by Steve Nelsen
WHAT SHOULD BE
Of all the genuses, Psathyrella might have had the most convoluted
wrangling about what species belong in it, although there are several rival candidates
among the dark-spored mushrooms. Kauffman, probably the most respected American
mycologist of his day, said in 1918 that as an Agaricus subgenus, Psathyrella
Fries (1838) refers to a tiny genus having the type species Ps. disseminata
Fr. It was characterized by being Coprinus-like, but non-deliquescing,
having black spores, a membranous, plicate to sulcate pileus, slender stem, vanishing
veil, and gills that do not become variegate-dotted (to distinguish it from Panaeolus
and some species of Coprinus). Kauffman says that Peck described 12 species,
but that he had personally only seen one other, cretata (Lasch Fr.), which
I have not seen mentioned more recently either in the U.S. or Europe, so a different
name is apparently used now. This concept of Psathyrella lasted in the
popular U.S. literature through Graham (1944), but has now been replaced by a
completely different one. Everybody calls the new genus Psathyrella (Fr.
Quél.). Singer says (1986) that Quélet decided in Champ. Jura Vosg.
(1872-3) that Fries had gotten the type species of Psathyrella Fr. wrong
because this species "did not fit the synopsis" that Fries published (although
it appears to me to fit the one Kauffman says Fries published). He therefore rejected
Fries' type species and replaced it with Psathyrella gracilis (Fr. Quél.).
That seems reasonable enough; gracilis fits what Kauffman lists as the
synopsis for Psathyrella just fine (as does disseminata). I have
not figured out what Kauffman called gracilis yet, but it is common enough
that he must have had a name for it.
By a logical process I do not pretend
to understand, other authors expanded Psathyrella (Fr. Quél.) into
a huge genus, incorporating part of Hypholoma (Fr. Quél.) and all
of Psathyra (Fr. Quél.). The original Psathyrella type species
disseminata, after spending a while in Pseudocoprinus (Kummer),
has been reabsorbed into Coprinus in most modern books. This appears to
demonstrate again that current mycologists don't really care about macroscopic
features, since the principal reason Fries separated Ps. disseminata in
the first place was that it does not deliquesce. Why they would completely change
the definition of a Friesian genus (and pretend that the Rules of Nomenclature
somehow justify it) escapes me. If one can both change the type species and eliminate
the original one from a genus, the new genus can have no overlap with the old
one, as occurred for Psathyrella. How could people want it to have the
same name, and why would they devise Rules that would make this happen?
Psathyrella has spores that vary from blackish through brown, purplish
brown, and pinkish gray (so spore color doesn't help much to tell if a species
is a Psathyrella), but apparently browns in the yellow-to-orange range
are excluded. Other characteristics are a veil that varies from absent to copious
and persistent (so veil characteristics don't help either), a cap surface that
varies from smooth to grooved all the way to hairy (also of no use), a stem that
is "usually" fragile and whitish, and complex combinations of microscopic features.
This genus concept obviously has nothing whatsoever to do with what Quélet
was talking about in 1872-3 (he recognized both Hypholoma and Psathyra),
so calling the genus Quélet's modification of Fries's name without giving
a reference is only confusing. It solved some other nomenclatural problems: Hypholoma
was hopelessly diverse, containing both the species people want to call Naematoloma
now, and ones that are Psathyrella. I have not seen what was wrong with
the name Psathyra. Quélet used Drosophila for a united genus
in 1886, but it was later declared to be an invalid name, although no one I have
read has bothered to say why. I hesitate to think that the fact that it is used
for a genus of fruit flies has anything to do with it.
Smith published a monograph financed by the National Science
Foundation on North American Psathyrella in 1972, describing over 400 species,
divided into ten subgenera, although a few have only one species each. Most of
the large number of new species reported in it do not appear to have been used
by anyone else since. The majority of them were found either near Ann Arbor, Michigan
(Smith's home), or Lodgepole, Idaho (where he spent summers with his wife, who
both was from there and was working on a paleontology degree in the area). Smith
comments that a good year for Psathyrella occurs only once about every
10--15 years, and that in a good year not only are there an order of magnitude
more individuals than in a bad year, there are also an order of magnitude more
species (hmmm. . .). His monograph finishes with a large appendix containing many
new species, and the comment that a third good year had occurred after the main
part was finished. I have heard a professional mycologist state that, in his opinion,
Smith had a tendency to describe aberrant individuals as if they were species.
Although the greater than 400 species number is sometimes repeated, the only general
book I have seen that describes even as many as 9% of this number is Smith, Smith
and Weber (1979), who cover 35. Singer (1986), who likes a lot of species as well
as most, only recognizes 74 Psathyrella species worldwide, acknowledging
that many more have been named, and mentioning that species concept in Psathyrella
is "not settled" (which appears to most often mean "people still refuse to use
my concept"). Smith's monograph has made it improbable for any amateur to conclusively
name almost any Psathyrella, because Smith's species are based on extremely
detailed microscopic features that I certainly don't know how to demonstrate.
Psathyrellas are the quintescence of "little brown mushrooms" to
most people, and I don't think any are considered good to eat. David Arora (1986)
believes that "They constitute an immense, monotonous, metagrobolizing multitude
of dull, whitish, buff, graying, or brownish mushrooms. . . ." (he does have a
tendency to get caught up in his own rhetoric, doesn't he?). Although Psathyrellas
are quite common, most mushroom books ignore them as completely as possible. Only
five are illustrated in Bessette, Bessette, and Fischer, and twelve in Phillips
(all rather ugly-looking fellows). I agree that many Psathyrellas are quite
ignorable, especially in age, but think that when they are young enough, some
have a quite striking appearance.
Here are photographs of three striking
The cap of this species is a rich reddish brown and the beautiful
corrugations arehighlighted with white, possibly remains of the white veil. It
was found in Baxters' Hollow (Sauk County on an October 1st, and I am calling
it Ps. deliniata (with relatively little hope of it actually being this
This species has deeper brown cap color and a contrasting,
abundant bright white veil that clings to the margin in gobs. The stem is rather
thick for a Pathyrella, and also a bright white. It is also from Baxters'
Hollow, on a September 5th.
I call this Psathyrella the "spring dot-cap" because I still
don't know exactly what it is. We often find it in May, instead of the Morels
we're hunting. It grows on well-rotted wood, has rather persistent hair-like remnants
of a veil on both cap and stem, and is slightly hygrophanous. The picture was
taken at Ferry's Bluff, Sauk County, on a May 13th, and I don't believe that I
have seen this species after the first of June.
Psathyrella species (whose names I naturally am not completely
sure of) provide excellent examples of some technical terms used in books describing
mushrooms. One of the irritating things to a beginner is the frequency with which
people make up special jargon to conveniently describe things about plants. The
jargon words usually don't mean anything in the absence of knowing exactly what
they were made up to designate without wasting a lot of time explaining in real
English words. All areas of science do this; it is necessary to prevent wasted
effort. Nevertheless, using jargon sure makes it harder to "join the club" and
figure out what the descriptions mean.
floccose: Covered with flakey looking scales,
usually arising from tearing of a fragile veil as growth occurs. This Psathyrella,
found on an October 27th at La Riviera City Park, Prairie du Chien (Crawford County),
has both floccose cap and stem (usually called the pileus and stipe by mycologists),
and is probably in Smith's Subgenus Pannucia, Section Pannucia,
which has many floccose species, although it is larger (5 cm) and shaggier than
illustrations I have seen of P. pannucia itself.
hygrophanous: Having sufficiently loosely packed
hyphae in the cap so that water is soaked up like a blotter, producing very large
differences in color between "soaked" and dry conditions. The figure to the right
shows Psathyrella hydrophylla, taken on a May 30th in the UW Arboreteum
in Madison (Dane County), and you can see completely dried as well as completely
hydrated caps close to each other. This is one of the commonest species, illustrated
in most books. You clearly must be very careful about matching colors to ones
in photographs for hygrophanous mushrooms, as their appearance changes greatly
with time, even within a couple of hours after picking. This makes photographs
of strongly hygrophanous species that were carried to a laboratory to photograph,
as for example in Phillips' books, look rather different than they did in the
woods. Most Psathyrella species are at least slightly hygrophanous, although
the degree varies a lot.
evanescent: Fragile and disappearing. The
fragile, wispy grayish veil still remaining on the turned over mushroom cap is
quite evanescent. No trace of such a veil was seen on any of the other caps in
this cluster. It is common not to be able to find traces of evanescent veils and
rings on any of several mushrooms, making you wonder whether there ever was one.
Nevertheless, whether there is a veil is frequently a branch point in keys (and
one of the reasons they are hard to use; if it appears high in the key, you often
have to follow both pathways all the way to their end, or get the wrong answer).
The cap is also hygrophanous, and unusually, dries in streaky patches starting
between the margin and the center. I have no idea what this Psathyrella
is. The picture was taken at Whitefish Dunes State Park in Door County, on a September
virgate: Sometimes `Englished' as "lined",
this means that although the surface of the cap is smooth, it is semi transparent
when hydrated, so darker regions are seen above the gills. When dehydrated, the
cap usually becomes opaque, and the "lines" can no longer be seen (so virgate
mushrooms also change greatly with age). I am not absolutely certain that the
mushroom shown on the right is a Psathyrella. The stems are rather thin
and fragile, but not white. It has a Mycenoid aspect, that is, it looks like some
Mycena species. The photograph was taken on an October 1st at Rowan Creek,
This species is strongly hygrophanous (which would not be
obvious if I had not set one cap that was dried out near the others), clearly
has an evanescent white veil covering the cap (most obvious as the tiny flecks
on the margin of the smallest cap), and rather unusually, also has a persistant
membranous ring, formed from a partial veil between the margin of the cap and
the stem. All of this has unfortunately not allowed me to tell what species of
Psathyrella it actually is. Taken on a September 11th at at Whitefish Dunes
S.P. in Door Count
This Psathyrella species is probably in Subgenus Pannucia,
has persistant white veil remants that hang on the pileus margin in triangular
tufts as well as very fine "hair like" remants on the rest of the cap, and is
faintly virgate. I have seen it several times, both in the spring and fall, but
this picture is from Baxters' Hollow on a June 30th.
This Psathyrella has a cap that is virgate all the
way to its broad umbo and the surface is tacky when wet, as it was in this picture.
I am trying to convince myself that this is Ps. verna. The photo was taken
on a May 17th, at Wyalusing S.P. in Grant County.
This Psathyrella borders on Aurora's "monotonous multitude",
but is seen to have a generous amount of white veil when it is young. The veil
is evanescent and disappears from most specimens the the time the cap is fully
expanded. The fragile, whitish stipe is typical of the genus. The picture was
taken on a September 20th at the Madison School Forest in Verona, Dane County.
This mushroom has a mycenoid aspect but dark spores. Its
pileus (cap) is campanulate (bell shaped) and strongly virgate (transparent when
hydrated) all the way to the umbo (center of the cap when it is raised) revealing
rather close (closely spaced) lamellae (gills), and is strongly hygrophanous ,
drying from the umbo toward the margin (edge of the cap). The cuticle (skin) of
the pileus has very large, swollen cells, which occurs in some species of Psathyrella,
but I am not even certain of the genus of this mushroom. It was photographed on
an October 26th in Wyalusing S.P., Grant County.
AND LIST OF SPECIESPsathyrella (Fr.)Quel. [= Drosophila Quel.
em. K&R (Mos. 3.7.7]: stipe fragile, usually mottled whitish to silvery; pileus
usually pale clay to dark brown when moist, becoming whitish to tan; spores cocoa
to chocolate-black, dull brick red, or purplish gray; in H2SO4
more or less violet to slate gray (distinguishing them from the spores of Paneolus,
which remain black in sulfuric acid) >400 species described from North America,
57 are listed in Gt.Brtn. (The following list of species is arranged according
to Smith's subgenera: A.H. Smith , Mem. N.Y. Bot. Garden, 1972,
24. I do not have this book and have transcribed it from notes it took;
there may be plenty of mistakes! I dropped the names of all species that I have
not seen mentioned elsewhere, which makes the list a lot shorter!)
Lachrymaria =Lachrymaria Pat.: spores verrucose, ornamentation
darkens more in KOH than wall, gills +/- mottled at maturity
- velutina (Fr)Sing.;
- velutina var. boughtoni (Pk.)AHSm.;
- rigidipes (Pk)AHSm;
- subcinnamomea sp.nov. AHSm.
Paneolina: (now many people transfer these species to Paneolus):
brown basidioles in hymen., no veil, spores ornamented (oil lens).
- castaneifolia (Murr)AHSm.
- Subgen. Psathyroides:
(Hypholoma, Drosophila subgen. Pluteopsis): fibril. to squam.
with colored fibrils; spores often angular/subangular
- lachrymabunda (Fr.)Sing. = cotonea(Quel.)KŁhn.
Mycophila: parasitic on Coprinus commatus and atramentarius
- albolutacea; maculata (Parker)AHSm.;
- duchesnayensis AHSm.;
- epimyces (Pk.)AHSm.=Stropharia coprinophyla Pk.
Pseudostropharia: annulus; pileus glabrous or with soon evanescent
fibrils (see also Pannucia sect. app.)
- caputmeduseae (Fr)K&M;
- annulata AHSm.;
- sphagnicola (Maire)Favre;
- kaufmannii sp.nov.AHSm.;
- longistriata AHSm.
- Subgen. Candolleana: no pleurocystidia
and margin first appendiculate (see also Flocculosae of Pannucia)
- incerta (Pk.)AHSm.;
- hymenocephala (Pk)AHSm.;
- candolleana (Fr.)Maire;
- longipes (Pk)AHSm.;
- huronensis snAHSm.;
- waltersii snAHSm.;
- bipellis Quel.=barleae (Bres.)Sm.
- Subgen. Pannucia
(Karst.)AHSm.: outer and/or partial veil well developed appendiculate margin;
- Sect. Appendiculatae: membranous to submembranous
- semivestita (B&B)AHSm.;
- hydrophila (Fr)Mairea
=puiliformis(Bull.:Fr.)Ort. a."Henrici: hydrophila is now puiliformis and the
smaller one that used to be it is laevissima: FldMyc'02,3(1)23;" [Sigh, the
renamers are at it again]
- Subsect. Largae:
- subagraria (Atk.)AHSm.;
- septentrionalis snAHSm.;
- Subsect. Subacutae:
- tsugae snAHSm.;
- Sect. Pannucia: young pileus with well defined
outer veil leaving copious remains; inner veil +/- fibrillose
- typhae (Kalch)P&D;
- vestita (Pk.)AHSm.;
- bifrons (Berk.)AHSm.;
- squamosa (Karst)Mos.;
- gossypina (Bull.:Fr.)Pears.
not in Smith, but is in US;
- carbonicola AHS =pennata(Fr.)Pers.&Dennis
- senex (Pk.)AHSm.
- Subgen. Homophron: incrusting
material (revived in KOH), thick walled pleurocystidia at neck or apex.
- sublaterita; conissans (Pk.)AHSm.;
- saccharinophila (Pk.)AHSm.;
- cernua (Vahl:Fr.)Mos.= papyracea(Pers.:Fr.)
- olympiana AHSm.;
- atomatoides (Pk.)AHSm.
- Subgen. Psathyrella:
veil very slight to absent, various cheiloscystidea
- minima Pk.;
(Fr.)Quel. fma. prona;
- hirta Pk = Psathyra coprobia;
- circellatipes Benoist;
- umbonata (Pk)AHSm.;
- atomata (Fr.)Quel.;
- gracilis (Fr)Quel
= corrugis (Pers.:Fr.)K&R caudata (Fr)Quel;
- ammophila (Dur.&Lev.)Ort.;
- ovatispora Sm. = paneoloides (R.Mre)Arn.;
- spadiceogrisea (Fr)Mre;
- multipedata (Pk) AHSm.,
- microsperma (Pk)AHS;
- agrariella (Atk.)AHSm.
- Subgenus Cystopsathyra:
grannulose veil: Subgenus
Conocybella: cystoderma have big knob at end (lecythiform); no veil:
- kellermanii (Pk.)Sing. OH (only type)
More and More
Psathyrella delineataA crop of what I believe to
be Psathyrella delineata turned up on woodchips at the Madison School Forest (on
the south edge of town of Verona, off Highway 59) this September. This is the
only Psathyrella I have ever seen that could be described as "big" and "stiking"
in appearence. It is an American species described by Peck (I have not yet found
as genus, but since Peck had Coprinus disseminatus as the type species of Psathyrella,
he certainly would not have put this there), and moved to Psathyrella by A. H.
Smith. The mature ones we found were at the high end of the quoted size range
of 3-10 cm., and the stems were up to 10 x 2-2.5 cm. wide, which is actually bigger
than the quoted range (4-10x1-1.6 cm), but the stems are very fat in buttons,
and slim down as they elongate. The caps are extremely rugose, having radiate
grooves that are especially visible in the mature caps, where the color difference
between the peaks and valleys gets large. There is a thick, white veil on the
buttons, but it tends to disappear in age, and more of it ends up on the cap margin
than on the stem, so they are described as lacking a ring. I looked on the internet,
and not surprisingly, found pictures of three clearly different species called
delineata; several smaller Psathyrellas have varying amounts of radiating grooving,
and I had found one earlier that I tried to make into delineata too.
More Psathyrella delineataThis is the only Psathyrella
I have ever seen that could be described as large and substantial. I have not
located the original desription by Peck, nor what genus he put it in. Both Peck
and Kaufmann in his 1918 book have Psathyrella centered upon Coprinus disseminatus
(unless you are behind the times enough to still use Pseudocoprinus, or up to
date enough to break Coprinus into its three new species), so he could not have
called it a Psathyrella. Adrienne and I found them in wood chips at the Madison
School Forest, at the south edge of Verona Township on October 12, 2002, during
a dry period when hardly anything else was out. The fully developed ones were
10 cm. wide, at the high end of the reported range (3-10 cm.) and the stems were
to 10 by as much as 2.5 cm. wide, which is wider than the quoted range (4-10 x
1-1.6 cm.). The fattest stems occur for the buttons, which have ollow but thick-walled
stems, and which narrow as they extend. The caps are strikingly radially grooved,and
especially stricking when older, and the ridges get lighter than the pits. I have
called smaller species that are radiately rugose "delineata" previously, but had
never seen this species before. Looking on the internet, posted pictures of at
least three different species are identified as delineata.
Psathyrella velutinaThis European and American species
that was already described by Fries is often removed to a separate genus, but
Singer and Smith, who never went out of their way to agree on anything, both want
it to be a Psathyrella. The spores are minutely wrinkled, but this is hard to
see without an oil immersion lens, and the very fibrillose cap and veil remnants
that stick tightly to the margin are easier ways to see it. The picture is taken
near Gunflint Lake in northeastern Minnesota.
Psathyrella conopileaThis rather mycenoid-shaped
Psathyrella was found on a WMS foray on Sept. 22, 1995. The caps are to 3 cm wide
and 2 high, and the spores 15-16x7 microns. The most characteristic feature is
not visible in the photograph: tiny dark brown "hairs" on the cap cuticle, to
about 1/4 mm long. These appear to grow more clustered than Smith's description
of conopilea, but fit everything else.
Psathyrella hymenocephalaRather large but thin-capped
clustered Psaths that have a rusty tan color at first. They appeared in the same
place where an elm was removed, in 1984 and 1987, at the end of May.
Psathyrella arenulinaThese are from a Lake Michigan
beach near Jacksonport, Door County. Peck described it as Agaricus (Psilocybe)
arenulina, found in 1876 at West Albany, New York. I do not know how to distinguish
Peck's species from that of Durieu (1796-1878) and L veill (1796-1870), ammophila,
which was described from North Africa earlier. Smith says that both occur in the
same habitat, growing on roots of beach grass
Psathyrella spp.This rather species has a rather
broad umbo and flaring bell shape at the margin, fine whitish veil remnants on
the cap, which was 3.5 cm wide in the larger specimen, which had a stem is 6 cm
long, and 0.35 wide. It has a cellular cap cuticle and 9x4.7 microns spores. I
have no idea what it should be called. Far too many of my pictures of Psathyrellas
fall into this class of unknown species. The picture was taken at Walking Iron
Park in Dane County.
Psathyrella 'rugose'This species is considerably
more delicate than delineata. The largest one had a stem 3x0.4 cm., and the spores
were 6.5 microns long. Its delicately outlined veining pattern on the young caps
is striking. I have no idea what it should be called. It is from Baxters' Hollow,
Sauk County on October 1st, 1988.