Fungal Systematics and Evolution
VOLUME 4
DECEMBER 2019
PAGES 97–170
doi.org/10.3114/fuse.2019.04.08
A revision of the genus Psathyrella, with a focus on subsection Spadiceogriseae
P. Voto1, F. Dovana2, M. Garbelotto3
Via Garibaldi 173, 45010 Villadose, Italy
Department of Life Sciences and Systems Biology, University of Torino, Viale P.A. Mattioli 25, I-10125, Torino, Italy
3
Department of Environmental Science, Policy, and Management; University of California, Berkeley, CA 94720, USA
1
2
*Corresponding author: matteog@berkeley.edu
Key words:
key
molecular phylogenetics
new taxa
systematics
taxonomy
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal
E-mail: p.crous@westerdijkinstitute.nl
Abstract: Specimens belonging to taxa traditionally assigned to the subsection Spadiceogriseae of the genus
Psathyrella were analyzed both morphologically and molecularly. Samples included mainly European collections,
selected GenBank accessions, and specimens of various North American taxa described by Smith (1972) and
deposited at the Herbarium of the University of Michigan (MICH). Three additional taxa from Africa and Central
America were also included. Bayesian and Maximum Likelihood analyses of two loci (ITS and Tef-1α) independently
and together supported the monophyletic nature of the subsection Spadiceogriseae, and identified nine statistically
supported clades within the subsection. North American and European species often fell within the same clade,
suggesting a relatively recent origin of the subsection or human induced intercontinental movement. While this
study determines for the first time that the presence of a white veil is diagnostic for the entire subsection, very few
morphological traits were associated with individual clades, but clades were often distinctively different in terms
of habitat association, suggesting that trophic interactions may have driven the evolution of this group of fungi.
Combined, morphological and DNA analyses revealed both expected and unexpected synonymies. The new
combinations P. vesiculosa, P. ochrofulva and P. sanjuanensis are proposed, and the new species P. rogersiae is
described. New information is provided on the taxonomic status and distribution of several species including P.
P. 85167,
albescens,
P. atrifolia,
P. bivelata, P. fatua, P. kauffmanii, P. aff. kauffmanii, P. incondita, P. infida, P.
Biodiversity agrariella,
Institute, P.O. Box
3508 AD Utrecht,
The Netherlands.
nitens, P. niveobadia, P. phegophila, P. pseudocorrugis sensu Kits van Waveren, P. subnuda. In total, 13 synonymies
were proposed. Based on DNA data, five species of uncertain validity were confirmed as valid, while six species may
be ambiguous and may require an in-depth re-analysis. The information gathered in this study was used to generate
a key to the species of the subsection Spadiceogriseae.
Effectively published online: 21 June 2019.
INTRODUCTION
In the morphology-based infrageneric classification of the
genus Psathyrella by Kits van Waveren (1985), the section
Spadiceogriseae (subgenus Psathyra), type species P.
spadiceogrisea, includes taxa with spores of medium dimension,
pleurocystidia not muricate and mainly utriform with rounded
to obtuse apex, and a veil neither granulose nor strongly
fibrillose-squamulose. In the same 1985 study, Kits van Waveren
further split the section in the two subsections Spadiceogriseae
and Lutenses. Species with gill edges predominantly lined
by large numbers of conspicuous sphaeropedunculate to
clavate cells, here called paracystidia, amidst a minority of
pleurocystidioid cells were placed in the first subsection. Based
on this study (see discussion), we include a white veil as an
additional morphological trait relevant for species within the
subsection.
As we demonstrate in this paper, Kits van Waveren’s
subsection Spadiceogriseae is a well constructed taxonomic
unit which has the important advantage to receive full
recognition by the molecular evidence, even when applied on
a worldwide scale. This subsection only needs to be integrated
with P. ammophila which, evidently because of its large
and dark spores, the Dutch author placed in the subgenus
Psathyrella, keeping and emending Morgan’s (1907) taxon
Deconica Ammophilae [section Ammophilae (Morgan) Kits van
Wav. emend., monospecific].
In the earlier work by Romagnesi (1944) the species of this
homogenous group are placed in section Fatuae (characterized
by colorless gill trama) subsection Spadiceogriseae Romagn.
(type species Drosophila spadiceogrisea), but not all of them
as Drosophila frustulenta sensu Romagnesi which, fide Kits van
Waveren (1985), corresponds to P. clivensis, is the type species
assigned to section Hydrophilae (defined on pigmented gill
trama) subsection Frustulentae. Therefore, Romagnesi (1944)
only partially realized the importance of the parameter of
dominating paracystidia (which he used for the delimitation of
subsection Spadiceogriseae inside section Fatuae) placing it at a
lower level than that of the gill trama pigmentation.
Recently, Örstadius & Knudsen (2012) and Örstadius et al.
(2015) have published important studies on the genus Psathyrella
including a modern revision of some species belonging to
subsection Spadiceogriseae. However, they have not treated this
subsection directly; in their keys formal supraspecific taxa are
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© 2019 Westerdijk Fungal Biodiversity Institute
97
Voto et al.
ignored and dichotomic steps are primarily based on partitions
of spore length.
When we started this study, a number of ambiguities about
European species morphologically belonging to subsection
Spadiceogriseae, many of which present in Kits van Waveren’s
monograph (1985), and not resolved by the two modern
works aforementioned, were still awaiting a definitive formal
recognition by the scientific community (e.g. P. agraria, P.
casca, P. niveobadia, P. phaseolispora, P. phegophila, and P.
pseudocorrugis sensu Kits van Waveren).
More complex and unresolved is the taxonomy of North
American species within the subsection Spadiceogriseae, mostly
because the taxonomic work by Smith (1972) on North American
specimens predated that of Kits van Waveren (1985) by over a
decade. Predominance of paracystidia had no recognition at
any supraspecific level in his systematic plan. He used the taxon
subgenus Psathyrella section Fatuae for species with spores of
medium length and pleurocystidia utriform to broadly rounded
at apex, and used the subsection Fatuae only to delimit the
terrestrial species. Psathyrella kauffmanii, for instance, is in
subgenus Pseudostropharia because of being annulate. The
formal species descriptions themselves show that Smith was
not aware of the taxonomical importance of the morphological
feature of the dominance of paracystidia later used by his
European colleague to define the subsection Spadiceogriseae.
Hence, while he noted the presence of both types of cystidia
on gill edges, he did not define their exact frequency ratio,
save for those cases when clavate cystidia were exceedingly
prevalent. The first aim of this work, therefore, was to search
through Smith’s monograph for those descriptions revealing
a possible inclusion of North American taxa in the subsection
Spadiceogriseae.
As should be expected, the differences in taxonomic
approach by the two systematists resulted in the definition of
species that may be difficult to compare based on their original
description. Thus, the second aim of this study was to provide a
unified classification of a good selection of North American and
European taxa.
Finally, our study attempts to clarify the taxonomic position
of some of the species described by Smith, either characterized
by ambiguously shaped pleurocystidial apices, a trait that makes
uncertain their assignment within sections Spadiceogriseae
or Pennatae, or characterized by unclear gills edge patterns,
a trait that makes uncertain their assignment to subsections
Spadiceogriseae or Lutenses.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
MATERIALS AND METHODS
Samples analyzed
Table 1 lists all of the specimens described included in the study.
In brief, morphological and molecular analyses were attempted
on a total of 98 samples, including 60 North American, one
Central American, 35 European and two African specimens. A
total of 47 holotypes and five paratypes were included in this
study.
We note that due to limited sample size, types vouchers of
the following taxa could not be obtained from the Herbarium
at the University of Michigan (MICH): P. ambusta, P. flexuosipes,
P. incondita, P. carolinensis, P. lactobrunnescens, P. latispora, P.
praecox and P. subcinerascens. Paratypes of P. flexuosipes, P.
98
incondita and P. praecox were loaned to us. Unfortunately, the
type vouchers of some European species, e.g. P. niveobadia,
P. phaseolispora, P. phegophila, and of the South American
species P. macquariensis could not be obtained. Finally, although
we obtained a type for the following species, they failed to
produce a DNA sequence: P. affinis, P. almerensis, P. ovaticystis,
P. psammophila, P. rugulosa, P. solheimii, P. solheimii var.
sanjuanensis, P. tenacipes, P. velibrunnescens, P. yaoundeana.
Table 2 lists all additional sequences used in the ITS phylogenetic
analysis and retrieved both from GenBank or UNITE.
Morphology
Spore color was assessed in water suspensions. Dried material
was first imbibed with either 5 % KOH or 10 % NH4OH. A 5 %
KOH stain was used to determine pigmentation of the marginal
cystidia and of the underlying cells. A 10 % NH4OH dilution
was used to determine traits of pleurocystidia including wall
thickness, presence of incrustation, and pigmentation. Congo
red was used for staining and picture-documentation of
microscopical features of interest.
A number of words in Smith’s monograph that are not
used in the European terminology were interpreted as follows.
Inequilateral (spore shape in profile) = amygdaliform. Terete (spore
shape) = bidimensional, in contrast to flattened tridimensional
spores which need to be described in length, width in side view,
and width in profile. Acute, obtuse, rounded and truncate (spore
pole and cystidial apex): we maintain Smith’s description of four
levels of rotundity, from sharp (acute) through midway (obtuse)
to broadly curved (rounded) or flat (truncate). Distant, close and
crowded (frequency of gills): we substitute the term ‘close’ with
the expression ‘moderately crowded’.
When observed in 10 % NH4OH, pleurocystidia may
show an irrelevant greenish tint, only yellowish to brownish
pigments are considered relevant. The cystidial cells on the
gill edge in Psathyrella are often of two types; cheilocystidia
and paracystidia. We call cheilocystidia those resembling
pleurocystidia, while we call paracystidia, sensu Huijsman (1955),
those having an apically undifferentiated shape (that is clavate,
spheropedunculate, obpyriform, sometimes slender and more
or less cylindrical); the term “dominating” in the text is used to
indicate the predominance on gills edges of either one of the
two types of cells described above. However, two ambiguous
cases can sometimes occur, i.e. either some marginal cells
may have an intermediate shape with a subtle differentiation
at the apex, or the pleurocystidia themselves may also be
undifferentiated in shape, e.g. ellipsoid to clavate. In both cases,
a clear-cut distinction between the two types of marginal cells
is not possible. The shape of cheilocystidia is usually omitted
in our descriptions because it generally mirrors that of the
pleurocystidia, except for a tendency to bear somewhat smaller
dimensions. Sometimes, they may be partially described if they
conspicuously deviate from the main pleurocystidial pattern;
however, cheilocystidia may be characterized by a narrower
apex or a slenderer neck than pleurocystidia.
Presence of a germ pore is assessed in KOH, while its shape
is assessed in water, as it may tend to swell when observed
in alkali. The letter Q indicates the quotient of bidimensional
spores, that is the ratio length/breadth, but when the spores
are tridimensional, the symbols fQ and pQ are used instead to
indicate the quotient in front and, respectively, in profile view
The term avQ stands for the average value of Q.
© 2019 Westerdijk Fungal Biodiversity Institute
Table 1. Data of specimens used in this study. Names previously proposed by other authors are identified by an asterisk.
Species
Coll. ID/Origin
Revised proposed name (*) Ecology
ITS/LSU
Tef 1-α
Drosophila pallidispora Murrill → P. neotropica
A.H. Sm.
Earle572 (NY00649148) type/CU
In soil in a garden
−
−
Psathyrella abieticola A. H. Sm.
Smith58673 (MICH5337) type/US
Under spruce and fir
KC992891
−
P. aff. kauffmanii
PRM897750/CZ
On very rotten trunk of Fagus
AM712260
−
P. affinis A.H. Sm.
MICH11857 type/US
On wet leaves in swamp
−
−
P. agraria Endere ad int.
KR0030008 type/DE
In a mossy cornfied
KU307507
−
? P. velibrunnescens
GenBank Accession Numbers
MICH47933/US
P. albescens
On sandy humus
MF326008
MF521824
MICH65241/US
? P. albescens
In moist soil and moss under Salix
MF325951
−
P. agrariella
MICH47939/US
=?
On sawdust pile
MF325950
−
P. ochrofulva comb. nov.
P. agrariella var. ochrofulva A.H. Sm.
MICH11858 type/US
On sandy humus in cut-over hardwoods
MF325987
−
P. albanyensis A.H. Sm.
MICH11862 type/US
On soil under aspen and conifers
MF325952
MF521823
P. albescens Hesler & A.H. Sm.
MICH11863 type/US
On decaying chip
−
MF521817
P. albescens
MCVE29106/IT
In a park
MF325953
MF521822
P. albescens
MCVE29107/IT
Among brambles and Robinia
MF326009
MF521821
P. albescens
MCVE29111/IT
On humus of Quercus ilex
−
MF521820
P. albescens
MCVE29113/IT
On humus at margin of wood
MF325954
MF521819
P. albescens
MCVE29116/IT
Among hardwood trees
MF325955
MF521818
P. almerensis Kits van Wav.
J. Daams (L) type/NL
On remnants of culms of Typha, Phragmittes and
Cirsium
−
−
P. thujina*
P. almerensis
LÖ31-04/SE
P. thujina*
On moist soil
KC992874
−
P. almerensis
LÖ379-06/SE
P. hellebosensis*
On remnants of Phragmites
KC992873
KJ732768
P. alnicola A.H. Sm.
MICH11868 type/US
Under alder
MF325957
MF521815
P. amarella A.H. Sm.
MICH11869 type/US
On leaf mold in hardwood forest
MF325958
−
P. ammophila (Durieu & Lév.) P.D. Orton
LÖ169-01/SE
On sand dune
KC992871
−
P. ammophila
LÖ359-11/SE
On moist soil
−
KJ732767
P. spadiceogrisea
MICH11873 type/US
On debris in a clearing
MF325959
−
MICH65256 paratype/US
Habitat unknown
−
MF521814
P. atomatoides (Peck) A. H. Sm.
LÖ249-82/SE
Moist on wood in a forest
KC992930
−
P. atrifolia (Peck) A.H. Sm.
MICH32732/US
P. praetenuis
Among cottonwood tree
MF325960
MF521813
P. atrifolia (Peck) A.H. Sm.
MICH65288/US
P. praetenuis
Habitat unknown
−
−
P. atrifolia (Peck) A.H. Sm.
MICH65287/US
=?
Habitat unknown
−
−
P. bivelata Contu
MCVE29104/IT
On grassy soil
MF325961
MF521812
P. bivelata
MCVE29117/IT
On soil in urban flowerbed
MF325962
MF521811
P. candolleana (Fr.: Fr.) Maire
LÖ38-00/SE
In a rich deciduous forest
DQ389720
−
P. psammophila
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
99
P. angusticystis A.H. Sm.
P. argillacea A.H. Sm.
A revision of Psathyrella, subsection Spadiceogriseae
© 2019 Westerdijk Fungal Biodiversity Institute
P. agrariella (G.F. Atk.) A.H. Sm.
P. agrariella
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
100
Table 1. (Continued).
Species
Coll. ID/Origin
P. candolleana
EB990221/IT
P. carinthiaca Voto
MCVE25611 type/AT
P. carminei Örstadius & E. Larss.
LÖ5-09 type/IT
P. clivensis (Berk. & Broome) P. D. Orton
Revised proposed name (*) Ecology
Among debris in clearing of pinewood
GenBank Accession Numbers
ITS/LSU
Tef 1-α
KF281384
−
MF325963
−
KC992880
KJ732773
LÖ182-03/SE
On the great alvar
DQ389683
KJ732774
P. corrugis(Pers.: Fr.) Konrad & Maubl.
LÖ171-01/SE
Attached to buried wood
−
KJ732757
P. corrugis
MCVE29121/IT
On litter of pine-needles
MF325964
−
P. cortinarioides P.D. Orton
MCVE28713/IT
On soil in chestnut and birch wood
MF326010
MF521810
P. cortinarioides
LÖ77-00/SE
Moist, rich with Alnus
KC992936
−
P. ellenae var. yubaensis Thiers & A.H. Sm.
MICH11919 type/US
On humus in conifer woods
MF326011
−
P. ephemera A.H. Sm.
MICH11921 type/US
On black muck in a swamp
MF325965
−
P. fagetophila Örstadius & Enderle
LÖ210-85 (M) type/SE
Woodland among leaves
KC992902
−
P. fatua (Fr.) Konrad & Maubl.
LÖ231-08/SE
On dry sandy soil
−
KJ732772
P. fatua
LÖ132-97 neotype/SE
On nitrophilous soil
DQ389681
−
P. fatua
MCVE29105/IT
On litter of Quercus ilex
MF325966
MF521809
P. fatua
MCVE29108/IT
On buried roots
MF325967
MF521808
P. fatua
MCVE29109/IT
On soil at the top of a ditch bank
−
MF521807
P. fatua
MCVE29110/IT
On litter of Quercus ilex
−
MF521806
P. fatua
MCVE29115/IT
In grass
−
MF521805
P. fatua
MCVE29122/IT
On soil in a park
MF325968
MF521804
P. fatua
MCVE29123/IT
On humus at margin of wood
−
MF521803
P. fatua
MCVE29124/IT
On grassy ground at the top of a river bank
−
MF521802
P. flexuosipes A.H. Sm.
MICH32961 paratype/US
On debris
MF325969
−
P. fragrans A.H. Sm.
MICH5347 type/US
On conifer debris
MF325970
MF521801
P. fulvoumbrina A.H. Sm.
MICH5833 type/US
P. fusca (J.E. Lange) A. Pearson
LÖ287-04/SE
Under aspen and fir
MF417625
MF521800
On calcareous soil
KC992892
KJ732779
P. fusca
Smith75099 (MICH48491)/US
In wet soil
MF325971
MF521799
P. gordonii (Berk. & Broome) A. Pearson & Dennis
MCVE28708/IT
On grassy soil
MF325972
MF521798
P. gordonii
MCVE28710/IT
On horse-manured soil
MF325973
MF521797
P. griseopallida Thiers & A.H. Sm.
MICH11939 type/US
On sandy soil near an oak log
MF325974
MF521796
P. hellebosensis Deschuyteneer & Melzer
MCVE29114/IT
On soil by a river
MF325956
MF521816
P. hirta Peck
LÖ142-00/SE
Cow dung
−
KJ732800
P. incondita A.H. Sm.
MICH36450 paratype/US
On elm wood
MF325975
−
P. incondita
MICH36451 paratype/US
Under pines
MF325976
MF521795
P. infida Quél.
MCVE29119/IT
On moist soil with deciduous trees
MF325977
MF521794
P. umbrosa
P. kauffmanii
=?
Voto et al.
© 2019 Westerdijk Fungal Biodiversity Institute
On humus in spruce forest
On soil in Pinus forest
P. owyheensis
Table 1. (Continued).
Species
Coll. ID/Origin
Revised proposed name (*) Ecology
GenBank Accession Numbers
ITS/LSU
Tef 1-α
P. kauffmanii A.H. Sm.
MICH11962 type/US
On hardwood debris
−
MF521792
P. kauffmanii
MCVE29101/ES
In beech leaf litter
MF325978
MF521793
P. leucotephra (Berk. & Broome) P.D. Orton
MCVE28705/ES
On soil under Populus and Platanus
MF325979
MF521791
MICH11965 type/US
On wet soil along stream
MF325980
−
LÖ201-02/SE
In a copse
−
KJ732778
P. lubrica A.H. Sm.
MICH11957 type/US
P. spadiceogrisea
On muck
−
MF521771
P. thujina*
P. lutulenta Esteve-Rav. & M. Villarreal
AH21379 type/ES
P. magnispora Heykoop & G. Moreno
24929 (AH) type/ES
P. mazzeri A.H. Sm.
MICH11963 type/US
P. albescens
On muddy soil
KC992875
KJ732769
In chalk grassland
KC992863
−
In a pasture
MF325981
MF521790
© 2019 Westerdijk Fungal Biodiversity Institute
P. microrhiza (Lasch: Fr.) Konrad & Maubl.
LÖ185-02/SE
In a rich deciduous forest
−
KJ732765
P. nitens A.H. Sm.
MICH11983 type/US
On debris
MF325982
MF521789
P. nitens
MICH33251/US
On debris
MF325985
MF521787
P. nitens
MICH33252/US
On alder debris
MF325986
MF521786
P. nitens
MICH33249/US
=?
On humus
MF325983
MF521788
P. nitens
MICH33250/US
=?
On humus
MF325984
−
P. niveobadia (Romagn.) M.M. Moser
MCVE30076/IT
On humus around a stump of Fagus
MK400421
MK408675
P. niveobadia
MCVE29102/IT
Around a stump of Fagus
MF325996
MF521780
P. obtusata (Pers.: Fr.) A. H. Sm.
LÖ88-01/SE
In moist soil in forest
DQ389711
−
P. obtusata
MICH49136/US
On hardwood logs
−
−
P. obtusata
MICH65721/US
P. atomatoides
Habitat unknown
MF326013
−
P. obtusata
MICH33261/US
=?
On debris of cedar slash
MF326012
−
P. olivaceopallida A.H. Sm.
MICH11990 type/US
P. kauffmanii
On humus in hardwoods
MF325988
−
P. oregonensis A.H. Sm.
MICH5840 type/US
On conifer wood
MF325989
−
P. ovaticystis Pegler
Pegler1433 (K(M)196311) type/UG
Among leaf litter
−
−
P. owyheensis A.H. Sm.
MICH5357 type/US
In grass near cow dung
MF325990
MF521785
P. pennata (Fr.) A. Pearson & Dennis
LÖ206-03/SE
P. phegophila Romagn.
Tassi97076/FR
P. piluliformis (Bull.: Fr.) P. D. Orton
MCVE29118/IT
P. piluliformis
LÖ162-02/DE
P. praecox A.H. Sm.
MICH49251 paratype/US
P. praetenuis A.H. Sm.
P. pratensis A. H. Sm.
P. kauffmanii
On burnt soil
−
KJ732813
On leaves and debris in calcareous Fagus wood
−
−
On stump of frondose tree
MF325991
MF521784
On stump of Fagus
DQ389699
KJ732808
On soggy leaf mold near edge of pond
MF325992
MF521783
MICH5361 type/US
On debris of cottonwood tree
MF325993
MF521782
MICH12009 type/US
In pasture on sod near cow manure
MF325994
−
Smith67836 (MICH12012) type/US
P.D. Orton (K) type/GB
P. gordonii*
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
101
P. psammophila A. H. Sm.
P. pseudogordonii Kits van Wav.
P. spadiceogrisea
In sandy soil
KC992856
−
On stumps of Fagus
−
−
A revision of Psathyrella, subsection Spadiceogriseae
P. limicola var. subpectinata A.H. Sm.
P. longicauda P. Karst.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
102
Table 1. (Continued).
Coll. ID/Origin
Revised proposed name (*) Ecology
GenBank Accession Numbers
ITS/LSU
Tef 1-α
P. pseudogordonii
Pérez-De-GregorioPG121105
(MCVE28705)/ES
P. leucotephra
On soil under Populus and Platanus
MF325979
MF521791
P. pseudolimicola A.H. Sm.
MICH5366 type/US
On mud in a cow pasture
−
MF521781
P. rogersiae Voto, Garbelotto & Chiarello
MCVE29120 type/IT
In moist plain with hygrophilous plants
MF325995
−
P. rostellata Örstadius
LÖ 228–85 type/SE
Deciduous wood
DQ389693
−
P. rugulosa A.H. Sm.
MICH12040 type/US
On a spruce log
−
−
P. seminuda A. H. Sm.
Smith34091 (MICH12043) type/US
On soil
KC992907
−
P. septentrionalis A.H. Sm.
MICH12045 type/US
On hardwood
MF326014
−
P. septentrionalis var. vesiculosa A.H. Sm.
MICH12044 type/US
P. vesiculosa comb. nov.
On hardwood logs
MF326015
−
On moist soil under aspen
−
−
P. sanjuanensis comb. nov.
Under aspen
−
−
On clayey, mossy soil
KC992877
KJ732770
P. solheimii McKnight & A.H. Sm. Mem.
MICH12054 type/US
P. solheimii var. sanjuanensis A.H. Sm.
MICH12055 type/US
P. sp
NL-0631/HU
P. spadiceogrisea (Schaeff.) Maire
LÖ102-98/SE
Among leaves in a pasture
−
KJ732771
P. spadiceogrisea
LÖ92-01/SE
In a deciduous forest
DQ389682
−
P. spadiceogrisea
Tassi04005 (MCVE29103)/FR
In a park
MF325997
MF521779
P. kauffmanii
P. spadiceogrisea
Smith74295 (MICH49606)/US
Debris
MF325998
−
P. sphaerocystis P. D. Orton
LÖ126-99/SE
Horse dung
−
KJ732803
P. sphagnicola (Maire) J. Favre
LÖ233-99/SE
On Sphagnum with Picea
KC992937
−
P. squamosa (P. Karst.) A. H. Sm.
LÖ194-96/SE
On a sandy roadside
KC992940
−
P. subargillacea A.H. Sm.
MICH12058 type/US
P. psammophila
On rotten wood
MF325999
−
P. subnuda (Karsten) A.H. Sm.
MICH33421/US
P. abieticola
On burned soil
MF326000
MF521778
P. subvinacea A.H. Sm.
MICH12085 Type/US
P. kauffmanii
On underside of logs
MF326001
MF521777
P. sulcatotuberculosa (J. Favre) Einhell.
MCVE29112/IT
On moist soil by a rice field
MF326002
−
P. tenacipes A.H. Sm.
MICH12091 type/US
On drying muck and small sticks in ash swamp
−
−
P. tenera Peck
LÖ81-83/SE
In a moist Alnus forest
KC992849
−
P. thujina A.H. Sm.
Smith66720 (MICH12097) type/US
On black muck
KC992876
MF521776
P. umbrosa A.H. Sm.
MICH12102 type/US
On soil among rotten wood
MF326003
MF521775
P. velatipes A.H. Sm.
MICH12106 type/US
P. ammophila
In a parking strip
MF326004
MF521774
P. velibrunnescens A.H. Sm.
MICH12107 type/US
On Alnus debris
−
−
P. subnuda var. velosa A.H. Sm.
MICH5376 type/US
P. abieticola
On debris of cottonwood tree
MF326005
MF521773
P. velosa
MICH33441/US
Habitat unknown
MF326006
−
P. vesiculocystis A.H. Sm.
MICH5380 type/US
On conifer duff
MF326007
MF521772
P. warrenensis A. H. Sm.
Smith70162 (MICH5382) type/US
P. abieticola
On grassy soil
KC992906
−
P. yaoundeana Mossebo & Pegler
Mossebo42B (K(M)55648) type/CM
Among grass
−
−
Voto et al.
© 2019 Westerdijk Fungal Biodiversity Institute
Species
A revision of Psathyrella, subsection Spadiceogriseae
Table 2. List of sequences used in the ITS analysis without matching morphological analysis, including original Collection ID, origin when available,
and GenBank or UNITE accession number.
Species
Coll. ID/Origin
GenBank/UNITE Accession Numbers
Psathyrella ammophila
O-F-253115/NO
UDB037642
Psathyrella ammophila
LO99/NO
UDB017672
Psathyrella ammophila
AH:947/ES
KY350223
Psathyrella ammophila
SZMCNL 3398
FN396112
Psathyrella ammophila
AH24456
KY350224
Psathyrella ammophila
LO169-01/SE
KC992871
Psathyrella ammophila
AH947
MF966506
Psathyrella ammophila
AH24456
MF966486
Psathyrella casca
AM1814/DE
MF668177
Psathyrella cascoides
AV080419/DE
MK045734
Psathyrella cascoides
DW170430/DE
MK026744
Psathyrella cf. gracilis
SOC997/US
FJ235146
Psathyrella clivensis
AM1913/FR
MK129249
Psathyrella fatua
DD8849/BE
MK182811
Psathyrella fatua
DW160426/DE
MH997496
Psathyrella fatua
SZMC-NL-0603
FN396142
Psathyrella fatua
AM1726/DE
MK072832
Psathyrella fatua
DD0223/BE
MK182825
Psathyrella fatua
AH33718
MF966492
Psathyrella fatua
AH:33718/ES
KY350222
Psathyrella fatua
DD2601/BE
MK182826
Psathyrella fatua
DD2602/BE
MK182822
Psathyrella fatua
DD0222/BE
MK182821
Psathyrella fatua
AH22059
MF966485
Psathyrella fatua
LO231-08
KC992879
Psathyrella fatua
AM445/DE
MK045704
Psathyrella jacobssonii
LO256-92/SE
KC992855
Psathyrella mammifera
HMJAU21908/CN
MG734734
Psathyrella mammifera
HMJAU 37882/CN
MG734740
Psathyrella marquana
AM1693/DE
MF668178
Psathyrella microrhiza
TU106657/EE
UDB011850
Psathyrella microrhiza
TU106614/EE
UDB011827
Psathyrella phegophila
AH45940/ES
KY350219
Psathyrella phegophila
HMJAU 28267/CN
MG734728
Psathyrella phegophila
BRNM705637
AM712277
Psathyrella phegophila
HMJAU 37848/CN
MG734738
Psathyrella prona
303322/US
MK607583
Psathyrella pseudogracilis
TU106565/EE
UDB011802
Psathyrella spadiceogrisea
TU118645/EE
UDB018041
Psathyrella spadiceogrisea
TU106529/EE
UDB011718
Psathyrella spadiceogrisea
TU106528/EE
UDB011717
Psathyrella spadiceogrisea
TU106574/EE
UDB011808
Psathyrella spadiceogrisea
TENN:068273/US
KY744152
Psathyrella spadiceogrisea
AM1244/DE
MK045702
Psathyrella spadiceogrisea
BRNM705637/CZ
AM712276
Psathyrella spadiceogrisea
GENT JR3565/BE
MK045679
Psathyrella spadiceogrisea
AM1242/DE
MK045696
© 2019 Westerdijk Fungal Biodiversity Institute
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
103
Voto et al.
Table 2. (Continued).
Species
Coll. ID/Origin
GenBank/UNITE Accession Numbers
Psathyrella spadiceogrisea
LO102-98/SE
KC992878
Psathyrella spadiceogrisea
AM1671/DE
MK045698
Psathyrella spadiceogrisea
HIAS120608/DE
MK045697
Psathyrella spadiceogrisea
SZMC-NL-0440
FM878024
Psathyrella spadiceogrisea
AM1581/ES
MK045703
Psathyrella spadiceogrisea
CBS 171.47/PT
MH856203
Psathyrella striatoannulata
INB3978642/CR
KY350221
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Psathyrella striatoannulata
INB4162132
KY350220
Psathyrella sublatispora
AH40381/ES
MF966502
Psathyrella sublatispora
LO190-97/SE
KC992854
Psathyrella subspadiceogrisea
HMJAU 35992 type/CN
KY678465
Psathyrella subspadiceogrisea
HMJAU 35996/CN
KY678466
Psathyrella thujina
AM1656
KY680791
Psathyrella thujina
AM1849
KY680792
Pictures of pleurocystidia and marginal cells are used to
show their shape and sometimes their color or incrustations
in ammonia or KOH; otherwise, they are shown in Congo red.
The most relevant spore shapes are illustrated in the Figures,
while spore color is better described in writing, due to possible
distortions of the digital imagery or to the small number of fully
ripe spores.
All species studied and described have clamps and 4-spored
basidia, therefore these two features are generally omitted.
Material studied
Personal collections and duplicates of some of the material
received as a gift are deposited in the fungarium of Museo di
Storia Naturale di Venezia (The Venice Museum of Natural
History, MCVE). All of Smith’s types, paratypes and other
collections used in this study were on loan from the University
of Michigan Fungarium (MICH). When the small size of a type
specimen prevented a loan, the collection manager selected for
us an available paratype from other collections cited by Smith
(1972). Other collections were received on loan from the Royal
Botanic Garden at Kew (K), the Naturalis Biodiversity Center
(former National Herbarium of Leiden, L), the Fungarium of
the New York Botanical Garden (NY), and from private fungaria.
Unfortunately, we were not able to obtain the types of P.
niveobadia, P. phegophila, P. phaseolispora and P. macquariensis.
Table 1 reports all samples studied, including the specimens
sequenced, the GenBank accession numbers of the sequences
generated, host and state of collection, and, when appropriate,
the name assigned to them following our revision.
DNA extraction, PCR and sequencing
DNA was extracted from all dried sporocarps as described by Swo
et al. (2004). Briefly, samples were ground to a fine dust using
a bead mill, suspended in a CTAB extraction buffer before being
frozen using dry ice and then thawed using a heating block set at
70 °C, for three freeze-thaw cycles. Samples were then incubated
at 70 °C for 30 min before being fully submerged in phenol :
104
chloroform : isoamyl alcohol (25:24:1) and centrifuged for 15
min at 13 000 g. Finally, DNA was purified from the supernatant
using the GeneClean Turbo kit (QBiogene, Inc). Samples that
failed to amplify using this extraction procedure were extracted
using the NaOH protocol described by Osmundson et al.
(2013). Briefly, 200 µL of 0.5 N NaOH were added to the ground
lyophilized tissue. Samples were briefly vortexed, then 5 µL were
transferred to a new tube containing 495 µL of 100 mM Tris, pH
8.0; 1 µL of this mixture was used as template DNA in a 25 µL
PCR reaction.
Two genomic loci were selected to conduct the phylogenetic
analyses. The markers were the nuclear ribosomal Internal
Transcribed Spacer (ITS) and the translation elongation factor
1-alpha (Tef-1α). Primers used for the amplification and
sequencing of ITS were 5’-CTTGGTCATTTAGAGGAAGTAA-3’
(ITS1f; forward) and 5’-TCCTCCGCTTATTGATATGC-3’ (ITS4;
reverse) (Gardes & Bruns 1993), while those used for Tef-1α
were 5’-GCYCCYGGHCAYCGTGAYTTYAT-3’ (983 forward) and
5’-ACHGTRCCRATACCACCRATCTT-3’ (1567 reverse) (http://
www2.clarku.edu/faculty/dhibbett/Protocols_Folder/
Primers/Primers.pdf). Amplification reactions were performed
in the following 25 µL reaction mixture: 5× buffer, 0.2 mM
dNTPs, 1.25 U/ µL of GoTaq polymerase (Promega Corp.
Madison, WI), 2.0 mM MgCl2 (Invitrogen Corp. Carlsbad,
CA, USA), 0.50 uM each non-degenerate primer or 0.64 uM
each degenerate primer, and approximately 20 ng of DNA.
The PCR program was: 95 °C for 3 min, followed by 35 cycles
of 95 °C for 40 s, different annealing temperature (53 °C for
ITS, and a touchdown between 66 °C and 56 °C with a 1 °C
decrease at each cycle for Tef-1α) for 55 s, 72 °C for 55 s, and an
extension step at 72 °C for 7 min. The purified PCR amplicons
of ITS and Tef-1α were Sanger-sequenced in-house at the
Forest Pathology and Mycology Laboratory (Berkeley, USA).
All amplicons were forward and reverse sequenced with the
related primers, and consensus sequences were generated
by using the Geneious software, v. 9.0.5 (Biomatters, Ltd).
Chromatograms of each sequence were analyzed by using both
Geneious and SnapGene® Viewer. The minimum acceptable
phred score considered per base was 20.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Molecular analyses
Analyses were performed on the four following datasets: 1)- the
first included 160 ITS sequences, 68 generated by this study, 83
obtained from GenBank and nine from UNITE; 2)- the second
included 72 Tef-1α sequences, 56 generated by this study and
16 selected from GenBank; 3)- the third included both ITS and
Tef-1α sequences for 52 taxa, 43 generated by this study, and
nine selected from GenBank; 4)- the fourth included ITS and Tef1α sequences for 224 taxa, both sequenced in this study and
previously analyzed by Örstadius et al. (2015). The sequences
of Bolbitius vitellinus (AFTOL730) and Mythicomyces corneipes
(AFTOL972) were selected as outgroups in all of the analyses.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
P. thujina AM1656 KY680791
P. lutulenta AH21379 ES Type KC992875
1/87
P. agraria KR0030008 DE KU307507
P. thujina Smith66720 (MICH12097) type US KC992876
1/77
P. almerensis LO31-04 SE KC992874
P. fatua SZMC NL 0603 FN396142
P. rogersiae MCVE29120 IT Type MF325995
P. albescens MCVE29113 IT MF325954
P. spadiceogrisea BRNM705637 CZ AM712276
P. spadiceogrisea GENTJR3565 BE MK045679
1/P. spadiceogrisea AM1671 DE MK045698
P. spadiceogrisea HIAS120608 DE MK045697
P. spadiceogrisea AM1242 DE MK045696
P. spadiceogrisea CBS 17147 PT MH856203
1/74
P. agrariella MICH47933 US MF326008
P. mazzeri MICH11963 US Type MF325981
1/85
P. spadiceogrisea TENN068273 US KY744152
P. albescens MCVE29107 IT MF326009
P. albescens MCVE29116 IT MF325955
P. spadiceogrisea AM1581 ES MK045703
P. spadiceogrisea AM1244 DE MK045702
1/85
P. agrariella MICH65241 US MF325951
P. ochrofulva MICH11858 US Type MF325987
1/95
P. albescens MCVE29106 IT MF325953
0.98/87 P. ammophila AH947 ES MF966506
P. ammophila AH947 ES KY350223
1/P. ammophila AH24456 KY350224
P. ammophila AH24456 MF966486
P. ammophila LO99 NO UDB017672
1/71
P. ammophila OF253115 NO UDB037642
P. ammophila LO169 01 SE KC992871
P. ammophila SZMCNL3398 FN396112
-/87
P. velatipes MICH12106 US Type MF326004
0.99/75
P. hellebosensis MCVE29114 IT MF325956
P. thujina AM1849 KY680792
P. almerensis LO379-06 SE KC992873
1/83
P. fatua AH22059 MF966485
P. fatua AH33718 MF966492
0.98/74
P. fatua LO132-97 SE Neotype DQ389681
P. fatua DW160426 DE MH997496
P. fatua MCVE29122 IT MF325968
P. fatua DD0223 BE MK182825
P. fatua DD8849 BE MK182811
P. fatua AM445 DE MK045704
P. fatua AM1726 DE MK072832
1/86
P. fatua MCVE29105 IT MF325966
P. fatua DD0222 BE MK182821
P. fatua DD2602 BE MK182822
P. fatua MCVE29108 IT MF325967
P. fatua LO231-08 KC992879
P. fatua DD2601 BE MK182826
P. fatua AH33718 ES KY350222
P. carminei LO5-09 IT Type KC992880
0.97/76
P. cf. gracilis SOC997 US FJ235146
P. owyheensis MICH5357 US Type MF325990
-/81
/ spadiceogrisea group
/ agraria
0.03
Fig. 1. Bayesian phylogram obtained from the general nrITS sequence alignment of Psathyrella spp. Bolbitius vitellinus was used as outgroup taxon.
Only BPP values ≥ 0.95 and MLB values ≥ 70 % are given above clade branches.
© 2019 Westerdijk Fungal Biodiversity Institute
105
Voto et al.
P. spadiceogrisea TU118645 EE UDB018041
P. cascoides AV080419 DE MK045734
P. spadiceogrisea LO92-01 SE DQ389682
P. cascoides DW170430 DE MK026744
P. spadiceogrisea LO102-98 SE KC992878
P. spadiceogrisea SZMC-NL-0440 FM878024
P. casca AM1814 DE MF668177
-/74
P. spadiceogrisea TU106529 EE UDB011718
P. amarella MICH11869 US Type MF325958
P. praecox MICH49251 US Paratype MF325992
0.99/P. spadiceogrisea MCVE29103 FR MF325997
1/99
P. clivensis AM1913 FR MK129249
P. clivensis LO182 03 SE DQ389683
P. spadiceogrisea TU106528 EE UDB011717
P. niveobadia MCVE29102 IT MF325996
P. alnicola MICH11868 US Type MF325957
P. oregonensis MICH5840 US Type MF325989
P. striatoannulata INB3978642 CR KY350221
P. striatoannulata INB4162132 KY350220
P. albanyensis MICH11862 US Type MF325952
P. fusca MICH48491 US MF325971
P. olivaceopallida MICH11990 US MF325988
P. kauffmanii MCVE29101 ES MF325978
P. spadiceogrisea MICH49606 US MF325998
P. subvinacea MICH12085 US Type MF326001
P. phegophila BRNM705637 AM712277
P. phegophila HMJAU28267 CN MG734728
P. phegophila AH45940 ES KY350219
P. phegophila HMJAU37848 CN MG734738
1/100
P. mammifera HMJAU21908 CN MG734734
P. mammifera HMJAU37882 CN MG734740
P. subspadiceogrisea HMJAU35992type CN KY678465
1/100
P. subspadiceogrisea HMJAU35996 CN KY678466
1/99 P. spadiceogrisea TU106574 EE UDB011808
P. marquana AM1693 DE MF668178
1/99 P. atrifolia MICH32732 US MF325960
P. praetenuis MICH5361 US Type MF325993
P. griseopallida MICH11939 US Type MF325974
P. sp. NL 0631 HU KC992877
P. fulvoumbrina MICH5833 US Type MF417625
P. umbrosa MICH12102 US Type MF326003
P. prona 303322 US MK607583
P. sublatispora AH40381 ES MF966502
P. sublatispora LO190 97 SE KC992854
P. jacobssonii LO256 92 SE KC992855
1/98
P. ellenae var. yubaensis MICH11919 US Type MF326011
P. rostellata LO228 85 SE DQ389693
1/84
/ sphagnicola
P. flexuosipes MICH32961 US Paratype MF325969
0.96/P. incondita MICH36450 US MF325975
P. sphagnicola LO233 99 SE KC992937
2x
1/100 P. aff. kauffmanii PRM897750 CZ AM712260
/ septentrionalis
P. vesiculosa MICH12044 US Type MF326015
1/78
P. septentrionalis MICH12045 US Type MF326014
2x
1/100
P. gordonii MCVE28708 IT MF325972
/ gordonii
P. gordonii MCVE28710 IT MF325973
1/100
P. cortinarioides LO77 00 SE KC992936
/ cortinarioides
P. cortinarioides MCVE28713 IT MF326010
1/100
P. atomatoides LO249 82 SE KC992930
/ atomatoides
P. obtusata MICH65721 US MF326013
P. limicola var. subpectinata MICH11965 US MF325980
0.97/73
P. fagetophila LO210-85 M SE Type KC992902
-/93
P. warrenensis Smith70162 MICH5382 US KC992906
/ agrariella
1/95
1/P. agrariella MICH47939 US MF325950
1/98
P. seminuda Smith34091 MICH12043 US Type KC992907
P. pratensis MICH12009 US Type MF325994
P. obtusata MICH33261 US MF326012
1/98
P. incondita MICH36451 US MF325976
/ incondita
P. squamosa LO194 96 SE KC992940
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
0.99/-
1/-
/ atrifolia
1/100
0.03
Fig. 1. (Continued).
106
© 2019 Westerdijk Fungal Biodiversity Institute
/ spadiceogrisea group (continued)
1/98
/ albanyensis
1/76
A revision of Psathyrella, subsection Spadiceogriseae
P. subnuda var. velosa MICH33441 US Paratype MF326006
P. vesiculocystis MICH5380 US Type MF326007
P. ephemera MICH11921 US Type MF325965
0.98/1/89
P. abieticola Smith58673 MICH5337 US Type KC992891
0.97/74
P. subnuda var. velosa MICH5376 US Type MF326005
1/98
P. subnuda MICH33421 US MF326000
P. fusca LO287-04 SE KC992892
0.95/76
P. carinthiaca MCVE25611 AT Type MF325963
P. fragrans MICH5347 US Type MF325970 /
1/99
P. piluliformis MCVE29118 IT MF325991
1/P. piluliformis LO162-02 DQ389699
1/99 P. nitens MICH11983 US Type MF325982
P. nitens MICH33251 US MF325985
0.95/100
0.98/99
P. nitens MICH33252 US MF325986
P. nitens MICH33249 US MF325983
1/86
P. nitens MICH33250 US MF325984
/ nitens
0.96/P. angusticystis MICH11873 US Type MF325959
1/99
P. obtusata LO88-01 SE DQ389711
P. psammophila Smith67836 MICH12012 US Type KC992856
P. subargillacea MICH12058 US Type MF325999
P. magnispora 24929-AH ES Type KC992863
3x
0.96/1/100 P. microrhiza TU106614 EE UDB011827
P. microrhiza TU106657 EE UDB011850
1/99
P. infida MCVE29119 IT MF325977
/ infida
P. tenera LO81 83 SE KC992849
P. pseudogracilis TU106565 EE UDB011802
0.97/85
P. corrugis MCVE29121 IT MF325964
P. bivelata MCVE29104 IT MF325961
P. candolleana EB990221 IT KF281384
P. bivelata MCVE29117 IT MF325962
/ candolleana
1/99
P. candolleana LO38 00 SE DQ389720
P. sulcatotuberculosa MCVE29112 IT MF326002
P. leucotephra MCVE28705 ES MF325979
3x
Bolbitius vitellinus isolate AFTOL 730
Mythicomyces corneipes AFTOL 972
0.97/-
6x
1/96
/ abieticola
carinthiaca
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
0.03
Fig. 1. (Continued).
Sequences of each dataset were first aligned using MAFFT v.
7.017 (Katoh et al. 2002) using default conditions for gap openings
and gap extension penalties. Finally, alignments were manually
adjusted using Geneious v. R 11.1.4 (https://www.geneious.
com, Kearse et al. 2012). The model of sequence evolution
GTR +GAMMA+I was selected with the program Modeltest
v. 3.7 (Posada & Crandall 1998). A partition homogeneity test
was performed with PAUP v. 4.0a (build 159) (Swofford 2003)
to assess character congruence between ITS and Tef-1α data
sets. All datasets were analyzed using Bayesian inference (BI)
and maximum likelihood (ML) criteria. The ITS and Tef-1α data
sets were partitioned. BI analysis was performed using MrBayes
v. 3.2 (Ronquist et al. 2012) with four incrementally heated
simultaneous Monte Carlo Markov Chains (MCMC) run for 10
M generations in datasets 1,2,3 and for 20 M generations in
dataset 4. Trees were sampled every 1 000 generations and the
first 25 % trees were discarded as “burn-in” . ML was performed
with RAxML v. 7.2.8. (Stamatakis 2006) in Geneious v. R 11.1.4
(Kearse et al. 2012), using a total of 1 000 bootstrap replicates
(Felsenstein 1985) to assess the relative robustness of the
branches of maximum likelihood in RAxML. Only BPP values
≥ 0.95 and MLB values ≥ 70 % were reported in phylogenetic
trees (Figs 1, 2). Pairwise % identity values of ITS and Tef-1α
sequences were calculated using MEGA v. 10.0.4 (Kumar et al.
2018). BI analyses of the fourth dataset were run on the CIPRES
Science Gateway (Miller et al. 2010).
Molecular data were used to contrast or support the extensive
morphology-based taxonomy, and to resolve uncertainties that
could not be resolved solely based on morphology. Nucleotide
homology calculated using BLAST (Altschul et al. 1990) and
positioning of sequences on phylogenetic trees were both used
to infer presence or absence of molecular conspecificity. When
both ITS and Tef-1α were >98 % homologous, and the taxa from
which both sequences were derived fell as immediate neighbors
on a BI tree, this evidence was regarded as strong support of
conspecificity. ITS and/or Tef-1α homology < 97 % was used to
support lack of conspecificity, while > 98 % homology of the ITS
only was regarded as inconclusive. If homology levels between
two taxa were 97–98 %, the phylogenetic positioning of the taxa
bearing the sequences was analyzed. Additionally, in these cases,
it was analyzed whether heterozygosities or indels were located
in introns or exons, and whether they were synonymous or not.
Non-synonymous exonic heterozygosities and indels in exons
were taken as indication of higher divergence between taxa.
Taxonomy
1. Psathyrella kauffmanii A.H. Sm., Mem. N. Y. Bot. Gard. 24: 69.
1972. Figs 4, 5.
Molecular heterotypical synonyms and misapplied names
proposed here: Psathyrella olivaceopallida A.H. Sm., Mem. N.
Y. Bot. Gard. 24: 138. 1972; Psathyrella subvinacea A.H. Sm.,
Mem. N. Y. Bot. Gard. 24: 124. 1972; Psathyrella fusca sensu
A.H. Smith, Mem. N. Y. Bot. Gard. 24: 355. 1972, p.p.; Psathyrella
spadiceogrisea sensu A.H. Smith, Mem. N. Y. Bot. Gard. 24: 354.
1972, p.p.
Morphological heterotypical synonyms proposed here: Psathyrella carolinensis A.H. Sm., Mem. N. Y. Bot. Gard. 24: 142. 1972;
© 2019 Westerdijk Fungal Biodiversity Institute
107
Voto et al.
P. albescens MICH11863 US Type MF521817
P. agrariella MICH47933 US MF521824
P. albescens MCVE29107 IT MF521821
P. mazzeri MICH11963 US Type MF521790
/ albescens
1/100 P. albescens MCVE29116 IT MF521818
P. albescens MCVE29111 IT MF521820
P. albescens MCVE29113 IT MF521819
1/78
P. albescens MCVE29106 IT MF521822
1/95 P. thujina MICH12097 US Type MF521776
/ thujina
P. lutulenta AH21379 ES Type KJ732769
1/76
P. argillacea MICH65256 US Paratype MF521814
1/71
P. hellebosensis MCVE29114 IT MF521816
/ hellebosensis
1/100
P. almerensis LÖ379-06 SE KJ732768
P.
velatipes
MICH12106
US
Type
MF521774
1/100
/ velatipes
P. ammophila LÖ359-11 SE KJ732767
P. fatua MCVE29124 IT MF521802
P. fatua MCVE29122 IT MF521804
1/89
0.99/93 P. fatua MCVE29108 IT MF521808
P. fatua MCVE29105 IT MF521809
/ fatua
P. fatua MCVE29123 IT MF521803
1/100
P. fatua LÖ231-08 SE KJ732772
P. fatua MCVE29115 IT MF521805
1/P. fatua MCVE29110 IT MF521806
P. fatua MCVE29109 IT MF521807
P. owyheensis MICH5357 US Type MF521785 / owyheensis
1/100
P. carminei LÖ5-09 IT Type KJ732773
1/100 P. praetenuis MICH5361 US Type MF521782
/ praetenuis
P. atrifolia MICH32732 US MF521813
-/94
P. griseopallida MICH11939 US Type MF521796
/ griseopallida
0.99/93 1/100
P. pseudolimicola MICH5366 US Type MF521781
P. umbrosa MICH12102 US Type MF521775
/ umbrosa
1/100 1/100 P. fulvoumbrina MICH5833 US Type MF521800
1/98
P. sp. NL 0631 HU KJ732770
P. lubrica MICH11957 US Type MF521771
P. praecox MICH49251 US Paratype MF521783
-/72
P. spadiceogrisea MCVE29103 FR MF521779
1/87
/ spadiceogrisea
P. spadiceogrisea LÖ102-98 SE KJ732771
1/96
P. niveobadia MCVE29102 IT MF521780
1/96
P. clivensis LÖ182-03 SE KJ732774
P. subvinacea MICH12085 US Type MF521777
P. kauffmanii MICH11962 US Type MF521792
1/99 P. kauffmanii MCVE29101 ES MF521793
/ albanyensis
P. fusca MICH48491 US MF521799
P. alnicola MICH11868 US Type MF521815
P. albanyensis MICH11862 US Type MF521823
P. microrhiza LÖ185-02 SE KJ732765
P. nitens MICH33252 US MF521786
1/100
P. nitens MICH33251 US MF521787
/ nitens
1/99
P. nitens MICH11983 US Type MF521789
P. nitens MICH33249 US MF521788
1/98 P. vesiculocystis MICH5380 US Type MF521772
-/76
P. subnuda var. velosa MICH5376 US Type MF521773 / fusca
1/100
P. subnuda MICH33421 US MF521778
P. fusca LÖ287-04 SE KJ732779
1/100 P. piluliformis MCVE29118 IT MF521784
/ piluliformis
1/100
P. piluliformis LÖ162-02 DE KJ732808
P. fragrans MICH5347 US Type MF521801
0.96/P. hirta LÖ14-00 SE KJ732800
P. cortinarioides MCVE28713 IT MF521810
P. sphaerocystis LÖ126-99 SE KJ732803
P. pennata LÖ206-03 SE KJ732813
1/P. incondita MICH36451 US MF521795
1/100 P. gordonii MCVE28710 IT MF521797
/ gordonii
P. gordonii MCVE28708 IT MF521798
P. corrugis LÖ171-01 SE KJ732757
1/99
P. longicauda LÖ201-02 SE KJ732778
P. infida MCVE29119 IT MF521794
1/100 P. bivelata MCVE29117 IT MF521811
0.97/93
/ bivelata
P. bivelata MCVE29104 IT MF521812
P. leucotephra MCVE28705 ES MF521791
Bolbitius vitellinus AFTOL 730
Mythicomyces corneipes AFTOL 972
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
/ spadiceogrisea group
0.03
Fig. 2. Bayesian phylogram obtained from the general Tef-1α sequence alignment of Psathyrella spp. Bolbitius vitellinus was used as outgroup taxon.
Only BPP values ≥ 0.95 and MLB values ≥ 70 % are given above clade branches.
108
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
P. mazzeri MICH11963 US Type
P. agrariella MICH47933 US
P. albescens MCVE29113 IT
/ albescens
1/100 P. albescens MCVE29116 IT
P. albescens MCVE29107 IT
1/97
P. albescens MCVE29106 IT
1/100
P. thujina MICH12097 US Type
/ thujina
-/85
P. lutulenta AH21379 ES Type
1/100
1/94
P. hellebosensis MCVE29114 IT
/ hellebosensis
P. almerensis LÖ379-06 SE
P. velatipes MICH12106 US Type
1/99
1/100
P. fatua MCVE29122 IT
P. fatua MCVE29108 IT / fatua
1/85
P. fatua MCVE29105 IT
P. owyheensis MICH5357 US Type / owyheensis
1/100
P. carminei LÖ5-09 IT Type
1/100
P. praetenuis MICH5361 US Type
-/81
/ praetenuis
P. atrifolia MICH32732 US
0.99/80
P. griseopallida MICH11939 US Type
1/100
P. umbrosa MICH12102 US Type
1/98
/ umbrosa
P. fulvoumbrina MICH5833 US Type
1/100
P. sp. NL-0631 HU
P. subvinacea MICH12085 US Type
0.97/86
P. kauffmanii MCVE29101 ES
1/99
P. fusca MICH48491 US
/ albanyensis
P. alnicola MICH11868 US Type
P. albanyensis MICH11862 US Type
0.98/78
1/81
P. praecox MICH49251 US Paratype
0.99/96
1/80
P. spadiceogrisea MCVE29103 FR
/ spadiceogrisea
1/99
P. niveobadia MCVE29102 IT
P. clivensis LÖ182-03 SE
P. nitens MICH33252 US
1/100
P. nitens MICH33251 US
/ nitens
1/100
P. nitens MICH11983 US Type
P. sp. MICH33249 US
0.99/1/100 P. vesiculocystis MICH5380 US Type
1/99
P. subnuda var. velosa MICH5376 US Type
/ fusca
1/100
P. subnuda MICH33421 US
P. fusca LÖ287-04 SE
P. incondita MICH36451 US
1/91
0.99/100 P. piluliformis MCVE29118 IT
1/100
/ piluliformis
P. piluliformis LÖ162-02 DE
P.
fragrans
MICH5347
US
Type
0.96/1/100 P. gordonii MCVE28710 IT
/ gordonii
P. gordonii MCVE28708 IT
P. cortinarioides MCVE28713 IT
P. infida MCVE29119 IT
1/100 P. bivelata MCVE29117 IT
1/97
/ candolleana
P. bivelata MCVE29104 IT
P. leucotephra MCVE28705 ES
Bolbitius vitellinus AFTOL 730
Mythicomyces corneipes AFTOL 972
1/88
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
/ spadiceogrisea group
1/100
0.03
Fig. 3. Bayesian phylogram obtained from combined nrITS and Tef-1α sequence alignment of Psathyrella spp. Bolbitius vitellinus was used as outgroup
taxon. Only BPP values ≥ 0.95 and MLB values ≥ 70 % are given above clade branches.
Psathyrella kauffmanii var. exannulata A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 71. 1972; Psathyrella phegophila Romagn., Persoonia
Suppl. vol. 2: 282. 1985.
Pileus 20–80(–90) mm broad, in early stages ovate to obtusely
conical, more or less umbonate campanulate, finally expanding
through convex to plane and more or less wavy to lobed and
rugulose, margin at first inflexed; when young dark, chestnut-
brown to cinnamon-brown or honey-brown, possibly with
violaceous shades, when mature dark greyish brown to tobaccobrown, hygrophanous, discoloring through warm ochraceous,
olivaceous ochraceous, ochraceous brown, ochraceous greybrown, greyish ochre, cinnamon-buff, finally fading to greyish,
buff, greyish ochraceous, pale olivaceous ochraceous, sometimes
with pinkish to pale violaceous grey tints; faintly translucently
striate when moist. Veil white to whitish, when young strongly
© 2019 Westerdijk Fungal Biodiversity Institute
109
Voto et al.
connecting margin of pileus and stipe, when mature leaving
usually fugacious fibrils around the margin of pileus and appressed
fibrils to floccules on lower half of stipe, also appendiculate at
margin, sometimes as a more or less membranous, more or less
incomplete annulus with striate upper surface on stipe. Lamellae
adnate, sometimes with a decurrent tooth; narrow to moderately
broad in age; crowded or moderately crowded; when young
whitish to greyish or brownish greyish, finally brownish grey, dark
brownish grey to purple-brown or dark reddish brown; edge white
fimbriate or not. Stipe 40–120 × (2.5–)3–10(–13) mm, sometimes
gradually broadened towards base, extreme base sometimes
laterally narrowed to a short pseudorhiza; white, not or little
discoloring brownish in age over the basal area; apex pruinose to
sometimes striate-sulcate, sometimes fibrillose to flocculose on
lower portion. Context fragile, thin at margin to moderately thick
at centre in pileus, fragile to firm in stipe, pale to concolorous with
surface. Odor indistinctive to mild. Taste mild to slightly bitter.
Spore deposit dark purple-brown. Spores (6.2–)6.8–10.3(–11) ×
(3.8–)4–5.7(–6) µm, on average 7.5–8.9 × 4.5–5.3 µm, Q (1.4–)
1.5–2(–2.3), avQ 1.6–1.82; in front view elliptic, oval, oblong,
sometimes irregularly angular-oval, irregularly angular-oboval,
angular-oblong, base truncate to subtriangular, sometimes
rounded, in profile adaxially flattened to subamygdaliform or
subphaseoliform, sometimes elliptic; dark reddish brown to dark
red; germ pore little to moderately distinct, mainly more or less
truncate to depressed, 1.4–1.8 µm broad. Basidia 4(2)-spored,
15–34 × 7–11 µm. Pleurocystidia (22–)26–56(–70) × 9–24 µm;
(broadly) utriform to (broadly) ellipsoid or clavate, sometimes
lageniform; apex rounded to obtuse, sometimes subobtuse, often
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
capitate to subcapitate, forked or not, incrusted or not; walls
sometimes or rarely weakly yellowish; numerous to scattered.
Cheilocystidia 19.5–55(–70) × 7.5–20(–25) µm, rarely attenuate
to subacute, pigment like pleurocystidia, scattered, sometimes
moderately numerous or locally numerous. Paracystidia 9–40
× 4–20 µm, sometimes cylindrical-elongate, walls sometimes
inconspicuously yellowish pigmented, rarely incrusted, numerous,
dominating. Cells below marginal cystidia rarely weakly yellowish
pigmented and incrusted. Clamps present.
Habitat: Caespitose to gregarious or scattered, on or connected
to wood or woody debris. The host tree of MCVE29101 and
the types of P. kauffmanii var. exannulata and P. phegophila is
European beech (Fagus sylvatica). The substrates of the types of P.
kauffmanii, P. olivaceopallida and P. subvinacea are indicated only
in general terms, as hardwoods and logs; however, habitat where
types of these species were collected has a dominance of sugar
maples (Acer saccharum), American beeches (Fagus grandifolia),
and northern red oaks (Quercus rubra) or other oaks (Quercus
spp.). The holotype of P. carolinensis was from forest humus at
1 220 m. a.s.l. Kits van Waveren (1985) considers Drosophila
fusca sensu Malençon & Bertault 1970 a misinterpretation of
P. phegophila, and in this case the host tree is a Quercus ilex in
Morocco at an elevation of almost 2 000 m. a.s.l.
Collections examined from MICH: Psathyrella kauffmanii A.H. Sm.,
holotype, Smith 66602; Psathyrella kauffmanii var. exannulata A.H. Sm.,
holotype, Smith 6285; Psathyrella olivaceopallida A.H. Sm., holotype,
Smith 66685; Psathyrella subvinacea A.H. Sm., holotype, Hoseney
Fig. 4. Psathyrella kauffmanii. MCVE2910. Photo M.À. Pérez-De-Gregorio.
110
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 5. Psathyrella kauffmanii. A.H. Smith 66602, MICH11962; MC Marginal Cystidia in KOH; PC Pleurocystidia; S1 Spores in water; S2 Spores in KOH.
641; Psathyrella fusca sensu A.H. Smith, Smith 75099; Psathyrella
spadiceogrisea sensu A.H. Smith, Smith 74295.
Other material examined: France, Île-de-France, Yvelines, forêt de
Saint-Germain, gregarious on woody debris in calcareous beechwood,
16. Nov. 1997, G. Tassi 9776, printed description in Tassi (2010). Spain,
Girona, Santa Pau, la Fageda d'en Jordà, 560 m. a.s.l., gregarious in
beech leaf litter, 7 Oct. 2000, J. Carbó & M.À. Pérez-De-Gregorio JC20001007.11 (duplicate MCVE29101), Fig. 4, Tef-1α MF521793, ITS
MF325978, printed description in Pérez de Gregorio et al. (2010).
Notes: In old exsiccata it often happens that only a limited
number of dark spores may be found; however, the dark color
and not very distinct germ pore of the spores, together with
pleurocystidia shape, are diagnostic when comparing this
species with the other lignicolous species of the subsection. The
sometimes annular veil and the habitat usually represented by
Fagus can also be diagnostic. However, the use of host or habitat
to corroborate the diagnosis should be done with caution,
because Fagus is a known host also for the closely related
species P. niveobadia.
© 2019 Westerdijk Fungal Biodiversity Institute
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Voto et al.
Because we could not examine the original material of P.
phegophila, we have selected a Spanish collection (MCVE29101)
that matches it satisfactorily, and that is fully described by both
text and pictures of the sporocarps in their natural habitat (Pérez
de Gregorio et al. 2010). DNA sequences of the two loci studied
and the morphology are a perfect match with sequences and
morphology of the holotype of P. kauffmanii. Another collection
we consider a good European representative of this taxon comes
from France and was described and illustrated by Tassi (2000),
as voucher 97076. Thanks to Tassi’s courtesy, this sample was
loaned to us, but, unfortunately, DNA extraction from this
specimen failed. It is to be noted that until now, European reports
always lack the annulus, like P. kauffmanii var. exannulata, one
of the synonyms ascertained in this study.
We could not obtain the holotype of P. carolinensis, however,
judging from its original description, and considering all the
variability observed within P. kauffmanii during the course of
this study, we consider it to be another synonym of P. kauffmanii.
Together with all the taxa regarded as conspecific, P.
kauffmanii also creates a compact cluster with three other species
described below: P. albanyensis, P. alnicola, P. oregonensis.
Psathyrella albanyensis has tridimensional and peculiarly
cordiform to pentagonal spores. Psathyrella alnicola is typically
associated with alders. Psathyrella oregonensis is characterized
by high ITS molecular homology with both P. kauffmanii and
P. alnicola (ITS homology was 100 % with P. alnicola and 99 %
with all other comparable material of P. kauffmanii), however,
based on one collection available to us, it can be stated that
this species differs from them because it is found in coniferous
habitats and because it has broader tridimensional spores. Thus,
based on available evidence, we conclude that P. oregonensis is
an autonomous taxon, maybe to be regarded as an intermediate
species between P. kauffmanii and P. albanyensis. It has been often
stated that high homology of ITS sequence among closely related,
albeit distinct, taxa is not necessarily indicative of conspecificity,
and may be due to interspecific horizontal gene transfer followed
by allelic sweeps that effectively eliminate those ITS alleles that
were originally present in the “receiver” fungal species (discussed
in Bergemann et al. 2009). In order to determine the exact
phylogenetic relationships among the above taxa, sequence
information from additional loci will be needed.
Psathyrella striatoannulata, recently described from
central America, is also strikingly near P. kauffmanii, based
both on molecular (approximately 99 % ITS homology) and
morphological data. Both taxa share the following: a striate
membranous annulus, pleurocystidia often undifferentiated and
possibly pigmented and incrusted, the general habit and colors
of the basidiomes. However, its authors report that incrustations
at the apices of the pleurocystidia distinctively stain reddish
brown in NH4, although this feature tends to gradually disappear
in old specimens. A further molecular comparison is needed to
resolve the relationship between these two species. We believe
the descriptions of Psathyrella spec. 3 by Ludwig (2007a) and of
P. obtusata var. aberrans sensu Breitenbach & Kränzlin (1995)
may represent P. kauffmanii.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
1a. Psathyrella kauffmanii A.H. Sm., Mem. N. Y. Bot. Gard. 24:
69. 1972. Fig. 5.
Typus: Psathyrella kauffmanii A.H. Sm. USA, Michigan, Huron
Mts. Marquette Co., Big Bay, 16 Jun. 1963, A.H. Smith 66602
(MICH11962), on hardwood debris. Tef-1α MF521792.
112
Loan of four sporocarps in good condition and a slice of pileus
for micro-morphology investigation.
Spores (6.2–)6.8–8.5(–9.3) × (3.8–)4.3–5.2(–5.7) µm, Q 1.5–1.7;
in front view elliptic to suboval, sometimes (narrowly) oval to
angular-oval, base often truncate to subtriangular, in profile
adaxially flattened to subphaseoliform; few dark red; germ pore
subdistinct, more or less truncate, ± 1.5 µm broad. Pleurocystidia
(22–)33–55 × 13–23 µm, utriform to ellipsoid or clavate; apex
obtuse, sometimes subobtuse, often capitate, sometimes
forked, rarely rostrate, often incrusted; walls sometimes weakly
pigmented; moderately numerous. Cheilocystidia 19.5–49 ×
9.5–18 µm, scattered. Paracystidia 10–35 × 5–19(–24) µm, often
cylindrical-elongate, numerous, dominating.
1b. Psathyrella kauffmanii var. exannulata A.H. Sm., Mem. N. Y.
Bot. Gard. 24: 71. 1972.
Typus: Psathyrella kauffmanii var. exannulata A.H. Sm. USA,
Michigan, Oakland Co., Highland Recreation Area, Haven Hill, 11
Jun. 1937, A.H. Smith 6285 (MICH11961), in a beech wood.
Loan of one sporocarp in good condition and a half pileus for
micro-morphology investigation.
Spores 6.9–9.1 × 4.3–5.6(–6), Q 1.4–1.9; in front view elliptic
to oval, sometimes oblong, (narrowly) angular-oval, base
often truncate to subtriangular, in profile adaxially flattened,
sometimes subphaseoliform; mostly orangish to reddish brown,
very few dark reddish brown to dark red; germ pore subdistinct,
mainly truncate, 1.5–1.8 µm broad, Pleurocystidia 35–47 ×
12–19 µm; utriform, sometimes clavate, often or sometimes
irregular in ventral part; apex rounded, subcapitate or not, often
or sometimes incrusted; walls sometimes weakly yellowish,
numerous. Cheilocystidia 28–42 × 10–18 µm, scattered.
Paracystidia 9–35 × 5–18 µm, walls sometimes yellowish
pigmented, numerous, dominating.
Notes: Our attempts to extract and amplify DNA failed, however
by morphological comparison we think this material can be
viewed as conspecific with P. kauffmanii. We note that Smith
(1972) reports spores color in KOH as ‘cocoa color and remaining
so a long time’, therefore not dark; however, we had difficulty in
finding dark spores in many old exsiccata of P. kauffmanii and
conspecific materials.
1c. Psathyrella olivaceopallida A.H. Sm., Mem. N. Y. Bot. Gard.
24: 138. 1972.
Typus: Psathyrella olivaceopallida A.H. Sm. USA, Michigan,
Cheboygan Co., Burt Lake, Colonial Point, 20 Jun. 1963, A.H.
Smith 66685 (erroneously reported as 4991) (MICH11990),
scattered to gregarious on humus in hardwoods. ITS MF325988.
Loan of one sporocarp lacking a slice of pileus for micromorphology investigation.
Spores (7–)7.3–9.2(–10) × 4.1–5(–5.1), Q 1.55–1.9(–2) µm;
in front view elliptic, oblong to narrowly oval, sometimes
oblong- to angular-oval, base often truncate to subtriangular,
in profile adaxially flattened, sometimes subphaseoliform;
few dark reddish brown; germ pore moderately distinct, often
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
more or less truncate, 1.4–1.5 µm broad. Pleurocystidia 29–43
× 10–22 µm; broadly utriform, utriform, ellipsoid, sometimes
lageniform, clavate; apex rounded to obtuse, subcapitate or not,
often forked, sometimes incrusted; walls sometimes weakly
yellowish, numerous. Cheilocystidia 28–43 × 10–20 µm, in part
(broadly) lageniform with short to elongate neck and subobtuse
to subacute apex, pigmented like pleurocystidia, scattered.
Paracystidia –28 × –15 µm, often weakly yellowish, numerous,
dominating. Cells below marginal cystidia hyaline.
Notes: Although the molecular evidence provided by the ITS
sequence alone is inconclusive as stated above, we retain this
species a synonym of P. kauffmanii. Smith may have found its
only collection when the sporocarps were already discolored:
the pale olive tint implied by the name and the greyish and
pinkish tones on fading reported in Smith’s (1972) description
belong to the chromatic set of the pileus of P. kauffmanii.
1d. Psathyrella subvinacea A.H. Sm., Mem. N. Y. Bot. Gard. 24:
124. 1972.
Typus: Psathyrella subvinacea A.H. Sm. USA, Michigan,
Washtenaw Co., Waterloo State Recreation Area, 31 Aug. 1967,
F. Hoseney 641 (MICH12085), on underside of logs on wooded
side of sand hill. Tef-1α MF521777, ITS MF326001.
Loan of three sporocarps in good condition for micro-morphology
investigation.
Spores 6.8–9.2(–10.5) × 4.2–5.2(–5.7), Q 1.4–1.8(–2.2); in front
view elliptic, oval to oblong-oval, sometimes (narrowly) angularoval, base rounded, truncate to subtriangular, in profile elliptic
to subamygdaliform, sometimes subphaseoliform; almost all
spores pale to very pale, mostly hyaline to very pale purplish
brown, some violaceous orange, very few dark violaceous brown;
germ pore indistinct. Pleurocystidia 32–50 × 10–24 µm; (broadly)
utriform; apex often subcapitate, sometimes forked, sometimes
incrusted; walls often weakly yellowish below apex. Cheilocystidia
26–42 × 12–19 µm, pigmented like the pleurocystidia, scattered.
Paracystidia 9–40 × 6–18 µm, walls often weakly yellowish
pigmented, numerous, dominating. Cells below marginal cystidia
weakly yellowish pigmented, sometimes incrusted.
Notes: Apart from the very pale pigmentation of the spores,
which was likely due to delayed maturation, both molecular and
morphological characters identify this taxon as conspecific with
P. kauffmanii.
1e. Psathyrella spadiceogrisea sensu A.H. Smith p.p.
Material examined: USA, Michigan, Jackson Co., Munith woods, 13 May
1967, A.H. Smith 74295 (MICH49606), debris. ITS MF325998.
Loan of three full sporocarps and a pileus, in good condition for
micro-morphology investigation.
Spores 7.5–10.3(–11) × 4.2–5.3(–5.6) µm, on average 8.9 × 4.7
µm, Q (1.4–)1.6–2(–2.3), avQ 1.73–1.83; in front view oblong,
elliptic, narrowly oval, rarely oblong-oval or angular-oval, base
rounded, rarely truncate to subtriangular, in profile adaxially
flattened to subamygdaliform, sometimes subphaseoliform;
reddish brown to dark red; germ pore moderately distinct,
mostly more or less truncate to subdepressed, ± 1.5 µm broad.
Pleurocystidia 28–51 × 13–22 µm; (broadly) utriform, sometimes
ellipsoid; apex rounded, often subcapitate, often forked,
sometimes incrusted; walls hyaline; scattered. Cheilocystidia
30–41 × –15(–18) µm, scattered. Paracystidia 10–30 × 5–14 µm,
often yellowish, numerous, dominating. Cells below marginal
cystidia scarcely yellowish pigmented or incrusted.
Notes: We have attempted to understand Smith’s concept of P.
spadiceogrisea by examining a Smith’s collection selected for
this purpose by MICH staff. For this material, we have obtained a
molecular sequence only for the ITS, which places it in the same
subclade as P. kauffmanii. Its terete spores exclude conspecificity
with P. albanyensis and P. oregonensis. Nonetheless, we have
compared it with P. kauffmanii and found a good correspondence
between the two. Because the only relevant difference between
the two is the slightly greater spore length and, consequently,
the slightly greater spore quotient, we retain this collection as a
synonym of P. kauffmanii. Unfortunately, we could not discover
any significant details about the habitat in which these samples
were collected. This suggested synonymy does not necessarily
apply to the other vouchers Smith deposited in MICH under this
name, which should be considered and evaluated separately.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
1f. Psathyrella fusca sensu A.H. Smith p.p.
Material examined: USA, Michigan, Oakland Co., Proud Lake, 3 Sep.
1967, A.H. Smith 75099 (MICH48491), in wet soil. Tef-1α MF521799,
ITS MF325971.
Loan of three sporocarps in good condition and a fragment of
pileus for micro-morphology investigation.
Spores 6.9–8.1 × 4.3–5.4 µm, in front view oval to elliptic, often
angular-oval, base often truncate to subtriangular, in profile
adaxially flattened, sometimes subphaseoliform; dark reddish
brown to dark red; germ pore moderately distinct, often more
or less truncate, 1.4–1.7 µm broad. Pleurocystidia 36–56 × 14–
19 µm; utriform; apex rounded, often subcapitate, sometimes
forked, no incrustation seen; walls hyaline; moderately
numerous. Cheilocystidia 30–55 × 11–20 µm, scattered,
sometimes locally numerous. Paracystidia 12–31 × 6–20 µm,
often weakly yellowish, sometimes incrusted, numerous,
dominating.
Notes: We have attempted to understand Smith’s concept
of P. fusca by examining a Smith’s collection selected for
this purpose by MICH staff. It has turned out to correspond,
both morphologically and molecularly, to P. kauffmanii. This
result does not necessarily apply to the other vouchers Smith
deposited in MICH under this name.
2. Psathyrella alnicola A.H. Sm., Mem. N. Y. Bot. Gard. 24: 176.
1972. Fig. 6.
Typus: Psathyrella alnicola A.H. Sm. USA, Idaho, Idaho Co.,
French Creek Grade, Salmon River, 6 Sep. 1964, A.H. Smith
70222 (coll. K.A. Harrison) (MICH11868), under alder. Tef-1α
MF521815, ITS MF325957.
Loan of one sporocarp in good condition and a half pileus for
micro-morphology investigation.
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Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 6. Psathyrella alnicola. A.H. Smith 70222, MICH11868; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
Spores 7.9–10 × 4.6–5.7(–6.2) µm (8–11 × 5–6 µm in Smith
1972), Q 1.45–2; in front view elliptic, oval to oblong,
sometimes (narrowly) angular-oval, base rounded to truncate,
sometimes subtriangular, in profile adaxially flattened, elliptic
to subamygdaliform, sometimes subphaseoliform; dark
reddish brown to dark red; germ pore subdistinct, more or less
truncate, 1.5–1.7 µm broad. Pleurocystidia (37–)45–67(–83)
× (10–)13–20(–32) µm; utriform to conical-lageniform, rarely
subcylindrical; apex obtuse to rounded, obscurely subcapitate or
not, sometimes forked, sometimes incrusted; walls sometimes
weakly pigmented, moderately numerous. Cheilocystidia 35–62
× 15–17 µm, scattered. Paracystidia 8–48 × 4–25 µm, sometimes
cylindrical-elongate, sometimes weakly pigmented, rarely
incrusted, numerous, dominating.
Notes: The holotype and other five collections determined by
Smith are undoubtedly associated with alder (Alnus). Collection
of the holotype was made from under a single alder, even if
many Ponderosa pines (Pinus ponderosa) and Douglas-firs
(Pseudotsuga menziesii) coexisted in the same site. The 99
% identity of both ITS and Tef-1α sequences with all other P.
kauffmanii studied here clearly identifies a very close relationship
between the two taxa, however, the hygrophilous Alnus habitat
is clearly distinct from the mesophilous habitat characteristic of
P. kauffmanii.
3. Psathyrella solheimii McKnight & A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 67. 1972. Fig. 7.
114
Typus: Psathyrella solheimii McKnight & A.H. Sm. USA, Wyoming,
Albany Co., Laramie Mts., Pole Mountain, Happy Jack Winter
Sports Area, 27 Jun. 1944, W.G. Solheim 2178 (MICH12054), on
moist soil under aspen, especially around beaver dams.
Loan of two portions of pilei for micro-morphology investigation.
Spores (7.4–)7.5–9.6(–10) × 4.9–5.6 µm (broader, 8–10 × 5–6.2
µm, in Smith 1972), Q (1.4–)1.5–1.9, avQ 1.66; in front view oblong,
elliptic, narrowly to cylindrical- oval, sometimes subangularoval, base broadly rounded, truncate to subtriangular, in profile
adaxially flattened to subelliptic, sometimes subamygdaliform,
rarely subphaseoliform; dark red-brown; germ pore moderately
distinct, truncate to subrounded, 1.1–1.7 µm. Pleurocystidia
(24–)30–45(–52) × (10–)11.5–21 µm, ellipsoid-utriform,
fusiform-utriform, utriform, (broadly) clavate, sublageniform,
ellipsoid, sometimes ellipsoid mucronate, obconical; apex
broadly rounded to obtuse, rarely subacute, sometimes forked,
rarely subcapitate, not incrusted; scattered. Cheilocystidia 17–
33(–38) × 7–18 µm, sometimes lageniform-mucronate, hyaline,
scattered to moderately numerous. Paracystidia 10–22(–25) ×
5.5–16(–18) µm, numerous, dominating; intermediate cells with
slightly bulging to submucronate apex, 17–32 × 11.2–17.5 µm,
scattered to moderately numerous towards the stipe. Marginal
cystidia and cells below them hyaline or very pale, not incrusted.
Notes: Unfortunately, DNA extraction of this material failed,
but based on the morphological evidence, we consider it to
belong to the subsection Spadiceogriseae. Smith (1972) keys it
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 7. Psathyrella solheimii. W.G. Solheim 2178, MICH12054; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
just beside P. kauffmanii, stating that the latter differs in longer
and often forked pleurocystidia, and in slightly larger spores.
This was possibly true when comparing P. solheimii to Smith’s
description of P. kauffmanii only, but when we take into account
all of the materials we have studied identified as P. kauffmanii,
we see those divergences disappear. A difference we can suggest
is that the pleurocystidia apices often tend to have a tapering
shape in P. solheimii, while in P. kauffmanii they tend to have
a more rounded, often capitate, shape. Moreover, the habitat
under aspen on moist soil, especially around beaver dams, is
in contrast with the mainly dry and beech dominated habitat
associated with P. kauffmanii. We have also checked P. solheimii
against P. alnicola, given they both share alder as habitat, but
their respective pleurocystidia show a marked disparity: on
average they are shorter and thicker in P. solheimii, while they
are longer and slenderer, and often with an elongate apex, in
P. alnicola. Moreover, gill edge cells are often mucronate in P.
solheimii and never so in P. alnicola. Until new molecular data
are made available, we opted to retain P. solheimii as a separate
species.
4. Psathyrella sanjuanensis (A.H. Sm.) Voto, Dovana & Garbel,
comb. nov. MycoBank MB831131. Fig. 8.
Basionym: Psathyrella solheimii var. sanjuanensis A.H. Sm., Mem.
N. Y. Bot. Gard. 24: 68. 1972. A.H. Smith 51837, MICH12055.
Typus: Psathyrella solheimii var. sanjuanensis A.H. Sm. USA,
Colorado, San Miguel Co., Ophir, 4 Aug. 1956, C. Barrows & A.H.
Smith (A.H. Smith 51837) (MICH12055), under aspen.
Loan of two small fragments of pilei for micro-morphology
investigation.
Spores (8–)8.2–9.8(–10.7) × (5–)5.2–5.8(–6) µm, on average
9.3 × 5.5 µm, Q (1.4) 1.5–1.9(–2.15), avQ 1.7; in front view
elliptic, sometimes cylindrical, narrowly oval, oblong, base
broadly rounded to subtruncate, sometimes obtuse, in profile
subamygdaliform to adaxially flattened, rarely subphaseoliform;
dark red-brown; germ pore distinct, rounded to subtruncate,
sometimes slightly eccentric in profile, 1.3–1.7 µm.
Pleurocystidia 33–50 × 10–16 µm, utriform to lageniform, often
elongate in upper part; apex rounded, often subcapitate, rarely
capitate, sometimes forked with up to 20 µm long protrusions;
scattered. Cheilocystidia 24–40(–45) × 7.5–13(–15) µm, in part
fusiform-lageniform with obtuse apex, apex rarely subcapitate
or forked, numerous to abundant and dominating from mid
gill towards the stipe, scattered to absent towards the pileus
margin. Paracystidia 12–25(–30) × 7–13(–16,5) µm, absent or
inconspicuous towards the stipe, numerous and dominating
towards the pileus margin. Marginal cystidia and cells below
them not pigmented, not incrusted.
Notes: The longest strip of gill edge we could examine was 4.5
mm long. Starting from the pileus margin towards the stipe
end, we found that paracystidia went from being numerous
and dominating (in 2 mm long gill segment) to being absent
or inconspicuous. Conversely, cheilocystidia showed an
inverse trend and went from being absent or rare to being
numerous, even abundant, and dominating (in 2.3 mm long
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Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 8. Psathyrella solheimii var. sanjuanensis. A.H. Smith 51837, MICH12055; PC Pleurocystidia; MC1 Marginal Cystidia towards the pileus margin;
MC2 Marginal Cystidia towards the stipe; S1 Spores in water; S2 Spores in KOH.
gill segment). Paracystidia were small and had approximately
the same dimensions as basidia [20–31 × 8–12(–15) µm], even
where they were dominating. Although we lack the support of
molecular evidence, we assume this collection does not belong
to the subsection Spadiceogriseae, rather it better fits into the
sister subsection Lutenses. Based on such cumulative evidence,
we see no reason to maintain its affiliation with P. solheimii,
and propose to elevate it to the species rank. Interestingly,
Smith (1972) too had suggested this taxon could ‘eventually
deserve recognition as an autonomous species’, a statement he
made based mainly on the shape of pleurocystidia and on the
consistency and colour of the annulus.
5. Psathyrella oregonensis A.H. Sm., Mem. N. Y. Bot. Gard. 24:
157. 1972. Fig. 9.
Typus: Psathyrella oregonensis A.H. Sm. USA, Oregon, Clackamas,
Beaver Creek, Mt Hood National Forest, 24 Oct. 1947, A.H. Smith
28182 (MICH5840), on conifer wood. ITS MF325989.
Loan of one sporocarp with incomplete pileus for micromorphology investigation.
Spores 7–10(–10.7) × 5–6.7 × 4.4–5.5 µm, fQ 1.3–1.7, avQ
1.5, pQ 1.4–1.9, avQ 1.7; in front view oval, elliptic, angularto triangular-oval, base mainly truncate, in profile elliptic,
subamygdaliform, subphaseoliform; reddish brown to dark
red; germ pore little distinct, 1.4–1.7 µm broad. Pleurocystidia
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(31–)37–66(–76) × 11.5–17 µm; slenderly utriform, sometimes
thickly utriform, sublageniform, subcylindrical, subconical; apex
rounded to obtuse, often subcapitate, sometimes forked in
Smith’s description, sometimes elongate, sometimes incrusted;
walls hyaline; numerous. Cheilocystidia 30–43 × 15–20 µm,
scattered. Paracystidia –39 × –21 µm, sometimes rhomboid,
almost hyaline, numerous, dominating.
Notes: Psathyrella latispora is one of the holotypes MICH
could not loan, however, based on its description, it belongs
to subsection Spadiceogriseae. Its type was collected cespitose
from unspecified coarse wood debris, and shares a number
of similarities with P. oregonensis from which it is mainly
differentiated for having terete spores.
6. Psathyrella rugulosa A.H. Sm., Mem. N. Y. Bot. Gard. 24: 158.
1972. Fig. 10.
Typus: Psathyrella rugulosa A.H. Sm. USA, Tennessee, Sevier Co.,
Great Smoky Mts. National Park, near Clingman’s Dome, 24 Jun.
1945, L.R. Hesler 17027 (MICH12040), on a spruce log.
Loan of one sporocarp in good condition for micro-morphology
investigation.
Spores (6.2–)7.5–8.8(–9.8) × (4–)4.3–5.2(–5.3) µm, Q (1.4–)
1.5–1.8(–2.3), avQ 1.7; in front view elliptic to suboval, base
mainly obtuse to rounded, in profile adaxially flattened to
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 9. Psathyrella oregonensis. A.H. Smith 28182, MICH5840; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
Fig. 10. Psathyrella rugulosa. L. R. Hesler 17027, MICH12040; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
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subphaseoliform, sometimes narrowly subamygdaliform or with a
faint suprahilar depression, rarely phaseoliform; pale, few at most
dull orangish; germ pore little distinct (callus), mainly subtruncate.
Pleurocystidia 38–65 × 9–17 µm, utriform to subcylindrical, often
with elongate-cylindrical neck, rarely subellipsoid or ventricoseconical; apex (4–)6–10 µm broad, rounded, sometimes obtuse to
subacute, often (laterally) subcapitate, rarely forked, sometimes
incrusted; numerous. Cheilocystidia up to 60 × 15 µm, scattered.
Paracystidia –30 × –20 µm, numerous, dominating.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: Smith (1972) reported spores with distinct and narrowly
truncate germ pore. Unfortunately, DNA extraction failed on
this material; however, general morphological evidence places
it into subsection Spadiceogriseae. Due to its pale spores and
indistinct germ pore it resembles P. clivensis, however, because
of its appendiculate veil and conifer habitat, Smith (1972) placed
it near P. oregonensis, from which it differs because of its pale
and terete spores.
7. Psathyrella albanyensis A.H. Sm., Mem. N. Y. Bot. Gard. 24:
140. 1972. Fig. 11.
Morphological heterotypical synonyms proposed here:
Psathyrella subcinerascens A.H. Sm., Mem. N. Y. Bot. Gard. 24:
172. 1972.
Typus: Psathyrella albanyensis A.H. Sm. USA, Wyoming, Albany
Co., Laramie Mts., drainage at the northeast base of Pole
Mt., 27.V.1956, W.G. & R. Solheim 4438 (MICH11862), on soil
under aspen and conifers at 2 438 m a.s.l. (8 000 feet). Tef-1α
MF521823, ITS MF325952.
Loan of three sporocarps in good condition and a pileus for
micro-morphology investigation.
Spores (6.2–)6.7–8.3 × 4.5–6.3 × 4.2–5.2 µm, (7–9 × 5–6 × 4–5 µm
in Smith’s description), fQ (1.2–)1.3–1.5(–1.6), pQ 1.45–1.7(–2.2);
in front view oval, elliptic to subcylindrical, often subtriangular or
cordiform to subpentagonal, sometimes subglobose, base often
truncate, apex sometimes attenuate and snout-like projected, in
profile elliptic, sometimes subamygdaliform; reddish brown to
dark red; germ pore indistinct, sometimes distinct but very small,
at most 1 µm. Pleurocystidia 31–45 × 12–17(–19) µm; utriform,
ellipsoid-rhomboid, sometimes fusiform, subcylindrical, apical
part often attenuate to conical or submucronate, apex obtuse
to sometimes subobtuse or subrounded, not incrusted, walls
hyaline; moderately numerous. Cheilocystidia 19–40 × 14–
15(–21) µm, scattered. Paracystidia 10–33(–36) × 6–16(–23)
µm, with walls weakly yellowish pigmented and sometimes
incrusted, numerous, dominating. Cells below marginal cystidia
weakly yellowish pigmented and with incrustations.
Notes: The molecular analysis nests this taxon within the P.
kauffmanii complex; morphologically it is characterized by often
tridimensional, often pentagonal-like spores, both pleuro- and
cheilocystidia showing a tendency to an attenuate and conical
Fig. 11. Psathyrella albanyensis. W.G. & R. Solheim 4438, MICH11862; PC Pleurocystidia; MC Marginal Cystidia in KOH; S1 Spores in water; S2 Spores
in KOH.
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A revision of Psathyrella, subsection Spadiceogriseae
apex, small and slender sporocarps with reddish rusty tints on
the pileus. Its habitat occurs at very high elevations (> 1 500 m
a.s.l.), in association with conifers and occasionally with aspen
(Populus tremuloides). Psathyrella oregonensis appears to be its
closest relative, based on its tridimensional spores and habitat
represented by conifers.
Psathyrella subcinerascens is represented only by the
holotype, and was one of the Smith’s species MICH could not
loan because of limited sample size. It was collected on muck
near beaver ponds in a mountain setting, thus suggesting that
arboreal debris may be a common habitat for this species. We
have noticed many morphological similarities with P. albanyensis
and propose the conspecificity of these two species.
8. Psathyrella ovaticystis Pegler, Kew Bulletin Additional Series
VI: 428. 1977. Fig. 12
Typus: Psathyrella ovaticystis Pegler. Uganda, Buganda Province,
Mengo Distr., Zika Forest, 12 Jun. 1968, D.N. Pegler 1433 (K(M)
196311), among leaf litter.
Loan of one sporocarp lacking a slice of pileus for micromorphology investigation and molecular analysis.
Spores 6.2–8.8(–9.1) × 4.2–5.1 µm, on average 7.8 × 4.9 µm, Q
1.5–1.8, avQ 1.6; in front view oval to cylindrical oval, irregularly
to angularly cylindrical-elliptical, irregularly to angularly oboval,
base broadly rounded to sometimes truncate, in profile adaxially
flattened to subphaseoliform; few spores darkish reddish brown
and with little to moderately distinct, mostly truncate, 1.4–1.7
µm broad germ pore. Pleurocystidia 20–32 × (9–)11–16 [apex
8–12.5(–15) µm broad when differentiated], cylindrical-utriform,
clavate-utriform, ellipsoid, clavate, ovoid, sometimes utriform;
apex rounded to almost truncate, sometimes subcapitate, not
forked, not incrusted, often collapsed in exsiccate material;
walls not pigmented; moderately numerous to scattered.
Cheilocystidia 18–28 × 9–16 µm, scattered to locally moderately
numerous. Paracystidia 11–25(–30) × 7.5–15(–19) µm,
dominating.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Notes: Pegler (1977) reports the dimension of the pileus to be
10 mm and the stipe’ s dimension to be 40 × 1.5 mm, therefore it
can be deduced that his collection consists in one only specimen.
Some characters of this taxon recall P. kauffmanii: color (Pegler
1977: ‘fuscobrunneae … dark fuscous brown’ without indication
of the liquid of observation) and shape of spores, and shape of
cystidia. However, it has distinctly shorter (20–26 × 9–12 µm
in Pegler 1977), bulkier on average, oftener undifferentiated
pleurocystidia. The carpophore has a reduced, slender habitus
and is devoid of veil. Although the DNA analysis failed, the
general micro morphological features seem to place it into the
subsection Spadiceogriseae, possibly close to the P. kauffmanii
complex.
9. Psathyrella spadiceogrisea (Schaeff.) Maire, Mém. Soc. Sci.
Nat. Maroc. 45: 113. 1937. Figs 13 (as P. spadiceogrisea), 14 (as
P. lubrica), 15 (as P. praecox), 16 (as P. amarella).
Fig. 12. Psathyrella ovaticystis. D.N. Pegler 1433, K(M) 196311; PC Pleurocystidia; MC Marginal Cystidia; S2 Spores in KOH.
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Habitat: gregarious or cespitose; on woody debris, in leaves, on
muck; in deciduous woods, in parks, on paths.
Collections examined from MICH: Psathyrella amarella A.H. Sm.,
Ammirati 2925; Psathyrella lubrica A.H. Sm., holotype, Smith 15019;
Psathyrella praecox A.H. Sm., paratype, Potter 4490.
Other material examined: France, Essonne, Mennecy, Parc de Villeroy, ?
Apr. 2004, gregarious, on humus, G. Tassi 04005 (duplicate MCVE29103,
Tef-1α MF521779, ITS MF325997), Fig. 13.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 13. Psathyrella spadiceogrisea. MCVE29103. Photo G. Tassi.
Molecular heterotypical synonyms proposed here: Psathyrella
amarella A.H. Sm., Mem. N. Y. Bot. Gard. 24: 353. 1972; Psathyrella
lubrica A.H. Sm., Mem. N. Y. Bot. Gard. 24: 144. 1972; Psathyrella
praecox A.H. Sm., Mem. N. Y. Bot. Gard. 24: 352. 1972.
Pileus 15–55 mm broad, in early stages obtusely conical with
inflexed margin, finally conical-convex to plane, obtusely
umbonate or not, sometimes radially rugulose in old sporocarps;
when young dark reddish brown sometimes with a violaceous
tinge, discoloring to brown, dark brown, cinnamon brown, rust
brown, ochraceous brown, brownish orange, yellow brown,
dark yellow brown, grey brown, yellow red, sometimes shiny,
hygrophanous, fading cream to alutaceous or whitish, often
with greyish tint; translucently striate when moist. Veil white,
fugacious, when young on pileus with flocci or fibrils around
margin, sometimes fringed at margin, on stipe with appressed
fibrils. Lamellae adnate, moderately broad in age, crowded to
moderately crowded; when young pale alutaceous to hazelbrown, finally purplish brown; edge white. Stipe 40–80×1.5–8
mm, generally equal; white to whitish; apex pruinose, base
sometimes fugaciously felted. Context thick in the centre of
pileus and thin towards the margin, fragile to moderately
fragile in stipe, pale. Odor indistinctive. Taste mild to bitterish.
Spore deposit dark purple-brown. Spores (6–)7–10.1(–10.7) ×
(3.5–)3.8–5.5(–5.8) µm, on average 7.7–9 × 4.3–5.2 µm, Q (1.4–)
1.6–2.2, avQ 1.75–2; in front view elliptic, narrowly oval, oblong,
subcylindrical, sometimes narrowly angular-oval, base often more
or less truncate, in profile subphaseoliform to adaxially flattened,
rarely amygdaliform; reddish brown; germ pore distinct, more or
less truncate, 1.3–1.7 µm broad. Basidia 4-spored, 18–24(–30) ×
7–9(–10) µm. Pleurocystidia 30–64 × 9–20 µm, mainly utriform,
also fusiform to lageniform, sometimes broadly utriform,
subcylindrical, obtusely subconical; apex rounded to subobtuse,
rarely acute, rarely subcapitate, sometimes forked, sometimes
incrusted; walls sometimes thickened, sometimes yellow brown
pigmented; numerous, rarely scattered. Cheilocystidia 25–45
(–50) × 10–18 µm, rare to scattered, rarely absent or rather
numerous. Paracystidia 8–33(–42) × 5–17(–20) µm, sometimes
rhomboid, rarely incrusted; walls sometimes thickened and
lightly pigmented (overall towards base); numerous, dominating.
Cells below marginal cystidia rarely incrusted, rarely slightly
pigmented. Clamps present.
120
Notes: As the holotype material of P. spadiceogrisea seems not
to be existing anymore we refer, for its molecular concept, to
Örstadius et al. (2015) and, for its morphological concept, to the
German description of the basionym Agaricus spadiceogriseus by
Schäffer (1771) (later followed by a latin description in Schäffer
1774) which includes a color drawing in table 237 (iconotypus).
Our morphological concept also matches that by Örstadius et al.
(2008) but for some little differences as they, probably, included
also P. niveobadia in their description.
Schäffer (1800) later authored a reprinted version of its
original table displaying a different pileus coloration including
distinct violaceous tints, and dirty pinkish for the smallest
sporocarps. The first color can also be found in P. fatua (see Figs
36, 38, 40) and P. albescens (see Fig. 27) when undergoing the
drying process; the distinct pink color, instead, is only known for
P. fatua (see Figs 37, 39). More comments on Schäffer’s table
and description are included in the notes about P. albescens, but
-based on the points above- it is likely that Schäffer’s reprinted
drawings may also include P. fatua collections.
The features of P. spadiceogrisea show a range of
variability. The initial strong tints of the pileus can linger into
the mature stage, until the pileus remains fresh, and may
discolor into greyish tints. The pleurocystidia are with or
without the following: incrustations, pigmentation or forked
apex. Morphological variability and the unrecognized presence
in Europe of the American taxon P. albescens (see below), at
least until now, have led to misinterpretations, superfluous
synonyms, and contradicting interpretations. Many subspecific
taxa have been reduced to synonymy: P. spadiceogrisea f.
mammifera, P. spadiceogrisea f. phaeophylla and Drosophila
spadiceogrisea f. russifolia by Ludwig (2007); P. spadiceogrisea
f. vernalis both by Ludwig (2007) and by Örstadius (2007). Also
Ludwig (2007) synonymized P. spadiceogrisea f. exalbicans to
P. fatua. Some of these synonymies, particularly the last two,
may be incorrect as their propounder did not know about the
presence of P. albescens in Europe; this however would not
change the main point that none of them are to be regarded
as good species.
The four vouchers we have analyzed in this study display
some degree of slight intraspecific variability in their molecular
identity too. However ITS sequence homology values were
always higher than 98.0 % among the type of P. amarella, the
paratype of P. praecox, and samples LÖ92-01 and MCVE29103
of P. spadiceogrisea. Similarly, Tef-1α sequence homology values
were always higher than 99.0 % among the type of P. lubrica, the
paratype of P. praecox, and samples LÖ102-98 and MCVE29103
of P. spadiceogrisea.
In MCVE29103 (G. Tassi’s voucher 04005), composed of
both young and mature specimens, the pilei are date brown,
ochraceous in the center, and encircled in the primordial and
young stages by a conspicuous band of white veil (see Fig. 13).
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
These macro morphological traits are similar to those described
for P. niveobadia; however the micro-morphological features of
this material overlap with those of P. spadiceogrisea.
The holotype of P. amarella shows some pleurocystidial
apex with a thickened and pigmented wall. All of the material
examined has shown a sparse presence of little incrustations on
pleurocystidia apex, but never a nucous covering.
The lack of strong morphological differences and the
similarity of DNA sequences all suggest these collections
represent the same species.
Psathyrella marquana and P. cascoides are two very recently
described species. The former has characters that, although close
to those of P. spadiceogrisea, are different enough to represent
its own species, and it is included in our key. The latter instead,
has characters fully consistent with those of P. spadiceogrisea,
and therefore it has to be regarded as a synonym.
Descriptions of P. casca in the literature are mostly
misidentifications of P. spadiceogrisea or P. niveobadia. Kits van
Waveren’s (1985) interpretation was reported by Örstadius et al.
(2008) as synonym to P. spadiceogrisea. Actually, we suspect it,
and also P. casca sensu Tassi (2000), to represent P. niveobadia
instead. We consider P. casca a dubious name.
For comparison with P. niveobadia see below.
9a. Psathyrella lubrica A.H. Sm., Mem. N. Y. Bot. Gard. 24: 144.
1972. Fig. 14.
Typus: Psathyrella lubrica A.H. Sm. USA, Michigan, Oakland Co,
Kent Lake, near New Hudson, 28 May 1940, A.H. Smith 15019
(MICH11957), on muck. Tef-1α MF521771.
Loan of two sporocarps in good condition and a portion of pileus
for gross morphology investigation.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Spores 7.5–9 × 3.8–5.5 µm, Q (1.46–)1.61–1.9(–1.93), avQ 1.75;
in front view elliptic, narrowly oval, subcylindrical, sometimes
narrowly angular-oval, base rounded to more or less truncate,
in profile adaxially flattened to subphaseoliform; reddish
brown; germ pore moderately distinct to distinct, more or
less truncate,1.3–1.7 µm broad. Pleurocystidia 45–62 × 10–18
µm; utriform to fusiform-utriform or lageniform, sometimes
subcylindrical; apex obtuse to subobtuse, sometimes rounded
to subacute, rarely subcapitate, sometimes forked, sometimes
incrusted; walls hyaline; numerous. Cheilocystidia 25–40 ×
10–15 µm, scattered. Paracystidia –33 × –13 µm, sometimes
rhomboid, numerous, dominating. Cells below marginal cystidia
hyaline, not incrusted.
Fig. 14. Psathyrella lubrica. A.H. Smith 15019, MICH11957; PC pleurocystidia; S1 Spores in water; S2 Spores in KOH.
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Editor-in-Chief
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E-mail: p.crous@westerdijkinstitute.nl
Fig. 15. Psathyrella praecox. V. Potter 4490, MICH49251; PC1 Pleurocystidia in NH4OH; PC2 Pleurocystidia in Congo red; MC Marginal Cystidia in Congo
red; S1 Spores in water; S2 Spores in KOH.
9b. Psathyrella praecox A.H. Sm., Mem. N. Y. Bot. Gard. 24: 352.
1972. Fig. 15.
Paratype: Psathyrella praecox A.H. Sm. USA, Michigan, Gratiot
Co., Ithaca, Schovence’s woods, 18 May 1948, V. Potter 4490
(MICH49251), on soggy leaf mold near edge of a pond. Tef-1α
MF521783, ITS MF325992.
Notes: Smith (1972) reports narrower spores, (6–)7–9 × 3.3–3.8
µm. Both morphology and DNA reveal that this paratype is a later
synonym of P. spadiceogrisea. This result does not necessarily
apply to the holotype, which could not be obtained due to
scarcity of material; however, from the general description of
the species we conclude that P. praecox is to be considered a
synonym of P. spadiceogrisea.
Loan of various sporocarps in good condition for gross
morphology investigation.
9c. Psathyrella amarella A.H. Sm., Mem. N. Y. Bot. Gard. 24:
353. 1972. Fig. 16.
Spores 7.5–10.1 × 3.8–5.1 µm, Q 1.7–2.2, avQ 2; in front view
elliptic, cylindrical to oval, sometimes narrowly angular-oval,
base often more or less truncate, in profile adaxially flattened to
subphaseoliform, sometimes phaseoliform or subamygdaliform;
reddish brown; germ pore distinct, often truncate, ± 1.6 µm
broad. Pleurocystidia 32–43 × 10–17 µm; utriform, sometimes
fusiform-utriform, subcylindrical; apex rounded to obtuse, not
subcapitate, sometimes forked, sometimes incrusted; walls
sometimes pigmented; numerous. Cheilocystidia 31–36 × 11–
12 µm, rare. Paracystidia 8–31 × 5–17 µm, rarely incrusted,
walls sometimes thickened and weakly pigmented, numerous,
dominating.
Typus: Psathyrella amarella A.H. Sm. USA, Michigan, Baraga Co.,
Silver River Area, Dynamite Hill Road, 7 Feb. 1969, J.F. Ammirati
2925 (MICH11869), gregarious on leaf mold in a hardwood
forest. ITS MF325958.
122
Loan of one pileus in good condition for gross morphology
investigation.
Spores 7–9.4 × 4–5 µm, Q 1.6–2.1; in front view elliptic,
subcylindrical, (narrowly) oval, base often more or less truncate,
in profile subphaseoliform to adaxially flattened; reddish
brown; germ pore distinct, subtruncate to rounded, ± 1.2–1.6
µm broad. Pleurocystidia 30–48 × 10–15(–20) µm; utriform,
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 16. Psathyrella amarella. J.F. Ammirati 2925, MICH11869; MC Marginal Cystidia in KOH; PC Pleurocystidia in NH4OH; S1 Spores in water; S2 Spores
in KOH.
© 2019 Westerdijk Fungal Biodiversity Institute
123
Voto et al.
ellipsoid-utriform, sometimes clavate-utriform to obtusely
fusiform; apex rounded, sometimes subacute, not subcapitate,
often forked to irregular, sometimes incrusted; walls sometimes
thickened, sometimes reddish brown pigmented; moderately
numerous. Cheilocystidia 20–33(–42) × 8–13 µm, scattered and
visible in mass. Paracystidia 11–31(–40) × 5–17(–25) µm, often
cylindrical, walls sometimes thickened and pigmented with a
very slightly ochraceous hue (detectable in mass); numerous,
dominating. Cells below marginal cystidia slightly pigmented,
sometimes incrusted.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: The molecular comparison of this species with the other
examined material of P. spadiceogrisea shows that the ITS
nucleotide homology ranges between 98 % and 99 % identity
with 1 to 14 mismatches. Unfortunately, we could not add the
evidence of a second locus; however, apart from the bitter taste,
we think its morphological traits fit well into the concept of P.
spadiceogrisea.
Fig. 17. Psathyrella niveobadia. MCVE29102 Photo B. De Ruvo.
10. Psathyrella niveobadia (Romagn.) M.M. Moser, Kleine
Kryptogamenflora die Röhrlinge und Blätterpilze (Agaricales)
2b/2: 276. 1978. Figs 17–22.
Pileus 15–72 mm broad, in early stages obtusely conical
to paraboloid with inflexed margin, then hemispherical,
finally through convex to plane, obtusely umbonate or not,
often radially rugulose in old sporocarps; when young dark,
violaceous brown to reddish brown or date-brown, possibly still
dark when mature, discoloring to ochraceous brown at centre,
more brown to greyish brown in periphery, hygrophanous,
fading cream to grey-cream or grey-beige, sometimes with
olivaceous tints; not translucently striate or only at extreme
margin. Veil white, fugacious, when young of fibrils on margin
and connecting margin to stipe. Lamellae 3–6 mm broad,
adnate to broadly adnate, crowded to moderately crowded;
when young grey brown, finally more or less dark brown;
edge white. Stipe 30–130(–200) × 3.5–10 mm, cylindrical,
sometimes very broad at apex (up to 17 mm) and then tapering
downwards, sometimes a little broadening towards the base,
often with a short to 10 cm long rooting base; hollow since
when young; white, discoloring pale brownish with age; apex
pruinose. Context fairly thick and tenacious in the pileus,
distinctly thick and rigid in the stipe cortex, only fragile in the
rooting stipe base portion. Odor weak to indistinctive. Spore
deposit purplish brown. Spores 6.9–9.5(–10) × (3.9–)4.2–5
µm, on average 7.4–8.3 × 4.5–4.7 µm, Q (1.5–) 1.55–2(–2.25),
avQ 1.7–1.8; in front view elliptic to cylindrical or narrowly
oval-cylindrical, base truncate to broadly rounded, in profile
phaseoliform, subphaseoliform or adaxially flattened, rarely
indented above the apiculus; orange-brown to reddish orange
brown (darkish reddish brown in NH4); germ pore distinct,
rounded, 1.0–1.5 µm broad. Pleurocystidia (26–)35–60(–70)
× (10–)14–18(–22) µm, utriform, lageniform, clavate, clavatemucronate, fusiform or irregularly tapering towards the
apex, sometimes cylindrical-utriform, ellipsoid-utriform; apex
broadly rounded to narrowly obtuse, sometimes truncate or
subacute, subcapitate or not, sometimes distinctly capitate,
sometimes forked, sometimes incrusted, often with grossly
mucous masses which gradually disappear in exsiccata and,
perhaps, also with age of sporocarps; walls often or sometimes
slightly thickened especially at apex and yellow brown
pigmented; base sometimes with one or two septa; scatter to
124
Fig. 18. Psathyrella niveobadia. MCVE29102. Photo B. De Ruvo.
numerous. Cheilocystidia 21–35(–45) × 12–22 µm, scattered.
Paracystidia 11–30(–40) × 7–18 µm; walls sometimes slightly
thickened and pale brown; numerous, dominating. Cells below
marginal cystidia generally not incrusted and not pigmented.
Habitat: gregarious to caespitose; often growing in spring,
lowlands to mountains; on broadleaves woody debris or around
stumps, dry to moist.
Material examined: France, Aisne, près de Russy-Bémont, vallée de
Russy, forêt de Retz, at the margin of the wood, gregarious on soil and
on debris, 14 Feb. 1993, G. Tassi 93014 (9314). Italy, Teramo, Rocca
Santa Maria, loc. Ceppo, 1 350 m a.s.l., caespitose around stump of
Fagus, 4 May 2003, B. de Ruvo, Figs 17, 18, (duplicate MCVE29102,
Tef-1α MF521780, ITS MF325996, more pictures of this collection
in
http://www.ambmuggia.it/forum/topic/478-psathyrella-fatua/);
Pescara, Lettomanoppello, Parco nazionale della Majella, 1 300 m a.s.l.,
gregarious to fasciculate among debris of leaves and needles or around
a Fagus stump in a calcareous mixed wood of Fagus, Picea, Abies and
Pinus), 13 May 2018, 18 May 2018 and 20 May 2018, A. Micucci, Figs
19–22, (MCVE30076, Tef-1α MK408675, ITS MK400421).
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 19. Psathyrella niveobadia. MCVE30076. Photo A. Micucci.
Fig. 20. Psathyrella niveobadia. MCVE3007. Photo A. Micucci.
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Voto et al.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 21. Psathyrella niveobadia. MCVE30076. Photo A. Micucci.
Notes: The case of P. niveobadia remains a little ambiguous.
We could not examine the French holotype of its basionym,
Drosophila niveobadia, therefore, we have studied two
French collections kindly provided to us by G. Tassi. The gross
morphology of voucher 04005 (duplicate MCVE29103) recalls
that of P. niveobadia, but decidedly belongs to P. spadiceogrisea
(see discussion above). The other voucher, 93014, has
microscopical characters in line with those of P. niveobadia
(shape of pleurocystidia, some pleurocystidia apex thick-walled,
dimension and orange-brown color of spores, small germ pore)
but, regretfully, DNA extraction failed.
Two other specimens from central Italy (collectors Micucci,
MCVE30076, and De Ruvo, MCVE29102), show morphological
characters that match perfectly those of P. niveobadia described
by Kits van Waveren (1985). They possess, and this is particularly
true for Micucci’s specimen, all the main expected diagnostic
features for this species, including: the thickness and rigidity
of the stipe cortex and, to a lesser extent, of the pileus, which
is scarcely, if at all, striate; the possibly rooting stipe base; the
orange-brown spores which are on average 7.4–8.3 × 4.5–4.7
µm large; the distinctly thickwalled pleurocystidia apex which
can be subcapitate or particularly narrow to tapered. From our
own observations we can add grossly mucous coverings (in
sufficiently fresh specimens at least) of the pleurocystidia apex
which is not reported in Kits van Waveren’s (1985) description.
126
We regard other characters in Kits van Waveren’s (1985)
description as less relevant, not exclusive or unstable; thus, for
instance, identifications based mainly on pileus color and veil
development, as emphasized in Kits van Waveren’s (1985) notes,
could actually confuse this species with P. spadiceogrisea, while
based mainly on rhomboid shape of paracystidia the species in
question could actually be P. fatua.
Molecular comparisons between the two Italian specimens
and the P. spadiceogrisea samples included in this paper
have produced somewhat ambiguous results, though. Tef-1α
nucleotide homology levels ranged between 98.14 % and 98.67
%, suggesting a possible conspecificity, but ITS nucleotide
homology values ranged between 96.77 % and 99.02 %,
only partially corroborating conspecificity. Because Micucci’s
sequences were obtained in late 2018, they were not used when
constructing the phylogenetic trees, thus their exact placement
in the phylogeny of the subsection is unknown. However, taking
into consideration the morphological differences listed above
and the presence of some, albeit limited, genetic divergence
between P. spadiceogrisea and P. niveobadia in the combined
phylogenetic tree (Fig. 3), we have decided to keep the species
P. niveobadia separate from P. spadiceogrisea. Of course,
we also acknowledge the specimens belonging to these two
species may instead represent extremes of a rather variable
single species.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 22. Psathyrella niveobadia. A. Micucci, MCVE30076; PC1 Pleurocystidia in Congo red; PC2 Pleurocystidium in NH4OH; S1 Spores in water; S2
Spores in KOH.
Other images in color drawings are in Tassi (2000: 368, pl. 7) and
Ludwig (2007b, pl. 378, figs 98.38A–98.38B).
11. Psathyrella albescens Hesler & A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 133. 1972. Figs 23–32 (as P. albescens), 33 (as P.
mazzeri).
Molecular heterotypical synonyms proposed here: Psathyrella
agrariella sensu A.H. Smith, Mem. N. Y. Bot. Gard. 24: 411. 1972,
p.p.; Psathyrella mazzeri A.H. Sm., Mem. N. Y. Bot. Gard. 24: 356.
1972.
Pileus 10–70 mm broad, in early stages hemispheric to convex,
then campanulate to convex, finally convex to plane and rarely
slightly depressed at centre, umbonate or not, sometimes
radially rugulose, sometimes wavy, margin for some time
more or less inflexed, finally straight to sometimes reflexed;
when young, dark yellowish brown to cinnamon-brown; when
mature, cinnamon-brown, yellowish brown, brownish orange,
light to medium Dresden brown, rarely pinkish buff, often moist
or shiny, hygrophanous, discoloring to white with ochraceous
centre for a time, finally drying from margin to greyish or
greyish violaceous; translucently striate when moist. Veil white,
when young with fibrils or flocci around the margin of pileus
and connecting it to stipe, when mature as fugacious fibrils
around the margin of pileus or appendiculate or folded back on
the gills, on stipe usually with few appressed fibrils, once as a
fragile submembranous annulus. Lamellae (broadly) adnate to
slightly decurrent, moderately broad, not or slightly ventricose,
crowded; when young whitish to pale pinkish buff or dull brown,
finally purplish brown to blackish brown; edge white, fimbriate.
Stipe 20–100(–140) × 1.5–6(–12) mm, often broadening
towards base or towards both apex and base, extreme base
sometimes bent, subrooting or characterized by a short to 50
mm long pseudorhiza; watery white to whitish, often discoloring
brownish; apex pruinose, sometimes subsquamulose or striate,
base often felted to subtomentose. Context pale, fragile to
rather rigid, in pileus thin. Odor and taste indistinctive to mild.
Spore deposit purplish black to blackish brown. Spores (6.3–)
6.7–10.2(–11.7)×(3.5–)3.7–5.4(–5.5) µm, on average 7.4–9
× (4.1–)4.5–4.9 µm, Q (1.5–)1.6–2.4(–2.6), avQ (1.70–)1.80–
1.90(–2.16); in front view oblong to elliptic or oval to narrowly
oval, sometimes (sub) angular-oval to cylindrical or irregular,
rarely obscurely narrowly oboval or with a median constriction,
base broadly rounded to truncate, sometimes triangular, in
profile adaxially flattened to subphaseoliform, phaseoliform
or indented above the apiculus, sometimes subamygdaliform;
orange-brown to orangish reddish brown; germ pore distinct,
more or less truncate, 1.3–1.8 µm broad. Basidia 4-spored,
16–24(–29) × 6–9(–10) µm. Pleurocystidia (28–)32–68(–70) ×
(8–)10–23(–28) µm, utriform to broadly utriform, subcylindrical,
fusiform to ellipsoid-rhomboid, rarely lageniform or clavate; apex
rounded to obtuse, sometimes subtruncate, rarely subobtuse,
subcapitate or not, forked or not, incrusted or not; walls
sometimes or often thickened, sometimes brownish pigmented;
numerous, sometimes moderately numerous. Cheilocystidia
21–50 (–58) × (7–)10–18(–25) µm, similar in incrustations and
pigment to the paracystidia, rare to scattered or sometimes
locally numerous. Paracystidia 8–48(–56) × 4–28(–30) µm,
sometimes incrusted, walls sometimes or often thickened and
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Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 23. Psathyrella albescens. MCVE29113. Photo P. Voto.
Fig. 24. Psathyrella albescens. MCVE29113. Photo P. Voto.
Fig. 25. Psathyrella albescens. MCVE29107. Photo P. Voto.
Fig. 26. Psathyrella albescens. MCVE29107. Photo P. Voto.
Fig. 27. Psathyrella albescens. MCVE29107. Photo P. Voto.
Fig. 28. Psathyrella albescens. Voto 20160313. Photo P. Voto.
128
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 29. Psathyrella albescens. MCVE29116. Photo M. Maletti.
Fig. 30. Psathyrella albescens. MCVE29111. Photo P. Voto.
Fig. 31. Psathyrella albescens. P. Voto, MCVE29113; PC1 Pleurocystidia in Congo red; PC2 Pleurocystidia in NH4OH; MC Marginal Cystidia; S1 Spores
in water; S2 Spores in KOH.
yellowish to brownish pigmented; very numerous, dominating.
Cells below marginal cystidia sometimes incrusted, sometimes
locally yellowish pigmented. Clamps present.
Habitat: Solitary to gregarious or subcespitose; on woody debris,
in leaves, in grass, on sandy humus; in parks, at margin of woods
or in wood clearings or in wood paths, in pastures, at roadsides.
Collections examined from MICH: Psathyrella agrariella sensu Smith,
Smith 25099 and McKnight F949; Psathyrella albescens Hesler & A.H.
Sm., holotype, Hesler 18895; Psathyrella mazzeri A.H. Sm., holotype,
Mazzer 6023.
© 2019 Westerdijk Fungal Biodiversity Institute
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Voto et al.
Other material examined: Italy, Belluno, Nevegal, about 1 000 m a.s.l.,
gregarious on debris at margin of forest, 24 Sep. 2010, P. Voto, Figs
23, 24, 31 (MCVE29113, Tef-1α MF521819, ITS MF325954); Rovigo,
Sant’Apollinare, 6 m a.s.l., 18 Nov. 2006, gregarious by a roadside
among litter of brambles and Robinia pseudoacacia, P. Voto, Figs 25–27
(MCVE29107, Tef-1α MF521821, ITS MF326009); Rovigo, Crespino, 3
m a.s.l., solitary in a park, 31 Oct. 2006, P. Voto (MCVE29106, Tef-1α
MF521822, ITS MF325953); Rovigo, Villadose, 3 m a.s.l., gregarious in
a park under Tilia platyphyllos, 13 Mar. 2016, P. Voto 20160313, Fig.
28; (Pesaro Urbino, Monti delle Cesane, botanical garden, gregarious
among hardwood trees, 29 Apr. 2012, M. Maletti, Fig. 29 (MCVE29116,
Tef-1α MF521818, ITS MF325955 ); Venezia, Chioggia, S. Anna, Nordio
forest, 2 m a.s.l., 12 Dec. 2007, solitary on humus of Quercus ilex, P.
Voto, Fig. 30 (MCVE29111, Tef-1α MF521820); at same location, 31 Oct.
2011, gregarious on woody debris, P. Voto 20111031.1; at same location,
5 Nov. 2015, gregarious on woody debris, P. Voto (MCVE29332).
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: Psathyrella albescens is difficult to distinguish from P.
spadiceogrisea based on micro morphological characters. We cite
the occurrence of two traits: possibly capitate pleurocystidia that
are absent or seldom found in P. spadiceogrisea, and spores with a
greater tendency to be phaseoliform or indented above the apiculus.
More consistent differences can be found in the gross morphology
and are listed below. The young pilei of P. spadiceogrisea have a
reddish brown (in agreement with Schäffer’s, 1771 table nr. 237)
to dark reddish brown or violaceous brown color, often retained
until maturity, while P. albescens pilei are at most dark yellowish
brown at first, and quickly become discolored to brownish orange,
never attaining a dark brown or reddish tint. Furthermore, the
discoloring pilei of P. spadiceogrisea usually fade to cream or
alutaceous tints, while those of P. albescens usually fade to white.
The veil of P. spadiceogrisea is fugacious and conspicuously located
on the margin of the pileus where it is usually no longer present in
mature sporocarps (in agreement with Schäffer’s, 1774 description
of ‘velo et annulo destitutus’), rarely and exclusively in early stages
of development it is in form of appendiculate flocci. Conversely,
in favorable climatic conditions, the veil of P. albescens can be still
appendiculate from the margin of the pileus or folded back on the
gills or, rarely, annulate on the stipe in mature sporocarps. Often,
P. albescens has a stipe base that is more or less swollen and can
be subrooting or characterized by a pseudorhiza up to 50 mm
long. Conversely, this trait is not reported by Schäffer (1774) for P.
spadiceogrisea, which has a stipe base at most slightly attenuated.
Finally, P. spadiceogrisea is only found on woody material, while P.
albescens can also be found in grasslands.
The features that can differentiate P. albescens from P. fatua
are slight and often they can do so only partially. The first has a
veil at times still appendiculate or folded back on the gills, rarely
annular, in mature specimens, while the second has a quickly
evanescent veil on the margin. The first species has a discoloring
pileus without distinct pinkish shades, at most and rarely pinkish
buff, and with a disk never attaining vivid colors: at most it
may only have a dull ochraceous tint. The second species has
discoloring pilei that may attain pinkish shades on the margin, and
vivid yellow tints on the disk. The gill edge is not pigmented in the
first species, but it can be red in the second. Pleurocystidia can be
brownish, forked, or incrusted in the first, while in the second they
are never brownish, and rarely forked or weakly incrusted. Spores
of P. albescens are orange-brown to orangish reddish brown and
often phaseoliform to indented above the apiculus, while those
of P. fatua are brown to dark red, rarely phaseoliform or indented
above the apiculus, sometimes subphaseoliform.
130
We expect this species to be present in herbaria of European
mycologists under the different names P. fatua, P. pseudocorrugis
sensu Kits van Wav., P. niveobadia, and P. spadiceogrisea.
Based on our evaluation, at least one voucher determined
as P. agrariella by Smith is actually P. albescens, thus rendering
Smith’s concept of P. agrariella dubious (see descriptions of
Smith 25099 and Mc Knight F949 below).
11a. Psathyrella albescens Hesler & A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 133. 1972. Fig. 32.
Typus: Psathyrella albescens Hesler & A.H. Sm. USA, Tennessee,
Knox, Farragut, 20.II.1949, L. R. Hesler 18895 (MICH11863), on
decaying chip pile. Tef-1α MF521817.
Loan of a portion of pileus in good condition for micromorphology investigation.
Spores 6.7–8.5(–10) × 3.7–4.4(–4.9) µm, Q (1.4–)1.6–2(–2.3),
avQ 1.88; in front view oblong to elliptic, sometimes irregular,
sometimes obscurely narrowly oboval or with median
constriction, rarely angular-oval, base rounded, sometimes more
or less truncate, in profile adaxially flattened to subphaseoliform,
sometimes indented above the apiculus; orange-brown to
orangish reddish brown; germ pore distinct, mostly more or
less truncate, about 1.6–1.7 µm broad. Pleurocystidia 35–52
× 13–17 µm, utriform, sometimes subcylindrical, ellipsoidrhomboid; apex rounded to obtuse, sometimes subcapitate,
forked or incrusted; walls sometimes thickened and pigmented;
numerous. Cheilocystidia small, scattered. Paracystidia 8–48
× 6.5–28 µm, often with yellowish incrustations, walls often
thickened, numerous, dominating. Cells below marginal cystidia
conspicuously incrusted.
11b. Psathyrella mazzeri A.H. Sm., Mem. N. Y. Bot. Gard. 24:
356. 1972. Fig. 33.
Typus: Psathyrella mazzeri A.H. Sm. USA, Michigan, Allegan
Co., Vivian Baker Farm near Otsego, 16 May 1970, S.J. Mazzer
6023 (MICH11963), scattered to gregarious in open pasture with
Phleum pratense and Dactylis glomerata. Tef-1α MF521790, ITS
MF325981.
Loan of two sporocarps in good condition for micro-morphology
investigation.
Spores 6.8–9.3(–10) × 3.8–5.2(–5.5) µm, Q (1.45–)1.65–1.9(–
2.1), avQ 1.79; in front view elliptic, oblong to narrowly oval,
rarely oval to subangular-oval, base rounded, rarely truncate,
in profile subphaseoliform to adaxially flattened, sometimes
phaseoliform to indented above the apiculus; orange-brown;
germ pore distinct, mostly more or less truncate, more or less
1.7–1.8 µm broad. Pleurocystidia 32–51 × 10–16 µm; utriform,
rarely lageniform; apex rounded, often subcapitate; incrustations
and pigment not seen; more or less numerous. Cheilocystidia
21–35 × 10–14 µm, scattered. Paracystidia 10–35 × 4–10 µm,
moderately incrusted, walls hyaline, numerous, dominating.
Cells below marginal cystidia hyaline, moderately incrusted.
12. Psathyrella agrariella (G.F. Atk.) A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 411. 1972.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 32. Psathyrella albescens. L.R. Hesler 18895, MICH11863; MC Marginal Cystidia in KOH; PC Pleurocystidia in NH4OH; S1 Spores in water; S2 Spores
in KOH.
Smith (1972) transferred Psilocybe agrariella to Psathyrella but,
apparently, without personally revising the original material; in
fact among the material examined he indicated only a number
of other collections that he deemed to represent that species.
MICH staff has provided us with three of those vouchers and they
have come out to represent at least two different taxa. Smith
28713 has dominating cheilocystidia, and molecularly is close to
P. seminuda and clearly out of subsection Spadiceogriseae. Smith
25099 has dominating paracystidia and matches P. albescens
both in morphology and DNA sequence. McKnight F949 also
resembles P. albescens based on macro and micro-morphology,
but the molecular evidence based only on the ITS sequence is
inconclusive.
In Atkinson’s basionym description of Psilocybe agrariella,
the pileus is pale reddish brown to pale rufous, the stipe
is concolorous to the pileus at the base, the carpophores
dimension is small, and the pleurocystidia are defined lanceolate
to subventricose. These parameters above are shared with
P. agrariella var. ochrofulva. The cheilocystidia are described
as broadly ventricose and shorter than the pleurocystidia,
sometimes very broad and bluntly mucronate: therefore it is
unclear whether true paracystidia are present or not, although
clavate to pyriform cells are reported at stipe apex intermixed
with pleurocystidioid cells. We conclude that the true identity
of Psilocybe agrariella remains dubious until the holotype is
morphologically revised and sequenced. We also suggest that
Smith’s concept of Psathyrella agrariella should be rejected both
because of Smith’s vague description of the gill edge cystidia,
which are reported as broadly ventricose, mucronate or saccate,
without an indication of the ratio between them, and because of
the lack of homogeneity in his vouchers.
12a. Smith 25099, MICH47933.
= Psathyrella albescens Hesler & A.H. Sm.
USA, Michigan, Oakland Co., Proud Lake, 26 May 1947, coll. Lange
& Smith, on sandy humus. Tef-1α MF521824. ITS MF326008.
Loan of a dozen and more sporocarps in good condition for
micro-morphology investigation.
Spores 6.3–9.1 × 4–5.2 µm, Q 1.5–1.8, avQ 1.7; in front view
elliptic to oval or angular-oval, sometimes subtriangular,
irregular, base generally truncate, in profile subphaseoliform
to adaxially flattened; orange-brown, rarely orangish reddish
brown; germ pore distinct, ± 1.5 µm broad. Pleurocystidia 28–50
× 10.5–16.5 µm; utriform; apex obtuse to rounded, sometimes
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Voto et al.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 33. Psathyrella mazzeri. S.J. Mazzer 6023, MICH11963; PC1 Pleurocystidia in NH4OH; PC2 Pleurocystidia in Congo red; MC Marginal Cystidia in
Congo red; S1 Spores in water; S2 Spores in KOH.
or often subcapitate, sometimes forked, sometimes incrusted,
walls hyaline or almost so; moderately numerous. Cheilocystidia
27–37 × 11.5–18 µm, moderately numerous. Paracystidia –30
× –22 µm, often incrusted, numerous, dominating. Cells below
marginal cystidia hyaline or almost so.
Notes: The material received was composed of solitary to
clustered sporocarps with stipes progressively slightly broadened
towards the base and at that point bent and attached to woody
debris; in many pilei the margin had a greyish tint. ITS and Tef-1α
sequences put it in synonymy with P. albescens, the morphology
is coherent with this result.
12b. Mc Knight F949, MICH65241
?= Psathyrella albescens Hesler & A.H. Sm.
USA, Utah, Duchesne Co., two miles south of Moon Lake, 9 Jul.
1955, coll. K.H. Mc Knight & R. Duke, in moist soil and moss
under Salix. ITS MF325951.
Loan of numerous sporocarps in good condition for micromorphology investigation.
132
The notes from the collector report the following: pileus 1.5–5.5
cm across, buffy brown to bistre when moist and a fibrillose
annulus; in the exsiccata, we saw long stipes, up to 90 × 3 mm,
and distinct fibrillose veil remains at the stipe apex.
Spores 7–10.1(–12) × 4.4–5.4(–5.7) µm, Q 1.5–2.1(–2.3), avQ
1.76–1.86; in front view elliptic to oval, sometimes irregular,
base rounded to truncate, in profile subphaseoliform to
adaxially flattened, sometimes subamygdaliform; orangebrown to orangish reddish brown; germ pore distinct, round
to rarely truncate, 1.5–1.8 µm broad. Pleurocystidia 34–53 ×
7.7–11.5 µm; utriform, subcylindrical, sometimes cylindrical,
flexuous; apex 6–12 µm broad, rounded to more or less
truncate, sometimes or often subcapitate, often forked,
often incrusted, walls often slightly thickened, sometimes
pigmented; numerous. Cheilocystidia 30–45 × 9.5–11 µm,
scattered to sometimes somewhat clustered. Paracystidia
–32 × –17 µm, often pigmented, hardly incrusted, numerous,
dominating. Cells below marginal cystidia locally slightly
pigmented.
Notes: The ITS sequence of this voucher has a high homology
with sequences of the various P. albescens collections, but its
homology is even higher with the sequence of P. agrariella var.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
ochrofulva. Although we lack here the corroborating evidence of
a second locus, the micro morphological traits are very similar to
those of P. albescens, and the possible presence of an annulus
is already demonstrated for that species by our Italian material
MCVE29113. Moreover, the bistre color of the fresh pileus
corresponds to the dark yellow brown color described for P.
albescens by Smith.
45) × 7–11(–13) µm, numerous, inconspicuously (because small
and often recurved to repent) dominating. Paracystidia 11–25(–
30) × 6–11(–15) µm, often incrusted, walls yellowish pigmented,
numerous. Cells below marginal cystidia forming a yellowish
discontinuous band approximately 25 µm broad towards the
stipe, becoming pale fulvous and up to approximately 100 µm in
width towards the pileus margin, generally not incrusted.
12c. A.H. Smith 28713, MICH47939.
Notes: The material received was composed of solitary to
clustered sporocarps with most stipes progressively slightly
broadened towards base and there attached to woody debris;
the pilei had a greyish to greyish violaceous margin; most gills
had a purplish edge partly visible by a lens. The phylogenetic
evidence places this material very close to P. seminuda (DNA
nucleotide homology: ITS= 99 % with a single mismatch) but we
have found some morphological divergences, such as narrower,
terete spores, distinctly smaller pleuro- and cheilocystidia and
the presence of numerous, though small, paracystidia not cited
for P. seminuda. As for the slightly pigmented gill edge it is to be
noticed that P. seminuda is described with cheilocystidia often
rusty brownish colored.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
=?
USA, Michigan, Washtenaw Co., Dexter, 18.V.1948, on sawdust
pile. ITS MF325950.
Loan for micro-morphology
sporocarps in good condition.
investigation
of
numerous
Spores (7.2–)7.5–8.7(–9) × 4.2–5 µm, Q 1.6–2, avQ 1.7–
1.8; in front view elliptic, sometimes oval, base obtuse, in
profile adaxially flattened to subamygdaloid, sometimes
subphaseoliform; reddish to brownish red; germ pore generally
distinct, generally truncate, sometimes eccentric in profile,
±1.5 µm broad. Pleurocystidia 27–52 × 9–15 µm; utriform,
sublageniform, sometimes subcylindrical; apex obtuse to rarely
rounded, (4–)5–7(–10) µm broad, rarely subcapitate, rarely
forked, often incrusted; numerous. Cheilocystidia (20–)23–34(–
13. Psathyrella ochrofulva (A.H. Sm.) Voto, Dovana & Garbel,
comb. nov. MycoBank MB831128.
Basionym: Psathyrella agrariella var. ochrofulva A.H. Sm., Mem.
N. Y. Bot. Gard. 24: 411. 1972. A.H. Smith 33628, MICH11858.
Fig. 34.
Fig. 34. Psathyrella ochrofulva. A.H. Smith 33628, MICH11858; PC1 Pleurocystidia in NH4OH; PC2 Pleurocystidia in Congo red; MC Marginal Cystidia
in Congo red; S1 Spores in water; S2 Spores in KOH.
© 2019 Westerdijk Fungal Biodiversity Institute
133
Voto et al.
Typus: Psathyrella agrariella var. ochrofulva A.H. Sm. USA,
Michigan, Cheboygan Co., Colonial Point Hardwoods near Burt
Lake, 13 Sep. 1949, A.H. Smith 33628 (MICH11858), gregarious
on sandy humus in cut-over hardwoods. ITS MF325987.
Loan of one pileus in good condition for micro-morphology
investigation.
Spores (7.5–)8–10.1 × 4.5–5.6(–6) µm [7–9(–10) × 3.8–4.5(–5)
µm in Smith, 1972], on average 9 × 5–5.3 µm, Q 1.4–2.2, avQ
1.7–1.8; in front view elliptic to oval, sometimes cylindrical,
base rounded to truncate, in profile adaxially flattened to
subelliptic, sometimes subphaseoliform; reddish brownish;
germ pore distinct, rounded to subtruncate, about 1.4–1.8(–2)
µm broad. Pleurocystidia 30–50(–53) × 11–17 µm, the smaller
ones towards gill edge; utriform, conical, fusiform, lageniform,
sometimes ellipsoid, subcylindrical, submucronate; apex
obtuse, sometimes rounded or subacute, sometimes forked
or truncate, sometimes incrusted, walls sometimes slightly
pigmented; moderately numerous. Cheilocystidia 23–36 × 11–
13 µm, inconspicuous, scattered. Paracystidia 9–30 × 4.5–16
µm, sometimes incrusted, walls sometimes or often slightly
thickened and yellowish pigmented, numerous, dominating.
Cells below marginal cystidia with pale pigment, incrusted.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: The ITS sequence places this taxon as an immediate
relative of P. albescens, but this sample has very variable
pleurocystidia, and, in accordance with Smith’s description,
a relevant number of them are subfusiform, progressively
attenuate towards the apex, resulting in an undifferentiated
transition between the ventricose and the apical part. This is a
pattern not usually found in P. albescens. Moreover, differently
from P. albescens, we have only found some subphaseoliform
spores, none distinctly phaseoliform or indented above the
apiculus. The pileus is yellowish fulvous when young, and
dull reddish cinnamon when mature, but red pigmentation in
the pileus of P. albescens is currently unknown. Given that we
have demonstrated how Smith’s concept of P. agrariella var.
agrariella is not univocal, we believe P. ochrofulva stands alone
as an autonomous species.
Psathyrella lactobrunnescens, described only from the
type collection on hardwood debris, is a species that could
not be loaned for scarcity of material. Judging from its
description, it belongs to subsection Spadiceogriseae, and,
based on a provisional morphological analysis, it resembles P.
ochrofulva, with whom it shares some peculiar characters such
as little differentiated, apically attenuate and never capitate
pleurocystidia; small and fragile sporocarps; and pigmented
paracystidia. On the contrary, its pilei are described as conical
and hardly expanding, while they are obtuse to convex or plane
in P. ochrofulva. The chromatic evolution of the pileus, initially
white and striate, then greyish brown, and finally faded white
again, is unusual and may suggest an albino form.
14. Psathyrella fatua (Fr.) Konrad & Maubl., Encycl. mycol. 14
(Agaricales 1): 125. 1948. Figs 35–43.
Pileus 8–52(–62) mm broad, in early stages broadly conical to
paraboloid, then expanding to convex, finally plane, with straight
to reflexed margin, subumbonate or not, sometimes rugulose
to wavy in old sporocarps; when young reddish brown, when
mature already discolored through brown to ochraceous brown,
134
Fig. 35. Psathyrella fatua. MCVE29105. Photo P. Voto.
Fig. 36. Psathyrella fatua. MCVE29109. Photo P. Voto.
yellowish brown, brownish orange, pale to medium Dresden
brown, pale pinkish chestnut brown, yellow red, sometimes
shiny, hygrophanous, quickly fading to whitish, cream,
alutaceous, pale to rich pink, pale greyish to pinkish violaceous,
with centre pale greyish ochraceous, pale ochraceous, yellowish
to vivid yellow, whitish, pale pinkish ochre, finally drying from
margin to greyish violaceous; translucently striate when moist.
Veil white, quickly evanescent, when young of flocci or fibrils
on stipe (mainly on lower half) and on the margin of the pileus
or appendiculate from it. Lamellae broadly adnate and usually
with a subdecurrent tooth, moderately broad, not or slightly
ventricose, crowded to moderately crowded; when young clay
to pale brownish grey, then through pinkish clay, lilaceous greybrown or purplish grey to finally dark purplish brown; edge
white, sometimes reddish brownish pigmented. Stipe (10–)
20–110(–125) × (1–)2–5.5(–7) mm, straight to flexuous, equal
to slightly progressively broadened towards base, sometimes
bent towards base, extreme base sometimes attenuate or
characterized by a pseudorhiza; white, sometimes discoloring
pale brownish; apex pruinose and sometimes striate, base
sometimes felted to substrigose. Context rather rigid in the stipe
that usually snaps when bent, whitish to pale, sometimes more
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 37. Psathyrella fatua. MCVE29122. Photo P. Voto.
Fig. 38. Psathyrella fatua. MCVE29124. Photo P. Voto.
Fig. 39. Psathyrella fatua. MCVE29124. Photo P. Voto.
Fig. 40. Psathyrella fatua. MCVE29124. Photo P. Voto.
Fig. 41. Psathyrella fatua. Voto 20181105.2. Photo P. Voto.
Fig. 42. Psathyrella fatua. MCVE29115. Photo M. Maletti.
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Voto et al.
Editor-in-Chief
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E-mail: p.crous@westerdijkinstitute.nl
Fig. 43. Psathyrella fatua. Voto, MCVE29109; MC Marginal Cystidia in KOH; PC1 Pleurocystidia in Congo red; PC2 Pleurocystidia in NH4OH; S1 Spores
in water; S2 Spores in KOH.
pigmented in stipe base. Odor and taste indistinctive. Spore
deposit purplish black. Spores (6.3–)6.8–10.5(–11.7) × 4–5.6
µm, on average 7.7–9 × 4.5–5.2 µm, Q (1.4–)1.5–2.2(–2.3),
avQ 1.6–2; in front view elliptic, oval, oblong, subcylindrical,
sometimes irregular, rarely angular-oval to cylindrical-oboval,
base truncate, rounded to obtuse, in profile adaxially flattened
to subamygdaliform, rarely or sometimes subphaseoliform,
rarely phaseoliform or indented above the apiculus; brown to
dark red; germ pore distinct, rounded to truncate, 1.2–2(–2.4)
µm broad. Basidia 4-spored, 15–25 × 7.5–10 µm. Pleurocystidia
(21–)30–60(–65) × 9–19(–21) µm, broadly to narrowly utriform,
fusiform, conical, lageniform, sometimes submucronate,
rarely clavate, ellipsoid, subcylindrical to rhomboid; apex
rounded to obtuse, rarely subobtuse, sometimes or often
subcapitate, rarely forked or incrusted; walls hyaline or very
slightly yellowish but never brownish, sometimes thickened
below apex; numerous to scattered. Cheilocystidia 20–45(–60)
× 8–18 µm, rare to scattered, sometimes (locally) moderately
numerous. Paracystidia 8–40(–45) × 5–23(–35) µm, sometimes
incrusted; walls often thickened and yellowish to reddish brown
pigmented (overall towards base); numerous, dominating. Cells
below marginal cystidia often incrusted, pale yellow to yellowish
brown with pigment often more concentrated at septa and in
discontinuous wall traits. Clamps present.
136
Habitat: Solitary, gregarious to cespitose; in foliar and on woody
debris in broadleaf wood stands, in parks, on paths, in grass; dry;
on calcareous and nitrophilous soil.
Material examined: Italy, Venezia, Chioggia, S. Anna, Nordio forest, 2 m
a.s.l., 29 Oct. 2006, gregarious on litter of Quercus ilex, P. Voto, Fig. 35
(MCVE29105, Tef-1α MF521809, ITS MF325966); at same location, 31
Oct. 2007, gregarious on litter of Quercus ilex, P. Voto (MCVE29110, Tef1α MF521806); Rovigo, Crespino, 1 m a.s.l., 28 Nov. 2006, gregarious on
soil at the top of a ditch bank, P. Voto, Figs 36, 43 (MCVE29109, Tef-1α
MF521807); Rovigo, Villadose, 3 m a.s.l., 19 Nov. 2013, gregarious on
dry grassy soil in a wooded park, P. Voto, Fig. 37 (MCVE29122, Tef-1α
MF521804, ITS MF325968); at same location, river Adigetto, 16 Nov.
2014, gregarious on often moist grassy ground at the top of the river
bank, P. Voto, Figs 38–40 (MCVE29124, Tef-1α MF521802); at same
location on a different spot, 5 Nov. 2018, gregarious, P. Voto 20181105.2,
Fig. 41; at same location on a different spot, 16 Nov. 2015, gregarious
and subcespitose, P. Voto (MCVE29333); Pesaro Urbino, Fano, loc.
Roncosambaccio, gregarious on buried roots, 24 Nov. 2006, M. Maletti
1867 (duplicate MCVE29108, Tef-1α MF521808, ITS MF325967); Pesaro
Urbino, Monte Giano, loc. Rio Secco, 14 Oct. 2011, gregarious among
grass on disturbed soil, M. Maletti 3035, Fig. 42 (duplicate MCVE29115,
Tef-1α MF521805).
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Notes: One of our aims was to provide molecular evidence
supporting the presence of pilei with pinkish tints and of
reddish gills’ edges within this species. Ludwig (2007b) was, to
our knowledge, the first to report a possible red underlining
of the gill edge. A number of collections we have examined
show these two parameters can be variously combined, even
in materials collected in the same place. Cross comparisons
of the sequences originated from various samples show that
all of our materials share a minimum homology value of 99.7
% at the ITS locus and an average 99 % Tef-1α identity, with
the only exception of MCVE29124 which shares a 97 % to 98
% identity with all others. However molecular evidence does
not provide substantial support to split this single voucher
from the others. In conclusion, our molecular identification
of P. fatua includes material with (see Figs 35, 37, 39) or
without a more or less evident pinkish peripheral half of
the discoloring pileus, and with (see Fig. 37) or without a
pigmented gill edge.
Elements differentiating P. fatua from P. spadiceogrisea
are listed below. Spores are less frequently phaseoliform and
rarely indented above the apiculus. Pleurocystidia are never
brown, are more varied in shape (utriform, lageniform, fusiform,
conical), and sometimes are more or less capitate, rarely forked
or incrusted. Gill edges are possibly pigmented. Sporocarps are
smaller and with a rapidly discoloring pileus that is never dark
colored when mature, while it may be pinkish. For differences
between P. fatua and P. albescens, see notes provided for the
latter.
15. Psathyrella tenacipes A.H. Sm., Mem. N. Y. Bot. Gard. 24:
440. 1972. Fig. 44.
Typus: Psathyrella tenacipes A.H. Sm. USA, Michigan, Cheboygan
Co., Wolf Bog, 13 Jun. 1948, A.H. Smith 28718 (incorrectly
28717 in original description) (coll. Singer, DiGilio & Smith)
(MICH12091), described as gregarious on muck and small sticks
partly burned in protologue, described as gregarious on drying
muck in ash swamp in Smith’s handwritten field notes.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Loan of various sporocarps in good condition for micromorphology investigation.
Spores 7–9.5(–10.1) × 4.6–5.7 µm (7–9 × 4.5–5.5 in Smith
1972), Q (1.45) 1.6–1.85(–1.9), avQ 1.7; in front view suboval
to elliptic, sometimes subangular to suboblong, base rounded
to subtruncate, in profile adaxially flattened, sometimes
subamygdaliform to subphaseoliform, orangish reddish brown;
germ pore distinct, truncate, 1.2–1.7 µm broad. Pleurocystidia
28–50 × 9–14(–16) µm, (narrowly) utriform, sometimes
lageniform to subcylindrical; apex 4.5–8(–10) µm broad, rounded
to obtuse, often or sometimes subcapitate, not incrusted; walls
thin, hyaline; moderately numerous. Cheilocystidia 24–30 ×
10–15 µm, scattered. Paracystidia –30 × –17(–20) µm; walls
thickened and yellowish pigmented in larger ones, numerous,
dominating. Cells below marginal cystidia yellowish pigmented.
Unfortunately, DNA extraction failed for this species, which
otherwise is remarkable for the presence of some peculiar
Fig. 44. Psathyrella tenacipes. A.H. Smith 28718, MCVE12091; PC Pleurocystidia; MC Marginal Cystidia in KOH; S1 Spores in water; S2 Spores in KOH.
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characters, such as a scarce and very fugacious veil; a tubular,
pliant, fairly tough stipe; pallid brownish gill edges and bistre
but soon discoloring pileus color. Based on micro- and gross
morphology, we see some similarity between P. tenacipes and P.
fatua, possibly suggesting conspecificity. However, because of
the lack of molecular evidence of P. fatua from any American
collections included in this study, we are not yet ready to recognize
this common European species as being present in North America.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
16. Psathyrella owyheensis A.H. Sm., Mem. N. Y. Bot. Gard. 24:
174. 1972. Fig. 45.
Molecular heterotypical synonyms proposed here: Psathyrella
carminei Örstadius & E. Larss., Mycol. Prog. 14: 18. 2015.
Typus: Psathyrella owyheensis A.H. Sm. USA, Idaho, Owyhee
Co., Sands Basin, 27 Apr. 1969, E. Trueblood 3135 (MICH5357),
gregarious in native grass near cow dung. Tef-1α MF521785, ITS
MF325990.
Loan of two sporocarps in good condition for micro-morphology
investigation.
Spores (7.5–)8.8–11.1(–12.5) × (3.8–)4.5–5.7(–6.5) µm, on
average 9.4 × 5.2 µm, Q (1.4–)1.6–2(–2.5), avQ 1.8–1.9; in
front view oval, elliptic, oblong, sometimes subcylindrical,
sometimes or often slightly irregular between median portion
and base, base obtuse to broadly rounded, in profile subelliptic
to subamygdaliform, sometimes subphaseoliform; dull
orange; germ pore distinct, often truncate, 1.5–2 µm broad.
Pleurocystidia 37–59 × 12–16 µm, utriform to lageniform,
sometimes subcylindrical; apex obtuse, sometimes subcapitate,
rarely forked or incrusted; walls sometimes weakly yellowish
pigmented; more or less scattered. Cheilocystidia 22–35 × 9–12
µm, scattered. Paracystidia –23 × –15 µm, walls hyaline or
almost so, numerous, sometimes incrusted, dominating.
Notes: The molecular sequences of the holotype of P. carminei
deviates by a few bases from the holotype of P. owyheensis
(DNA nucleotide homology: ITS = 99 % identity, Tef-1α = 98 %
identity), we consider this to be a good double locus evidence
of conspecificity; micro-morphology matches well too. We
could not ascertain the exact trophism of the holotype (and only
collection) of P. owyheensis: it could be the nitrogen-rich soil, due
to the nearby dung, or grassy material, we cannot even exclude
sagebrush (Artemisia tridentata) debris as this plant is widely
distributed in the Sands Basin. Psathyrella carminei is reported
on soil in a forest with pines but there are a number of species
in subsection Spadiceogriseae growing both with and without
association to woody material, e.g. P. fatua, P. clivensis and P.
albescens. Psathyrella carminei is said to smell of fish while P.
owyheensis has indistinct odor, but we know of other Psathyrella
Fig. 45. Psathyrella owyheensis. E. Trueblood 3135, MICH5357; PC1 Pleurocystidia in NH4OH; PC2 Pleurocystidia in Congo red; MC Marginal Cystidia;
S1 Spores in water; S2 Spores in KOH.
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A revision of Psathyrella, subsection Spadiceogriseae
species, e.g. P. bipellis and P. saponacea, which may or may not
possess a peculiar odor.
17. Psathyrella hellebosensis Deschuyteneer & Melzer, Bull
AMFB 10: 5. 2017. Figs 46, 47.
Spores (6.5–)7–8.5(–11) × 4.7–5.3(–5.6) µm, Q (1.4–)1.5–1.8, in
front view oval to angular-oval, with a base often subtruncate,
in profile rarely subphaseoliform, reddish brown, with distinct
germ pore. Pleurocystidia, 43–58(–64) × 11–15(–17) µm,
utriform, often forked to three-lobed. Paracystidia sometimes
incrusted and locally weakly yellowish pigmented, dominating.
thujina, however it has spores mainly and distinctly phaseoliform,
and its habitat is not specifically defined wet or moist soil. It
remains a dubious European taxon. Psathyrella phaseolispora,
whose type could not be studied, is a species described based
on a single dilapidated old specimen found in a manured PoaLolietum grassland, and resembles P. clivensis but with distinctly
phaseoliform and somewhat more pigmented spores, and a
somewhat more defined germ pore; it too remains dubious.
However, pending more morphological descriptions or convincing
molecular evidences, we decided to insert these two species in
our key to facilitate their identification from more collections.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Material examined: Italy, Pesaro, loc. Villa Ceccolini, near river Foglia,
26 Mar. 2011, M. Maletti 2768 (MCVE29114), gregarious on soil in an
unkempt open riparian site. Tef-1α MF521816, ITS MF325956.
Notes: The colors and habitus of this Italian collection are in
perfect agreement with the description of this species. The
molecular evidence of the Tef-1α DNA region clearly isolates this
species from neighboring allied taxa, and places it confidently
among species of the subsection Spadiceogriseae that are not
found in association with woody material.
Two other species should be mentioned here because of
their non-lignicolous habitat. Psathyrella agraria (Enderle 1996),
reported from mossy soil in a cornfield (Zea mays, Poaceae),
shares a 99 % ITS identity with the types of P. lutulenta and P.
Fig. 46. Psathyrella hellebosensis. MCVE29114. Photo M. Maletti.
Fig. 47. Psathyrella hellebosensis. P. Voto, MCVE29114; MC Marginal Cystidia in KOH; PC Pleurocystidia; S1 Spores in water; S2 Spores in KOH.
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18. Psathyrella rogersiae Voto, Garbelotto & Chiarello, sp. nov.
MycoBank MB831126. Figs 48, 49.
Typus: Italy, Veneto, Vicenza, Trissino, river Guà, about 100 m
a.s.l., 28 Mar. 2013, O. Chiarello (MCVE29120). ITS MF325995.
Pileus 30–40 mm broad, soon convex with inflexed margin,
when mature slightly wavy and sometimes slightly umbonate;
when young dark brown to dark grey-brown, when mature
brown to dark yellowish brown, hygrophanous, fading
through various tints of yellowish brown (greyish, dull,
medium, bright), finally pale grey; faintly translucently striate
when moist. Veil white, as a scarce and fugacious fringe on
margin of pileus. Lamellae adnate or with a decurrent tooth;
moderately narrow to somewhat ventricose in age; about
Pileus 30–40 mm latus, mox convexus sed margine inflexa, per
mature leviter undulatus et interdum leviter umbonatus; primo
umbrinus vel murinus, per mature brunneus, deinde spadiceus,
hygrophanus, in sicco cinereus; humido leviter striatus. Velum
album, fibrillosum, pauper, fugax. Lamellae adnatae, modice
ventricosae, subdistantes; initio griseae, deinde purpureae, acie
alba. Stipes 35–50 × 3.5–6 mm, cylindraceus, albus. Odor ut sapor
indistincti. Sporae (6.8–)7.5–10 × 4.6–5.2(–5.5) µm, oblongae,
ellipsoideae, subovoideae, nonnullae subphaseoliformes, in
aqua observatae aurantiacae-brunneae, poro germinativo
distinctae. Basidia 4-sporigera. Pleurocystidia 34–54(–59) × 13.3–
16.5 µm, utriformia vel sublageniformia, apice obtusa, interdum
vel saepe subcapitata, raro furcata, numerosa. Cheilocystidia
28–45 × 7.5–15 µm, pleurocystidiis similia vel fusoidea, dispersa
sed modice numerosa ad stipitem. Paracystidia 15–35 × 7.5–21
µm, interdum leviter flavida ut crassitunicata, numerosa. Fibulae
adsunt. Gregaria in udo prati.
Fig. 48. Psathyrella rogersiae. MCVE29120. Photo O. Chiarello.
Etymology. The epithet is a dedication to Patricia Rogers, a
collection manager in the University of Michigan Herbarium
who helped us navigate the collections at the MICH herbarium.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 49. Psathyrella rogersiae. O. Chiarello, MCVE29120; PC Pleurocystidia; MC Marginal Cystidia in KOH; S1 Spores in water; S2 Spores in KOH.
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A revision of Psathyrella, subsection Spadiceogriseae
28 when mature; when young greyish, finally purplish; edge
white. Stipe 35–50 × 3.5–6 mm, cylindrical, occasionally bent
towards apex or base; white, a little discoloring brownish over
the basal area; apex pruinose, naked or with very scarce velar
fibrils. Odor and taste indistinctive. Spores (6.8–)7.5–10 × 4.6–
5.2(–5.5) µm, on average 8.2–9.2 × 4.8–5 µm, Q (1.5–)1.53–
1.88 (–1.96), avQ 1.68–1.75; smooth; in front view elliptic to
suboval, sometimes sub angular-oval, oblong, base rounded
to subtruncate, in profile adaxially flattened, sometimes
subphaseoliform, rarely distinctly phaseoliform; dull orange
to brownish orangish; germ pore distinct, rounded, 1.3–1.7
µm broad. Basidia 4-spored, 17–28 × 8–11 µm. Pleurocystidia
34–54(–59) × 13.3–16.5 µm, with quotient 2.7–3.8(–4.2), on
average 3.42; utriform to sublageniform, with often elongate
and sometimes eccentric neck, apex rounded to obtuse,
often or sometimes subcapitate, rarely forked; numerous.
Cheilocystidia 28–45 × 7.5–15 µm, sometimes varying to
obtusely subfusiform, scattered at margin of pileus to
moderately numerous towards stipe. Paracystidia 15–35 ×
7.5–21 µm, sometimes elongate-ellipsoid, walls sometimes
thickened and yellowish pigmented, numerous, dominating.
Cells below marginal cystidia incrusted and weakly yellowish
pigmented. Pileipellis of clavate to subglobose, 13–30(–40)
µm broad cells; underlying hyphae with incrusted pigment.
Clamps present.
Habitat: Gregarious; in sandy grassland near a river bank, the area is
a moist, often flooded, permeable alluvial plain with Salix alba trees
and various hygrophilous plants, such as Carex spp., Phragmites
australis, Typha latifolia, Scirpus spp., Ranunculus aquatilis.
Notes: The ITS sequence places this fungus in the subclade P.
albescens. The differences against P. albescens are many and
evident, ranging from not thickset utriform pleurocystidia and
somewhat less phaseoliform spores to distinctly dark grey brown
colors of pileus, scarce veil and hygrophilous habitat; P. ochrofulva
is a tiny, differently colored species with different pleurocystidia
and habitat. Because of its hygrophilous habitat it can be compared
to the American taxon P. thujina, which Örstadius et al. (2015)
proposed as a prior name for the European P. almerensis and
P. lutulenta, but its smaller spores and slenderer pleurocystidia
represent a distinctive combination of characters. It is also
comparable to P. hellebosensis which differs mainly because of its
more pigmented, largely ovoid to angular-ovoid spores.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
19. Psathyrella argillacea A.H. Sm., Mem. N. Y. Bot. Gard. 24:
367. 1972. Fig. 50.
Paratype: Psathyrella argillacea A.H. Sm. USA, Wyoming, Albany
Co., Laramie River, North Fork River, 27 Jul. 1950, A.H. Smith
35220 (MICH65256). Tef-1α MF521814.
Fig. 50. Psathyrella argillacea. A.H. Smith 35220, MICH65256; PC1 Pleurocystidia in NH4OH; PC2 Pleurocystidia in Congo red; MC Marginal Cystidia in
KOH; S1 Spores in water; S2 Spores in KOH.
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Voto et al.
Loan of three pilei and some entire or broken stipes for gross
morphology investigation.
Spores 7.3–11 × 4.4–5.6(–6.2) µm, Q 1.5–2.5, avQ 1.8–2; in front
view elliptic to (narrowly) oval, base rounded to subtruncate, in
profile adaxially flattened, subcylindrical to subamygdaliform,
rarely subphaseoliform; dull orangish brown to brown; germ
pore distinct, rounded to subtruncate, about 1.8 µm broad.
Pleurocystidia 38–55 × 12–15 µm; utriform, fusiform-utriform;
apex obtuse to rounded, rarely subacute, often forked, often
incrusted, walls sometimes brownish pigmented; moderately
numerous. Cheilocystidia 25–45 × 12–15 µm, scattered.
Paracystidia 12–30 × 5–19.5 µm, sometimes ellipsoid, often
incrusted, walls often thickened and yellowish pigmented,
numerous, dominating. Cells below marginal cystidia slightly
incrusted and pigmented.
Lauricolae expressly characterized by lignicolous habitat. The
lack of macro characters in the description of this paratype
prevents a thorough comparison with P. owyheensis, a very
similar species based on micro-morphology (see key). Yet, the
two are surely different based on DNA evidence.
20. Psathyrella umbrosa A.H. Sm., Mem. N. Y. Bot. Gard. 24:
391. 1972. Fig. 51.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: Best match when blasting the sequence of this material
is with the holotype of P. lutulenta but with a weak identity
that undoubtedly excludes conspecificity. No field notes are
attached to this paratype and on careful examination of the
whole exsiccata no fragments of non-fungus material could be
found for a clue to the habitat; however the holotype, which we
could not study for scarcity of material, and all other collections
examined by Smith (1972) are reported on hardwood debris and
litter, moreover Smith (1972) placed the species in subsection
Molecular heterotypical synonyms proposed here: Psathyrella
fulvoumbrina A.H. Sm., Mem. N. Y. Bot. Gard. 24: 413. 1972.
Typus: Psathyrella umbrosa A.H. Sm. USA, New Mexico, Rio
Arriba Co, Sangre de Cristo Mountains, Aug. 1963, C.A. Barrows,
1416 (MICH12102), densely gregarious on soil filled with rotten
wood. Tef-1α MF521775, ITS MF326003.
Loan of three sporocarps in good condition for micro-morphology
investigation.
Spores (6.7–)7.1–9.3 × 4.3–5.7 µm, Q 1.4–2.1; in front view
suboblong to broadly oval, rarely subangular-oval, base mainly
broadly rounded, rarely subtruncate, in profile subelliptic
to adaxially flattened or subphaseoliform, sometimes
subamygdaliform, apex truncate in front view and in profile;
orangish to vaguely reddish brown; germ pore distinct, truncate,
Fig. 51. Psathyrella umbrosa. C.A. Barrows 1416, MICH12102; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH; GM
Greenish masses/incrustations in hymenium.
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A revision of Psathyrella, subsection Spadiceogriseae
1.5–1.8 µm broad. Pleurocystidia 31–51 × 10.5–16 µm, utriform
to sublageniform or subfusiform, sometimes ventricose-conical;
apex obtuse to subcapitate; walls hyaline; moderately numerous.
Cheilocystidia 23–40 × 8–14 µm, scattered. Paracystidia mainly
9–12 × 5–7 µm, some 22–32 × 9–13 µm, dominating among
numerous basidia and basidiola.
20a. Psathyrella fulvoumbrina A.H. Sm., Mem. N. Y. Bot. Gard.
24: 413. 1972.
Typus: Psathyrella fulvoumbrina A.H. Sm. USA, Idaho, Owyhee
Co, Boulder Creek, South Fork, 4 Jun. 1969, E. Trueblood 3167
(MICH5833), gregarious under aspen and fir. Tef-1α MF521800,
ITS MF417625.
Loan for micro-morphology investigation of four sporocarps in
good condition.
Spores 7–8.7(–9.5) × 4.4–5.7 µm, Q 1.3–1.9; in front view elliptic
to oval, base broadly rounded, in profile subelliptic to adaxially
flattened or subamygdaliform, rarely subphaseoliform; redbrown; germ pore distinct, mostly truncate, 1.5–1.8 µm broad.
Pleurocystidia 33–55 × 10–22 µm, narrowly utriform, narrowly
sublageniform, narrowly fusiform-utriform, broadly ventricose-
conical; apex subcapitate or not, sometimes (irregularly)
capitate, sometimes subacute, sometimes forked; walls hyaline;
moderately numerous. Cheilocystidia 22–27 × 9–12 µm, scattered.
Paracystidia mainly small, some 20–30 × 10–13 µm, dominating
among numerous, up to 10 µm broad basidia and basidiola.
Note: There is full correspondence, both molecular and
morphological, between P. fulvoumbrina and P. umbrosa
therefore they are synonyms.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
21. Psathyrella praetenuis A.H. Sm., Mem. N. Y. Bot. Gard. 24:
110. 1972. Fig. 52.
Molecular heterotypical synonyms proposed here: Psathyrella
atrifolia sensu A.H. Smith, Mem. N. Y. Bot. Gard. 24: 146. 1972,
p.p.
Typus: Psathyrella praetenuis A.H. Sm. USA, Idaho, Bonner Co.,
Upper Priest River, 1 Oct. 1968, A.H. Smith 76781 (MICH5361),
gregarious on debris under cottonwood trees. Tef-1α MF521782,
ITS MF325993.
Loan of three sporocarps in good condition for micro-morphology
investigation.
Fig. 52. Psathyrella praetenuis. A.H. Smith 76781, MICH5361; PC Pleurocystidia in NH4OH; MC Marginal Cystidia in KOH; S1 Spores in water; S2 Spores
in KOH.
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Spores (7.5–)8–11(–11.8) × 4.5–6 µm, Q (1.5–)1.7–2.1(–2.2),
avQ 1.9; in front view elliptic to oblong, sometimes suboval,
base rounded, in profile subelliptic to subamygdaliform,
sometimes subphaseoliform; sordid brown with a reddish
orangish tinge; germ pore distinct, mostly truncate, 1.6–1.7 µm
broad. Pleurocystidia 33–55 × 11.5–17 µm, utriform, sometimes
subcylindrical; apex obtuse, sometimes forked, sometimes
(irregularly) subcapitate, often incrusted; walls sometimes
brownish pigmented; more or less moderately numerous.
Cheilocystidia 22–50 × 11–22 µm, scattered. Paracystidia 9–39(–
49) × 7–22 µm, sometimes submucronate, walls often thickened
and pale yellowish pigmented; numerous, dominating. Cells
below marginal cystidia faintly yellowish pigmented and
incrusted.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: Smith (1972) defines this species as devoid of any trace
of veil but places it in subgen. Pannucia, section and subsection
Appendiculatae on the assumption that sporocarps may start
with an appendiculate veil. For further comments see next
species.
Psathyrella atrifolia* (Peck) A.H. Sm., Contr. Univ. Mich. Herb.
5: 38. 1941.
*The original name ‘atrofolia’ should be correctly spelled
‘atrifolia’.
Notes: Smith (1972) has proposed a description of this taxon based
on numerous collections but he did not report having revised
the holotype of the basionym, Hypholoma atrofolium Peck. We
have attempted to understand Smith’s concept of P. atrifolia by
examining three of his collections selected for this purpose by
MICH staff. There is a conspicuous correspondence, both molecular
(DNA nucleotide homology: ITS= 100 %) and morphological,
between P. praetenuis and Smith 73945 (MICH32732) therefore
they are conspecific; it is to be noticed that both materials were
found among cottonwood tree (Populus deltoides) debris. Smith
87038 (MICH65288), found under aspen (Populus spp.), is likewise
morphologically identical but failed to produce a sequence;
however, we consider it conspecific too. DNA extraction failed
also with Smith 86968 (MICH65287), habitat unknown, but in this
case we have found some differences in the spore parameters and
it may not be identical to the other vouchers, indeed it recalls P.
argillacea. Of course these results do not necessarily apply to the
other vouchers Smith deposited in MICH under this name, nor can
we be assured of the correct identity between Smith’s concept
and Peck’s basionym as Peck described only, and partially too,
the spores among the microscopical characters. Here follows our
revision of Smith 73945.
21a. A.H. Smith 73945, MICH32732.
= Psathyrella praetenuis A.H. Sm.
Material examined: USA, Idaho, Bonner Co., Priest Lake, Tule Bay, 10
Oct. 1966, in a cottonwood flat. Tef-1α MF521813, ITS MF325960.
Loan of many sporocarps in good condition for micromorphology investigation.
Spores (7.4–)8.3–9.7(–11.1) × (3.9–)4.5–5.7(–6.3) µm, on average
9 × 5.1 µm, Q (1.5–)1.7–1.9(–2.2), avQ 1.85; in front view elliptic,
144
suboval to oblong sometimes angular-oval, base rounded to
subtruncate, in profile adaxially flattened to subellipsoid, rarely
or sometimes subphaseoliform, sordid brown with a reddish
orangish tinge; germ pore distinct, mostly truncate, 1.6–1.8 µm
broad. Pleurocystidia 30–54 × 10–20(–21.5) µm, utriform; apex
obtuse to rounded, rarely subacute, sometimes forked, rarely
incrusted; walls hyaline; moderately numerous. Cheilocystidia
26–44 × 10–15 µm, rare. Paracystidia 9–38 × 6–19.5 µm,
sometimes submucronate, walls often thickened and pale
yellowish brownish pigmented, scarcely incrusted; numerous,
dominating. Cells below marginal cystidia often pale yellowish
brownish pigmented, scarcely incrusted.
22. Psathyrella griseopallida Thiers & A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 417. 1972. Fig. 53.
Typus: Psathyrella griseopallida Thiers & A.H. Sm. USA, California,
San Diego Co, Lake Henshaw, San Luis Rey Campground, 10
Mar. 1970, H.D. Thiers 25079 (MICH11939), Smith’s protolog:
gregarious on sandy soil near an oak log, Thiers’s field note:
gregarious in vicinity of dead oak logs. Tef-1α MF521796, ITS
MF325974.
Loan of one entire pileus and a narrow slice of pileus for micromorphology investigation.
Spores (7.3–)8.85–11.9(–13.4) × (4.3–)4.8–5.7(–6.2) µm (8–10 ×
4–5 µm in Smith, 1972), on average 10.2 × 5.29 µm, Q (1.6–)
1.7–2(–2.2); in front view oblong, oval, subtriangular, base
rounded to sometimes truncate, in profile adaxially flattened
to faintly subphaseoliform, sometimes subamygdaliform,
rarely phaseoliform; red-brown; germ pore distinct, truncate
to broadly rounded, 1.7–2 µm broad. Pleurocystidia (28–)32–
60 × 10–19 µm, utriform, subcylindrical, conical, ventricoseconical, sometimes subellipsoid to sublageniform; apex obtuse
to rounded, sometimes or often subcapitate, sometimes
incrusted; walls hyaline; (moderately) numerous. Cheilocystidia
22–50 × 10–17 µm, mainly lageniform to attenuate, apex mainly
obtuse to sometimes subacute, scattered to locally moderately
numerous. Paracystidia 8–28(–30) × 5–17 µm, apex often
incrusted, walls thickened and at most very faintly pigmented in
larger ones, otherwise thin and hyaline, numerous, dominating.
Note: Fragments of leaves are clearly recognizable scattered
among the exsiccata and connected to some stipe bases,
confirming the association with arboreal material.
23. Psathyrella pseudolimicola A.H. Sm., Mem. N. Y. Bot. Gard.
24: 262. 1972. Fig. 54.
Typus: Psathyrella pseudolimicola A.H. Sm. USA, Idaho, Idaho
Co, Seven Devil’s Mountains, Nezperce National Forest, Papoose
Creek, 10 Aug. 1962, A.H. Smith 65850 (MICH5366), gregarious
on mud in a cow pasture (mud and manure present). Tef-1α
MF521781.
Loan of three sporocarps in good conditions for micromorphology investigation.
Spores 8.5–10.7(–11.8) × 4.9–5.9(–6.7) µm, on average 9–10.5
× 5–5.5 µm, Q (1.6–)1.7–1.9(–2); in front view (narrowly)
oval to elliptic-oval, base obtuse to rounded, sometimes
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A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 53. Psathyrella griseopallida. H. D. Thiers 25079, MICH11939; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in
water; S2 Spores in KOH.
Fig. 54. Psathyrella pseudolimicola. A.H. Smith 65850, MICH5366; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in
water; S2 Spores in KOH.
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Voto et al.
truncate, in profile (narrowly) adaxially flattened-ellipsoid to
subphaseoliform, sometimes phaseoliform; reddish brown;
germ pore distinct, rounded to subtruncate, 1.5–1.9 µm broad.
Pleurocystidia 33–53 × 10–16(–19) µm, utriform, sometimes
ellipsoid-utriform, lageniform; apex rounded to obtuse,
subcapitate or not; walls sometimes faintly pigmented; scattered
to moderately numerous. Cheilocystidia 25–43 × 12–13 µm, often
(broadly) lageniform, sometimes cylindrical-utriform (recalling
the broadly cylindrical-utriform pleurocystidium sketched by
Smith 1972), scattered. Paracystidia up to 35 × 15 µm, hyaline,
numerous, dominating. Cells below gill edge hyaline.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: Smith (1972) reports the spores 8.5–11 × 4.5–5 (up to 12 ×
6) µm as if he had found only a few broader spores. The molecular
evidence places it beside the previous species, P. griseopallida,
but at a consistent distance (DNA nucleotide homology: Tef-1α =
95 %). Smith (1972) commented having a doubt as to whether
this species is coprophilous or not as ‘it was not found on cow
dung though ample opportunity was at hand’; on examination of
the exsiccata multiple loosen dried leaves and a small amount of
crumbled unidentifiable plant material were found in the box with
the dried fungi but none attached at stipe bases. We presume the
habitat to be either herbicolous or on nitrogen-rich soil.
24. Psathyrella velatipes A.H. Sm., Mem. N. Y. Bot. Gard. 24:
442. 1972. Fig. 55.
= Psathyrella ammophila (Durieu & Lév.) P.D. Orton
Typus: Psathyrella velatipes A.H. Sm. USA, Michigan, Washtenaw
Co., Ann Arbor, Liberty St., 30 Sep. 1971, A.H. Smith 80623
(MICH12106), gregarious-cespitose in a parking strip. Tef-1α
MF521774, ITS MF326004.
Loan for gross morphology investigation of four sporocarps,
some of which broken.
Spores (8.2–)8.7–10.5(–12) × 5–5.8 µm, on average 9.9 × 5.5 µm,
Q (1.5–)1.7–2.1(–2.2), avQ 1.8; in front view cylindrical-oblong
to elliptic, base obtuse, in profile subamygdaliform, sometimes
subphaseoliform, base often with a suprahilar depression;
reddish brown; germ pore distinct, sometimes eccentric in
profile, 1.8–2(–2.5) µm broad. Pleurocystidia 28–40 × 11–15
µm; narrowly utriform, lageniform, sometimes fusiform-conical
to flexuous, rarely ovoid-utriform to thickly utriform, often
emerging for only 10 µm length or less, neck often elongate;
apex obtuse to rounded, sometimes subcapitate, rarely forked;
scattered. Cheilocystidia 28–55 × 10–15 µm, sometimes
submucronate, pigment as in the paracystidia, rare to scattered
towards margin of pileus, moderately numerous towards stipe.
Paracystidia –37 × –21 µm, often incrusted, walls often thickened
and yellowish pigmented moreover towards base, numerous,
dominating. Cells below marginal cystidia often incrusted and
yellowish pigmented.
Fig. 55. Psathyrella velatipes. A.H. Smith 80623, MICH12106; MC Marginal Cystidia in KOH; PC Pleurocystidia; S1 Spores in water; S2 Spores in KOH.
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Notes: There is morphological and molecular evidence that
this species corresponds to P. ammophila. The distinctly
pigmented marginal cells recall the description of P. ammophila
f. marginata by Bon (1988). Smith (1972) was particularly
impressed by the ‘distinct fibrillose scurfy’ covering of the
stipe, but we know that P. ammophila has initially a dense velar
coating possibly leaving wooly layers on the stipe too. The
habitat in Ann Arbor town, far away from the sea shore and
sand dunes, is not an obstacle for recognition of this species
as sand from river or lakeshore caves is often used in making
the foundation of urban green areas; moreover coarse soil,
probably including grains of sand, was found attached to the
stipe base. The spores of P. ammophila are usually described
as larger by European mycologists, but Heykoop et al. (2017)
report collections with smaller spores, similar in size to those
of P. velatipes. Actually this species spores have a very wide
range, both in dimension and in quotient: (7.5–)8.5–14 × (4.7–)
5–8.5 µm, avQ 1.5–1.9.
Smith (1972) considered P. arenulina the American
counterpart of P. ammophila and Kits van Waveren (1985)
treated it as a synonym. We have not studied this material
but, basing on Smith’s (1972) description, it shares almost all
characters, including scarcity of pleurocystidia, indeed none
reported (‘normally very scarce and easily overlooked’ in
Kits van Waveren, 1972 for P. ammophila). Only the pileus is
unusually ‘widely and irregularly striate’ while the spores in KOH
are ‘cocoa-color slowly becoming darker’ which, as we explain
further, generally corresponds to a medium, not dark, color in
water. Overall we accept this synonymy.
Psathyrella dunarum, created on the base of a solitary
sporocarp found on a mossy coastal sand dune, has a strongly
striate pileus, scarce subobtusely lageniform pleurocystidia
and orange-brown, 8–9 × 4.5–5.5 µm, on average 8.7 × 5.4 µm,
spores. We see a strong correspondence between P. dunarum
and P. ammophila therefore, unless molecular evidence should
demonstrate convincingly the existence of an autonomous
species, we consider them synonyms.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
25. Psathyrella neotropica A.H. Sm., Mem. N. Y. Bot. Gard. 24:
108. 1972. Fig. 56.
Basionym: Drosophila pallidispora Murrill, Mycologia 10(2): 64
(1918). Typus: Cuba, Herradura, 17 Aug. 1907, F.S. Earle 572 (NY
00649148), in soil in a garden.
Loan for gross morphology investigation of fragments of pilei
and stipes.
Spores (7–)8–12.2(–13.8) × 5–7.5(–8.5) µm [9–12(–13)×5–6.5
µm in revision of Smith, 1972], on average 10.25 × 6.4 µm, Q
1.3–1.9(–2.05), avQ 1.6; wall often irregular in shape or with
protrusions, swellings or discontinuities, in front view suboval
to elliptic, sometimes cylindrical, base rounded to truncate, in
profile adaxially flattened, amygdaliform, subamygdaliform,
elliptical, phaseoliform to subphaseoliform; yellowish; germ
Fig. 56. Psathyrella neotropica. F. S. Earle 572, NY 00649148; Spores in NH4OH.
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pore generally absent to indistinct (callus), sometimes as a
protruding bubble in alkali, rounded, 1.5–1.8 µm broad when
discernible. Hymenial cystidia, basidia and clamps could not be
observed because the material revived very badly.
Typus: Psathyrella septentrionalis A.H. Sm. USA, Michigan,
Chippewa Co., Tahquamenon Falls State Park, near Paradise,
2 Jul. 1951, D. Palmer 36553 (MICH12045), on hardwood. ITS
MF326014.
Notes: Unfortunately, DNA sequencing failed because of
contamination and likewise our morphological revision too was
strongly hampered by poor rehydration of the sample. However
we are able to report an unpublished private revision, included
in the holotype box, Pegler made in 1986 in which he reported
spores 8.5–12×5.5–7 µm, on average 9.5–10.5 × 5.7–6.3 µm, Q
1.4, very pale, pleurocystidia ventricose-fusoid, cheilocystidia
spheropedunculate. Pegler’s notes seem to clearly implicate
that paracystidia are dominating on the gill edge; this and
the description by Smith (1972) of broadly rounded apex of
pleurocystidia confidently places this taxon into subsection
Spadiceogriseae. Beyond the anomalous characteristics of the
spore outline, which we tend to view as a casual sufferance
and not as a taxonomical attribute, we can remark on the short
pleurocystidia, 28–37 × 10–14 µm, as reported by Smith (1972).
Although we cannot know its phylogenetic position, for some
traits this material is reminiscent of the European P. clivensis
which we have not found in any of the American materials
studied.
Loan of two sporocarps and a pileus in good condition for micromorphology investigation.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
26. Psathyrella septentrionalis A.H. Sm., Mem. N. Y. Bot. Gard.
24: 134. 1972. Fig. 57.
Spores (7–)7.3–8.7(–9) × 4.1–5(–5.2) µm, Q 1.6–2(–2.1); in
front view elliptic to narrowly oval, base obtuse, sometimes
rounded, in profile adaxially flattened to subphaseoliform,
sometimes distinctly phaseoliform; orangish to reddish brown;
germ pore distinct, 1–1.5(–1.7) µm broad. Pleurocystidia 35–53
× (7–)9.5–20(–22) µm, utriform to obtusely ventricose-conical,
sometimes cylindrical-ellipsoid, conical to sublageniform; apex
obtuse to rounded, rarely subacute, sometimes subcapitate
or somewhat irregular, rarely capitate, rarely incrusted;
moderately numerous to numerous. Cheilocystidia 24–50 × 13–
22(–25) µm, numerous to abundant, dominating. Paracystidia
8–15 × 6–10 µm and numerous, 15–20 × 10–15 and moderately
numerous.
Notes: This is one of those species where we could not know, by
means of Smith’s (1972) vague description only, whether or not
it had the attributes of subsection Spadiceogriseae; molecular
evidence and abundant, dominating cheilocystidia clearly
exclude it.
Fig. 57. Psathyrella septentrionalis. D. Palmer 36553, MICH12045; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
148
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 58. Psathyrella vesiculosa. A.H. Smith 63485, MICH12044; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
27. Psathyrella vesiculosa (A.H. Sm.) Voto, Dovana & Garbel,
comb. nov. MycoBank MB831130. Fig. 58.
Basionym: Psathyrella septentrionalis var. vesiculosa A.H.
Sm., Mem. N. Y. Bot. Gard. 24: 136. 1972. A.H. Smith 63485,
MICH12044.
Typus: Psathyrella septentrionalis var. vesiculosa A.H. Sm., USA,
Michigan, Chippewa Co, Tahquamenon River, White House
Landing, 30 Jun. 1961, A.H. Smith 63485 (MICH12044), cespitose
on hardwood logs. ITS MF326015.
Loan of two sporocarps and a pileus in good condition for micromorphology investigation.
Spores 6.9–9.9(–10.6) × 3.5–5.5 µm, Q 1.4–2.1(–2.6), avQ
1.8–1.9; in front view oblong to elliptic, base obtuse to
attenuate, in profile adaxially flattened or with a faint suprahilar
depression, sometimes amygdaliform, subamygdaliform to
subphaseoliform; pale orange-brown, pale dull orange; germ
pore indistinct (callus), ± 1,5–1,7 µm broad. Pleurocystidia 29–50
× (10–)13–17 µm, utriform, fusiform, conical, clavate-mucronate
to lageniform, sometimes ellipsoid; apex obtuse to subacute,
sometimes subcapitate; very numerous. Cheilocystidia 30–45
× 9–15 µm, apex often subacute, scattered. Paracystidia 7–37
× 4–20 µm, apex sometimes obscurely mucronate to obscurely
acutely papillate, hyaline, numerous, dominating.
Notes: Smith (1972) described the spores of P. septentrionalis
var. vesiculosa as ‘in KOH chocolate color, gradually fading
and with an ochraceous tinge (in about three hours)’ and
with a distinct germ pore, but we found them distinctly pale
immediately and the germ pore indistinct; he also detected
longer pleurocystidia, up to 60 µm. Psathyrella septentrionalis
and P. vesiculosa show a molecular proximity (DNA nucleotide
homology: ITS = 99 % with five mismatches) but there are a
number of micro morphological elements that distinguish
P. vesiculosa, such as the spores (distinctly paler, somewhat
slenderer, often attenuate at base, scarcely subphaseoliform),
the gill edge dominated by the paracystidia, and partly
the pleurocystidia which are somewhat slenderer and less
rounded at apex. In macro morphology, we also notice that
the pileus of P. septentrionalis is described as obtuse to
convex, expanding from broadly convex to plane (Smith’s
plates 43b, 46a and 46b clearly confirm this), while the pileus
of P. vesiculosa is described as obtusely conical becoming
campanulate. Predominance of paracystidia and, partly, shape
of pleurocystidia may induce to consider P. vesiculosa a possible
member of subsection Spadiceogriseae, but it has a colored
veil and, as we anticipated in the introduction, this parameter
has to be given a relevant discriminating importance. This is
one of those species evidencing that dominating paracystidia
alone are not exclusive of subsection Spadiceogriseae.
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28. Psathyrella velibrunnescens A.H. Sm., Mem. N. Y. Bot. Gard.
24: 128. 1972. Fig. 59.
Typus: Psathyrella velibrunnescens A.H. Sm. USA, Washington,
Jefferson Co., Bottom, Clearwater River, 9 May 1939, A.H. Smith
13240 (MICH12107), on Alnus debris.
Loan of a slice of a pileus for micro-morphology investigation.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Spores 7.7–9.5(–10) × 4.5–5.1(–5.4) µm, Q 1.6–2.1(–2.2), avQ
1.8; in front view oblong to elliptic, sometimes oval, base mainly
obtuse, in profile adaxially flattened to subphaseoliform; dull
orangish or paler; germ pore indistinct (callus). Pleurocystidia
40–79 × 9–17 µm, utriform to subcylindrical, often elongatecylindrical in upper part; apex mainly rounded, sometimes or
often (irregularly) subcapitate; very numerous. Cheilocystidia
27–51 × 9–12(–19) µm, scattered. Paracystidia 9–41×5–23 µm,
numerous, dominating. No pigment or incrustation seen on any
type of cystidia.
Notes: Unfortunately DNA extraction failed on this material,
but, based on morphological evidence, we consider P.
velibrunnescens to pertain to the P. septentrionalis complex. We
note that Kits van Waveren (1985), on the contrary, regards it as
an American subspecies of P. chondroderma (= P. pertinax). For
further comment see notes to the following species.
29. Psathyrella aff. kauffmanii, Czech Republic, Bohemia, Volary,
Černý Kříž, Mount Šumava, Jelení vrch area, 870 m. a.s.l., 13 Jul.
2000, J. Holec 61/2000 (PRM897750, erroneously registered in
GenBank as BRNM:705626), gregarious on very rotten trunk of
Fagus sylvatica in a mixed forest with also Picea abies and Abies
alba. ITS AM712260.
?= Psathyrella velibrunnescens A.H. Sm.
(Description produced from data and images received from
Vašutová).
Pileus 15–40 mm, campanulate, red-brown, hygrophanous,
distinctly radially rugulose. Veil yellowish whitish at margin
of pileus. Lamellae pale violet-brown. Stipe 40–70 × 7–9 mm,
cylindrical, whitish, brownish towards the base, innately white
marbled. Smell rubber-like. Spores 7.5–9 × 4–5 µm, Q±1.7–2.1,
avQ±1.8, in front view elliptic, in profile subphaseoliform,
rarely subamygdaliform to adaxially flattened, light to medium
brown, germ pore little distinct (callus). Pleurocystidia (40–)
46–60(–72) × 12–14 µm, utriform to sublageniform, with apex
obtuse to rounded, rarely subacute. Cheilocystidia 30–40 × 11–
15 µm, scattered. Paracystidia 20–34 × 9–13 µm, numerous,
dominating.
Notes: When blasting the ITS sequences of P. septentrionalis
and P. vesiculosa in GenBank we have discovered that they
allied with P. aff. kauffmanii from Europe synthetically
Fig. 59. Psathyrella velibrunnescens. A.H. Smith 13240, MICH12107; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
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A revision of Psathyrella, subsection Spadiceogriseae
described in Vašutová et al. (2008). Upon our request,
Vašutová has provided us with a more detailed description, has
shown us some images of microscopical characters, and has
discussed with us the molecular outcome. Various characters
link P. aff. kauffmanii to P. vesiculosa: the gill edge dominated
by paracystidia, the pale spores, the little distinct germ pore,
the slender pleurocystidia, the yellowish veil at the margin of
a campanulate pileus; on the contrary the often phaseoliform
spores with rounded, not attenuate base seem to differentiate
them; moreover P. aff. kauffmanii has very thickset, 7–9 mm
broad, stipes, while P. vesiculosa has slender, 1.5–5 mm broad
stipes. A more interesting comparison, though not based on
phylogenetic data, can be made with the unsequenced P.
velibrunnescens. A very strong correspondence can be noticed
between the two, both in gross and micro characters as well
as in the lignicolous habitat and, but for the lack of molecular
support, we would suggest their conspecificity. However, while
we wish Vašutová to find and describe it again in more detail,
surely P. aff. kauffmanii has been shown to be a European
species on its own right. It only remains to understand
whether it may be autonomous from or conspecific with any
of the American species of the phylogenetic cluster to which
it belongs.
Psathyrella conica, recently described from China, shares
many morphological characters with the last three species here
described, namely P. vesiculosa, P. velibrunnescens and P. aff.
kauffmanii. Like them, it has paracystidia dominating, utriform
pleurocystidia, pale spores with an indistinct or absent germ
pore, campanulate to conical pilei and a lignicolous habitat.
Moreover, like them it is molecularly outside of spadiceogrisea
clade and has a veil which its authors have noticed not to be
merely white (‘white to dirty white’). We strongly suspect that,
just as P. aff. kauffmanii is a European representative of P.
vesiculosa group, P. conica could be instead an Asian member
of the same group.
30. Psathyrella obtusata (Fr.) A.H. Sm., Contr. Univ. Mich. Herb.
5: 55. 1941, sensu Smith, Mem. N. Y. Bot. Gard. 24: 385. 1972.
We have attempted to understand Smith’s (1972) concept of P.
obtusata by examining three Smith vouchers, Ammirati 2811,
Harrison 8023 and Smith 73998, selected for this purpose by
MICH staff. Our revision of these materials has revealed that
they represent three different species: Harrison 8023 has the gill
edge dominated by the cheilocystidia, Ammirati 2811 and Smith
73998 have the gill edge lined with numerous paracystidia often
mucronate to attenuate at the apex.
30a. J.F. Ammirati 2811, MICH49136.
= Psathyrella obtusata (Fr.) A.H. Sm.
Material examined: USA, Michigan, Baraga Co., Huron River, Skanee
Road, 25 Jun. 1969, on hardwood log.
Loan of two sporocarps in good condition for micro-morphology
investigation.
The original gross description by the collector reports pilei color
ranging from watery sayal brown to dark brown, fading to clay
color at centre and tan at margin; stipe more or less up to 6
mm broad at apex, white fibrillose, sordid brownish in age; odor
slight, taste slightly raphanoid. Spores (5.5–)6.7–8.3(–9.1) × 4.3–
5.6(–6) µm, Q 1.4–1.7; in front view elliptic to oval, sometimes
somewhat irregular, base generally rounded, in profile adaxially
flattened to subphaseoliform; dull orangish to reddish orangish;
germ pore little distinct, small. Pleurocystidia (35–)40–63 ×
(7.5–)8–16(–17.5) µm; narrowly utriform, narrowly fusiform,
(flexuously) conical, sometimes sublageniform; apex obtuse,
sometimes subacute, rarely subcapitate to submucronate, often
incrusted; walls sometimes slightly thickened and pale yellowish
below apex; numerous. Cheilocystidia 28–43 × 8.5–15.5 µm,
locally scattered to numerous, locally dominating. Paracystidia
13–40 × 9.5–20(–27) µm, often mucronate to attenuate,
yellowish, often incrusted towards base, numerous, locally
dominating. Cells below marginal cystidia yellowish pigmented
and incrusted.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Note: We failed to sequence this material but it has the typical
characters of P. obtusata.
30b. K.A. Harrison 8023, MICH65721. Fig. 60.
= Psathyrella atomatoides (Peck) A.H. Sm.
Material examined: Canada, Nova Scotia, Victoria Co., Forks Baddeck,
22 Sep. 1968, densely cespitose on sawdust pile. K.A. Harrison 8023,
MICH65721. ITS MF326013.
Loan of numerous sporocarps in good condition for micromorphology investigation.
The original gross description by the collector reports pilei up
to 3.6 cm broad, slightly conic to subumbonate, sometimes
expanding to nearly convex, hygrophanous, when young dark
brown, when dried tan, radially striate, veil connecting pileus
and stipe when young then fibrillose on margin; gills up to 2
mm broad, close, dark smoky brown; stipe 30–35 × 3–4 mm,
covered by fibrils. Spores 6.8–8.5(–9.1) × 3.8–4.8 µm, Q 1.5–
1.9(–2.1); in front view oblong, elliptic to oval, base generally
obtuse, in profile amygdaliform, subamygdaliform to adaxially
flattened, sometimes subphaseoliform to narrowly elliptic or
subfusiform; strong orange-brown to red-orange; germ pore
indistinct to little distinct, small. Pleurocystidia 45–65 × 10–15(–
17) µm; lageniform to conical, sometimes subutriform, fusiform,
irregularly cylindrical-mucronate; apex subobtuse to subacute,
rarely subcapitate, often incrusted; walls often thickened and
yellowish to yellowish brownish below apex; numerous to
moderately numerous. Cheilocystidia 32–60 × 10–12 µm, more
often fusiform-conical, scattered near pileus margin, numerous
elsewhere, dominating. Paracystidia 22–28 × 9–10 µm, often
incrusted, more or less scattered, numerous near pileus margin.
Cells below marginal cystidia pigmented near pileus margin,
hyaline elsewhere.
Notes: This material is in good molecular concordance, though
with evidence provided by only one locus (DNA nucleotide
homology: ITS= 99 % with 1 mismatch), with P. atomatoides;
morphology too is very similar except for the distinct reddish
pigmentation of the gill edge in a short, 3 mm long, segment
near the pileus margin, in correspondence with the dominance
zone of the paracystidia.
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Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 60. Psathyrella obtusata. K. A. Harrison 8023, MICH65721; GE Gill edge at pileus margin; PC Pleurocystidia in NH4OH; S1 Spores in water; S2
Spores in KOH.
30c. A.H. Smith 73998, MICH33261. Fig. 61.
=?
Material examined: USA, Idaho, Bonner co., Priest River, Priest River
Experimental Forest, 6 Oct. 1966, on debris of cedar slash, A.H. Smith
73998, MICH33261. ITS MF326012.
Loan of five, entire or broken, sporocarps in good condition for
micro-morphology investigation.
The original gross description by the collector reports pilei watery
tawny when moist, striate half way; veil of white particles at
margin; gills broad, adnate, when young brownish, when mature
near hair brown; stipe pallid, soon honey below, thinly fibrillose.
Spores 7–8.5(–9.2) × 4.2–5 µm, on average 7.9 × 4.55 µm, Q 1.5–
2, avQ 1.7; in front view elliptic to oval, base rounded to obtuse,
in profile amygdaliform, subamygdaliform to somewhat adaxially
flattened, sometimes subphaseoliform; reddish brown; germ pore
distinct, rounded to truncate, 1.4–1.7 µm broad. Pleurocystidia
29–45(–52) × 10–16 µm; variable, ellipsoid, fusiform, conical,
lageniform, sometimes narrowly utriform, sublageniform,
flexuous; apex subacute to obtuse, sometimes acute, rarely
minutely rounded subcapitate, sometimes incrusted; numerous
152
to moderately numerous. Cheilocystidia 21–33 × 8.5–12(–14) µm,
moderately numerous to locally rare. Paracystidia 14–23(–26) ×
8–12.5 µm, often mucronate to subpapillate, hyaline, numerous.
Cells below marginal cystidia hyaline.
Notes: This voucher corresponds neither to any European
species, nor to any American material sequenced in this work;
however phylogenetic analysis, though based on one locus
only, clearly places it outside subsection Spadiceogriseae.
Smith (1972) has described some species with Pennatae-type
pleurocystidia and a conspicuous presence of paracystidia on
the gill edge so it should be placed among, and compared with,
them, which is outside this paper’s purpose.
31. Psathyrella limicola var. subpectinata A.H. Sm., Mem. N. Y.
Bot. Gard. 24: 416. 1972.
Typus: Psathyrella limicola var. subpectinata A.H. Sm. USA,
Wyoming, Carbon Co., Lower French Creek, Medicine Bow
Mountains, 30 Jun. 1950, A.H. Smith 34407 (MICH11965),
gregarious on wet soil along stream. ITS MF325980.
Loan for gross morphology investigation of three sporocarps in
good condition.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 61. Psathyrella obtusata. A.H. Smith 73998, MICH33261; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
Spores 6.8–9.3 × 3.8–5 µm, Q (1.6–)1.8–2.1(–2.3), avQ 1.9; in
front view (narrowly) oval to elliptic, base rounded, in profile
subamygdaliform to subphaseoliform, sometimes adaxially
flattened; reddish brown; germ pore indistinct to moderately
distinct, sometimes distinct, rounded to truncate, 1.2–1.6
µm broad. Pleurocystidia 29–41 × 10–13 µm; utriform to
fusiform-utriform; apex subrounded, sometimes subcapitate,
sometimes weakly incrusted, walls slightly pigmented;
moderately numerous. Cheilocystidia 20–30(–37) × 8–10 µm,
walls sometimes thickened and weakly pigmented in lower
half, generally numerous, generally dominating. Paracystidia
8–20(–28) × 6–13 µm, sometimes irregular, walls in larger ones
sometimes thickened and weakly pigmented, numerous, locally
dominating where larger ones are crowded. Cells below marginal
cystidia sometimes with thickened and weakly pigmented walls.
Notes: This material has a 99 % ITS nucleotide homology with
a number of species outside of subsection Spadiceogriseae: P.
warrenensis, P. seminuda, P. fagetophila, and Smith 28713, one
of the Smith vouchers under the name of P. agrariella.
32. Psathyrella affinis A.H. Sm., Mem. N. Y. Bot. Gard. 24: 232.
1972. Fig. 62.
Paratype: Psathyrella affinis A.H. Sm. USA, Michigan, Oakland
Co., Highland Lake, 14 Jun. 1945, A.H. Smith 20414 (MICH11857),
gregarious to scattered on wet leaves in swamp.
Loan for gross morphology investigation of two pilei.
Spores (6.8–)8–10.4(–12.5) × (4.6–)5.4–6.2(–6.8) µm, Q (1.4–)1.5–
1.6(–2.1); in front view (broadly) oval, sometimes subtriangular
to elliptic, base often subtruncate, in profile adaxially flattened,
elliptic, subphaseoliform; orange-brown; germ pore distinct to
indistinct, sometimes eccentric in profile, rounded, 1.3–1.6 µm
broad. Basidia 8–13 µm broad, some 2-spored. Pleurocystidia 25–
40 × 10–17(–20) µm; ventricose-lageniform, sometimes utriform
to lageniform; apex 4.5–6(–7.5) µm broad, obtuse, sometimes
subobtuse to subrounded, sometimes subforked; more or less
moderately numerous. Cheilocystidia 27–38 × 11–15 µm, more or
less scattered to locally absent. Paracystidia 7.5–31(–36) × 7–17(–
22.5) µm, often incrusted towards base, walls sometimes faintly
pigmented, numerous, dominating.
Notes: Unfortunately we failed to sequence this material, so we
have no molecular information about its phylogenetic position.
The pleurocystidial shape is particular: the obtuse, never
rounded nor acute to subacute, apex is somewhat ambiguous
but the neck is much slenderer than the ventricose portion,
looking on the whole unfit for section Spadiceogriseae.
33. Psathyrella nitens A.H. Sm.
Because of scarcity of material, the holotype could be obtained
for DNA sequence only. The MICH staff has selected four other
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Voto et al.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Fig. 62. Psathyrella affinis. A.H. Smith 20414, MICH11857; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
collections from the same site by Smith and deposited under
the same name for micro-morphology investigation. Two of
them, Smith 30241 and Smith 30239, correspond molecularly
to the holotype and morphologically to the description given
by Smith (1972) therefore they are confidently assigned to the
typical P. nitens. Smith 29642 deviates significantly from the
holotype (DNA nucleotide homology: ITS= 99 %, Tef-1α = 94 %)
and we have found that the spores are appreciably smaller,
the pleurocystidia and paracystidia are somewhat smaller and
the apex of the pleurocystidia are more obtuse. Smith 30175
is distinctly closer to Smith 29642 (DNA nucleotide homology:
ITS= 99 % with 1 mismatch) than to the other three vouchers
(DNA nucleotide homology: ITS= 99 % with 8 to 9 mismatches)
and it too displays the same differences exactly as found in
Smith 29642 therefore they are conspecific; we consider them
a separate taxon from P. nitens and report their morphological
revision in a unified description. All these vouchers fall outside
of subsection Spadiceogriseae and nest beside P. obtusata;
their pleurocystidia are never thickly utriform and the apex is
sometimes more or less acute but never rounded, the gill edge
has often intermediate, mucronate to rhomboid, cells.
33a. Psathyrella nitens A.H. Sm., Mem. N. Y. Bot. Gard. 24: 320.
1972.
Typus: Psathyrella nitens A.H. Sm. USA, Washington, Pierce
Co., Mt. Rainier National Park, Lower Tahoma Creek, 21 Aug.
154
1948, A.H. Smith, 30388 (coll. D.E. Stuntz) (MICH11983), on
debris; only for molecular sequence. Tef-1α MF521789, ITS
MF325982.
Loans for micro-morphology investigation: USA, Washington,
Pierce Co., Mt. Rainier National Park, Lower Nisqually, 18
Aug. 1948, A.H. Smith 30239 (MICH33251), on debris, Tef-1α
MF521787, ITS MF325985; USA, Washington, Pierce Co., Mt.
Rainier National Park, Lower Tahoma, 18 Aug. 1948, A.H. Smith
30241 (MICH33252), on alder debris, Tef-1α MF521786, ITS
MF325986.
Spores 8.3–11.2(–12) × 5–6.5(–6.8) µm, Q 1.4–1.7(–1.9); in
front view oval to elliptic, base obtuse to broadly rounded, in
profile suboval, amygdaliform to subamygdaliform, sometimes
subphaseoliform; reddish brownish; germ pore distinct, mainly
truncate to subtruncate, 1–1.7 µm broad. Pleurocystidia 42–
75(–87) × 11–17.5 µm; lageniform, conical, subutriform, usually
with elongate neck; apex acute to subobtuse, sometimes
subcapitate or forked, incrusted or not; moderately numerous.
Cheilocystidia 25–45(–49) × 11–12.5(–16) µm, mainly with
neck very short, looking mucronate, to medium long (2–10
µm), locally numerous and in tufts. Paracystidia 16–36 × 8–14
µm, walls sometimes or often thickened, yellowish pigmented,
numerous, dominating. Cells below marginal cystidia with
yellowish pigment.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
33b. A.H. Smith 29642, MICH33249; A.H. Smith 30175;
MICH33250. Fig. 63.
basidia and basidiola. Cells below marginal cystidia mostly or only
locally with yellowish pigment, sometimes incrusted.
=?
34. Psathyrella psammophila A.H. Sm., Mem. N. Y. Bot. Gard.
24: 357. 1972.
Loans for micro-morphology investigation: USA, Washington,
Pierce Co., Mt. Rainier National Park, Lower Tahoma, 30 Jul.
1948 (Smith 29642), Tef-1α MF521788, ITS MF325983; at same
location, 16 Aug. 1948, coll. D.E. Stuntz & B. Knowles, on humus
(Smith 30175), ITS MF325984.
No description was attached to either voucher by the collectors.
Spores (7.5–)8.1–9.6(–10.5) × (4.5–)5–5.6(–5.8) µm, on average
8.9 × 5.2 µm, Q (1.44) 1.52–1.88(–2), avQ 1.70; in front view
elliptic, sometimes suboval, base rounded, in profile adaxially
flattened, sometimes amygdaliform to subamygdaliform; reddish
brownish; germ pore distinct, mainly truncate, 1.1–1.5 µm broad.
Pleurocystidia 40–60 × 8.8–16 µm; narrowly utriform to lageniform,
sometimes fusiform to conical, sometimes flexuous, neck variably
long; apex obtuse, rarely subacute, sometimes incrusted;
moderately numerous to numerous. Cheilocystidia (18–)31–45
× 7–15 µm, apex sometimes incrusted, scattered to numerous.
Paracystidia 11–24(–38) × 6–12(–17) µm, walls sometimes or
often thickened and slightly yellowish pigmented towards base,
moderately numerous to numerous and intermixed with many
Molecular heterotypical synonyms proposed here: Psathyrella
angusticystis A.H. Sm., Mem. N. Y. Bot. Gard. 24: 383. 1972;
Psathyrella subargillacea A.H. Sm., Mem. N. Y. Bot. Gard. 24:
386. 1972.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Typus: Psathyrella psammophila A.H. Sm. USA, Michigan,
Washtenaw Co., Pinckney Recreation Area, 7 May 1964, A.H.
Smith, 67836 (MICH12012), on sandy soil. ITS KC992856.
Loan for micro-morphology investigation of a slice of a pileus in
good condition.
Spores 7.5–9.3(–11) × (4.4–)4.7–5.5(–5.7) µm, on average 8.2
× 5 µm, Q 1.5–1.7(–2), avQ 1.6; in front view broadly oval to
elliptic, base rounded, in profile subphaseoliform to adaxially
flattened, sometimes distinctly phaseoliform; brownish orange,
sometimes with a vague reddish shade; germ pore mainly
indistinct and small. Pleurocystidia 40–60 × 10–16(–18) µm;
utriform, rarely cylindrical-ellipsoid to clavate; apex obtuse,
sometimes rounded, sometimes mucronate to subcapitate,
Fig. 63. Psathyrella nitens. A.H. Smith 29642, MICH33249; PC1 Pleurocystidium in NH4OH; PC2 Pleurocystidia in Congo red; MC Marginal Cystidia; S1
Spores in water; S2 Spores in KOH.
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rarely forked, often incrusted; walls often weakly thickened
and yellowish pigmented; numerous. Cheilocystidia 25–40(–50)
× 8–15 µm, sometimes subacute, moderately numerous to
scattered. Paracystidia –20 × –15 µm, sometimes mucronate,
often incrusted, numerous.
Notes: On accurate observation of the holotype material it can
be noticed that some debris of little twigs or grass stems are
attached at the stipe base. We have found the ITS sequence in
GenBank, unfortunately we have failed to add the Tef-1α region
sequence.
Psathyrella psammophila is very close to the two following
species, P. subargillacea and P. angusticystis (DNA nucleotide
homology: ITS = 99 % with 4 and 1 mismatches respectively).
A number of characters join these three taxa together such as
small sporocarps, rudimentary veil, fragile context, pale colored
and campanulate to broadly conical, never expanded, mature
pileus, moderately crowded to distant (not distinctly crowded)
gills, spore parameters in general including the little distinct germ
pore, cheilocystidia generally sparse, paracystidia numerous but
generally small, general feature of pleurocystidia. In all three
cases we lack a second locus evidence; however, as we have
found no relevant morphological difference, we consider both
taxa conspecific with P. psammophila, the slightly paler spores
of P. angusticystis being the only different element we could
possibly point out in objection.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
34a. Psathyrella subargillacea A.H. Sm., Mem. N. Y. Bot. Gard.
24: 386. 1972.
Typus: Psathyrella subargillacea A.H. Sm. USA, Michigan, Luce
Co., Tahquamenon Falls State Park, near Paradise, 23 Jun. 1963,
A.H. Smith, 66717 (coll. A.H. Smith & S. Brough) (MICH12058),
on rotten wood. ITS MF325999.
Loan for micro-morphology investigation of one sporocarp and a
half cut lengthwise, both in good condition.
Spores (7.3–)7.7–9.3(–10) × 4.4–5.5(–6.2) µm, Q (1.45–)1.6–1.9(–
2.05), avQ 1,74; in front view oval to elliptic, sometimes irregular,
base rounded, in profile subphaseoliform to subamygdaliform;
pale dull orangish; germ pore indistinct (callus). Pleurocystidia
(30–)38–63 × 7–17 µm; narrowly utriform, fusiform, conical,
sometimes cylindrical to ellipsoid, rarely thickly utriform; apex
obtuse to subacute, rounded, rarely acute, often subcapitate,
sometimes forked, rarely rostrate; often incrusted; walls
sometimes weakly pigmented; numerous. Cheilocystidia 25–55
× 8–17 µm, often with subacute apex, incrustations not seen,
scattered, towards margin of pileus numerous. Paracystidia 10–
28 × 6–20 µm, incrusted, numerous, often dominating.
35. Psathyrella incondita A.H. Sm., Mem. N. Y. Bot. Gard. 24:
405. 1972.
Since the holotype could not be obtained because of its
scarcity, the MICH staff has selected two other collections Smith
determined and deposited under this name. Our revision has
evidenced they are unequivocally two different taxa: Harrison
10471 comes molecularly near P. flexuosipes and could
represent the true P. incondita, Hoseney 1728 is distant from it.
Both materials fall outside of section Spadiceogriseae.
35a. K.A. Harrison 10471, MICH36450.
?= Psathyrella incondita A.H. Sm.
Material examined: USA, Michigan, Marquette Co., Ives Lake, Loop Rd.,
21 Jul. 1971, det. Smith, gregarious on elm wood, K.A. Harrison 10471,
MICH36450. ITS MF325975.
Loan for micro-morphology investigation of one almost entire
pileus in good condition.
Loan for micro-morphology investigation of two sporocarps and
some fragments of pileus.
Spores (7.2–)7.5–9.5(–11.2) × 4.3–5.3(–6.2) µm, on average
8.5 × 4.9 µm, Q (1.55–)1.65–1.95(–2.15), avQ 1.75; in front
view oval to elliptic, base subtruncate, rounded to obtuse
(tapered in Smith’s sketches), in profile adaxially flattened to
subphaseoliform, sometimes subamygdaliform to phaseoliform;
brownish to reddish orange; germ pore small and indistinct
(callus). Pleurocystidia (30–)46–68 × (8–)10–17 µm; conical,
subcylindrical, fusiform, sometimes ellipsoid, ellipsoidmucronate, narrowly utriform, lageniform; apex 4–8 µm broad,
obtuse, sometimes rounded, rarely subacute, sometimes
subcapitate, rarely forked, sometimes weakly incrusted;
walls sometimes weakly thickened below apex; moderately
numerous. Cheilocystidia 25–48 × 7.5–17 µm, scattered to
moderately numerous. Paracystidia –23 × –20 µm, sometimes
rhomboid to mucronate, walls often pigmented and thickened,
numerous. Cells below marginal cystidia weakly pigmented.
Spores (6.2–)7–8.7 × 3.3–4.4 µm, Q 1.8–2.45(–2.6), avQ 2–2.1;
in front view cylindrical to oblong, sometimes irregular, base
obtuse, in profile mainly subamygdaliform to subphaseoliform;
pale reddish brown to brownish orangish; germ pore distinct,
mainly truncate, ±1.5 µm broad. Pleurocystidia 33–64 × 10–
16.5 µm; lageniform, ventricose-fusiform, conical, utriform,
ellipsoid; apex 3.5–6.5 µm broad, subacute to obtuse,
sometimes irregular to bent, often incrusted; numerous.
Cheilocystidia (25–)35–50(–52) × 9–15 µm, often incrusted,
moderately numerous, dominating. Paracystidia –25(–31) ×
–11 µm, more or less numerous. Cells below marginal cystidia
hyaline.
34b. Psathyrella angusticystis A.H. Sm., Mem. N. Y. Bot. Gard.
24: 383. 1972.
Material examined: USA, Michigan, Livingston Co., Hell, Fralick’s
Tree Farm, 28 Oct. 1970, det. Smith, under pines, F.V. Hoseney 1728,
MICH36451. Tef-1α MF521795, ITS MF325976.
Typus: Psathyrella angusticystis A.H. Sm. USA, Michigan, Emmet
Co., Wilderness Point, Wilderness State Park, 24 Sep. 1953, A.H.
Smith, 43146 (MICH11873), gregarious on debris in a clearing.
ITS MF325959.
156
35b. F.V. Hoseney 1728, MICH36451. Fig. 64.
=?
Loan of various sporocarps entire and in fragments for micromorphology investigation.
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 64. Psathyrella incondita. F. V. Hoseney 1728, MICH36451; PC Pleurocystidia; MC Marginal Cystidia at centre of gill; S1 Spores in water; S2 Spores
in KOH.
The notes of gross morphology by the collector report pileus 4.7
cm broad and stipe up to 5 mm thick at apex.
However micro morphological characters do not fully correspond
to those of P. squamosa.
Spores (6.1–)6.8–8.1(–9.5) × (3.7–)4–4.8(–5.7) µm, on average
7.7 × 4.4 µm, Q 1.5–1.9(–2.05), avQ 1.7; in front view oval,
sometimes oval-sublimoniform to triangular, base rounded,
in profile subelliptic to adaxially flattened, amygdaliform to
sublimoniform, sometimes oval; orange-brown; germ pore
moderately distinct to indistinct, 1–1.2 µm broad. Pleurocystidia
(23–)25–40 × (8–)10–15 µm; lageniform to broadly ventricoselageniform, sometimes subutriform, sometimes flexuous
towards apex; apex obtuse, sometimes subacute, submucronate,
subcapitate, often incrusted; walls sometimes sub hyaline;
scattered. Cheilocystidia 22–39 × (7.5–)9–15 µm, moderately
numerous at centre of gill edge, more scattered elsewhere.
Paracystidia –27 × –16 µm, sometimes rhomboid, hyaline,
moderately numerous at centre of gill edge, inconspicuous
elsewhere. Gill edge at centre with equal dominance of both
types of cells or with locally alternate predominance. Cells
below marginal cystidia hyaline.
36. Psathyrella flexuosipes A.H. Sm., Mem. N. Y. Bot. Gard. 24:
145. 1972.
Notes: The best reliable match when blasting the ITS region
is with LO194-96 and LO164-96, two vouchers representing P.
squamosa, with 99 % identity; when blasting the Tef-1α region
none is above 90 % identity but it must be noticed that Tef1α regions of the two aforesaid vouchers are not sequenced.
Paratype: Psathyrella flexuosipes A.H. Sm. USA, Oregon,
Douglas Co., Lake Tahkenitch, 21 Nov. 1935, A.H. Smith 3578
(MICH32961), on debris. ITS MF325969.
Loan for gross morphology investigation of one sporocarp with
a broken pileus.
Spores 7–9(–10) × 4.1–5.3 µm, Q (1.3–)1.5–2.1, avQ 1.7; in
front view oval to elliptic, base rounded, in profile adaxially
flattened, subamygdaliform, subphaseoliform; bright reddish;
germ pore distinct, truncate to subrounded, 1.5–1.8 µm broad.
Pleurocystidia 38–60 × 9–17 µm; fusiform-utriform, fusiform,
ventricose-fusiform to ventricose-conical, sometimes (slenderly)
utriform; apex obtuse, never acute or broadly rounded, often
(grossly) incrusted; numerous to moderately numerous.
Cheilocystidia 30–60 × 7.5–16 µm, sometimes minutely
incrusted, numerous and generally dominating. Paracystidia
9–35 × 7–18 µm, sometimes submucronate, sometimes
incrusted, walls sometimes or often thickened and yellowish
pigmented, numerous and locally dominating.
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Notes: We wanted to study this species because Smith (1972)
did not specify the ratio between cheilocystidia and paracystidia
but, unfortunately, MICH could not loan the holotype for
scarcity of material. However, this paratype does not belong
to subsection Spadiceogriseae and has no closely resembling
counterpart in Europe.
37. Psathyrella abieticola A.H. Sm., Mem. N. Y. Bot. Gard. 24:
348. 1972. Figs 65–66A (as P. subnuda var. velosa), 66B (as P.
vesiculocystis).
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Molecular heterotypical synonyms and misapplied names
proposed here: Psathyrella subnuda sensu A.H. Smith, Mem.
N. Y. Bot. Gard. 24: 138. 1972, p.p.; Psathyrella subnuda var.
velosa A.H. Sm., Mem. N. Y. Bot. Gard. 24: 139. 1972; Psathyrella
vesiculocystis A.H. Sm., Mem. N. Y. Bot. Gard. 24: 349. 1972.
Pileus 15–40 mm broad, in early stages obtusely conical to
convex with straight margin, expanding broadly conic to broadly
convex or sub plane, moist or not; dark cinnamon-brown, rusty
cinnamon, Mars brown, buckthorn brown, hygrophanous,
fading to dull clay color, dingy tan, cinnamon buff, somewhat
pale cinnamon-drab, brownish grey; translucently striate when
moist. Veil white, quickly evanescent, when young with few thin
patches of fibrils along the margin appendiculate fragments, thin
and fibrillose on stipe. Lamellae adnate to ascending-adnate;
narrow to moderately broad; moderately crowded; when young
brownish to dingy cocoa-color, finally violaceous fuscous to dark
vinaceous brown; edge white. Stipe 30–110 × 1–6 mm, equal or
nearly so; white, watery white to pallid, not discoloring; apex
pruinose. Context (very) fragile, thin in pileus, concolorous to
surface. Odor and taste indistinctive. Spore deposit fuscous
black. Spores (7–)7.7–10(–11.2) × (5–)5.5–7.2 × (4–)4.4–5.5
µm, fQ 1.3–1.6, pQ 1.5–2; in front view broadly oval, often
or sometimes (irregularly) angular-oval, sometimes broadly
elliptic, rarely subglobose, triangular, oboval or mitriform,
base obtuse to broadly rounded, sometimes truncate, apex
often attenuate to sometimes snout-like projected, in profile
subelliptic to adaxially flattened, rarely subamygdaliform, oval
or subphaseoliform; dusky brown with a reddish to orangered shade; germ pore distinct, mainly truncate, sometimes
somewhat oblique in profile, 1.6–2(–2.5) µm broad. Basidia
4-spored, 17–25 × 6–11(–11.8) µm. Pleurocystidia 33–60(–65) ×
9–20 µm, broadly utriform to subcylindrical, often or sometimes
ellipsoid-saccate to subclavate, sometimes sublageniform; apex
broadly rounded to obtuse, rarely subcapitate, rarely incrusted;
walls hyaline, sometimes thickened below apex; moderately
numerous, rarely numerous. Gill edge cystidia 8–33(–48) ×
4.5–19(–25) µm, a mixture of three types of cystidia: distinctly
utriform to sometimes sublageniform, 25–33 × 12–13 µm,
usually scattered; ellipsoid to rhomboid, 21–29 × 10–17 µm,
numerous; saccate to clavate, 9.5–30 × 6–17 µm, numerous.
Caulocystidia up to 140 × 20 µm, utriform to clavate. Clamps
present.
Fig. 65. Psathyrella subnuda var. velosa. F. P. Sipe, 1100, MICH33441; PC Pleurocystidia; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
158
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 66. A. Psathyrella vesiculocystis. A.H. Smith & H. D. Thiers 70089, MICH5380; Marginal Cystidia. B. Psathyrella subnuda. A.H. Smith 56936,
MICH33421; Marginal Cystidia.
Habitat: Gregarious or scattered; on humus of cottonwood or
conifer, once on burned soil.
Loan for micro-morphology investigation of very tiny fragments,
the largest measuring 2 mm.
Collections examined from MICH: Psathyrella subnuda sensu A.H. Smith,
Smith 56936; Psathyrella subnuda var. velosa A.H. Sm., holotype, Smith
73947; Psathyrella vesiculocystis A.H. Sm., Smith & Thiers 70089.
The notes from the collector report a pileus 3–4 cm broad,
conical, smooth, light cinnamon drab, expanded when older,
without surface structures; stipe white, fragile gills drab with
white edge; spores up to 10 × 7 µm, reddish, dark with violet
tints in deposit. Spores 8.2–10(–10.7) × 6–7.1 × 4.8–5.5 µm,
fQ 1.3–1.5, pQ 1.6–2; in front view oval, triangular, angularoval to mitriform, base truncate to broadly rounded, apex
often attenuate to sometimes snout-like projected, in profile
subelliptic to adaxially flattened, dusky brown with an orangered shade; germ pore distinct, mostly truncate, sometimes
somewhat oblique in profile, 1.6–2 µm broad. Pleurocystidia
33–55 × 12–20 µm, broadly to cylindrical utriform or ellipsoidsaccate, apex broadly rounded, sometimes obtuse; walls hyaline,
sometimes thickened; moderately numerous. Gill edge cystidia
10–33 × 7–16(–19) µm, a mixture of scattered distinctly utriform
cells, scattered obtusely fusoid cells and numerous, small to
large, saccate to clavate cells.
Notes: The full description above comprises our revision and
Smith’s (1972) description of all taxa we acknowledge as
conspecific, with exclusion of Smith’s (1972) description of P.
subnuda. The presence of many pleurocystidia not distinctly
utriform but subcylindrical, ellipsoid-saccate to subclavate
makes the gill edge cystidial pattern highly composite and a
neat distinction between cheilocystidia and paracystidia hardly
applicable. Sometimes the intermediate marginal cells are very
small and can be ambiguously confused with the surrounding
paracystidia, making these latter seem dominating.
We have not examined material of P. abieticola, but we have
found strong molecular correspondence between P. vesiculocystis,
P. subnuda var. velosa, and the holotype of P. abieticola present in
GenBank (accession nr. KC992891). According to Smith’s (1972)
description, we see a good overlap of the morphological data,
including the tridimensional, strongly oval to angular-oval spores
with truncate base and attenuate to snout-like projected apex.
For comments on P. subnuda sensu Smith see below.
Psathyrella fusca is a common, relatively similar European
species but it is clearly not conspecific based on strong molecular
evidence and very different spores.
37a. Psathyrella subnuda var. velosa A.H. Sm., Mem. N. Y. Bot.
Gard. 24: 139. 1972. Fig. 65.
Typus: Psathyrella subnuda var. velosa A.H. Sm. USA, Idaho,
Bonner Co, Tule Bay, Priest Lake, 5 May 1966, A.H. Smith 73947
(MICH5376), on debris in cottonwood flat; only for molecular
sequence. Tef-1α MF521773, ITS MF326005.
Paratype studied: Psathyrella subnuda var. velosa A.H. Sm. USA,
Oregon, Pruetts place, 20 Nov. 1949, F.P. Sipe, 1100 (det. Smith)
(MICH33441), habitat not reported. ITS MF326006.
Notes: We could only observe scattered little traits of gill edge
and we believe that most utriform and, overall, fusoid marginal
cells must have collapsed, at least in their apical part, given the
badly fragmented state of the material received.
37b. Psathyrella vesiculocystis A.H. Sm., Mem. N. Y. Bot. Gard.
24: 349. 1972. Fig. 66A.
Typus: Psathyrella vesiculocystis A.H. Sm. USA, Idaho, Idaho co,
French Creek Grade, Salmon river near Burgdorf, 4 Sep. 1964,
A.H. Smith & H.D. Thiers 70089 (MICH5380), gregarious on
conifer duff. Tef-1α MF521772, ITS MF326007.
Loan of one sporocarp and one pileus in good condition for
micro-morphology investigation.
Spores 7.8–9.8 × (5–)5.5–6.5(–7) × (4–)4.5–5.5 µm, fQ 1.3–1.55,
pQ 1.6–2; in front view oval to subangular-oval, base obtuse to
broadly rounded, sometimes truncate, apex often attenuate
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Voto et al.
to snout-like projected, in profile adaxially flattened; dusky
brown with an orange-red shade; germ pore distinct, mainly
truncate, sometimes somewhat oblique in profile, 1.6–1.8 µm
broad. Pleurocystidia 37–60 × 12–20 µm, broadly utriform,
subcylindrical to subclavate, apex broadly rounded; walls
hyaline; moderately numerous. Gill edge cystidia 8–48 × 4.5–18
µm, a mixture of some distinctly utriform cells and numerous,
dominating, small to large, saccate, clavate and subellipsoid cells
in variable number.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Notes: Smith (1972) detected some broader cheilocystidia, up to
25 µm broad, and reported occasional subechinulate protrusions
near the apex of pleurocystidia. These last however are only the
deceptive optical result of inwardly collapsed thin-walled apices
sometimes leaving a rugged or spiked crater; this occurrence has
been observed also in the following collection, Smith 56936 (P.
subnuda sensu A.H. Smith).
37c. Psathyrella subnuda (P. Karst.) A.H. Sm., Contr. Univ. Mich.
Herb. 5: 61. 1941.
We have attempted to understand Smith’s (1972) concept of
P. subnuda by examining the voucher Smith 56936 selected
for this purpose by MICH staff. We have found a sufficiently
good correspondence, both morphological and molecular,
with P. abieticola. This result does not necessarily apply to the
other vouchers Smith deposited in MICH under this name. The
basionymous name Psathyra subnuda P. Karst. sensu auct. Eur.
is generally associated to P. spadiceogrisea. Kits van Waveren
(1985), and Romagnesi, in personal communication to him,
considered it extremely dubious.
Smith 56936, MICH33421. Fig. 66(b).
= Psathyrella abieticola
Material examined: USA, California, Humboldt Co., Trinidad, 22 Dec.
1956, on burned soil, Smith 56936, MICH33421. Tef-1α MF521778, ITS
MF326000.
Loan of one sporocarp in good condition with pileus and stipe
separated for micro-morphology investigation.
The notes of gross morphology by the collector report pileus
mars brown, veil absent, stipe dull white and not darkening.
Spores (7.2–)7.7–10(–11.2) × 5.2–7.2 × 4.4–5.5 µm, fQ 1.3–1.6,
av. fQ 1.45, pQ 1.5–1.8 (1.9), av. pQ 1.6; in front view oval,
broadly elliptic, sometimes (irregularly) angular-oval, rarely
oboval, base obtuse to rounded, rarely subtruncate, apex
attenuate or not, rarely snout-like projected, in profile adaxially
flattened, rarely subphaseoliform; dusky reddish brown; germ
pore distinct, mainly truncate, sometimes somewhat oblique
in profile, 1.6–2(–2.5) µm broad. Pleurocystidia 33–55 × 11–17
µm; utriform to subcylindrical, sometimes sublageniform; apex
rounded, sometimes subcapitate; walls hyaline, sometimes
thickened; moderately numerous. Gill edge cystidia 9.5–33 ×
6–17 µm, a mixture of three types of cells: distinctly utriform
to sublageniform, 25–33 × 12–13 µm, scattered; ellipsoid to
rhomboid and 21–29 × 10–17 µm, numerous; saccate to clavate
and 9.5–30 × 6–17 µm, numerous.
160
38. Psathyrella infida Quél., Bull. Soc. bot. Fr. 23: 329. 1877. Figs
67–69.
Pileus 10–20 mm broad, in early stages conical-campanulate
and umbonate with inflexed margin, finally convex and
subumbonate or not, radially rugulose in old sporocarps; when
young dark to blackish brown, hygrophanous, discoloring
reddish to ochraceous brown, at centre remaining for a time
so then fading to ochre, in periphery fading to white, at margin
keeping for a time an ochre-brown color often with a pinkish
tint; translucently striate when moist. Veil white, already
reduced to scarce or absent on most pilei but still present
as small to large flocks on some young pilei (see Fig. 62, at
the centre) and as small tufts on lower half stipe. Lamellae
adnate, broad, not or little ventricose, distant (about 16–18);
greyish then purplish grey or with brownish tinge; edge pale,
reddish towards margin of pileus. Stipe 18–40(–55) × 1.8–2.5
mm, cylindrical, often flexuous; concolorous with pileus,
hygrophanous, discoloring white to translucent-hyaline in
upper half, pale to subconcolorous with pileus in lower half;
apex apparently not pruinose, base sometimes with appressed
mycelium. Spores (10.5–)11–13.7(–15) × (5.2–)6.2–7(–8) µm,
on average 12–12.5 × 6.4–6.6 µm, Q (1.7–)1.75–2(–2.2), avQ
1.85–1.9; in front view elliptic, base obtuse, in profile elliptic to
subamygdaliform, sometimes adaxially flattened to narrowly
limoniform, apex sometimes snout-like projected; dark brown;
germ pore distinct, truncate, 1.5–2 µm broad. Basidia 4-spored,
11.5–12.5 µm broad. Pleurocystidia 23–63 × 11.5–16(–22) µm,
fusiform to lageniform, rarely utriform or conical; neck short
and more or less rostrate to long, sometimes flexuous; apex
subacute, rarely obtuse; moderately numerous. Cheilocystidia
–50 × 11–15 µm, scattered to locally moderately numerous.
Paracystidia 15–40 × 9–20(–29) µm; walls often thickened,
hyaline to slightly pigmented towards base; numerous and
dominating. Cells below marginal cystidia yellow-brown to
pale yellow-brown pigmented. Clamps present.
Habitat: Gregarious to cespitose; among moist grass of
alluvial plain near river bank with Salix alba trees and various
hygrophilous plants.
Collection examined: Italy, Vicenza, Trissino, near the river Guà, on
moist soil with deciduous trees, 28 Mar. 2013, O. Chiarello (det. P.
Voto), MCVE29119. Tef-1α MF521794, ITS MF325977.
Notes: We base our concept of P. infida on Romagnesi as he
studied French material and is presumably the most reliable
in understanding Quélet’s short diagnosis. Therefore we refer
to Kits van Waveren’s (1985) description as he reports a full
description received from Romagnesi and adds one’s own
microscopical observations of an exsiccatum received by the
same French mycologist.
Kits van Waveren (1985) cites a description by Einhellinger
(1973) but that German material is reported as having larger
spores and 2(1)-spored basidia thus suggesting it may represent
P. prona.
The above description of our collection fits well with that
by Kits van Waveren (1985). The main features of this species
are large dark spores, fusiform to lageniform pleurocystidia,
small sporocarps, distant and broadly adnate to triangular
gills, strongly developed but evanescent flocculose veil, pileus
possibly drying with pinkish shades at margin, gill edge possibly
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 67. Psathyrella infida. MCVE29119. Photo O. Chiarello.
Fig. 68. Psathyrella infida. MCVE29119. Photo O. Chiarello.
Fig. 69. Psathyrella infida. O. Chiarello, MCVE29119; MC Marginal Cystidia in KOH; PC Pleurocystidia; S1 Spores in water; S2 Spores in KOH.
reddish pigmented, habitat associated to deciduous trees in
moist to dry soil.
From a morphological perspective, various species are very
similar, the closer allies among them are P. romagnesii, which has
more distant gills and smaller sporocarps, P. orbicularis, which
has the gill edge dominated by cheilocystidia, P. orbitarum,
which has smaller spores on average.
The ITS sequence of our sample is very similar to that of
P. tenera found in GenBank (DNA nucleotide homology: ITS
= 99 % with 7 mismatches). It shares the moist habitat and
many morphological characters (for micro-characters of P.
tenera we refer to Smith 1972, and to Örstadius et al. 2015
who both have revised Peck’s type). However, there are two
relevant elements of differentiation given by the spores which
are distinctly shorter and have consequently a lower quotient
and by the gill edge which is markedly dominated by small
to large paracystidia intermixed with generally scattered
cheilocystidia. Unfortunately only the ITS region sequence is
available in GenBank for P. tenera; however, until a second
locus contradicting evidence should emerge, we consider this
collection a consistent representative of P. infida and a good
species in its own right.
© 2019 Westerdijk Fungal Biodiversity Institute
161
Voto et al.
39. Psathyrella bivelata Contu., Bull. Soc. Mycol. Fr. 107: 86.
1991. Figs 70–73.
This species is very close to P. candolleana from which it
differs for having larger spores, (6.5–)7.5–12.5(–15) × (4.2–)
4.5–6.7(–7.5) µm, on average 9–9.5 × 5–5.5 µm, of a medium
to somewhat pale reddish brown color and with presence of
some 2-spored basidia. The habitat of P. bivelata is, at least
as up to now ascertained, herbicolous and possibly associated
to nitrogen-rich soil but we cannot exclude it could also be
detriticolous on woody humus outside of woods. However, two
peculiar characters bond both species together: the presence of
a strong stripped incrustation on the pileal general veil cells and
the broadly ellipsoid to globose shape of a number of the same
cells, this last character possibly more emphasized in P. bivelata.
In gross morphology these two characters produce a tendency
of the general veil to turn brownish, or even to be so already in
young specimens, and to reduce itself to little flocks.
After its original collection in Sardinia, Italy, in 1989 a long
period of oblivion followed, until it was found for a second
time in 2006, again in Italy, precisely in Veneto. Since then, we
have identified this species in other collections from the same
region. The first Veneto collection including images was fully
described by Voto (2007), although images of the type were
not included in the protologue. Due to the fact that CAG, where
the type is hosted, does not loan any material, we have now
provided a reference sequence for that collection and we have
re-deposited it as MCVE29104. More illustrations are in Voto
(2013) (under the name of P. candolleana, voucher D030208,
now MCVE29117) and Voto (2011). More images from the
Veneto collections are in this paper, including two additional
illustrations of MCVE29104.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Materials examined: Italy, Veneto, Rovigo, Villadose, 3 m a.s.l.,
gregarious in grass in garden, 10 Nov. 2005, P. Voto, Fig. 70, 71
(MCVE29104, Tef-1α MF521812, ITS MF325961); at same location, 6
May 2006; at same location, 28 May 2006, at same location, 6 Jun.
2006; Rosolina Mare, 0–1 m a.s.l., gregarious in grass in garden, 10 Nov.
2007, P. Voto, Figs 72, 73 (MCVE29330); Vicenza, 39 m a.s.l., caespitose
in grass in urban flower bed near a Platanus, 27 May 2012, E. Zanella
(MCVE29117, former Voto D030208, Tef-1α MF521811, ITS MF325962).
Fig. 70. Psathyrella bivelata. MCVE29104. Photo P. Voto.
Fig. 71. Psathyrella bivelata. MCVE29104. Photo P. Voto.
40. Psathyrella yaoundeana Mossebo & Pegler, Kew Bulletin 53:
1001. 1998. Fig. 74.
Typus: Psathyrella yaoundeana Mossebo & Pegler. Cameroon,
Yaoundé, Cité-Verte, 30 Oct. 1996, D.C. Mossebo 42B
(K(M)55648), among grass.
Loan of two entire sporocarps somewhat mouldy for micromorphology investigation.
The authors described this taxon as provided with frequent,
obtusely ventricose-lageniform to broadly fusoid pleurocystidia
(utriform with rounded to obtuse apex by their sketched
illustration and following our terminology) and with a mixture,
without frequency ratio, of obtusely clavate, utriform to
sublageniform cells on the gill edge. These data made it an
eligible species of subsection Spadiceogriseae and the authors
themselves commented that it most closely approached P.
spadiceogrisea. In our revision we could find no pleurocystidia
but we noticed that various cheilocystidia, dislodged by pressure
162
Fig. 72. Psathyrella bivelata. Voto20071109.2. Photo P. Voto.
on the coverslip, floated and casually anchored themselves on
the gill sides. Cheilocystidia are about 21–37 × 12–17(–22) µm
large, the gill edge has the typical pattern of that of P. candolleana
with absent to very inconspicuous paracystidia. Spores (6.2–)
6.5–8.5(–9.8) × (3.7–)4.2–5(–5.5) µm, Q (1.4–)1.6–2, avQ 1.7
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
Fig. 73. Psathyrella bivelata. Voto20071109.2; Cells of general veil on pileus in Congo red.
Fig. 74. Psathyrella yaoundeana. Mossebo 42B, K(M)55648; MC Marginal Cystidia; S1 Spores in water; S2 Spores in KOH.
© 2019 Westerdijk Fungal Biodiversity Institute
163
Voto et al.
[we have also found a number of broader spores, up to 5.7(–6.3)
µm with Q 1.3–1.5], in front view elliptic to oval, base rounded,
in profile mainly adaxially flattened to subphaseoliform,
sometimes phaseoliform to elliptic; pale brown with a vague
reddish shade; germ pore distinct, rounded, 1–1.7 µm broad.
We failed to obtain the ITS sequence and only obtained a
dirty unusable sequence of the Tef-1α region from this sample;
however, on the whole, we are convinced this material belongs
to section Spintrigerae.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Phylogeny
The ITS alignment included 673 characters, and contained
380 (56.5 %) variable sites; the Tef-1α alignment included
607 characters, and contained 335 (55.2 %) variable sites;
the concatenated ITS and Tef-1α alignment included 1 281
characters, and contained 679 (52.0 %) variable sites. The
partition of homogeneity tests found no conflicts between the
ITS and the Tef-1α alignments. Both Maximum Likelihood and
Bayesian analyses produced the same tree topologies. Only the
Bayesian trees with posterior probability and bootstrap values
are shown. The most significant differences between the ITS
(Fig. 1) and the Tef-1α (Fig. 2) trees were mostly due to variation
in the list of taxa covered by each, due to failed sequencing of
one of the loci for some samples. The concatenated ITS-Tef1α tree was characterized by higher statistical support of its
branches, and can be used to infer the overall phylogenetic
relationships among the Psathyrella samples analyzed. In
the concatenated tree, Psathyrella samples were all placed
in a monophyletic group containing several statistically
supported clades, including a clade representative of the
subsection Spadiceogriseae. The nitens clade is placed as
the closest relative to the spadiceogrisea clade, while the
candeollana clade appears as the most distantly related to the
spadiceogrisea group, at least among the taxa include in this
analysis. Refer to the tree in the Supplementary Fig. 1 for a
better coverage of species to identify the closest relatives to
the spadiceogrisea group. In an intermediate position between
the nitens and the candeollana clade, the analysis identified
a clade including the three subclades fusca, piluliformis, and
gordonii and the two taxa P. incondita and P. cortinarioides.
Psathyrella infida was a solitary taxon outside all of the clades
and subclades identified by this analysis.
The spadiceogrisea group was further subdivided in nine
strongly supported groups, identified in Fig. 3 by the following
subclades: albescens, thuijna, hellebosensis, fatua, owyheensis,
praetenuis, umbrosa, albanyensis, spadiceogrisea. These
nine subclades provide a strong and exhaustive framework
to understand the evolution of species within the section
Spadiceogriseae, and to differentiate between potential gross
misidentifications (taxa falling in different subclades), and
nomenclature issues caused by the close relatedness of sister
taxa (taxa falling within the same subclade) possibly due to
the lack of distinctive morphological traits. Besides the limited
morphological variation when comparing closely related taxa,
the composition of taxa within each subclade may be associated
with geographic range and/or with habitat.
Pairwise genetic distances within clades were in general
low [ITS (0.000)0.006(0.020), Tef-1α (0.000)0.007(0.020);
(min)average(max)], even for morphologically distinct
species, and five clades included both North American and
European species suggesting either a recent evolutionary
164
divergence, or a dynamic phylogeographic history resulting
in the intercontinental movement of species. No distinctive
morphological trait was associated with clades, except for the
albanyensis clade characterized by dark, often tridimensional
spores, and with a little to moderately distinct germ pore.
Clades though did differ in habitat association as follows. Clades
spadiceogrisea and albanyensis were found in association
with woody substrates, even in the case of P. clivensis, which
apparently grows in grasslands but is also associated with
woody debris. Clades umbrosa and praetenuis appeared to be
intermediate and include both lignicolous and non-lignicolous
species. All remaining clades seemed to be distinctively more
detached from woody substrates and included species growing
in grasslands or in “specialized” habitats such as wetlands
populated by hygrophilous plants (e.g. P. thuijna), and sandy
substrates (P. ammophila).
In spite of the fact that the bulk of the sequences of the
largest dataset came from the published study by Örstadius et
al. (2015), the concatenation of Tef-1α and ITS failed to meet the
requirements of the partition of homogeneity tests. Nonetheless,
we opted to provide the phylogenetic tree resulting from this
larger dataset in the supplementary materials (Supplementary
Fig. 1). The resulting tree has in fact the convenience of being
directly comparable with that of Örstadius et al. (2015), of
including a larger number of taxa within and outside the section
Spadiceogriseae, and of allowing for a relative placement of
the group within the genus. Once again, the spadiceogrisea
group results as monophyletic and includes nine well supported
clades. Three species, namely P. sublatispora, P. prona and P.
jacobssonii, made up an additional clade that - based on ITS
analysis - could be regarded either as part of the same group, or
as a sister clade to the subsection (Fig. 1).
Based on morphology and molecular analyses combined,
this study describes a new species (P. rogersiae), renames three
species (P. vesiculosa, P. ochrofulva, P. sanjuanensis), identifies
as present in Europe two taxa previously described only in
North America (P. albescens, P. kauffmanii), resolving the related
nomenclature issues, and suggests the possible presence
in Europe of a third American species (see notes about P.
velibrunnescens and P. aff. kauffmanii). Five species of uncertain
status were fully validated (P. bivelata, P. carinthiaca, P. fragrans,
P. infida, P. aff. kauffmanii).
The analysis of the holotypes of P. fusca, P. spadiceogrisea,
P. agrariella, P. atrifolia and P. subnuda is still missing, however
the combined DNA and morphological analysis of non-typical
vouchers (Smith 1972) reveals that Smith may have had an
ambiguous concept of these species. In particular, the last three
species remain dubious. Likewise, the analysis of the holotypes
is still missing for P. niveobadia, P. phaseolispora and P. agraria
ad int., three European taxa Smith never studied; the latter two
species remain dubious while we propose a provisional solution
for the first.
Finally, in this study we have identified 13 heterotypical
synonyms (see Table 1). All of the specific taxonomic contributions
of this paper are listed in Table 1. We have previously discussed
our approach with regards to how DNA sequence information
was used to confirm or contest species boundaries, especially
when morphology was insufficient to make that call. In brief,
while DNA homology < 97 % was used with confidence to
identify distinct taxa, highly homologous sequences (> 98 %)
cannot be used by themselves to exhaustively determine
conspecificity, because of possible transfer of alleles between
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
species, especially if closely related. High DNA homology at
one locus can though be used to support conspecificity if such
conspecificity is further suggested by homology at another locus,
morphology, habitat, and geographic range. We have regarded
homology levels of 97 % and 98 % as border-line, and looked
for non-synonymous polymorphisms in exonic regions (DNA
variation resulting in a variation in aminoacids transcribed) or
for indels (insertion or deletions) in intronic regions to suggest
the evolutionary significance of this relatively small variation in
DNA sequences. Individual species description drew on such
DNA-based comparisons when needed.
DISCUSSION
A first clue of monophyly of the subsection Spadiceogriseae
supported by molecular evidence was provided in Vašutová
et al. (2008), though based only on three vouchers. The
parsimonious tree in Örstadius et al. (2015) show that such
monophyly is confirmed when applied to a wider European
range of species. The results obtained through the present work
confirm the monophyly of a joint clade spadiceogrisea in Europe
and America, with four cases that need be discussed in detail.
P. umbrosa has paracystidia which, although dominating
the gill edge, are mainly unusually small and intermixed
with numerous basidia and basidiola;
b) Psathyrella sp. (NL-0631) is described as having scattered
cheilocystidia and very scattered paracystidia;
c) P. ochrofulva (and P. lactobrunnescens, if indeed it belongs
to the clade spadiceogrisea) has often fusiform to conical
pleurocystidia equipped with highly variable, mainly obtuse,
apex, which is unusual in comparison to usually utriform to
lageniform cystidia with mainly rounded apex;
d) P. aff. kauffmanii and P. vesiculosa are outside of the clade
spadiceogrisea but are morphologically associated to the
subsection Spadiceogriseae. However, it can be noted that
they, together with the unsequenced P. velibrunnescens,
are characterized by a distinctly pigmented veil, which
is otherwise unknown in this group. From this last cited
case we infer that a white veil must be included among
the morphological parameters defining the subsection
Spadiceogriseae.
a)
Based on morphological analysis and on sequence data from both
ITS and Tef-1α, nine statistically well-supported clades comprise
the subsection Spadiceogriseae. ITS data alone identified an
additional clade including P. prona, P. sublatispora, and P.
jacobssoni that, although rather distantly related to the other
clades, could be regarded as being included in the subsection.
However, these three taxa were not included in this study, and
therefore no direct comparative morphological data or Tef-1α
sequence information is available to confirm their taxonomic
placement. Due to the distinct positioning of this clade within
an expanded - yet still monophyletic - P. spadiceogriseae
subsection, morphology-based taxonomic positioning may also
be more ambiguous than for the other taxa beloging to a more
narrowly defined subsection.
The paramount character of the subsection Spadiceogriseae,
i.e. the dominance of paracystidia on the gill edge, can be
present also in other subdivisions of the genus and, therefore,
this character must have originated independently multiple
times in the evolution of Psathyrella. Yet, the set of characters
that together define the subsection Spadiceogriseae is
morphologically unambiguous.
The revision, based on both morphological and molecular
data, of a number of samples and holotypes, has resolved some
problematic taxonomic situations, the more relevant of which
are listed immediately below.
The European P. phegophila has been shown to be a solid
species with a transcontinental range, though its correct and
prior name has to be the Smith epithet P. kauffmanii; this result
is only morphology-based as we lack the type sequence and
morphological revision of P. phegophila, however the European
collection we selected as a reference point is perfectly adherent
to its concept.
Previously unknown in Europe, the American species P.
albescens is discovered and documented in the old continent;
this taxon was until now repeatedly misidentified in Europe.
Although our conclusions were not based on the
sequencing of the type, nor on its morphological examination,
we propose to accept the autonomy of the European species
P. niveobadia, after analyses of some collections fully matched
its morphological concept. We were also able to describe the
differences with its closest relative, i.e. P. spadiceogrisea. Our
molecular comparisons revealed there is some evolutionary
divergence between the two species, however, because this
divergence is modest and ambiguous when comparing results
from the two loci, we expect future analyses based on multiple
loci will be necessary to definitely confirm or discard our
decision.
Based on the examination of some samples with pigmented
gill edge and pinkish pileus margin, we propose, with molecular
corroboration, that the taxon described with the informal name
of P. pseudocorrugis sensu Kits van Waveren (1985) should be
considered an intraspecific variation of P. fatua. The correct,
original interpretation, P. pseudocorrugis sensu Romagnesi, on
the contrary, is a species outside subsection Spadiceogriseae
because of its gill edge dominated by cheilocystidia (Örstadius
& Knudsen 2008).
The little-known Czech material present in GenBank under
the provisional name of P. aff. kauffmanii is more completely
described here, and we show that it nests within a small complex
of American species.
A critical examination of the clade spadiceogrisea in the
Tef-1α tree indicates that species within this clade may not
be distributed randomly within subclades, but there may be
trophism-based grouping. In subclades spadiceogrisea and
albanyensis, all species grow strictly associated with arboreal
material and humus, with the only partial exception of P.
clivensis growing usually in grassland but also found directly
attached to (buried) sticks (Tassi 2000). A second group,
containing Psathyrella sp. NL-0631 and subclades umbrosa and
praetenuis, has an intermediate situation with three lignicolous
species and two others not associated with arboreal material.
The subclades albescens, thujina, hellebosensis, fatua,
owyheensis and the species P. ammophila, on the contrary,
only include one species, P. argillacea, collected consistently
in association with arboreal material. All other fungi of this
last grouping are generally found in open fields, grasslands,
pastures, etc. with the partial exceptions of P. fatua, P. albescens
and P. owyheensis which extend their trophism to both kinds
of habitat. Some species in this complex have developed strict
associations with specialized habitats, such as the association
© 2019 Westerdijk Fungal Biodiversity Institute
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
165
Voto et al.
with hygrophilous plants in moist to wet soil reported for P.
thujina and allied taxa, or the association with sandy habitats
reported for P. ammophila.
Spore color in the subsection Spadiceogriseae is broadly
assignable to two categories: (a) brown, pale reddish, orange
or paler, and, (b) red-brown or darker. Generally, and using
Smith’s terminology, spores of the first group in KOH have a
cocoa color or paler tones possibly turning slowly to chocolate
color. Those in the second group are from the beginning, or
rapidly turn into, a chocolate or a darker color when in KOH. Of
course, intermediate situations can occur. Because Smith (1972)
described spore color only in KOH, this observation can offer a
possible clue for deducing spore color in water of Psathyrella
species described by Smith and not yet revised.
Some species outside of subsection Spadiceogriseae have
been studied here as well. The holotypes of P. fragrans and P.
carinthiaca were sequenced and, based on their molecular and
morphological characters, they can be confirmed as close to,
but autonomous from, P. piluliformis. Collections of the littleknown European species P. infida and P. bivelata are respectively
described and commented upon and their phylogenetic
sequences have been generated to offer a molecular reference
point. Finally, MCVE28713 represents an interesting rare albinistic
occurrence of P. cortinarioides, and was also sequenced.
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
Key to subsection Spadiceogriseae
At the conclusion of this study, we provide an intercontinental
key of the subsection Spadiceogriseae. Note that besides all the
species previously discussed in this paper, the key also includes
an additional taxon, identified in GenBank as Psathyrella sp.,
voucher NL-0631, and belonging to the P. umbrosa complex.
Psathyrella affinis, P. basii, P. cortinarioides, P. dunensis,
Kauffmania larga (formerly P. larga), P. obtusata, P. pertinax,
P. psammophila and P. sanjuanensis though not belonging to
the clade spadiceogrisea, are included in the key because they
have, or occasionally may have, the gill edge lined by numerous
paracystidia and/or they may have utriform pleurocystidia.
The users of this key should take into consideration the limited
knowledge of variability in morphology and habitat for the
species described with few or only one collection. After each
species name, the continent where its occurrence is ascertained is
reported in brackets; in the most recurrent cases of America and
Europe the initial letters A and E respectively are affixed instead.
A document by Voto, containing a key to all European species
of Psathyrella and a list of the principal synonyms with reference
to their first propounders, is also published online in the website
of The Associazione Micologica ed Ecologica Romana - A.M.E.R.
(www.ameronlus.it/chiavi_micologia.php).
1.
Germ pore absent to indistinct; spores pale to orangish (light to medium brown in P. aff. kauffmanii,
(pale) yellow-brown in P. subspadiceogrisea) ........................................................................................................................ 2
Germ pore distinct and/or spores distinctly pigmented .......................................................................................................... 10
2.
2.
Spores 4.5–6.5 µm broad or more ............................................................................................................................................. 3
Spores 6.5–10 × 3.5–5.5 µm ...................................................................................................................................................... 5
3.
3.
Pileus 30–140 mm; pleurocystidia 40–90 × 10–24; associated to woody debris ............................. see Kauffmania larga (A, E)
Pileus within 40 mm broad ........................................................................................................................................................ 4
4.
4.
Pleurocystidia 28–37 × 10–14 µm; found on soil in garden .............................................................................. P. neotropica (A)
Pleurocystidia 35–60 × 10–18 µm; associated or not to woody debris ................................................................ P. clivensis (E)
5.
5.
Veil yellow to brown colored; pileus conical to campanulate; lignicolous
.................................................................. see P. vesiculosa (A), P. velibrunnescens (A), P. aff. kauffmanii (E), P. conica (Asia)
Veil white ................................................................................................................................................................................... 6
6.
6.
Spores on average 7–8 µm long ................................................................................................................................................. 7
Spores on average longer ........................................................................................................................................................... 9
7.
Pleurocystidia often yellow below the apex; spores not triangular, often phaseoliform; veil flocculose,
extending to halfway or centre of the pileus; smell indistinct ............................................................ see P. cortinarioides (E)
Pleurocystidia not pigmented; spores partly subtriangular, not or sometimes phaseoliform; veil scanty ................................ 8
1.
7.
8.
8.
Pileus –75 mm broad; stipe 30–90 × 5–12 mm; smell sometimes sweetish ................................................... see P. pertinax (E)
Pileus –23 mm broad, at start brown with darker centre; stipe 30–50 × 2.5–3 mm; smell indistinct;
spores (pale) yellow-brown .......................................................................................................... P. subspadiceogrisea (Asia)
9.
Pleurocystidia utriform to subcylindrical, often with elongate-cylindrical neck, rarely subellipsoid or ventricose-conical;
apex (4–)6–10 µm broad, mainly rounded, sometimes obtuse to subacute; paracystidia never mucronate
.......................................................................................................................................................................... P. rugulosa (A)
Pleurocystidia more varied, with apex mainly obtuse; paracystidia sometimes mucronate
............................................................................................................................... see P. obtusata (A, E), P. psammophila (A)
9.
10.
10.
166
Growing on wood, in woody debris, leaf litter, humus, in parks or forests .............................................................................. 22
Growing in open fields, grasslands, pastures, on herbaceous debris, among hygrophilous plants,
in nitrogen rich soil, arenicolous, on dry to wet soil ............................................................................................................ 11
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
11.
11.
Spores on average less than 9.3 µm long and 5.1 µm broad ................................................................................................... 12
Spores longer and/or broader .................................................................................................................................................. 16
12.
Spores often distinctly ovoid to angular-ovoid, reddish brown; pleurocystidia often capitate to forked;
on moist soil .............................................................................................................................................. P. hellebosensis (E)
Spores different ........................................................................................................................................................................ 13
12.
13.
Pileus usually still distinctly greyish brown when mature, not pink at margin; gill edge not pigmented;
spores rarely or sometimes phaseoliform; pleurocystidia not or rarely incrusted, forked or brownish .............................. 14
Pileus quickly discoloring, usually to brownish orange shades, sometimes with pinkish tints,
fading to white, drying greyish violaceous; gill edge sometimes pigmented; stipe at the base sometimes bent,
swollen, or equipped with a pseudorhiza [If pleurocystidia fusiform with apex in part mucous or subcapitate,
see P. dunensis (E)] ............................................................................................................................................................... 15
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
13.
14.
14.
15.
15.
Spores reddish brown; in moist, often flooded, permeable alluvial plain with Salix and hygrophilous plants
......................................................................................................................................................................... P. rogersiae (E)
Spores orangish; in open pastured ground or in forest soil ......................................................................... P. owyheensis (A, E)
Pleurocystidia (narrowly) utriform, sometimes fusiform, rarely lageniform or clavate, capitate or not,
forked or not, incrusted or not, sometimes brownish; spores often ± phaseoliform, to ± indented above
the apiculus, orangish brown to orangish reddish brown; veil mature sometimes still appendiculate at the
margin of the pileus, folded back on the gills or, rarely, annular on the stipe; gill edge rarely weakly pigmented;
pileus at most and rarely pinkish buff ......................................................................................................... P. albescens (A, E)
Pleurocystidia utriform, lageniform, fusiform, conical, rarely clavate, capitate or not, rarely incrusted or forked,
never brownish; spores rarely phaseoliform or indented above the apiculus, sometimes subphaseoliform,
brown to dark red; veil fugacious; gill edge sometimes red; pileus sometimes discoloring with distinct to
weak pinkish tints .................................................................................................................................................. P. fatua (E)
16.
16.
Spores mainly and distinctly phaseoliform .............................................................................................................................. 17
Spores at most sometimes subphaseoliform, not or rarely distinctly phaseoliform ................................................................ 18
17.
17.
Spores 9.7–11.6(–12.5) × (5.2–)5.5–6(–6.5), on average about 10.7 × 5.8 µm ............................................ P. agraria ad int. (E)
Spores 8–10 × 5–6 µm, on average 9 × 5.5 µm ............................................................................................ P. phaseolispora (E)
18.
18.
Spores pale, orangish to reddish yellow .................................................................................................................................. 19
Spores more pigmented, generally distinctly reddish brown ................................................................................................... 20
19.
19.
Stipe with a conspicuous white, woolly ring; on soil in a flood-plain, Hungary ............................... Psathyrella sp. NL-0631 (E)
Stipe without a ring; in open pastured ground or in forest soil ................................................................... P. owyheensis (A, E)
20.
Habitat arenicolous on coastal dunes or inland, not associated to moist soil or hygrophilous plants;
spores (7.5–)8.5–14 × (4.7–)5–8.5 µm, avQ 1.5–1.9 ................................................................................ P. ammophila (A, E)
Habitat different, associated to hygrophilous plants in moist soil or on mud in cow pastures ................................................ 21
20.
21.
21.
22.
22.
23.
Growing in wet to moist soil among hygrophilous plants (e.g. Typha, Phragmites, Cirsium, Epilobium);
sometimes clampless ...................................................................................................................................... P. thujina (A, E)
[If pleurocystidia (ventricose-) lageniform to obconical, see P. basii (E)]
Habitat different, on mud in a cow pasture (it is dubious whether this species is coprophilous/nitrophilous or not)
................................................................................................................................................................ P. pseudolimicola (A)
Spores tridimensional or broader than 5.5 µm in front view, strongly pigmented; germ pore little distinct;
stipe without a ring [if pleurocystidia mainly ventricose-lageniform with obtuse apex, see P. affinis (A)] ......................... 23
Spores on average < 5.5 µm broad, not or inconspicuously tridimensional; some species with a moderately
distinct germ pore [If spores (8–)8.2–9.8(–10.7) × (5–)5.2–5.8(–6) µm, dark, germ pore distinct,
on moist soil under aspen, see P. sanjuanensis (A)] ............................................................................................................. 25
23.
Spores often cordiform to pentagonal, 7–9 × 4.5–6.3 × 4–5 µm, fQ 1.3–1.5; on soil in a mixture of aspen
and conifers at about 2 438 m a.s.l. (8 000 feet) ......................................................................................... P. albanyensis (A)
Spores not cordiform, on average > 5.5 µm broad at least in front view ................................................................................. 24
24.
24.
Spores 9–12 × 5.5–7 µm; on unspecified sticks ................................................................................................... P. latispora (A)
Spores 7–10(–10.7) × 5–6.7 × 4.4–5.5 µm; on conifer wood .......................................................................... P. oregonensis (A)
© 2019 Westerdijk Fungal Biodiversity Institute
167
Voto et al.
25.
25.
26.
26.
Pleurocystidia 20–32 × (9–)11–16 µm long, with mainly undifferentiated apex; veil absent or scarce;
pileus 10 mm; stipe 40 × 1.5 mm; spores dark, oval, cylindrical oval, oboval, sometimes irregular or
angular, with broadly rounded to subtruncate base; germ pore little to moderately distinct ............... P. ovaticystis (Africa)
Pleurocystidia larger and/or more differentiated; different combination of characters .......................................................... 26
Pleurocystidia in NH4 often with reddish brown mucoid to granular coverings and granular contents,
utriform to clavate or cylindrical, sometimes slightly thick-walled and pigmented, 45–52 × 14–21 μm;
stipe with a striate, membranous ring; spores 7.5–9.5(–10) × 4–5.5 µm ............................................. P. striatoannulata (A)
Pleurocystidia apex generally not equipped with coverings and contents staining reddish brown in NH4 .............................. 27
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
27.
27.
28.
28.
29.
29.
Spores dark, often with a truncate to subtriangular base, sometimes (irregularly) angular-oval or angular-oblong;
germ pore moderately to little distinct; veil when mature often still distinct with floccules at the margin
or appendiculate or as a membranous ring; pleurocystidia forked or not, incrusted or not,
sometimes or rarely weakly pigmented ............................................................................................................................... 28
Different combination of characters; without a ring (occasionally in P. albescens); if spores dark then germ pore distinct
(see P. fatua) ........................................................................................................................................................................ 30
Pleurocystidia mainly slender, (37–)45–67(–83) × (10–)13–20(–32) µm, never clavate,
often with a tapering and elongate upper portion, scarcely subcapitate; gill edge cystidia never mucronate;
in association with Alnus .................................................................................................................................... P. alnicola (A)
Pleurocystidia shorter, (22–)26–56(–70) × 9–24 µm, sometimes clavate, with an apex often broad,
tapering or not in the upper portion; gill edge cystidia sometimes or often mucronate;
mainly in association with Alnus or Fagus ........................................................................................................................... 29
In mesophilous habitat, mostly with Fagus; cheilocystidia rarely attenuate to subacute at the apex;
pleurocystidia utriform to ellipsoid or clavate, sometimes lageniform, with an often capitate to
subcapitate apex, (22–)26–56(–70) × 9–24 µm ........................................................................................ P. kauffmanii (A, E)
In moist habitat under Alnus; cheilocystidia often mucronate; pleurocystidia more varying: utriform, clavate,
sublageniform, mucronate, obconical, with a rarely subcapitate apex, (24–)30–45(–52) × (10–)11.5–21;
spores (7.4–)7.5–9.6(–10) × 4.9–5.6 µm .......................................................................................................... P. solheimii (A)
30.
30.
Spores on average < 9.2 µm long and < 5.1 µm broad ............................................................................................................. 34
Spores on average longer and/or broader ............................................................................................................................... 31
31.
31.
Spores reddish orangish to dull orange; pileus sometimes fading to pinkish buff ................................................................... 32
Spores reddish brown .............................................................................................................................................................. 33
32.
Pleurocystidia utriform to lageniform, sometimes subcylindrical, apex rarely forked or incrusted;
paracystidia (sub) hyaline; spores in front view sometimes oblong to subcylindrical or slightly irregular;
context rigid; odor sometimes of fish ...................................................................................................... P. owyheensis (A, E)
Pleurocystidia utriform, apex often forked or incrusted; paracystidia often yellowish;
spores not so; context fragile ......................................................................................................................... P. argillacea (A)
32.
33.
33.
34.
34.
35.
168
Pileus clay color turning to greyish tints, 1–3 cm; stipe 20–30 × 2–4 mm, equal;
spores medium reddish brown; on oak debris .......................................................................................... P. griseopallida (A)
Pileus discoloring to cinnamon buff with a slightly redder centre, fading pale date brown, 3–5 cm;
stipe 60–120 × 3–6 mm, subclavate; spores sordid reddish brown; often on Populus debris ...................... P. praetenuis (A)
Paracystidia mainly 9–12 × 5–7 µm, some –32 × –13 µm, hyaline, among numerous basidia and basidiola;
pleurocystidia rarely to sometimes forked, hyaline, never incrusted or pigmented;
pileus pale dingy cinnamon brown turning dark brown when mature, fading to pallid, 1–3 cm broad;
stipe 20–50 × 1–3 mm; spores sometimes or rarely subphaseoliform, (6.7–)7–9.3(–9.5) × 4.3–5.7 µm
.......................................................................................................................................................................... P. umbrosa (A)
Paracystidia generally more conspicuous; different combination of characters ...................................................................... 35
Pleurocystidia sometimes brownish, forked or capitate, apex often with pigmented thickwalled traits
or covered with grossly mucous masses disappearing in old exsiccata; pileus not or scarcely striate,
when young with dark tones of violaceous brown to reddish brown, possibly still dark when mature,
15–72 mm; stipe 30–130(–200) × 3.5–10 mm, base often attenuate to strongly rooting;
spores on average 7.4–8.3 × 4.5–4.9 µm, orange-brown to reddish orange brown
(darkish reddish brown in NH4), often phaseoliform; context of pileus and especially of stipe cortex thick,
rigid, tenacious .............................................................................................................................................. P. niveobadia (E)
© 2019 Westerdijk Fungal Biodiversity Institute
A revision of Psathyrella, subsection Spadiceogriseae
35.
Pleurocystidia apex not mucous; stipe base at most subrooting or equipped with a pseudorhiza;
spores brown, reddish brown, dark red (see P. albescens and P. tenacipes with orangish reddish brown spores);
context not distinctly thick and rigid (but species with somewhat rigid stipe cortex can occur,
e.g. P. fatua, P. albescens, P. tenacipes) ................................................................................................................................ 36
36.
Pleurocystidia often little differentiated between ventricose and apical part, sometimes or often subcapitate,
not forked, not incrusted, not pigmented; paracystidia yellow and gill edge pallid brownish;
pileus becoming distinctly rugulose, not discoloring pinkish, 10–40 mm; spores orangish reddish brown,
7–9.5(–10.1) × 4.6–5.7 µm, sometimes subphaseoliform; stipe rather stringy, fibrous, cartilaginous-pliant,
30–60 × 2–3.5 mm; in moist to wet habitat (on muck and small sticks partly burned in the protologue,
on drying muck in ash swamp in the field notes) ............................................................................................ P. tenacipes (A)
Different combination of characters; in dry habitat ................................................................................................................. 37
Editor-in-Chief
Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Inst
E-mail: p.crous@westerdijkinstitute.nl
36.
37.
37.
38.
38.
39.
39.
40.
40.
41.
41.
Pileus –20 mm broad, striate; pleurocystidia often little differentiated, apex mainly obtuse or subacute,
rarely rounded, never capitate; stipe –30 × –2.5 mm; paracystidia sometimes or often with a pale pigment
(gill edge possibly pigmented?) [If pileus usually drying with pinkish tints, pleurocystidia apex in part mucous
or subcapitate, see P. dunensis (E)] ...................................................................................................................................... 38
Larger sporocarps or else different combination of characters ............................................................................................... 39
Pileus yellowish fulvous to reddish cinnamon, mature broadly convex to plane; pleurocystidia conical,
utriform-conical, fusiform, ellipsoid, also utriform, subcylindrical, lageniform, slightly pigmented,
apex sometimes mucronate, forked, truncate, incrusted; spores orange-brown
(in KOH bright cocoa slowly turning chocolate in Smith 1972) ..................................................................... P. ochrofulva (A)
Pileus white turning greyish brown when mature and fading white, obtusely conical and hardly expanding;
veil apparently absent; pleurocystidia mainly subfusoid to subutriform, not pigmented, not mucronate,
not forked, not incrusted; spores in KOH dark cocoa brown slowly turning darker in Smith (1972)
............................................................................................................................................................ P. lactobrunnescens (A)
Pileus not striate, hardly expanding, 25–30 mm broad, bronze brown, then gray- to clay-brown
or pale ochre to cream; stipe 50–70 × 3–4 mm; spores rarely subphaseoliform, reddish brown,
8–10.5 × 4.5–5 µm; germ pore small; pleurocystidia utriform to subutriform, scarcely subcapitate,
rarely yellowish brown incrusted ................................................................................................................... P. marquana (E)
Pileus striate when fresh, usually expanding; sporocarps of the same dimension or larger;
different combination of characters ..................................................................................................................................... 40
Pleurocystidia sometimes capitate, rarely incrusted or forked, never brownish; spores brown to dark red,
rarely phaseoliform, sometimes subphaseoliform; pileus reddish brown, quickly discolouring through
brown to brownish orange or paler, sometimes with distinct to weak pink tints, 8–50 (60) mm broad;
gill edge sometimes red; stipe (10–)20–110 × 1–5.5(–7) mm, base sometimes attenuate or with a pseudorhiza
................................................................................................................................................................................ P. fatua (E)
Pleurocystidia sometimes brownish, incrusted, forked; spores not dark red; gill edge generally not pigmented;
spores on average 7.4–9 µm long, usually often ± phaseoliform to ± indented above the base;
pileus rarely with vague pinkish shades (P. albescens) ......................................................................................................... 41
Pileus when young dark yellowish brown to cinnamon-brown, discolouring through shades of brownish orange
(rarely pinkish buff) to finally white, 10–70 mm; stipe 20–100(–140) × 1.5–6(–12) mm, base often attenuate,
subrooting or equipped with a pseudorhiza; veil mature sometimes still appendiculate or folded back on the gills,
rarely annulate; spores orangish brown to orangish reddish brown; pleurocystidia subcapitate or not
..................................................................................................................................................................... P. albescens (A, E)
Pileus dark (violaceous) reddish brown, when mature sometimes often still strongly pigmented,
usually not fading completely white, 15–55 mm; stipe 40–80 × 1.5–8 mm, base at most sub attenuate;
veil mature fugacious; spores reddish brown; pleurocystidia rarely capitate .................................... P. spadiceogrisea (A, E)
ACKNOWLEDGEMENTS
We are grateful to the curators of the following herbaria: K, MICH, NY,
and particularly wish to express our gratitude to the MICH’s Collection
Manager and correspondent, Patricia Rogers, for her collaboration in
answering our many questions about the MICH specimens studied.
We wish to thank Kathy Richmond, of the Southern Idaho Mycological
Association (SIMA), who verified a collection site in Idaho, Joaquim
Carbó, Ottorino Chiarello, Alessio Micucci, Miquel À. Pérez-De-Gregorio,
Bruno de Ruvo, Marco Maletti, Gerard Tassi, Renato Tizzoni, Evelina
Zanella and Marino Zugna who supplied us with gifts of collections
and pictures; thanks are also due to Giovanni Robich of the herbarium
MCVE who for a time assisted us in communicating with the herbaria.
Also, we are grateful to L. Örstadius and L. Nagy for providing to us
details about their Psathyrella sp. NL-0631, and to M. Vašutová for the
collaboration on P. aff. kauffmanii. The key also includes an additional
© 2019 Westerdijk Fungal Biodiversity Institute
169
Voto et al.
taxon, identified in GenBank as Psathyrella sp., voucher NL-0631,
which belongs to the P. umbrosa complex. This was possible thanks to
courtesy of L. Nagy, who collected it together with Z.S. Gorliczai, and to
L. Örstadius, who examined it microscopically.
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Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail: p.crous@westerdijkinstitute.nl
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Supplementary Material: http://fuse-journal.org/
Fig. S1. Bayesian phylogram obtained from combined nrITS and Tef-1α
sequence alignment of Psathyrella spp. Bolbitius vitellinus was used as
outgroup taxon. Only BPP values ≥ 0.95 and MLB values ≥ 70 % are
given above clade branches. In addition to sequences generated by this
study and to select GenBank accessions, most sequences published by
Örstadius et al. (2015) are included in this tree.
© 2019 Westerdijk Fungal Biodiversity Institute
Fig. S1, part 1
1/71
0.95/92
1/93
P. sphagnicola LO233-99
/ sphagnicola
P. vesterholtii JHP10-086 Type
P. flexuosipes MICH32961 US Paratype
P. incondita MICH36450 US
P. rostellata LO228-85 type
P. ellenae var. yubaensis MICH11919
/ ellenae
1/100
P. sabuletorum LO196-98 type
P. sabuletorum JV90 77
P. fibrillosa LO138-00
P. gordonii MCVE28708 IT
P. gordonii MCVE28710 IT
P. pervelata SZMC NL-1950
1/91
P. gordonii LO220-95
1/100
P. vestita SZMC NL-2346
P. sp LO96-11
P. atomatoides LO249-82
/ atomatoides
P. obtusata MICH65721
-/96
1/95
1/100
1/96
0.03
P. flexispora LO228-00
P. vesiculosa MICH12044
1/100
P. aff. kauffmanii P9 AM712260
/ septentrionalis
P. septentrionalis MICH12045
P. cortinarioides MCVE28713
P. cortinarioides LO77-00
P. spintrigeroides LO127-86
1/97
P. pennata LO206-03
/ lacuum
P. lacuum Huijsman KC992887
1/99
P. orbicularis LO210-04
0.99/93
P. orbicularis LO149-11
1/93
/ ichnusae
P. ichnusae Contu080106 Type
P. hololanigera Hausknecht071109
-/98
P. squamosa LO164-96
1/100
/ incondita
P incondita MICH36451 US
0.97/P. squamosa LO194-96
P. fimiseda LO56 96 type
P. hirta LO142-00
P. impexa LO162-03
P. dicrani LO270-04
0.96/98
/ dicrani
P. parva LO23-08
1/99
P. seymourensis LO42-87
P. laricina Smith64604 MICH type
P. siccophila LO417-06 type
P. pseudocasca LO17-04
P. umbrina LO235-04
P. kitsiana LO217-85 type
1/100
P. madida LO377-06 Type
1/97
P. madida LO369-06
/ madida
P. merdicola LO45-02 Type
P. sp. Contu071230
P. suavissima LO4-87
P. scatophila LO64-95 type
P. obtusata 33261
P. scanica LO183-09 Type
0.98/P. warrenensis Smith70162 type
P. limicola var. subpectinata MICH11965 US
P. rubiginosa LO107-98
P. fagetophila LO210-85 type
P.
senex
LO115-02
1/85
0.99/73
P. seminude Smith34091 MICH type
P. agrariella MICH47939 US
/ agrariella
1/100
P. fennoscandica LO95-96
0.95/78
P. fennoscandica LO484-05 Type
-/99
P. romagnesii LO267-04
1/91
1/100
P. romagnesii LO85-98
P. pratensis MICH12009 US Type
1/99
P. noli-tangere LO83-03
P. pseudocorrugis LO226-06
1/100
P. tenuicula LO37-04
P. tenuicula LO58-03
0.96/84
P. lyckebodensis LO301-11 Type
1/95
P. globosivelata Schumacher035
1/99
P.
albofloccosa
Sivertsen65-89
/ albofloccosa
1/98
P. kellermanii de Meulder11242
P. sphaerocystis LO126-99
P. fragrans MICH5347 US Type
P. obscurotristis Wilhelm489 Type
1/99
P. piluliformis MCVE29118 IT
/ carinthiaca
P. piluliformis LO162-02
P. carinthiaca MCVE25611 AT Type
0.98/79
P. pertinax LO259-91
1/97
P. mucrocystis LO103-98
P. echinata ZT12073
P. vesiculocystis MICH5380 US Type
P. velosa MICH33441 US Paratype
P. velosa MICH5376 US Type
P. abieticola Smith58673 type
P. subnuda MICH33421 US
P. fusca LO287-04
P. conferta GE02 007 PC type
-/85
/ abieticola
1/100
P. panaeoloides LO293-04
P. panaeoloides LO44-03
0.98/P. ephemera MICH11921 US Type
P. rybergii LO373-06 Type
1/98
P. pygmaea LO97-04
P. olympiana LO32-02
P. stridvallii LO104-98 Type
P. arenosa LO220 96 Type
P. epimyces WU19965
/ gordonii
P. albescens MCVE29116 IT
P. albescens MCVE29107 IT
P. agrariella MICH47933 US
P. mazzeri MICH11963 US Type
096/83
P. agrariella MICH65241 US
-/86
P. ochrofulva MICH11858 US Type
P. albescens MCVE29106 IT
P. albescens MCVE29113 IT
P. albescens MCVE29111 IT
0.98/86
P. albescens MICH11863 US Type
P. agraria KR0030008 KU307507
P. lutulenta AH21379 Type
-/89
P. argillacea MICH65256 US Paratype
0.96/79
P. thujina LO31-04
1/89
P. thujina Smith66720 MICH type
P. rogersiae MCVE29120 IT Type
0.98/81
/ agraria
P. ammophila LO169-01
1/100
0.97/P. ammophila LO359-11
P. velatipes MICH12106 US Type
1/97
P. almerensis MCVE29114 IT
P. almerensis LO379-06
1/100
P. carminei LO5 09 Type
P. owyheensis MICH5357 US Type
1/92
P. fatua LO132 DQ389681
P. fatua MCVE29105 IT
P. fatua MCVE29108 IT
P. fatua LO231-08
P. fatua MCVE29122 IT
P. fatua MCVE29124 IT
-/95
P. fatua MCVE29123 IT
P. fatua MCVE29115 IT
P. fatua MCVE29109 IT
P. fatua MCVE29110 IT
1/100
P. praetenuis MICH5361 US Type
-/91
P. atrifolia MICH32732 US
1/99
P. griseopallida MICH11939 US Type
1/88
/ atrifolia
P. pseudolimicola MICH5366 US Type
1/99
1/100
P. umbrosa MICH12102 US Type
P. fulvoumbrina MICH5833 US Type
P. sp NL0631
0.96/P. alnicola MICH11868 US Type
1/97
P. oregonensis MICH5840 US Type
P. subvinacea MICH12085 US Type
P. phegophila SZMC-NL-3527
P. olivaceopallida MICH11990 US
P. kauffmanii MCVE29101 ES
P. fusca MICH48491 US
P. spadiceogrisea MICH49606 US
1/87
P. kauffmanii MICH11962 US Type
/ albanyensis
P. striatoannulata KY350220
P. albanyensis MICH11862 US Type
-/86
P. spadiceogrisea LO92-01
1/77
P. spadiceogrisea LO102-98
P. praecox MICH49251 US Paraype
P. amarella MICH11869 US Type
P. spadiceogrisea MCVE29103 FR
-/92
P. clivensis LO182-03
P. lubrica MICH11957 US Type
P. niveobadia MCVE29102 IT
P. sublatispora LO190-97 type
P. jacobssonii LO256-92 type
P. nitens MICH33252 US
1/100
P. nitens MICH33251 US
0.98/96
P. nitens MICH11983 US Type
1/100
P. nitens MICH33249 US
/ angusticystis
1/98
P. nitens MICH33250 US
P. angusticystis MICH11873 US Type
1/100
P. psammophila Smith67836 MICH type
P. obtusata LO88-01
P. subargillacea MICH12058 US Type
1/100
P. cotonea LO136-00
1/75
P. duchesnayensis Smith61737 MICH type
0.96/P. caput-medusae LO36-94
/ caput-medusae
2x
P. maculata LO240-84
0.98/95
P. magnispora Barta Spittelberg
1/96
P. magnispora 24929-AH type
/ magnispora
P. mesobromionis Arnolds01-174 L type
P. microrhiza LO136-08 Type
P. microrhiza LO185-02
P. lutensis LO98-03
P. longicauda LO201-02
/ spadiceogrisea group
Fig. S1, part 2
1/74
P. longicauda LO254-91
P. tenera LO81-83
P. tenera LO382-89
P. potteri LO271-01
P.
tenera
AHS65853
/ infida
1/89
P. prona LO91-99
0.99/74
P. infida MCVE29119 IT
1/100
P. stercoraria Kytovuori Virrat 1991
P. stercoraria LO460-05 type
P. sp LO312-92
P. calcarea LO211-03
1/P. lilliputana LO130-09 Type
P. orbitarum LO257-90
-/75
P. riparia Brooks1600 MICH type
P. pratensis LO23-94
1/100
P. purpureobadia 9956 L type
-/75
/ efibulata
P.
vinosofulva
LO2-88
1/79
P. romellii LO240 01 UPS type
P. effibulata LO37-96 type
0.98/99
P. bipellis LO426-05
0.99/95
P. bipellis LO207-96
/ bipellis
P. bipellis LO50-04
P. amarescens Arnolds02-78 type
1/98
P. corrugis LO171-01
1/100
P. corrugis MCVE29121 IT
/ amarescens
0.96/99
P. pseudogracilis LO287-06
P. pseudogracilis LO172-02
1/98
P. multipedata LO237-04
/ multipedata
P. supernula LO250-04
P. badhyzensis 79478 TAA type92883
P. bivelata MCVE29104 IT
P. bivelata MCVE29117 IT
0.99/P. candolleana EB990221 KF281384
0.97/P. trinitatensis TL9035
P. candolleana LO38-00
/ candolleana
0.98/P. tuberculata ADK3564
P. efflorescens Pegler2133
0.96/P. tuberculata ADK4162
1/100
P. leucotephra MCVE28705 ES
P. leucotephra LO138-01
P. luteopallida Sharp20863 MICH type
P. sulcatotuberculosa MCVE29112 IT
P. typhae LO21-04
Mythicomyces corneipes AFTOL 972
-/77
1/83
1/100
1/1/88
Bolbitius vitellinus isolate AFTOL 730
0.03