species.
Back to Main Menu