A peer-reviewed open-access journal
MycoKeys 52: 89–102 (2019)
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
doi: 10.3897/mycokeys.52.31415
RESEARCH ARTICLE
http://mycokeys.pensoft.net
89
MycoKeys
Launched to accelerate biodiversity research
A new species of Psathyrella
(Psathyrellaceae, Agaricales) from Italy
Giovanni Sicoli1, Nicodemo G. Passalacqua2, Antonio B. De Giuseppe2,
Anna Maria Palermo1, Giuseppe Pellegrino1
1 Department of Biology, Ecology and Earth Science, The University of Calabria, 87036 Arcavàcata di Rende,
Cosenza, Italy 2 Museum of Natural History of Calabria and Botanical Garden, The University of Calabria,
87036 Arcavàcata di Rende, Cosenza, Italy
Corresponding author: Giovanni Sicoli (giovanni.sicoli@unical.it)
Academic editor: Bryn Dentinger | Received 6 November 2018 | Accepted 13 March 2019 | Published 16 May 2019
Citation: Sicoli G, Passalacqua NG, De Giuseppe AB, Palermo AM, Pellegrino G (2019) A new species of Psathyrella
(Psathyrellaceae, Agaricales) from Italy. MycoKeys 52: 89–102. https://doi.org/10.3897/mycokeys.52.31415
Abstract
Sporophores of a new Psathyrella species have been reported for the first time as growing at the base of
Cladium mariscus culms in the Botanical Garden of the University of Calabria, Rende, Cosenza, southern
Italy. The fungus was initially identified as P. thujina (= P. almerensis) by means of both ecology and macroand microscopic characteristics of the basidiomes, then referred to P. cladii-marisci sp. nov. after extraction,
amplification, purification and analysis of the rDNA ITS region. We came to this conclusion after comparing our specimen with the descriptions of the taxa available in the literature for the genus Psathyrella.
Keywords
Agaricomycetes, Basidiomycota, Fen-sedge, Marshes, southern Italy, Taxonomy
Introduction
Within the cosmopolitan fungal genus Psathyrella (Fr.) Quél. (Agaricales, Psathyrellaceae),
about one hundred species have traditionally been recognised in Europe, almost all saprotrophs and found in many and diverse environments. Either terrestrial or lignicolous,
they grow mainly on organic debris from various origins, such as dung, post-fire locations
and dead stems of larger herbaceous plants (Vesterholt and Knudsen 1992). Psathyrella
basidiomes are pileate, stipitated and exannulate or, at most, with a fugacious ring and
the hymenophore is gilled, pale pink when young, turning brown with age due to a dark
Copyright Giovanni Sicoli et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
90
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
spore print. Moreover, they have, as the etymology indicates, a very fragile and ephemeral
consistency. Despite these common macroscopic characters of the basidiomes, a recent
phylogenetic analysis revealed the extremely complex origin of this genus, recognising
species as belonging to a Psathyrella sensu stricto group or to P. sensu lato complex, the former including 19 clades and the latter involving eight genera (Coprinellus, Kauffmania,
Cystoagaricus, Typhrasa, Lacrymaria, Homophron, Coprinopsis, Parasola), thus consistently
widening the list of such “psathyrelloid” basidiomycetes (Örstadius et al. 2015).
During an investigation on the mycoflora of the Botanical Garden at the University
of Calabria (Rende, Cosenza, Italy), basidiomes of an apparently “psathyrelloid” fungus were detected at the base of a fen-sedge [Cladium mariscus (L.) Pohl (Cyperaceae)],
a cosmopolitan-distributed plant species (Lansdown et al. 2018) occurring in marshy
places of most Italian regions (Bartolucci et al. 2018), but rarely in southern Italy.
Based on records reported by Örstadius et al. (2015), nine clades of Psathyrella s.s.
include species associated with moist soils and marshy places: “spadiceogrisea” (four
species), “fibrillosa”, “noli-tangere” and “prona” (two species each), “candolleana”, “cystopsathyra”, “lutensis”, “obtusata” and “pygmaea” (one species each). Nevertheless, only
three species have been found to be growing on sticks or on remnants of hygrophilous
plants: P. lutensis (Romagn.) Bon, as a monospecific “lutensis” clade, P. thujina A.H.
Sm. (=P. almerensis Kits van Wav.) in the “spadiceogrisea” clade and P. typhae (Kalchbr.)
A. Pearson & Dennis in the “candolleana” clade.
The aim of this work was therefore to identify our basidiomes by using both
morpho-ecological and biomolecular tools. This was highly encouraged by the habitat
peculiarity and the close relationship with a plant species with which no species of
Psathyrellaceae had ever been found associated.
Materials and methods
Eight basidiomes of the above “psathyrelloid” fungus were observed and collected on
10 April 2018, as gregarious all around and at the base of Cladium mariscus cut culms
(Fig. 1). In 2012, that plant had been removed, together with the whole clump of mud
attached to its roots, from a natural marsh named Lago dell’Aquila (Laureana di Borrello, Reggio Calabria, southern Italy) and transplanted to the Botanical Garden at the
corner of a 90 × 37 cm-wide and 30 cm-deep concrete tank, which had permanently
been kept full to the brim with water. Since then, some leaves of water lily (Nymphaea
alba L.) have been introduced to float on the water surface inside the tank and the mud
mass has been increasing, while the C. mariscus plant has been expanding and producing new culms that are cut every year.
Morphology
The basidiomes were first macroscopically examined for features, colours, sizes, hymenophore
shape, pileus and stipe ornamentations, smell and taste. Then, the structures of the basidiome
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
91
Figure 1. A tuft of Cladium mariscus planted in a tank at the Botanical Garden of the University of
Calabria, southern Italy (A), and first-sight features of Psathyrella basidiomes at the base and in-between
of remnants of excised culms of the plant (B).
were microscopically inspected for cheilo- and pleurocystidia occurrence and features,
presence of clamp connections, basidia and spore features. These observations were carried
out under a light microscope (Axioplan 2 Imaging Microscope, Carl Zeiss, Germany) at
400 and 1,000 magnifications on fragments of pileipellis and gills placed on slides in 10%
92
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
NH4OH. The results were compared with those published in the morphological keys for the
Psathyrella species and, more specifically, with those species reported as the closest, according
to morphology and ecological site conditions, i.e. P. thujina, P. typhae and P. lutensis (Kits
van Waveren 1985, Vesterholt and Knudsen 1992, Christan et al. 2017, Henrici 2017).
DNA Extraction, Amplification and Sequencing
One of the basidiomes was dehydrated at room temperature and destroyed for molecular analysis: DNA extraction, amplification, purification and sequencing of the nuc
rDNA internal transcribed spacer region (ITS). DNA extraction was implemented
by using CTAB protocol (Doyle and Doyle 1987) and the ITS region was amplified
using the primer combination ITS1F/ITS4 (White et al. 1990). The polymerase
chain reaction (PCR) was performed in a 25-µl reaction volume containing 1.0 µl
DNA, 2.5 µl 10 × 5-Prime–MasterMix Buffer (Thermo Fischer Scientific, Waltham,
Massachusetts, USA) and 1.25 µl of each primer (10 µM/µl). The PCR was carried
out according to the following amplification programme: 3 min initial denaturation
at 94 °C, 35 cycles (30 s denaturation at 94 °C, 1 min annealing at 55 °C, 45 s extension at 72 °C) and a 10 min final extension at 72 °C. This programme was carried
out in a T1000 Thermocycler (Biometra, Goettingen, Germany). The PCR products were purified using a QIAquick PCR purification kit (Qiagen Inc., Valencia,
California, USA). Sequencing was performed by means of a Bigdye terminator cycle
sequencing kit (Applied Biosystems, Foster City, California, USA). The sequencing
reaction was run by BMR Genomics (Padua, Italy) on a 96-capillaries ABI 3730XL
DNA Sequencer.
Forward and reverse DNA fragment electropherograms were checked by means of
the CHROMAS 2.6.5 software (technelysium.com.au) for a complete reconstruction of
the ITS1, ITS2 and 5.8 gene fragments. Ambiguous regions at the start and the end of the
alignment were deleted and gaps were manually adjusted to optimise the alignment. The
sequence generated for this study is deposited in GenBank with the code MK080112.
Alignment and Phylogenetic Analysis
Consensus sequences were generated from both forward and reverse primer reads in the
BioEdit sequence alignment editor, version 7.2.5 (Hall 1999), then homology searches
were performed at the National Centre for Biotechnology Information (NCBI) Web
site using BLAST. This sequence was then compared with those of the Psathyrella species
deposited in GenBank on which the phylogenetic analysis had recently been performed
(Padamsee et al. 2008, Battistin et al. 2014, Örstadius et al. 2015, Yan and Bau 2018). A
total of 45 ITS sequences, including three Coprinellus spp. (Table 2) were aligned using
MAFFT with the L-INS-i option (Katoh et al. 2017). The aligned ITS dataset consisted
of 702 nucleotide sites (including gaps). FASTA alignments from MAFFT were loaded in
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
93
IQ-TREE 1.5.6 (Nguyen et al. 2014) to perform Maximum Likelihood Analysis. Clade
robustness was assessed using a bootstrap (BT) analysis with 1,000 replicates (Felsenstein
1985). Phylogenetic trees were visualised using the FigTree v1.3.1 (Rambaut 2009).
Results
Morphology
The macro- and micro-morphological features of the basidiomes collected at the base
of the fen-sedge plant in the Botanical Garden are shown in Figures 2, 3. At first sight,
by observing the macro-level characters, i.e. the small-medium size, the extreme fragility at handling and the brown-blackish spore print, the basidiomes were easily assigned
to the Psathyrella genus (Vesterholt and Knudsen 1992). Secondly, the occurrence of
sphaeropedunculate and clavate cells along the gill edge and the utriform shape of some
cheilo and pleurocystidia seemed to direct them to the Section Spadiceogriseae Kits van
Wav., subsection Spadiceogriseae (Romagn.) ex Kits van Wav. (Kits van Waveren 1985).
If we compare the morphological features of our specimens with those belonging
to the closest Psathyrella species, a number of differences emerge (Table 1). Our specimens appeared to be more similar to P. thujina (Henrici, 2017), previously described
as P. almerensis (Kits van Waveren 1985, Vesterholt and Knudsen 1992), except for
Table 1. Main differences between our Psathyrella sp. and the closest species, according to the morphological characteristics of basidiomes and mycelium, and ecology. (Differences from our specimen are in
bold characters).
Morpho-ecological
characteristics
Pileus diameter (cm)
Pileus colour
Psathyrella sp.
P. thujina
P. typhae
P. lutensis
3.5
Hazelnut brown,
then beige brown
2.5
Warm brown, then
beige brown
2.5
Pinkish-ochre brown,
then pale flesh brown
White with a
pruinose apex
White with a
pruinose apex
Whitish to pale brown
Spore size (µm)
7.2–11.8 x 4.3–6.0
9.0–11.5 x 4.5–6.5
Cheilocystidia
Versiform, chiefly
utriform
Utriform
Utriform
Utriform
7.5–11.5(12.0) x
5.5–8.0
Versiform, chiefly
utriform
Absent
4.0
Dark reddish
brown, then very
pale brown
White with a
pruinose apex,
brownish base
9.0–10.0 x 4.5–5.5
NO
Marshes, on cut
culms of Cladium
NO
Marshes, on cut
culms of Typha,
Phragmites, Cirsium,
Epilobium
Autumn to winter
NO
Marshes, on cut culms
of Typha, Epilobium,
Scirpus, Phragmites,
Rumex, Iris
Summer
Stem colour
Pleurocystidia
Mucoid deposits on cystidia
Habitat
Seasonal occurrence
Spring
Versiform, chiefly
utriform
Versiform, chiefly
utriform to
ventricose
YES
Deciduous forests,
on sticks in mud
Summer to autumn
94
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
Figure 2. Macro-morphological characteristics of the Psathyrella basidiomes: scales of velar origin on pilei
tops and margins, and beige-coloured gills (A); cylindrical, white and exannulate stems under a lateral
profile (B); colour-shading of a cap hygrophany and fibrillose details of velar-originated scales (C); gills
turning brown-purplish with spore maturation and a fibrillose surface of a stem base (D); a pruinose stem
apex bearing a mature hymenophore with white gill edge lines (E).
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
95
Figure 3. Micro-morphological characteristics of the Psathyrella mycelium: clavate and sphaeropedunculate
(A), and cylindric (B, C) cells at a gill edge; differently clavate (D, E) and utriform (F) cheilocystidia; variously utriform-shaped pleurocystidia (G, H, I); a fibulate hypha (J); a 4-spored basidium (K); basidiospores (L).
96
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
the pileus diameter reaching 3.5 cm in our specimens, but never exceeding 2.5 cm
in this species. Furthermore, our Psathyrella revealed versiform-shaped cheilocystidia,
while those reported for P. thujina are only utriform. P. typhae was also divergent for
the pileus diameter, not exceeding 2.5 cm, but even for pileus and stipe colours and
for lacking pleurocystidia. On the other hand, the mucoid deposits, characterising the
pleurocystidioid cheilocystidia of P. lutensis, were absent in our specimens. In addition,
the spore length range was wider in our specimens than in P. thujina and P. lutensis and
all the closest three species, which showed larger spores on average.
As for ecology, the plant genus Cladium Browne has never been reported as a substrate to any other Psathyrella, although P. thujina and P. typhae are commonly found
on the remnants of ecologically similar plants (Kits van Waveren 1985, Vesterholt and
Knudsen 1992, Örstadius et al. 2015, Henrici 2017). Furthermore, the genus Cladium
was not mentioned in the unique Italian report of P. thujina, which was found “in
open sites, close to any hygrophilous plants” (Voto 2016), in accordance with Henrici
(2017) who refers this species to reed-beds and generic damp marshy habitats. Finally,
our specimen was collected in the spring, whereas the above three other Psathyrella species seem to occur in other seasons.
DNA Analysis
The obtained nrDNA sequence was 702 bp long. By comparing it with those published
in GenBank, we obtained a data matrix composed of 44 taxa and 710 characters, 276
gap-free sites and 240 conserved sites. The highest homology (99%) was observed with
P. candolleana (Fr.) Maire, which was confirmed by the phylogenetic analysis (Fig. 4).
Indeed, the phylogenetic tree shows that our specimen falls into the “candolleana”
clade, such a heterogeneous group, including taxa from different morphology, ecology
and geographic provenance and, amongst them, the above-cited P. typhae (Battistin et
al. 2014, Örstadius et al. 2015, Yan and Bau 2018).
Discussion and conclusions
Based on results from both morphological and molecular analysis, our collection cannot be assigned to a known species. According to morphology, our Psathyrella should
be closer to P. thujina (Section Spadiceogriseae). By contrast, the DNA ITS sequence
would undoubtedly include it in the “candolleana” clade, where each species showed up
to a 99% ITS sequence similarity with our sample. The most widespread and known
species in this clade, P. candolleana and P. leucotephra (Berk. & Broome) P.D. Orton,
both commonly occurring in Europe, too, are however morphologically very different
from our specimen, by forming large pilei (diameter up to 8.0 cm) and lacking pleurocystidia; furthermore, the latter frequently even shows a torn annulus in the upper
part of the stem, which we did not observe in our Psathyrella (Kits van Waveren 1985,
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
97
Table 2. Species used for the phylogenetic analyses including GenBank Accession Numbers and published references.
GenBank accession No.
Reference
Psathyrella abieticola
Species
KC992891
Örstadius et al. 2015
P. almerensis
KC992874
Örstadius et al. 2015
P. almerensis
KC992873
Örstadius et al. 2015
P. ammophila
KC992872
Örstadius et al. 2015
P. candolleana
AB306311
Ogura-Tsujita and Yukawa 2008
P. candolleana
DQ389720
Larsson and Örstadius 2008
P. candolleana
MG734719
Yan and Bau 2018
P. candolleana
MG734720
Yan and Bau 2018
P. cladii-marisci
MK080112
This study
P. conferta
KC992890
Örstadius et al. 2015
P. conica
MG734713
Yan and Bau 2018
P. flexispora
MF966494
Heykoop and Moreno 2002
P. fusca
MF966503
Heykoop and Moreno 2002
P. impexa
KC992900
Örstadius et al. 2015
P. kellermanii
KC992920
Örstadius et al. 2015
P. luteopallida
MG734736
Yan and Bau 2018
P. lutensis
MG734748
Yan and Bau 2018
P. lutensis
DQ389685
Larsson and Örstadius 2008
P. lutulenta
KC992875
Örstadius et al. 2015
P. madida
KC992932
Örstadius et al. 2015
P. parva
KC992912
Örstadius et al. 2015
P. prona
KJ939634
Larsson and Örstadius 2008
P. pseudogracilis
KC992853
Örstadius et al. 2015
P. purpureobadia
NR_119670
Larsson and Örstadius 2008
P. romagnesii
DQ389716
Larsson and Örstadius 2008
P. saponacea
MH155965
Yan and Bau 2018
P. senex
MG734732
Yan and Bau 2018
P. singeri
MG734718
Yan and Bau 2018
P. squamosa
KC992939
Örstadius et al. 2015
P. squamosa
MG367206
Yan and Bau 2018
P. subsingeri
MG734714
Yan and Bau 2018
KJ138423
Battistin et al. 2014
P. sulcatotuberculosa
FJ899635
Frank et al. 2010
P. tenuicula
DQ389706
Larsson and Örstadius 2008
P. thujina
KC992873
Örstadius et al. 2015
P. thujina
KC992874
Örstadius et al. 2015
P. thujina
KY680791
Örstadius et al. 2015
P. thujina
KY680792
Örstadius et al. 2015
P. trinitatensis
KC992882
Örstadius et al. 2015
P. tuberculata
MH497604
Yan and Bau 2018
P. typhae
DQ389721
Larsson and Örstadius 2008
Coprinellus heterothrix
FM878018
Nagy et al. 2011
C. impatiens
FM163177
Nagy et al. 2011
C. silvaticus
KC992943
Örstadius et al. 2015
P. tenera
98
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
Figure 4. One of the most parsimonius trees from the phylogenetic analysis of Psathyrella spp. based on
nrDNA sequence data. Bootstrap values are shown above branches based on 1,000 replicates (values below
50 are not shown).
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
99
Vesterholt and Knudsen 1992, Consiglio 2005). The “candolleana” clade encompasses
two more European species according to two recent phylogenetic analyses (Nagy et al.
2011, Battistin et al. 2014): P. sulcatotuberculosa (J. Favre) Einhell., previously regarded
as a variety of P. typhae (Kits van Waveren 1985), which mainly differs from our Psathyrella and from P. typhae itself with a partially-sulcate and -tuberculate pileus surface,
and P. badiophylla (Romagn.) Bon which forms spores normally exceeding 10–11 µm
in length (Kits van Waveren 1985, Vesterholt and Knudsen 1992); in addition, both
also lack pleurocystidia, which was considered to be such a morphologically relevant
character to induce the establishment of the Section Spintrigerae within the subgenus
Psathyra (Fr.) Sing. ex Kits van Wav. (Kits van Waveren 1985). Moreover, except for P.
typhae, which is the only Psathyrella ecologically comparable to our collection, all the
above species are reported to grow in diverse site conditions, i.e. close to stumps of trees
or on branches, on moist ground, in grass, on mossy woods or on various other vegetable matter (Kits van Waveren 1985, Vesterholt and Knudsen 1992). Finally, as far as
we know, other species in the “candolleana” clade are even geographically more distant,
each colonising a different kind of organic debris (Padamsee et al. 2008, Örstadius et
al. 2015, Yan and Bau 2018).
Therefore, within this framework, the placement of our fungus into the “candolleana” clade, together with other species showing strong differences for geographic and ecologic reasons, should not prevent the recognition of a new Psathyrella species.
Anyhow, more and more scientific contributions are remarking that the genetic analysis of a fungus aiming at taxonomic purposes can alone generate artefacts, i.e. “false positive” or “chimeras”, especially when such analysis is implemented by using a unique gene
(Thines et al. 2018, Lücking et al. 2018). A polyphasic approach, i.e. based on the combination and integration of all the available informative data (Colwell 1970), is becoming
more and more desirable for taxonomic research in mycology, whereas the ITS rDNA
region is still considered as the universal genetic marker for fungi (Schoch et al. 2012).
On the basis of the outcomes deriving from the morphologic, ecologic and biomolecular characteristics which we have identified in this note, we are therefore inclined
to establish a new species of Psathyrella.
Taxonomy
Psathyrella cladii-marisci Sicoli, NG Passal., De Giuseppe, Palermo & Pellegrino,
sp. nov.
Figs 1–3
Etymology. The specific epithet derives from Cladium mariscus, the name of the plant
where it was first detected.
Diagnosis. Similar to P. thujina from which it differs by showing a larger pileus
(about 40% larger), a wider range of spore length, versiform cheilocystidia and basidiomes occurring in spring.
100
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
Holotype. Italy. Calabria, Cosenza, Rende, Orto Botanico Università della
Calabria. 39°21'25.05"N, 16°13'44.57"E, 220 m a.s.l., marsh at the base of cut
culms of a Cladium mariscus (L.) Pohl plant, transplanted from Lago dell’Aquila
(Laureana di Borrello, Reggio Calabria, southern Italy) at the corner of a concrete
tank maintained full of water, 10 April 2018, Antonio Biagio De Giuseppe &
Giovanni Sicoli (CLU F302).
Description. Habit psathyrelloid. Pileus up to 3.5 cm diam., conical-convex
when young, hemispheric to applanate at maturity, with a deeply striate margin,
hazelnut in colour, turning to pale beige when dry. Pileipellis with evident concentric
arachnoid fibrils of velar origin, whitish and easily removable, often exceeding the
cuticle margin. Lamellae distant, ventricose, adnate, intermingled with numerous
lamellulae, initially pale pink, then intensely brown-purplish. Lamella edge whitish
with numerous sphaeropedunculate cells. Stipe, very fragile, cylindrical, white, exannulate with a diffuse fibrillosity especially on the basal surface, apical surface pruinose. Basidiospores 7.2–11.8 × 4.3–6.0 µm (n = 100), ellipsoid to ovoid-ellipsoid, with
a thick and smooth wall, adaxially flattened with a central 2µm-wide germ pore and a
distinct hilar appendix. Spore-print dark brown. Basidia clavate, 4-spored. Cheilocystidia versiform, often utriform, seldom cylindrical to clavate. Pleurocystidia utriformshaped. Mycelium septate and clamped. Context with apparently no smell, taste mild.
Habit, habitat and distribution. In small groups (gregarious), on the culm remnants of Cladium mariscus. So far, known only from the type locality.
Conclusions
This probably rare and, apparently, never before detected species could occur more
commonly if further surveys confirmed a sort of preference for C. mariscus as a growing
substrate for the fungus. This plant was observed all over Italy (Bartolucci et al. 2018),
although becoming more and more scattered due to the progressive surface reduction
of its natural growing environment, i.e. marshes and wet sites quite close to the sea at
mid-low altitudes. These sites have been long subjected to draining and other forms
of anthropogenic land uses. Since human activities have been causing a deep influence and restriction on density and distribution of the spontaneous flora, including C.
mariscus, the gradual depletion of plant biodiversity in such sites could also result in
negative effects on fungal diversity, thus rendering even more scarce the occurrence of
basidiomes of such taxa as P. cladii-marisci in Italy.
Acknowledgements
We are very grateful to Pasquale A. Cicirelli and Nicola Fico for their precious advice
in the digital image processing.
A new species of Psathyrella (Psathyrellaceae, Agaricales) from Italy
101
References
Bartolucci F, Peruzzi L, Galasso G, Albano A, Alessandrini A, et al. (2018) An updated checklist of
the vascular flora native to Italy. Plant Biosystems – An International Journal Dealing with all
Aspects of Plant Biology 152(2): 179–303. https://doi.org/10.1080/11263504.2017.1419996
Battistin E, Chiarello O, Vizzini A, Örstadius L, Larsson E (2014) Morphological characterisation and phylogenetic placement of the very rare species Psathyrella sulcatotuberculosa.
Sydowia 66(2): 171–181. http://hdl.handle.net/2318/152677
Christan J, Hussong A, Dondl M (2017) Beiträge zur Familie Psathyrellaceae: Psathyrella spintrigeroides, Psathyrella supernula, Psathyrella typhae. Mycologia Bavarica 18: 35–58.
Colwell RR (1970) Polyphasic Taxonomy of the Genus Vibrio: Numerical Taxonomy of Vibrio
cholerae, Vibrio parahaemolyticus, and Related Vibrio Species. Journal of Bacteriology
104(1): 410–433. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC248227/
Consiglio G (2005) Contributo alla conoscenza dei Macromiceti dell’Emilia-Romagna. XXIII.
Famiglia Coprinaceae - Parte terza. Bollettino del Gruppo Micologico G. Bresadola – Nuova Serie BGMB 48(2): 7–22.
Doyle JJ, Doyle JL (1990) Isolation of plant DNA from fresh tissue. Focus 12: 13–15.
Felsestein J (1985) Phylogenies and comparative method. The American Naturalist 125(1):
1–15. https://doi.org/10.1086/284325
Frank JL, Coffan RA, Southworth D (2010) Aquatic gilled mushrooms: Psathyrella fruiting in the
Rogue River in southern Oregon. Mycologia 102: 93–10. https://doi.org/10.3852/07-190
Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium, Series 41: 95–98.
Henrici A (2017) Psathyrella: the state of play – including P. thujina new to Britain. Field Mycology 18(3): 87–91. https://doi.org/10.1016/j.fl dmyc.2017.07.007
Heykoop M, Moreno G (2002) Studies in the genus Psathyrella in Spain. IV. Psathyrella submicrospora sp. nov. and P. microsporoides nom. nov. Mycotaxon 83: 425–433.
Katoh K, Rozewicki J, Yamada KD (2017) MAFFT online service: multiple sequence alignment,
interactive sequence choice and visualization. Briefings in Bioinformatics: 1–7. https://doi.
org/10.1093/bib/bbx108
Kits van Waveren E (1985) The Dutch, French and British species of Psathyrella. Persoonia,
Suppl. Vol. 2: 1–300.
Larsson E, Örstadius L (2008) Fourteen coprophilous species of Psathyrella identified in the
Nordic countries using morphology and nuclear rDNA sequence data. Mycological Research 112: 1165–1185. https://doi.org/10.1016/j.mycres.2008.04.003
Lücking R, Kirk PM, Hawksworth DL (2018) Sequence-based nomenclature: a reply to Thines
et al. and Zamora et al. and provisions for an amended proposal “from the floor” to allow
DNA sequences as types of names. IMA Fungus 9(1): 185–198. https://doi.org/10.5598/
imafungus.2018.09.01.12
Nagy LG, Walther G, Házi J, Vágvölgyi C, Papp T (2011) Understanding the evolutionary
processes of fungal fruiting bodies: correlated evolution and divergence times in the Psathyrellaceae. Systematic Biology 60: 303–317. https://doi.org/10.1093/sysbio/syr005
102
Giovanni Sicoli et al. / MycoKeys 52: 89–102 (2019)
Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2014) IQ-TREE: a fast and effective
stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology
and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300
Ogura-Tsujita Y, Yukawa T (2008) High mycorrhizal specificity in a widespread mycoheterotrophic plant, Eulophia zollingeri (Orchidaceae). American Journal of Botany 95: 93–97.
https://doi.org/10.3732/ajb.95.1.93
Örstadius, L, Ryberg M, Larsson E (2015) Molecular phylogenetics and taxonomy in Psathyrellaceae (Agaricales) with focus on psathyrelloid species: introduction of three new genera and
18 new species. Mycological Progress 14:25. https://doi.org/10.1007/s11557-015-1047-x
Padamsee M, Matheny B, Dentinger BTM, McLaughlin DJ (2008) The mushroom family
Psathyrellaceae: Evidence for large-scale polyphyly of the genus Psathyrella. Molecular Phylogenetics and Evolution 46: 415–429. https://doi.org/10.1016/j.ympev.2007.11.004
Rambaut A (2009) FigTree version 1.3.1 [computer program] http://tree.bio.ed.ac.uk
Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA, Chen W, and Fungal
Barcoding Consortium (2012) Nuclear ribosomal internal transcribed spacer (ITS) region
as a universal DNA barcode marker for Fungi. PNAS 109(16): 6241–6246. https://doi.
org/10.1073/pnas.1117018109
Thines M, Crous PW, Aime MC, Aoki T, Cai L, Hyde KD, Miller AN, Zhang N, Stadler M
(2018) Ten reasons why a sequence-based nomenclature is not useful for fungi anytime
soon. IMA Fungus 9(1): 177–183. https://doi.org/10.5598/imafungus.2018.09.01.11
Vesterholt J, Knudsen H (1992) Psathyrella (Fr.) Quél. In: Nordic Macromycetes (Vol. 2),
Hansen N and Knudsen H (Eds) Nordsvamp, Copenhagen, 236–252.
Voto P (2016) Rare Agaricales in Polesine I: Psathyrella, Conocybe, Lepista. Rivista di Micologia
59(2): 163–174.
Yan JQ, Bau T (2018) The Northeast Chinese species of Psathyrella (Agaricales Psathyrellaceae).
MycoKeys 33: 85–102. https://mycokeys.pensoft.net/article/24704/
White TJ, Bruns T, Lee S, Taylor JW (1990) Amplification and direct sequencing of fungal
ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ
(Eds) PCR protocols: a guide to methods and applications. Academic Press Inc., New York,
315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1