Molecular phylogenetics and taxonomy in Psathyrellaceae (Agaricales) with focus on psathyrelloid species: introduction of three new genera and 18 new species

Abstract

Based on traditional morphology, sequence data, and phylogenetic analyses, 18 new species are here described: Coprinellus christianopolitanus, Coprinopsis musae, C. udicola, Psathyrella arenosa, P. carminei, P. fennoscandica, P. ichnusae, P. lilliputana, P. lyckebodensis, P. madida, P. rybergii, P. sabuletorum, P. scanica, P. siccophila, P. stridvallii, P. sublatispora, P. vesterholtii, and Typhrasa nanispora. Psathyrella hololanigera and P. tenera are reported as new to Europe and P. parva as new to the Nordic countries. A four-gene dataset on Psathyrellaceae were analyzed by Maximum Parsimony, Maximum Likelihood, and Bayesian methods. Constraint analyses were performed to determine limits of /Psathyrella, and to evaluate whether the /Coprinellus, /cordisporus, and /gossypina clades could be regarded with confidence as monophyletic clades outside of the clade /Psathyrella. This was not unambiguously supported. Based on the phylogenetic results, Kauffmania is proposed as a monotypic genus for the species P. larga and Typhrasa for P. gossypina and the new described species T. nanispora. The genus Homophron is formally validated and three combinations are proposed: H. spadiceum, H. cernuum, and H. camptopodum. The genus Cystoagaricus Singer is emended and the following new combinations are proposed: C. hirtosquamulosus, C. squarrosiceps, C. olivaceogriseus, and C. silvestris. Neotypes have been selected for seven species described by Fries, and ITS sequence data for these were generated. The following new combinations are proposed: Coprinopsis canoceps, C. cineraria, C. melanthina, C. submicrospora, C. uliginicola, and Typhrasa gossypina. Brief comments are given to other species of interest. Psathyrella ornatispora were found not to belong in Psathyrellaceae. A key to 106 psathyrelloid species in Northern Europe is provided.

Introduction

Psathyrella is a large genus in Agaricales with 1,004 records in Index Fungorum (www.indexfungorum.org) including synonyms, varieties, forms, and names currently linked to genera such as Lacrymaria or Coprinopsis. The number of recognized species in Europe is about 100 and in the Nordic countries 78 (Örstadius and Knudsen 2012). The species in Psathyrella are known to have a wide geographic distribution (e.g., Smith 1972; Pegler 1977, 1983; Kits van Waveren 1985; Singer 1969, 1978; Grgurinovic 1997; Bi et al. 1987; Hoashi 2008). As presently understood, the genus includes species with mostly fragile and non-deliquescent basidiomata. The cap size varies from small to large, is smooth to innately fibrillose, often hygrophanous, mostly with white to dark reddish brown colours, and a copious to scanty veil. The pileipellis is considered a hymeniderm. Clamps are present or absent. The size of the spores varies and hymenial cystidia are present. The species are mostly regarded as saprotrophic and commonly growing on wood, soil, dung, plant residuals, and in swamps, but other nutritional modes has been reported such as the association with mycoheterotrophic orchids (Selosse et al. 2010) and the mycoparasitism in P. epimyces (Smith 1972).

The major taxonomic treatments of the genus Psathyrella by Smith (1972), Romagnesi (1982), Kits van Waveren (1985), and Singer (1986) were mainly based on phenotypic characters although sometimes complemented by mating tests using haploid mycelia (Galland 1972; Jurand and Kemp 1972). The infrageneric classification differs among authors. Smith, Romagnesi, and Singer include Lacrymaria as a subgenus while Kits van Waveren treats it as a genus of its own. According to Singer (1986) the family Coprinaceae comprises subfamily Coprinoideae, with the single genus Coprinus, subfamily Psathyrelloideae with the genera Psathyrella and Macrometrula, and subfamily Panaeoloideae including the genera Panaeolina, Panaeolus, Copelandia, and Anellaria.

In recent years, molecular sequence data have improved our understanding of relationships among dark spored agarics (Hopple and Vilgalys 1999; Moncalvo et al. 2002; Matheny et al. 2006). As a result, major changes to the genus classification have been suggested (Redhead et al. 2001a, b; Gams 2002). Extended specimen sampling of the major lineages within Psathyrellaceae and analyses based on sequence data of the LSU region (Vašutová et al. 2008; Padamsee et al. 2008) or a combination of the ITS and LSU regions (Larsson and Örstadius 2008) showed that the limits of Psathyrella are unclear. These studies suffered from poor resolution of deeper nodes but they all recovered Coprinellus and Coprinus cordisporus nested with Psathyrella in a large supported clade.

Recently, several studies using newly developed comparative phylogenetic methods based on alignments of multi-gene datasets have been published, aiming to explore evolutionary aspects of the development of characters within Psathyrellaceae. These studies have shed light upon the evolution of autodigestion, species evolution, and divergence times, but also on the delimitation of taxa (Nagy et al. 2009, 2010, 2011). Several new species have been described, both within Coprinellus and Coprinopsis (Nagy et al. 2012, 2013a).

Nagy et al. (2013b) presented phylogenetic analyses of a four-gene dataset sampled with the intention to cover all major clades within Psathyrellaceae. In the paper, 14 clades are recognized, not all of them well supported, and the authors suggest that these clades form the basis for a splitting and reclassification of Psathyrella.

Because of the phylogenetic results, several species originally described in Psathyrella have been transferred to other genera such as Parasola and Coprinopsis (e.g. Larsson and Örstadius 2008; this paper). Despite their phylogenetic placement they still have the apperance and habit of a Psathyrella species, and we will refer to such species collectively as psathyrelloid.

The study by Larsson and Örstadius (2008) had a focus on the coprophilous species in Psathyrella. In this study, we have expanded the specimen sampling of Psathyrella further, to include the majority of species that occur in Europe in an effort to identify and characterize the species that occur in Northern Europe. To identify and get support for the description of new species, a combination of classical morphology and ITS sequence data were used. To confirm or discriminate between closely related and morphologically similar species described from Europe and other parts of the world, a large number of type specimens were studied and sequenced. For some species described by Fries, neotypes are selected.

A four-gene dataset was generated in order to inferring the phylogenetic relationships among species and genera within Psathyrellaceae. In addition to the ITS and LSU regions, sequence data of two protein-coding genes (β-tubulin and Tef-1α) were selected. These are the same regions as used by Nagy et al. (2013b). Based on the phylogenetic results, we propose several taxonomical novelties and nomenclatural changes.

Materials and methods

Morphology

Many species of Psathyrella are fragile and important characters are easily destroyed during handling. Therefore, all basidiomata were photographed before being collected. The aim was to describe the collection complete with notes on ecology already in the field. The presence of a pseudorhiza was checked. As most species are hygrophanous, it is necessary to note the colours of moist and striate caps before they are dried. Colour names follow the Munsell soil colour charts (Munsell 1975), cited as Mu. in the text. If present, the evanescent veil on both cap and stem was described. Sometimes the veil is present only in young basidiomata and seen as fibrils close to the cap margin or as dispersed fibrils on the stem surface. When well-developed, the veil can be observed as flocci, scales, or patches on the cap surface. In most species, the stem is pulverulent or pruinose at the apex with more or less evident veil remnants below. The upper part of the veil sometimes leaves an annulus or fugacious ring-zone that must be noted. This zone is rarely present at the lower part of the stem. If possible, all stages of basidiomata development were collected in order to cover changes in cap colours and veil features. Moreover, the cap and stem surface was examined for projecting hairs that are present in a few species (e.g., P. tenuicula). Before drying the material spore prints were taken and any green reaction of the gill edge in 10 % NH₄OH was observed in a microscope.

Micromorphological characters were observed using a Nikon Eclipse E200 light microscope equipped with phase contrast. Digital images were recorded with a Nikon Infinity 2 camera. For each collection, 10 to 20 mature spores were measured in water at ×1,000 magnification. Abnormally large or small spores were not considered. Other microscopic characters were studied in a 10 % NH₄OH solution and measured to nearest micron. To observe the hymenial cystidia, a complete lamella was cut off with a razor blade and soaked for a while. The gill edge was removed in order to check the cheilocystidia. The middle portion of the gill was cut out, crushmounted, and pleurocystidia, basidia, subhymenium, and hymenophoral trama studied. The layers of the pileus were observed halfway from the margin by cutting tangential to the pileus, called a ‘scalp’. The cells of the pileipellis and the hyphae of the pileitrama can then be observed. If the material admitted, a radial cut to the pileus was done instead, resulting in a better picture of the different layers. In addition, if a veil was present above the pileipellis it could be more satisfactory located. Finally, the veil tissue from cap margin and the presence of clamps were checked. As for the shape of spores and cystidia the terminology of Vellinga (1988) was followed. Spores were mounted in a solution of ammonia while cystidia and other cells were mounted in a solution of ammonia stained with Congo Red before capturing digital images. To get a better resolution, most microscopical features except spores were then uniformly coloured with the program Adobe Photoshop CS2. All scale bars in the figures represent 10 μm. Line drawings of the micro-morphology are presented in association with the species descriptions.

Collections are deposited in Herbarium GB, University of Gothenburg, if not otherwise indicated. Types and other collections were studied as loans from AH, AMNH, B, BAFC, BR, C, CBM, CUP, E, F, G, GB, GDGM, GH, H, K, L, LD, MICH, NY, NYS, O, PC, S, SGO, SZE, TAA, TROM, TURA, UPS, WAG-W, WBS, WU, XAL, and ZT. Material was also received as gifts or loans from several private herbaria. Data on type specimens and other specimens morphologically studied, but not sequenced are provided as Supplementary data S1.

Specimen sampling

Our sampling emphasized Psathyrella in Europe and its northern part, aiming to find and include representatives from all known or putative taxa of Psathyrella. To be more representative, the specimen sampling was extended to outside Europe, including also tropical regions and the Southern Hemisphere. Also, other genera of the Psathyrellaceae were to some extent taken into account.

The ITS and LSU sequence dataset from our study on coprophilous Psathyrella species (Larsson and Örstadius 2008) was extended and complemented. Sequences in this study were generated from 182 specimens, representing 132 species of Psathyrella sensu lato and, in total, 148 species in Psathyrellaceae, and includes 55 type specimens (Table 1). In addition, sequence data (ITS, LSU, β-tub, and Tef-1α) representing six Psathyrella, 11 Coprinellus, two Coprinus cordisporus, one Cystoagaricus, seven Coprinopsis, and four Parasola species were retrieved from GenBank and added to the dataset. The selection was based on previous molecular studies of Agaricales and Psathyrellaceae (Vellinga 2004; Padamsee et al. 2008; Vašutová et al. 2008; Nagy et al. 2011, 2012, 2013b). Based on results from earlier molecular phylogenetic studies of Agaricales (Moncalvo et al. 2002; Matheny et al. 2006), representatives of Bolbitius, Mythicomyces, Stagnicola, Agrocybe, and Conocybe were selected as the outgroup in the analyses.

Table 1 Data of specimens used in the phylogenetic analysis

For the neotypification of species originally described by Fries from Sweden, representative collections were selected and the ITS generated, if not already included in the large dataset.

Molecular work

Sequences from four regions were generated for the study: the complete ITS region and about 1,000 base pairs (bp) of the 5′ end of the LSU nuclear ribosomal DNA, about 1,000 bp of translation elongation factor subunit 1 alpha (Tef-1α), and about 500 bp of the β-tubulin gene. DNA extractions, PCR reactions, and sequencing were performed as described in Larsson and Örstadius (2008). Primers used to amplify the complete ITS region and the 5′ end of the LSU region were ITS1F (Gardes and Bruns 1993) and LR21, LR0R, and LR7 (Hopple and Vilgalys 1999); for Tef-1α we used EF983F and EF2218R (www.aftol.org/pdfs/EF1primer); for β-tub B36f and B12r (Nagy et al. 2011). Primers used for sequencing were ITS1, ITS4 (White et al. 1990), Ctb6 (http://plantbio.berkeley.edu/~bruns/) and Lr5 (Hopple and Vilgalys 1999), EF983F, EF2218R, and 1567Ra (www.aftol.org/pdfs/EF1primers), B36f and B12r. DNA extraction, PCR, and sequencing of old type specimens follow Larsson and Jacobsson (2004).

Phylogenetic analyses

Sequences were edited and assembled using Sequencher 4.1 (Gene Codes, Ann Arbor, MI, USA). Alignment of individual genes was performed using the L-INS-i strategy as implemented in MAFFT v. 7.017 (Katoh and Standley 2013). The alignment was adjusted manually using the data editor in PAUP* 4.0b12 (Swofford 2003). Sequences have been deposited in GenBank and accession numbers are listed in Table 1.

Maximum likelihood (ML) analysis was performed using RAxML 7.0.4 (Stamatakis 2006). It was tested for conflict between the genes by comparing supported clades. Support values for each gene were estimated by 1,000 rapid bootstraps and conflicts in clades with higher support than 70 was considered significant. Four different partitioning schemes were compared: all genes in one partition, each gene in separate partitions (three partitions), each gene in separate partitions and third position partitioned separately in the protein coding genes (five partitions), and each gene partitioned separately and each codon position partitioned separately (seven partitions). It was also compared if GTR plus gamma or GTR plus gamma and invariable sites was the best fit. All model comparisons were done with AIC. Support values for the catenated genes under the best fitting model were calculated from 10,000 rapid bootstraps. Seven specific constraints were implemented to test if the following could be rejected: 1) Psathyrella s.s. is monophyletic excluding /cordisporus, 2) Psathyrella s.s., /Coprinellus, and /candolleana are monophyletic excluding /cordisporus, 3) Psathyrella s.l. is monophyletic excluding /cordisporus, 4) Psathyrella s.s. including /gossypina is monophyletic, 5) Psathyrella s.s. including the /candolleana clade is monophyletic, 6) Psathyrella s.l. is monophyletic excluding /Coprinellus, 7) /Parasola is nested in Psathyrella s.l. with at least the /gossypina clade more basal (i.e., Psathyrella s.l. is not monophyletic). The constraint topologies were compared to the ML tree using the SH-test as implemented in RAxML.

Bayesian analysis was done in MrBayes 3.1.2 using the partitioning scheme and model selected in the ML analysis. Four parallel runs with four chains each (three heated) were executed. Each run had 200 million generations, sampling every 10,000th generation. Burn-in was determined using TRACER (Rambaut and Drummond 2007) and AWTY (Nylander et al. 2008). Constraints one to six above were compared using posterior odds calculated from the Bayesian MCMC run. Instead of constraint seven it was tested directly if Psathyrella sensu lato is monophyletic. Posterior odds are conservative compared to Bayes factors when assuming equal prior on all topologies (Bergsten et al. 2013).

Phylogenetic analyses were based on the concatenated gene alignments. Heuristic searches for the most parsimonious trees were performed using PAUP* (Swofford 2003). All transformations were considered unordered and equally weighted. Variable regions with ambiguous alignment were excluded and gaps were treated as missing data. Heuristic searches with 1,000 random-addition sequence replicates and TBR branch swapping were performed, saving at most 25 trees in each replicate. Relative robustness of clades was assessed by the bootstrap method using 1,000 heuristic search replicates with 100 random taxon addition sequence replicates and TBR branch swapping, the latter saving at most 25 trees in each replicate.

Results

Phylogenetic results

The aligned complete dataset, including 31 sequences downloaded from GenBank, consisted of 218 taxa and 3,604 characters. After exclusion of ambiguous regions from the ITS region, and from intron regions of the Tef-1α and β-tub genes, 2,566 characters remained for the analysis. Of these, 1,381 were constant, 263 were variable but parsimony uninformative, and 922 (36 %) were parsimony informative.

The maximum parsimony analysis yielded 300 equally parsimonious trees (length = 8,158 steps, CI = 0.2392, and RI = 0.6112). One of the trees is presented as a phylogram in Fig. 1a and b. The bootstrap analysis recovered Psathyrellaceae as monophyletic including six major supported clades, /Psathyrella s. l. (including /Coprinellus and /cordisporus, 89 %), /gossypina (90 %), /Lacrymaria (100 %), /Homophron (100 %), /Coprinopsis (82 %), and /Parasola (100 %). Within /Psathyrella s.l., /Psathyrella s.s. received no support, but 19 major and minor clades within /Psathyrella s.l. were weakly to strongly supported; these are marked as subclades in the phylogram in Fig. 1a and b. Some of the subclades more or less correspond to morphological species groups and taxa, and these are further described and discussed below.

Fig. 1

a and b One of the most parsimonius trees from the phylogenetic analysis based on sequences of a concatenated data set from four nuclear genes (ITS, LSU, β-tubulin and Tef-1α). Bootstrap, Bayes, and ML values are indicated for the major clades, thick lines indicate support of at least 50 %, 0.95, 50 %, respectively. Major and minor clades discussed in the text are indicated with scale bars and named

Fig. 2

Photos of six of the new described species, a Psathyrella madida, b P. stridvallii, c P. carminei, d Coprinopsis musae, e P. fennoscandica, f P. vesterholtii. Photos of all new species are provided in Supplementary data S3

Fig. 3

Photos of species in the new described genera. a Typhrasa gossypina TL2011-389713 (C) Photo T. Laessoe. b Cystoagaricus silvestris Sweden Skåne. Benestad, Örups almskog, 18.9.2008, I. Månsson & L. Örstadius 163–08. c Homophron spadiceum Sweden, Skåne, Kristianstad, Sa Lingenäset, 11.9.2003, L. Örstadius 75–03. d Homophron spadiceum Sweden. Skåne, Kristianstad, Sa Lingenäset, 11.9.2003, L. Örstadius 75–03. e Kauffmania larga Sweden Västergötland, Habo, N. of Sjöbol, 22.10.1999, E. Grundel, LÖ197-99

No conflict was found between genes. The model with seven partitions and GTR plus gamma and invariable sites was supported as the best by AIC. None of the constraints were significantly worse using the SH-test so in the ML framework we cannot reject that the /cordisporus and /Coprinellus clades, respectively, are outside Psathyrella s.s. or that the /gossypina and /candolleana clades, respectively, are part of /Psathyrella s.s. Neither can we reject that /Psathyrella s.l. is paraphyletic with respect to /Parasola. The ML bootstrap support values are shown in Fig. 1a and b.

The burn-in for the Bayesian analysis was set to 100 million generations. The monophyly of /Psathyrella s.l. is supported (posterior odds 32.3), but none of the other constraints received support (posterior probability to low to be measured). In a Bayesian context it was thus rejected that the /cordisporus and /Coprinellus clades are outside Psathyrella s.l., and that the /candolleana or /gossypina clades are within /Psathyrella s.s. In the Bayesian analysis, Psathyrella s.s. received a BPP value of 0.98. Bayesian supports are shown in Fig. 1a and b.

From our sampling and phylogenetic analysis we recognize 116 species as belonging to Psathyrella, and of these, 15 are new and described below. In addition, we also identified and described one species of Coprinellus and two of Coprinopsis as new to science.

Psathyrella ornatispora does not belong to Psathyrellaceae and should be moved to another genus in Agaricales. Best match in GenBank when blasting the ITS region is with Melanophyllum haematospermum, but with 93 % identity only. Ninety-two species were identified to occur in Northern Europe.

Discussion

In this study we find support for recognizing the clades, /Parasola, /Coprinopsis, /Homophron, /Lacrymaria, and /gossypina, as distinct lineages within Psathyrellaceae. /Coprinellus and /cordisporus fall within the larger supported clade we here named /Psathyrella s.l. The molecular support for the clade /Psathyrella s.s., that includes the majority of species described in Psathyrella, is low but with support from the morphology we suggest recognizing the clade as distinct.

The results are very much in congruence with our previous study (Larsson and Örstadius 2008) based on ITS and LSU sequence data alone, but with the major difference that the support values for the majority of clades are higher.

Several new species, with a focus on psathyrelloid taxa, were added to the data set and particular efforts were made to find representatives for the minor, less species rich clades. We believe that it has strengthened the results and our aim to evaluate morphological characters.

The phylogeny is based on the same regions of sequence data as used in Nagy et al. (2013b), but analysed in a more traditional way. To split Psathyrella into several smaller genera as proposed by Nagy et al. (2013b) was not supported by our analyses, and can neither be supported nor legitimized by finding distinct and segregating morphological characters that unambiguously circumscribe the supported clades. The situation with the paraphyletic Psathyrella is not solved, but we believe that a more conservative approach is to prefer until stronger evidence for an alternative solution is available.

Morphological characters

A morphological characterization of the clades is difficult. For several of the already formally described and circumscribed genera within Psathyrellaceae, the phylogenetic analyses now suggest many of the corresponding clades to include a morphologically heterogeneous assemblage of species. In Table 2 a summary of discriminating characters for each major clade recognized in this study is presented with some further remarks here. Terminology of the pileipellis follows Clémençon et al. (2004). Basidia: psathyrelloid species are ususally circumscribed as having monomorphic basidia, and such species are now also shown to occur in Coprinopsis. However, these species should be examined closer to see if some of the specimens may have both mono- and dimorphic basidia. Pseudoparaphyses: in coprinoid species basidia are surrounded by pseudoparaphyses (brachybasidioles) in a typical pattern (Uljé 2005, 59). Only rarely we find this pattern in psathyrelloid species. The psathyrelloid species Coprinopsis marcescibilis and C. pannucioides have been shown to belong in the genus Coprinopsis (Larsson and Örstadius 2008). In the present study, additional psathyrelloid species are combined or described as new in Coprinopsis. Coprinopsis canoceps, C. melanthina, C. submicrospora, and C. udicola that lack the mentioned pattern of pseudoparaphyses, while in C. musae and C. uliginicola it is present, however, somewhat diffuse in the latter. Pileipellis: the psathyrelloid species Coprinopsis cineraria, C. marcescibilis, C. melanthina, C. pannucioides, C. submicrospora, and C. udicola all have a cutis, hyphae of often inflated cells, that also occur in other Coprinopsis species. Coprinopsis canoceps has a cutis with transition to a paraderm, while the pileipellis of C. musae and C. uliginicola is difficult to assess. Psathyrella species have a hymeniderm, about 30 of them with a transition to a paraderm. By Psathyrella gordonii and P. epimyces the pileipellis is similar to a cutis.

Table 2 A summary of morphological characters used to circumscribe and discriminate the nine genera and C. cordisporus within Psathyrellaceae, recognized in this study

Psathyrella

In this study we focus on the genus Psathyrella and species with psathyrelloid characteristics. We describe 15 new species known from Europe, but many of them may have a wider distribution range. Most of the species are placed within Psathyrella s.s., but Typhrasa nanispora occurs as a sister species to P. gossypina, in the /gossypina clade, and the two species is suggested to be placed in the new describe genus Typhrasa, see taxonomy section.

In total, our dataset has ITS sequence data of 49 type specimens of species described in Psathyrella, including the newly described species. Four of the sequences were recovered in other clades, and these species are here transferred to corresponding genera within Psathyrellaceae; see taxonomy section. The sequence of the type of P. ornatispora fell outside of Psathyrellaceae and the species should be transferred to another family and genus in Agaricales. The species has a deviating morphology with a granular mealy-scaly or furfuraceous cap surface, crowded free gills, very small spores, and lacks hymenial cystidia.

On a proposal from Redhead et al. (2001b) the Committee for Fungi (Gams 2002) recommended conserving the name Psathyrella (Fr.) Quél. with Agaricus gracilis Fr. as type. However, P. gracilis is a younger synonym of P. corrugis [Agaricus corrugis Pers., Neues Mag Bot 1:104, 1794; Agaricus corrugis Pers.: Fr., Syst Mycol 1:298, 1821; Psathyrella corrugis Pers. Konrad and Maubl., Encyclop Mycol 14:123, 1949; = Agaricus gracilis Fr., Syst mycol 1:299, 1821 (non-Agaricus gracilis Pers., Syn meth Fung:425, 1801); Psathyrella gracilis (Fr.) Quél., Mém Soc Émul Montbéliard, Sér. II, 5:152, 1872.]. In this study, the name P. corrugis is, therefore, used and is also the traditional way of naming the species in Europe.

The ITS sequence data of some type specimens indicate the occurrence of synonyms. The ITS of P. corrugis and the type specimen of P. amarescens are identical. According to Arnolds (2003) P. amarescens should differ from P. corrugis in having a red-brown to purple-brown cap, a bitter to acrid taste, and a gill edge with dark dots. We found that the species is very variable in morphology, e.g., in cap size and colour, and in the shape of the cystidia.

Psathyrella vinosofulva and P. riparia also have identical ITS sequences. Apart from slightly larger spores by the former they also agree morphologically. In comparison with P. purpureobadia, the two differ in two base pairs and one deletion event in the ITS1 region. We examined 25 collections of P. purpureobadia, all growing on dung. The type collection grew “in extensively grazed grassland” (Arnolds 2003). Orton (1960) reported P. vinosofulva to grow on soil in ash wood while Smith (1972) stated “in sandy soil along a stream” for his P. riparia. We consider P. purpureobadia to be an independent species, and P. riparia a later synonym of P. vinosofulva.

ITS sequence data of the type specimens of P. lutulenta and P. thujina and two additional specimens determined as P. thujina (LÖ379-06 and LÖ31-04) were included. The four collections are similar in morphology except that clamps were not found in P. thujina LÖ31-04. A few nucleotide differences were found in the ITS region between P. thujina LÖ379-06 and the other three. All together, 20 collections of P. thujina and the type specimen of P. almerensis (L) were examined by morphology alone. They were all morphologically similar except for that some of them lacked clamps. Leiden (L) generally do not allow sequencing of type collections. However, we consider both P. almerensis and P. lutulenta to be later synonyms of P. thujina.

The ITS sequences of the type of P. magnispora and the Austrian collection named P. magnispora differ in two base pairs in the ITS1 region. We only succeeded in generating the ITS2 region for the type specimen of P. mesobromionis, but the region is identical for all three sequences. Morphologically, the Austrian collection deviates by the dominance of 2-spored basidia. We only recognize one species and the oldest name is P. magnispora.

A key to the 106 psathyrelloid species that we recognize in Northern Europe, amended from the key in Funga Nordica (Örstadius and Knudsen 2012), including most of the herein newly described species was constructed and is provided as supplementary data S2. Besides the presence of hymenial cystidia, the size, shape, and pigmentation of the spores are important characters for species identification. Data on the spore morphology is provided with the Supplementary data S2.

Coprinellus

In most phylogenetic studies of Psathyrellaceae, the species referred to Coprinellus form a strongly supported clade (Larsson and Örstadius 2008; Padamsee et al. 2008; Nagy et al. 2010, 2013b). However, often, as in this study, /Coprinellus is recovered within a larger paraphyletic clade /Psathyrella s.l., Fig. 1b. We recognize Coprinellus as a distinct genus and we here describe the new species C. christianopolitanus.

/cordisporus

Coprinus cordisporus, a species with a coprinoid aspect, has taken various positions in the phylogenetic trees presented through the years (Hopple and Vilgalys 1999; Moncalvo et al. 2002; Larsson and Örstadius 2008; Vašutová et al. 2008; Padamsee et al. 2008; Nagy et al. 2010). Its rectangular to pentagonal spores can be compared to the rectangular to triangular spores in some species from the /gossypina clade. The powdery veil found in C. cordisporus is made up of subglobose to ellipsoid cells that also are found in species in sect. Cystopsathyra. Most interesting are the lageniform shape of the cheilocystidia in C. cordisporus. Similar type of cystidia can be found in many species in both Psathyrella and Coprinellus, but they are not present in other genera of Psathyrellaceae in Europe, except for the psathyrelloid species Coprinopsis canoceps. Pseudoparaphyses are present in C. cordisporus, but rarely found in Psathyrella. The pileipellis is a cutis. This unique set of morphological characters may explain the uncertain position on a single branch that the species gets in the phylogenies.

Based on comprehensive phylogenetic and morphologic studies of the C. cordisporus/C. cardiasporus complex Keirle et al. (2004) demonstrated C. cardiasporus to be a later synonym of C. cordisporus. They stated that C. cordisporus represents a complex of closely allied taxa. Based on ITS data the C. cordisporus complex was placed close to Coprinellus as in the present study. Uljé (2005) recognized four species based on morphology. Our own observations is that C. cordisporus is a morphologically variable species. The pileipellis of the C. cordisporus taxa has a cutis (Keirle et al. 2004) while the species in Coprinellus has a hymeniderm.

/gossypina

The clade is divided into three supported subclades that correspond to three separate genera. The subclade including Cystoagaricus strobilomyces, Psathyrella hirtosquamulosa, P. squarrosiceps, P. olivaceogrisea, and P. silvestris is characterized by a unique combination of characters as a pigmented layer of scales, flocci, or bundles of fibrils on cap and stem and mitriform or angular spores. We propose the combination of the four Psathyrella species to Cystoagaricus.

Psathyrella larga is an ordinary looking Psathyrella lacking outstanding features and is difficult to separate from for example P. rostellata. The new genus Kauffmania is here described to reflect better the species phylogenetic position.

The subclade including Psathyrella gossypina and T. nanispora is recognized and characterized by the rostrate hymenial cystidia with oily drops. The non-European species P. canadensis, P. delineata, and P. fraxinophila are also reported to have oily drops in their cystidia (Smith 1972). The collection A.H. Smith 29994 of P. canadensis (MICH) and the holotype of Hypholoma delineatum (NYS) were examined. The two species agree satisfactory in morphology with P. gossypina. The genus Typhrasa is here described for this group of species, see taxonomy section below.

Lacrymaria

/Lacrymaria is a strongly supported clade that include species with almost identical ITS and LSU sequences. The sequence variation between the species seems to be higher in the β-tubulin gene that may be more suitable for disentangle and segregate between the species. In morphology the genus Lacrymaria is fairly easy to identify, with species having tomentose, brown to reddish-brown caps, mottled gills, verrucose spores and pleurocystidia in small fascicles, often capitate.

Homophron

Phylogenetic analyses based on molecular data have shown that species that share the unique combination of morphological characters, as lacking a veil and having cystidia with crystals or incrustations, form a strongly supported clade (Larsson and Örstadius 2008; Padamsee et al. 2008; Vašutová et al. 2008; Nagy et al. 2012). Britzelmayr (1883) published the new name Homophron as a subgenus under Agaricus, characterized by”Velum fehlt. Stiel straff oder steif. Sporen braun.” Homophron contained ten species among others Agaricus spadiceus. Kits van Waveren (1985:155) considered the name Homophron illegitimate referring to that Britzelmayr (1883:174) did not indicate the rank, but on page 181 the rank subgenus is clearly given. Singer (1951) combined Homophron as a subgenus under Psathyrella and selected “P. spadicea (Schaeff. ex Fr.) Sing.” as type. Cooke (1953) raised Homophron to the rank of genus and selected Agaricus particularis Britzelm. as type, one of ten species mentioned in the original description of Homophron (Britzelmayr 1883). However, as no basionym was cited the combination to genus was not validly published (Donk 1962; Horak 1968). Therefore, we here propose to raise subgenus Homophron to genus level to validate the name and genus Homophron for this group of species and propose the combination of the three species, H. spadiceum, H. cernuum, and H. camptopodum.

Coprinopsis

In our analyses the sequence data of the type specimens of Psathyrella submicrospora, P. cineraria and P. uliginicola came out in the /Coprinopsis clade together with sequences of P. canoceps and P. melanthina. The species are psathyrelloid in appearance and was therefore described in Psathyrella. As mentioned above some of them lack the typical pattern of pseudoparaphyses that is thought to be a morphological characteristic for the genus Coprinopsis.

We here propose the combination of the five species to Coprinopsis based on the molecular results and in addition we describe the two new species, C. musae and C. udicola (Fig. 2).

Nomenclatural Novelties

Coprinellus christianopolitanus Örstadius & E. Larss., sp. nov. [MB 811478] – Holotype: Sweden, Skåne, Kristianstad, Tivolipark, 10.IX.2008, L. Örstadius 141–08 (GB). Fig. 4. (S3: Fig. 3a–d.)

Fig. 4

Coprinellus christianopolitanus. L. Örstadius 141–08; Ch Cheilocystidia; Pi Pileocystidia; Basidioma ×3; Scale bars 10 μm

Etymology. The epithet refers to the Latin name of Kristianstad where the fungus was found.

Pileus primo semiglobatus, 3 mm latus, 3 mm altus, flavo-brunneus, demum expansus, 5 mm latus, ochraceus, pubescens, velum non visum. Lamellae distantes, adnexae, acie alba fimbriata. Stipes 6 × 0.5 mm, albus, pubescens. Odor nullus. Sporae 13–16.5 × 8–9 μm, ellipsoideae, oblongae, poro germinativo excentrico, inconspicuo. Basidia 4-sporigera. Cheilocystidia 32–52 × 9–25 μm, anguste lageniformia, subcapitata vel clavata, dispersa, cellulis clavatis immixta, in NH₄OH viridiscentia. Pleurocystidia nulla. Pileocystidia 27–62 × 9–14 μm. Fibulae adsunt. Ad terram in horti.

Cap when young 3 mm high and 3 mm broad, semiglobate, yellow brown, when mature expanding and 5 mm broad, ochraceous, paler towards margin, pubescent sub lente, sulcate, veil not seen. Gills adnexed, distant, L = c. 13, grey with white fimbriate edge, becoming grey black. Stem 6 × 0.5 mm, white, pubescent. Smell none. Taste not recorded.

Spores 13–16.5 × 8–9 μm, av. 14.7 × 8.4 μm, Qav. = 1.8, ellipsoid, oblong, sometimes slightly irregular or papillate, in water dark reddish brown (Mu. 2.5YR 2.5/4), with indistinct and eccentric germ pore. Basidia 4-spored, 22–25 × 11–12 μm, surrounded by pseudoparaphyses. Cheilocystidia of two types: A: 32–52 × 9–25 μm, narrowly lageniform, often subcapitate to clavate at apex, scattered, when fresh apex turning yellow or green in a 10 % solution of ammonia, a reaction not seen in dried material, B: 17–42 μm wide, clavate, ellipsoid to sphaeropedunculate, numerous. Pleurocystidia not seen. Pileipellis a hymeniderm. Pileocystidia 27–62 × 9–14 μm, similar in shape and reaction of ammonia solution to the cheilocystidia of type A, sometimes thick-walled at base. Clamps present.

Habitat and distribution: Growing in a small group in a lawn of a park. Known from the type locality in southernmost Sweden.

Notes - The species can be recognized by small basidiomata, small, lageniform cystidia with subcapitate to clavate apex turning yellow or green in a 10 % solution of ammonia, and large spores with indistinct germ pore. Van de Bogart (1975) the author of Coprinus simulans reported “pileocystidia and caulocystidia when fresh reacting to ammonia water by becoming a bright blue-green color”. A green reaction of cystidia is known also from some species of Psathyrella.

Coprinopsis musae Örstadius & E. Larss., sp. nov. [MB 811479] – Holotype: Denmark, E-Jylland, Randers, in Randers Rainforest, a tropical greenhouse, on debris of Musa sp., 11.VII.2006, J. Vesterholt 06–179 (C). Figs. 2d, 5. (S3: Fig. 4a–b, d.)

Fig. 5

Coprinopsis musae. J. Vesterholt 06–179; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. Named after its occurrence on Musa (bananas).

Pileus 4–11 mm latus, ovoideus, deinde conico-parabolicus, brunneolo-aurantiacus, hygrophanus, humido striatus, laevis, siccus, glaber, in sicco pallescens, velum non visum. Lamellae confertae, subangustae, fere liberae, adscendentes, cinereo-aurantiacae, acie albae. Stipes 12–34 × 0.8–1.5 mm, cylindraceus, albus, sursum pruinosus, deorsum innato fibrillosus, submarmoratus, velum nullum. Odore inconspicuo vel acido. Sapore nullo. Sporae 9–10.5 × 5.8–6.2 μm, ellipsoideae, oblongae, ovoideae, amygdaliformes, valde pallidae; poro germinativo nullo vel inconspicuo. Basidia 4-sporigera. Pleurocystidia nulla. Cheilocystidia 25–45 × 8–12 × 6–14 μm, anguste utriformia, numerosa. Fibulae adsunt. Ad lignum in sylva tropica.

Cap 4–11 mm broad, ovoid, then conical, rounded conical to paraboloid, low campanulate, brownish orange (Methuen 5C4-6D4), at margin greyish orange (5B3), hygrophanous, striate almost to centre, smooth, dry, not pubescent, drying to greyish orange, orange grey, yellowish white (5B3, 6B2, 4A2), no veil seen. Gills crowded, rather narrow, almost free, adscending, greyish orange (5B3) with paler, slightly flocculose edge. Stem 12–34 × 0.8–1.5 mm, cylindrical or gradually widened to 2 mm at base, not bulbous, pulverulent upper part, white, downwards innately fibrillose, no veil seen, slightly marbled. Smell very faint, acidic. Taste none.

Spores 9–10.5 × 5.8–6.2 μm, av. 9.7–9.8 × 5.9–6 μm, Qav. = 1.6–1.7, ellipsoid, oblong, ovoid, in profile sometimes amygdaliform, in water hyaline to very pale brown (Mu. 10YR 8/3-8/4); germ pore absent or with a faint pore. Basidia 4-spored, 18–28 × 8–10 μm; pseudoparaphyses often seen. Pleurocystidia absent. Cheilocystidia numerous, 25–45 × 8–12 (at base) × 6–14 (at apex) μm, narrowly utriform, no clavate cells seen. Scalp cap halfway from margin: the upper layer consists of narrow hyphae indicating the presence of a suprapellis or less likely a veil, below a layer of subglobose to ellipsoid, pale yellow, 20–38 μm wide cells; pileitrama of yellow pigmented hyphae. Clamps seen on stem hyphae.

Habitat and distribution: So far only known from a tropical greenhouse where it grew on woody remnants, branches, and leaves of Musa (bananas).

Notes - Coprinopsis musae is characterized by its small size, extremely pale spores, lacking pleurocystidia, lacking veil. This psathyrelloid Coprinopsis comes genetically close to C. marcescibilis from which it differs in having smaller and significantly paler spores and smaller basidiomata. The experienced collector, Jan Vesterholt, did not notice a veil. C. marcescibilis has characteristic dentate flocci at cap margin. Psathyrella typhae belonging to another genus, is similar in morphology, but differs in having a veil and in growing on plant debris of, e.g., Carex, Scirpus, and Typha in wet or moist places.

Additional specimen: Denmark, E-Jylland, Randers, in Randers Rainforest, a tropical greenhouse, on remnants of Musa, 11.VII.2006, J. Vesterholt 06–180.

Coprinopsis udicola Örstadius, A. Melzer & E. Larss., sp. nov. [MB 811480] – Holotype: Germany, Sachsen, Delitzsch, 7.X.2008, A. Melzer 1240 (GB). Fig. 6. (S3: Fig. 5a–d.)

Fig. 6

Coprinopsis udicola. A. Melzer 1240; Pl Pleurocystidia; Ch Cheilocystidia; Basidioma ×1,5; Scale bars 10 μm

Etymology. The epithet refers to the preference of moist habitat.

Pileus 20–25 mm latus, 15–20 mm altus, conicus vel conico-convexus, saepe undulatus, humido striatus, obscure fuscus, in sicco pallescens sine roseo; velum fibrillosum. Lamellae modice confertae, sinuatae, primo albidae, dein griseo-brunneae vel atro-brunneae; acie alba fimbriata. Stipes usque ad 80 × 2 mm, eradicatus, albus, modice fibrillosus, apice albopulverulentus. Sporae 11.5–14 × 6–7 μm, anguste ovoideae, subcylindricae, oblongae, amygdaliformes, citriformes, rubidae, poro germinativo distinctae. Basidia 4-sporigera. Cheilocystidia 30–80 × 7–18 μm, polymorpha, lageniformia, subcylindrica, anguste utriformia, interdum flexuosa, septata, furcata vel capitata, numerosa. Pleurocystidia 30–75 × 10–14 μm, cheilocystidiis similia, raro vel absunt. Cellulae veli 60–125 × 5–17 μm. Fibulae adsunt. Caespitosus, udus.

Cap conical to conico-convex, when young about 10 mm broad and 10 mm high, mature 20–25 mm broad and 15–20 mm high, often wavy or slightly inflexed at margin, moist striate, young and mature dull grey brown with a brown centre, pallescent to grey or whitish without pink; veil when young as white fibrillose remnants more than halfway from margin. Gills rather close, sinuate, unequal, at first whitish, becoming grey brown to black brown, with conspicuously micaceous, uneven, white edges similar to the ones of Coprinellus micaceous. Stem up to 80 × 2 mm, equal, often bent, no pseudorhiza, white, glossy, with sparsely fibrillose veil remnants, pulverulent at apex. Not deliquescent. Smell fungous. Taste not recorded.

Spores 11.5–14 × 6–7 μm, av. 12.5 × 6.2 μm, Qav. = 2.0, narrowly ovoid, subcylindrical, oblong, in profile amygdaloid to citriform, sometimes with a slight suprahilar depression, in water red (Mu. 2.5YR 4/8), germ pore distinct, central, rarely slightly eccentric. Basidia 4-spored, 17–25 × 10–11.5 μm. Cheilocystidia 30–80 × 7–18 μm, polymorphic, lageniform, subcylindrical, narrowly utriform, sometimes flexuous, septate, forked, or headlike at apex, abundant. Pleurocystidia 30–75 × 10–14 μm, similar to cheilocystidia, scattered to absent close to gill edge. Pileipellis a cutis with short, wide cells, 35–60 × 20–40 μm, pale brown. Pileitrama with pale pigmented hyphae. Veil cells 60–125 × 5–17 μm, pale. Clamps present on stem base hyphae.

Habitat and distribution: In fascicles of about twenty basidiomata, moist with Aegopodium podagraria, Urtica dioica, and Phalaris arundinacea. Only known from the type locality.

Notes - The species can be recognized by the shape and size of the cystidia, the pale cap, and the large, amygdaliform spores. The genetically close Coprinopsis canoceps has smaller spores and cystidia. C. marcescibilis differs in having smaller cystidia and characteristic denticulate flocci at cap margin.

Psathyrella arenosa Örstadius & E. Larss., sp. nov. [MB 811481] – Holotype: Sweden, Everöd, by the railway 2,3 km. N.E of the church, 19.X.1996, L. Örstadius 220–96 (GB). Fig. 7. (S3: Fig. 6a–d.)

Fig. 7

Psathyrella arenosa. L. Örstadius 220–96; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×2; Scale bars 10 μm

Etymology. The epithet refers to the growing on sandy places.

Pileus 4–15 mm latus, primo convexus, tum expanso-planus, luteolo-rubidus, striatus, hygrophanus, in sicco cremeus; velum fibrillosum dispersum. Lamellae distantes, cinereae; acie concolora vel albida. Stipes 20–35 × 1–3 mm, pallide brunneus, apice pulverulentus, deorsum fibrillosus. Odor indistinctus; sapor mitis. Sporae 7–9 × 4–5 μm, oblongae, ellipsoideae, ovoideae, subphaseoliformes, subamygdaliformes, luteolo-rubrae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 25–55 × 7–13 μm, lageniformia, subutriformia, dispersa vel numerosa. Cheilocystidia 20–40 × 7–13 μm, pleurocystidiis similia, modice numerosa, cellulis clavatis immixta. Cellulae veli 15–70 × 2–10 μm. Fibulae adsunt. Subgregarius ad terram arenosam.

Cap 4–15 mm broad, convex, then expanded to nearly plane, yellowish red, dark reddish brown (Mu. 5YR 4/6, 3/3), faintly to distinctly striate when moist, hygrophanous, drying to cream; veil as dispersed fibres halfway from margin. Gills distant, L = 11–25, adnate to adnexed, mature pinkish grey (Mu. 5YR 6/2) with white or concolorous edge. Stem 20–35 × 1–3 mm, slightly enlarged towards base, pale brown, pulverulent at apex, fibrillose lower part. Taste mild. Smell not distinctive.

Spores 7–9 × 4–5.5 μm, av. 7.3–8.5 × 4.4–5.2 μm, Qav. 1.6–1.8, oblong, ellipsoid, ovoid, in profile sometimes subphaseoliform or subamygdaliform, in water yellowish red (Mu. 2.5YR 5/8 – 5YR 6/8), germ pore distinct. Basidia 4-spored, 17–26 × 7–9 μm, sometimes partly intracellular pigmented. Pleurocystidia 25–55 × 7–13 μm, lageniform, subutriform, scattered to numerous, pale. Cheilocystidia of two types: A: 20–40 × 7–13(−20) μm, similar to the pleurocystidia, rather numerous, B: small, clavate, scattered, sometimes numerous towards cap margin. Scalp cap halfway from margin: pileipellis consists of ellipsoid, clavate to subglobose cells 10–25 μm wide, pale; pileitrama made up of strongly pigmented hyphae, sometimes incrusted. Hymenophoral trama moderately pigmented. Veil cells 15–70 × 2–10 μm, clamped.

Habitat and distribution: Growing solitary or in small groups among mosses in open places, on dry, sandy, or humus rich soil. Known from Denmark and the southernmost province Skåne in Sweden.

Notes - The species is recognized by its small size, scanty veil, size of spores and cystidia, and the habitat on sandy, calcareous soil. Psathyrella senex differs in having larger basidiomata, a veil with flocci when fresh, and a habitat on debris of wood or among fallen leaves or terricolous in forests. P. seymourensis can be separated by a floccose veil and acute cystidia. P. ichnusae has broader pleurocystidia, sometimes thick-walled, paler cap, and a habitat on burnt soil.

Additional specimens: Denmark, NE. Jylland, Skagen, Grenen, 19.VIII.2000, J. Vesterholt with sons JV00-229 (C); Sjaelland, Gilleleje Feriecenter, 21.V.2012, M. Sonniks MSO2012-445438 (C); Zealand, Naestved Øvelseplads, 19.V.2010, E. Kristensen 2010–94357 (C); Sweden, Skåne, Åhus, Älleköpinge, 7.X.1999, L. Örstadius 146–99; Everöd, by the railway 2,3 km. N.E of the church, 24.IX.1987, L. Örstadius 155–87; Ivetofta, Grödby, 31.VIII.2006, L. Örstadius 250–06; Löderup, 0,5 km. N. of the seashore, 19.X.1997, L. Örstadius 184–97; N.O. of Löderup, 2.XI.2011, L. Örstadius 329–11; Lyngsjö, 0,6 km. N. of the church, 9.X.2006, L. Örstadius 360–06; Hommentorpsvägen, 28.X.2001, L. Örstadius 330–01; Ystad, Klintholmen, 2.XI.2011, L. Örstadius 328–11.

Psathyrella carminei Örstadius & E. Larss., sp. nov. [MB 811482] – Holotype: Italy, Calabria, Cosenza, Serra Vurga, Longobucco, 22.IX.2008, C. Lavorato (GB). Figs. 2c, 8. (S3: Fig. 7a–d.)

Fig. 8

Psathyrella carminei. C. Lavorato Longobucco; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×0,5; Scale bars 10 μm

Etymology. In honour of Carmine Lavorato who discovered the fungus.

Pileus usque ad 35 mm latus, primo semiglobosus, tum expanso-planus, rugosus, rufo-brunneus, striatus, hygrophanus, in sicco pallescens; velum flocculosum. Lamellae adnatae, modice confertae, primo albidae, demum atro-brunneae. Stipes usque ad 55 × 4 mm, aequalis, ad basim incrassatus, albidus, apice pulverulentus, deorsum fibrillosus. Sapor mitis. Odor piscis similis. Sporae 9–11 × 5–5.5 μm, oblongae, subcylindricae, leviter irregulares, subphaseoliformes, rufo-flavae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 40–55 × 11–18 μm, utriformia, lageniformia, modice numerosa. Cheilocystidia 30–45 × 10–16 μm, pleurocystidiis similia, dispersa, cellulis clavatis immixta. Fibulae adsunt. In silva conifera (Pinus).

Cap up to 35 mm broad, at first semiglobate, then expanded plane, wrinkled, red brown, striate halfway when moist, hygrophanous, pallescent on drying, veil as white fugacious flocci. Gills adnate, medium spaced, when young whitish, becoming blackish brown with pale edge. Stem up to 55 × 4 mm, equal, inflated towards base, whitish, pulverulent at apex, with fibrillose veil remnants downwards. Taste mild. Smell of fish.

Spores 9–11 × 5–5.5 μm, av. 10 × 5.2 μm, Qav. = 1.9, oblong, subcylindrical, slightly irregular, in profile sometimes subphaseoliform, in water reddish yellow (Mu. 5YR 6/8), with distinct germ pore. Basidia 4-spored, 22–25 × 8–10 μm. Pleurocystidia 40–55 × 11–18 μm, utriform to lageniform, rather numerous, pale. Cheilocystidia of two types: A: 30–45 × 10–16 μm, similar to pleurocystidia, scattered to rather numerous, B: 9–16 μm wide, clavate, numerous. Pileipellis (scalp) of 15–45 μm wide, subglobose to ellipsoid, pale cells. Pileitrama with moderately pigmented hyphae. Clamps present.

Habitat and distribution: Growing on soil in a forest with Pinus laricio subsp. calabrica. The soil is slightly acidic, pH 5,5–6. USDA-classification: Humic Psammentic, Dystrupets, mixed mesic. Location of the forest, southeast. Montane Mediterranean climate. Known only from the type locality in Italy.

Notes: The spore size, and non-forked, non-incrusted utriform cystidia are important characters for Psathyrella carminei. It differs from the genetically close P. spadiceogrisea and P. fatua in having larger spores and a smell of fish.

Psathyrella fennoscandica Örstadius & E. Larss., sp. nov. [MB 811483] – Holotype: Sweden, Träne, Stavsbacka, in a pasture grazed by cows, 12.X.2005, L. Örstadius 484–05 (GB). Figs. 2e, 9. (S3: Fig. 8a–d.)

Fig. 9

Psathyrella fennoscandica. L. Örstadius 484–05; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. The epithet refers to the occurrence in Finland, Norway, and Sweden.

Pileus 15–40 mm latus, conico-convexus, raro fere planus, primo castaneus, humido striatus,

hygrophanus, in sicco pallescens; velum fibrilloso-floccosum ad dimidium ad medium, glabrescens. Lamellae subdistantes, ventricosae, adnatae, griseae, acie alba. Stipes 50–120 × 2–5 mm, modice rigidus, saepe ad basim bulbosus, albidus. Odor indistinctus; sapor mitis. Sporae 8–10.5 × 5–6 × 4.5–5.5 μm, ovoideae, subfusiformes, oblongae, interdum amygdaliformes, rubidae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 35–80 × 10–16 μm, anguste utriformia, lageniformia, conica, dispersa vel numerosa. Cheilocystidia 20–45 × 6–12 μm, pleurocystidiis similia, numerosa. Cellulae veli 20–100 × 2–8 μm. Fibulae adsunt. Solitaria vel gregaria in sylva saepe ad terram vel ad lignum.

Cap 15–40 mm broad, paraboloid, then expanded campanulate, obtusely conical, convex, rarely almost applanate, at first dark reddish brown (Mu. 5YR 3/2-3/4-4/3), when moist striate up to ¾ from margin, hygrophanous, when drying sometimes ochre-brown at centre (e.g., Mu. 5YR 6/8), then becoming pale brown or grey; veil in young stages as floccules halfway from margin and some fibres towards centre, glabrescent. Gills medium spaced, L = 24–28, ventricose, adnate, at first grey, pink (Mu. 7.5YR 7/4), then reddish grey (Mu. 5YR 5/2), with minutely white floccose edge. Stem 50–120 × 2–5 mm, rather stiff, equal, often with distinct basal bulb, whitish, very pale brown, pulverulent striate at apex, with fugacious fibres or floccules from veil downwards. Smell not distinctive. Taste mild. Spore print black with reddish tinge (Mu. 10R 2.5/1-2.5/2).

Spores 8–10.5 × 5–6 × 4.5–5.5 μm, av. 8.7–9.7 × 5.2–5.7 × 4.9–5.3 μm, Qav. = 1.6–1.7 (in front view), 1.7–1.9 (in profile), ovoid, subfusiform, oblong, slightly angled, in profile with a slight suprahilar depression or amygdaliform, in water red (Mu. 2.5YR 4/6), almost opaque, with distinct germ pore. Basidia 4-spored, 16–26 × 8–9 μm. Pleurocystidia 35–80 × 10–16 μm, narrowly utriform, lageniform, conical, scattered to numerous, pale. Cheilocystidia of two types: A: 20–45 × 6–12 μm, similar to pleurocystidia, numerous, B: small, clavate, of variable frequency. Pileipellis a hymeniderm of 15–55 μm wide cells. Pileitrama made up of coarsely incrusted hyphae. Hymenophoral trama pale to moderately pigmented. Veil cells 20–100 × 2–8 μm, cylindrical. Clamps present on veil and stem hyphae.

Habitat and distribution: Solitary to gregarious in coniferous and deciduous forests, pastures, with Alnus glutinosa, A. incana, Betula sp., Deschampsia cespitosa, Frangula alnus, Picea abies, Pinus silvestris, Populus tremula, Sphagnum sp., Vaccinium myrtillus, oligotrophic, moist, on mossy, decayed branches or logs, on other debris, on soil, on grass, among leaves. So far known from Finland, Norway, and Sweden.

Notes - Psathyrella fennoscandica is recognized by red brown cap, rather scanty veil, bulbous stem base, narrowly utriform to lageniform cystidia, dark subfusiform spores, and moist oligotrophic habitat. Psathyrella senex differs in having slightly smaller, paler, non-fusiform spores, and prefers richer, sometimes nitrophilous habitats.

Additional specimens: Finland, Uusimaa, Tuusula, Ruotsinkylä, 31.VIII.1993, I. Kytövuori 93-602a (H); Varsinais-Suomi, Kustavi, Rahi, Leonsaari, W. of Isoperä bay, 3.XI.1996, J. Vauras 12059 (TURA); Norway, Oppland, Søndre Land, lake Trevatna, Vassenden, 14.IX.1984, G. Gulden 336/84 (O, Psathyrella sp.); Sweden, Blekinge, Kyrkhult, Snövleboda, 9.IX.1993, A. Blom LÖ 189–93; Ronneby, Piskabacken, 27.IX.2001, A. Nyström LÖ 199–01; Medelpad, Borgsjö, Bergåsen, 13.IX.1995, I. Kytövouri LÖ 89–95; Höganäs, 30.VIII.1993, D. Broström LÖ 148–93; Julåsen, 3.IX.1993, J. Stålberg LÖ 171–93; 14.IX.1995, L. Örstadius 99–95; Tubbo, 28.VIII.1989, L. Örstadius 236–89; Skåne, Andrarum, Borgarehögen, 5.X.1991, L. Örstadius 245–91; Glimåkra, E. of Nygårdstorpet, 25.IX.1997, L. Örstadius 119–97; Hjärsås, N. of Immeln, 8.IX.1989, L. Örstadius 305–89; Hörröd, Rebbetuaröd, 9.IX.1989, L. Örstadius 307–89; N. Åkarp, Hillarp klint, 17.IX.1996, L. Örstadius 95–96; Näsum, Hagstad, 12.X.1995, L. Örstadius 204–95; Örkened, Örnanäs, 5.X.1996, L. Örstadius 161–96; Vinslöv, Åraslöv bog, 29.IX.2001, L. Örstadius 214–01; Småland, Femsjö, Yaberg, 17.IX.1987, L. Örstadius 115–87; Uppland, Vänge, Fiby urskog, 8.IX.1994, I. Kytövuori LÖ 46–94; Värmland, Fryksände, Gultberget, 5.X.1991, J. Stålberg 91–114; Hammarö, St. Skagene, 12.IX.1987, L. Örstadius 110–87; Högboda, 12.IX.1987, L. Örstadius 102–87; Övre Ullerud, Torsberget, 13.IX.1992, L. Örstadius 242–92, 243–92; Stora Kil, Apertin, Kil, 14.IX.1986, J. Stålberg 86:84; Västergötland, Blidsberg, Göpåsen, 19.IX.1992, H. Toresson LÖ 306–92; Fristad, Gingri, 14.IX.1990 Toresson & Örstadius 248–90; Södra Säm, Klevaberget, 16.IX.1992, L. Örstadius 261–92; Timmele, Blankered, 15.IX.1992, L. Örstadius 255–92; Tvärred, Kråkeboberget, 19.IX.1992, L. Örstadius 297–92; Västmanland, Surahammar, Kanalhagen, 25.IX.1996, L. Bsenko 96–1941; Lönnbro bog, 9.X.1997, L. Bsenko 97–5555; Väratorp West, 29.IX.1999, L. Bsenko 99–1536.

Psathyrella ichnusae Örstadius, Contu, E. Larss. & Vizzini, sp. nov. [MB 811484] – Holotype: Italy, Sardinia, Olbia-Tempio, Golfo Aranci, Golfo di Marinella, 6.I.2008, M. Contu (GB). Fig. 10. (S3: Fig. 9a–d.)

Fig. 10

Psathyrella ichnusae. M. Contu 080106; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. The epithet refers to Ichnusa, an old name for Sardinia.

Pileus 2–18 mm latus, campanulatus vel convexus, hygrophanus sed non striatus, initio obscure ochraceo-brunneus, deinde ochraceo-argillaceus; velum album fibrillosum. Lamellae distantes vel subconfertae, sinuato-adnatae, ochraceae, dein obscure brunneae, ad aciem albo-flocculosae. Stipes 12–32 × 1–2 mm, valde fragilis, albus vel ochraceotinctus, sericeus. Odor atque sapor nulli. Sporae in cumulo brunneae. Sporae 7.8–9.2 × 4.4–5 μm, oblongae, subcylindricae, anguste ovoideae, amygdaliformes, in aqua observatae aurantiacae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 32–48 × 10–18 μm, utriformia vel lageniformia, dispersa. Cheilocystidia 25–35 × 10–16 μm, pleurocystidiis similia, dispersa vel modice numerosa, cellulis clavatis immixta. Cellulae veli 5–10 μm latae. Fibulae adsunt. Inter graminis, ad terram loco deusto, gregaria.

Cap 2–18 mm broad, at first convex, campanulate-convex to conico-paraboloid, then convex, not seen expanded plane, without a distinct umbo, not deliquescent, hygrophanous, not striate, when young and fresh dark ochre-brown, then ochre-buff; veil in young stages covered by thin fibrillose white veil remnants, still present on the margin when mature. Gills distant to medium spaced, sinuate-adnate, ochre, becoming dark bister-brown, with a paler flocculose edge. Stem 12–32 × 1–2 mm, very fragile, white with ochre tinge, sericeous; veil in young stages forming a dense fleece connecting stem with margin of cap. Context thin and fragile. Smell and taste none. Spore print brown.

Spores 7.8–9.2 × 4.4–5 μm, av. 8.3 × 4.6 μm, Qav. = 1.8, oblong, subcylindrical, narrowly ovoid, in profile sometimes narrowly amygdaliform, in water yellowish red (Mu. 5YR 5/8), with distinct germ pore. Basidia 4-spored, 18–20 × 8–9 μm. Pleurocystidia 32–48 × 10–18 μm, utriform to lageniform, sometimes with thickened walls, scattered, pale. Cheilocystidia of two types: A: 25–35 × 10–16 μm, similar to pleurocystidia, scattered to rather numerous, B: small, clavate, rather numerous. Pileipellis similar to a hymeniderm; pileitrama with rather strongly pigmented hyphae. Veil cells 5–10 μm wide. Clamps present.

Habitat and distribution: Gregarious on burnt soil in a coastal grassland near the sea. So far only known from the type locality in Italy.

Notes - The species can be recognized by the small size, rather well-developed veil, spore size, cystidia shape, and habitat. Psathyrella ichnusae differs from P. arenosa in having broader sometimes pigmented pleurocystidia, rather numerous small clavate cheilocystidia, and a habitat on burnt soil.

Psathyrella lilliputana Örstadius & E. Larss., sp. nov. [MB 811485] – Holotype: Sweden, Halland, Harplinge, Mannarp, 18.IX.2009, L. Örstadius 130–09 (GB). Fig. 11. (S3: Fig. 10a–d.)

Fig. 11

Psathyrella lilliputana. L. Örstadius 130–09; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×4; Scale bars 10 μm

Etymology. The epithet refers to the very small basidiomata that are small enough to inhabit the land of Lilliput in Swift’s Gulliver’s Travels.

Pileus 1–4 mm latus, semiglobosus, convexus, rufo-brunneus, leviter striatus, hygrophanus; velum disperse fibrillosum. Lamellae distantes, late adnatae, brunneae; acie alba. Stipes 6–8 × 0.2–0.5 mm, albidus, sparsim fibrillosus, apice pulverulentus. Odor nullus. Sporae 9.5–11 × 5–6 μm, oblongae, subfusiformes, citriformes, amygdaliformes, interdum subrostellatae vel irregulares, luteolo-rubrae; poro germinativo valde distinctae. Basidia 4-sporigera. Pleurocystidia 25–42 × 6–12 μm, anguste lageniformia, utriformia, fusiformia, subcylindrica, dispersa. Cheilocystidia 25–38 × 5–11 μm, pleurocystidiis similia, numerosa. Fibulae adsunt. Habitat: Ad lignum, in pascuis.

Cap 1–4 mm broad, semiglobate to convex, reddish brown, faintly striate, hygrophanous, veil as dispersed fibrils. Gills distant, L = 7–8, rather broadly adnate, brown with white edge. Stem 6–8 × 0.2–0.5 mm, whitish, with sparsely fibrillose veil remnants, pulverulent at apex. Smell none. Taste not recorded.

Spores 9.5–11 × 5–6 μm, av. 10.1 × 5.3 μm, Qav. = 1.9, oblong, subfusiform, sometimes snout-like projected at apex or irregular in outline, in profile citriform, amygdaliform, or with a suprahilar depression, many small oil drops, in water yellowish red (Mu. 5YR 5/6-4/6); germ pore very distinct. Basidia 4-spored, 16–19 × 8–9 μm. Pleurocystidia 25–42 × 6–12 μm, narrowly lageniform, utriform, fusiform, subcylindrical, rarely forked, scattered, pale. Cheilocystidia 25–38 × 5–11 μm, similar to the pleurocystidia, numerous. Scalp cap halfway from margin: pileipellis made up of clavate to subglobose, ellipsoid to pyriform, 15–30 μm wide, pale cells; pileitrama of strong pigmented hyphae. Clamps frequently seen on septa on hyphae in the stipe.

Habitat and distribution: Growing in a small group on small remnants of deciduous wood in a pasture with Quercus. So far only known from the type locality.

Notes - The species can be recognized by tiny basidiomata, small cystidia, and snout-like projected spores. About 15 species in Europe are known with a cap size of 4 mm or less. Only Psathyrella orbitarum and P. purpureobadia have a mean-value of spore size matching P. lilliputana. The latter one can be separated by purple-tinged cap and lack of clamps. P. orbitarum is genetically rather closely related with P. lilliputana but differs in having acute cystidia and regularly shaped spores.

Psathyrella lyckebodensis Örstadius & E. Larss., sp. nov. [MB 811486] – Holotype: Sweden, Skåne, Trolle-Ljungby, Lyckeboda, on sandy soil in a pasture, 27.X.2011, L. Örstadius 301–11 (GB). Fig. 12. (S3: Fig. 11a–d.)

Fig. 12

Psathyrella lyckebodensis. L. Örstadius 301–11; Pl Pleurocystidia; Ch Cheilocystidia; Basidioma ×1; Scale bars 10 μm

Etymology. The name refers to Lyckeboda where the fungus was discovered.

Pileus 12 mm latus, conico-convexus, brunneo-fulvus, non-striatus, granulis omnino. Lamellae adnatae, distantes vel modice confertae, ventricosae, brunneo-fulvae. Stipes 40 × 1.8 mm, deorsum fibrillosus, apice pulverulentus. Odor nullus. Sporae 8–9.2 × 4–4.6 μm, oblongae, subcylindricae, brunneo-flavae; poro germinativo nullo vel inconspicuo. Basidia 4-sporigera. Pleurocystidia 55–80 × 12–22 μm, lageniformia, utriformia, modice numerosa. Cheilocystidia 30–75 × 9–22 μm, pleurocystidiis similia, numerosa, cellulis clavatis immixta. Fibulae adsunt. Habitat: Ad terram, in pascuis.

Cap 12 mm broad, conico-convex, brownish yellow (Mu. 10YR 6/8), not striate, entirely covered by grey white grains. Gills rather broadly adnate, medium spaced to distant, L = c. 14, ventricose, concolorous with cap, with a faint pruinose edge. Stem 40 × 1.8 mm, slightly paler than cap, white fibrillose, pulverulent at apex. Smell none. Taste not recorded.

Spores 8–9.2 × 4–4.6 μm, av. 8.6 × 4.4 μm, Qav. = 2.0, oblong, subcylindrical, sometimes with a wide upper part or a slight suprahilar depression, in water brownish yellow (Mu. 5YR 6/8); germ pore absent or rarely indistinct. Basidia 4-spored, 17–30 × 7.5–9 μm. Pleurocystidia 55–80 × 12–22 μm, lageniform to utriform, upper part rarely head-like or subacute, rather numerous. Cheilocystidia of two types: A: 30–75 × 9–22 μm, similar to pleurocystidia, abundant, B: small, clavate, scattered. Pileipellis an epithelium of loosely arranged often subglobose, pale cells, 15–45 μm wide; pileitrama with pale brown pigmented hyphae. True clamps and pseudoclamps seen.

Habitat and distribution: Solitary on sandy soil with some deciduous trees in a dry pasture grazed by cows. So far only known from the type locality in Sweden.

Notes - The species is distinguished by a granular cap surface, small size, and rather large hymenial cystidia. It belongs to section Cystopsathyra (Kits van Waveren 1985) recognized by a powdery-mealy cap surface. Less than ten species are known worldwide from the section, all of them rare. The species differs from Psathyrella albofloccosa in having slightly smaller, elongate spores and larger, sometimes subacute cystidia. P. sphaerocystis has very small cystidia, wider spores and grows on dung while P. globosivelata has smaller spores and often grows on remnants of Agaricales species.

Psathyrella madida Örstadius & E. Larss., sp. nov. [MB 811490] – Holotype: Sweden, Skåne, Åhus, Sånnarna, 12.X.2006, L. Örstadius 377–06 (GB). Figs. 2a, 13. (S3: Fig. 12a–d.)

Fig. 13

Psathyrella madida. L. Örstadius 377–06; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. The epithet refers to the preferences of moist habitat.

Pileus 7–33 mm latus, conico-convexus, interdum irregularis, spadiceus, fulvus, ad marginem leviter striatus, hygrophanus, in sicco pallescens, velum fibrilloso-floccosum. Lamellae modice confertae, adnatae, ventricosae, griseo-brunneae, acie alba. Stipes 10–35 × 1.5–3 mm, albus, apice pruinosus, deorsum fibrillosus. Odor nullus. Sporae 7–9.5 × 4–5 μm, ovoideae, oblongae, subfusiformes, amygdaliformes, subcitriformes, rubidae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 35–50 × 12–26 μm, clavata, sphaeropedunculata, utriformia, lageniformia, numerosa vel dispersa. Cheilocystidia 25–55 × 8–35 μm, pleurocystidiis similia, numerosa. Cellulae veli 40–190 × 6–40 μm. Fibulae adsunt. Habitat: In solo sabuloso udo, cum Phragmite.

Cap 7–33 mm broad, at first semiglobate, then conico-convex, convex, expanding to plane with deflexed margin, sometimes irregular, reddish brown, reddish yellow, fulvous (Mu. 5YR 5/4, 5/6, 6/8), with the darkest colours towards centre, faintly striate at margin, hygrophanous, pallescent on drying; veil from margin to centre with a fibrillose to floccose coating, evanescent. Gills medium spaced, L = 24–30, adnate, ventricose, at first white grey, becoming grey brown with white fimbriate edge. Stem 10–35 × 1.5–3 mm, whitish, pulverulent at apex, downwards with floccules from veil remnants. Smell none. Taste mild.

Spores 7–9.5 × 4–5 μm, av. 7.9–8.5 × 4.4 μm, Qav. = 1.8–1.9, narrowly ovoid to oblong, sometimes subfusiform, in profile narrowly amygdaliform, subcitriform or with a slight suprahilar depression, in water red to yellowish red (Mu. 2.5YR 4/6-3/6, 5YR 5/8); germ pore distinct. Basidia 4-spored, 18–26 × 8–9 μm. Pleurocystidia 35–50 × 12–26 μm, clavate, sphaeropedunculate, utriform, lageniform, numerous to scattered, pale. Cheilocystidia 25–55 × 8–35 μm, similar in shape to pleurocystidia, numerous. Scalp cap halfway from margin: pileipellis with subglobose, ellipsoid, 20–55 wide, pale cells, some cells also clavate reminding of a hymeniderm; pileitrama of pale pigmented hyphae. Veil cells 40–190 × 6–40 μm. Clamps frequent on stem base.

Habitat and distribution: Growing single or in small groups on wet to moist sandy soil close to remnants of Phragmites communis. The locality is often inundated during parts of the year. So far only known from the type locality.

Notes - The species is recognized by the rather pale cap, copious veil, large veil cells, cystidia shape, and the moist habitat. The two collections grew ten feet from each other. Specimen 369–06 differs from the holotype in having clavate to sphaeropedunculate pleurocystidia in addition to utriform and lageniform ones. The genetic differences, however, were small. The closely related P. merdicola, also with a copious veil, can be separated by smaller spores and growth on dung.

Additional specimen: Sweden, Skåne, Åhus, Sånnarna, on wet to moist sandy soil close to remnants of Phragmites communis,10.X.2006, L. Örstadius 369–06.

Psathyrella rybergii Örstadius & E. Larss., sp. nov. [MB 811491] – Holotype: Sweden, Skåne, Fjälkestad, Perstorp, 11.X.2006, Örstadius & Ryberg 373–06 (GB). Fig. 14. (S3: Fig. 13a–d.)

Fig. 14

Psathyrella rybergii. L. Örstadius 373–06; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×2; Scale bars 10 μm

Etymology. In honour of the mycologist Arne Ryberg for his contributions to a better knowledge of fungi in southern Sweden.

Pileus 5–11 mm latus, campanulatus vel convexus, rufo-brunneus, striatus, hygrophanus, in sicco pallescens; velum fibrillosum. Lamellae distantes, adnatae, pallide brunneae; acie alba fimbriata. Stipes 15–22 × 1–1.5 mm, pulverulentus, fibrillosus. Odor nullus. Sporae 8.5–9.5 × 5–5.5 μm, oblongae, interdum leviter irregulares, subhexagonae, sub microscopium rufae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 35–60 × 10–22 μm, utriformia, numerosa. Cheilocystidia 30–45 × 10–16 μm, modice numerosa, pleurocystidiis similia, cellulis parvis clavatis immixta. Fibulae adsunt. Habitat: In silva frondosa.

Cap 5–11 mm broad, campanulate to convex, reddish brown (Mu. 2.5YR 5/4), striate more than halfway from margin, hygrophanous, pallescent on drying; veil present as fibres close to margin. Gills distant, L = 12–14, adnate, light brown (Mu. 7.5YR 6/4), with white fimbriate edge. Stem 15–22 × 1–1.5 mm, pulverulent 2/3 from apex, decreasing in intensity downwards, fibrillose lower part. Smell none.

Spores 8.5–9.5 × 5–5.5 μm, av. 9.1 × 5 μm, Qav. 1.8, oblong, sometimes slightly irregular, subhexagonal, in profile sometimes subphaseoliform, in water red (Mu. 2.5YR 5/6), with distinct germ pore. Basidia 4-spored, 20–28 × 8–9 μm. Pleurocystidia 35–60 × 10–22 μm, utriform, numerous, pale yellow below apex. Cheilocystidia of two types: A: 30–45 × 10–16 μm, similar to pleurocystidia, rather numerous, B: small, clavate, scattered to numerous. Scalp cap halfway from margin: pileipellis with pale yellow, 20–55 μm wide, subglobose to ellipsoid cells; pileitrama and hymenophoral trama with strongly pigmented hyphae. Clamps seen at hyphae of stem base.

Habitat and distribution: On soil among gravel or sand on a path in a deciduous forest. Known from the type locality only.

Notes - The species is distinguished by its small basidiomata, rudimentary veil, utriform cystidia, habitat, and pulverulent stem. P. panaeoloides can be separated by often larger basidiomes, by the density of the gills (L = 20–30), and by more irregular spores. Moreover, it is not known to grow among gravel in forests.

Psathyrella sabuletorum Örstadius & E. Larss., sp. nov. [MB 811492] – Holotype: Sweden, Skåne, N. Åsum, A3 former military camp, 29.X.1998, L. Örstadius 196–98 (GB). Fig. 15. (S3: Fig. 14a–d.)

Fig. 15

Psathyrella sabuletorum. L. Örstadius 196–98; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. The name epithet refers to the habitat on sandy places.

Pileus 5–15 mm latus, conico-convexus, convexus, brunneus, striatus usque ad medium, hygrophanus, in sicco pallide brunneus; velum fibrillosum. Lamellae distantes, adnexae, pallide brunneae. Stipes 20–35 × 1–1.5 mm, fragilis, apice pulverulentus, deorsum fibrillosus. Odor nullus. Sporae 8–11.5 × 4.5–6 μm, ovoideae, oblongae, subcylindricae, interdum amygdaliformes, rufo-flavae; poro germinativo indistinctae. Basidia 4-sporigera. Pleurocystidia 35–55 × 8–16 μm, lageniformia, fusiformia, numerosa vel dispersa. Cheilocystidia 25–40 × 9–14 μm, pleurocystidiis similia, dispersa vel destituta, cellulis clavatis immixta. Fibulae adsunt. In locis siccis et apertis ad terram arenosam.

Cap 5–15 mm broad, conico-convex, convex, strong brown (Mu. 7.5YR 5/6) to reddish brown, distinctly striate to centre, hygrophanous, drying to pale brown; veil fibrillose at margin. Gills distant, L = 11–16, adnexed, pale brown, with fimbriate edge. Stem 20–35 × 1–1.5 mm, fragile, sometimes a fibrillose zone, pulverulent at apex, fibrillose downwards. Smell none.

Spores 8–11.5 × 4.5–6 μm, av. 8.5–10.9 × 4.9–5.4 μm, Qav. = 1.7–2, ovoid, oblong, subcylindrical, sometimes slightly irregular, amygdaliform or with a suprahilar depression, occasionally subphaseoliform, in water reddish yellow (Mu. 7.5-5YR 6/8); germ pore indistinct to seemingly absent. Basidia 4-spored, 16–23 × 8–10 μm. Pleurocystidia 30–55 × 8–16 μm, lageniform to fusiform, sometimes conical or with flexuous walls, numerous to scattered. Cheilocystidia of two types: A: 25–40 × 8–14 μm, similar to pleurocystidia, rather numerous to absent, B: 8–18 μm wide, clavate to obpyriform, several cells deep from margin and halfway inwards, decreasing in number and then numerous to scattered. Scalp cap halfway from margin: pileipellis consists of subglobose to ellipsoid cells 15–50 μm wide and pale; pileitrama made up of faintly to moderately pigmented hyphae. Veil cells 15–60 × 5–10 μm. Clamps found on stem hyphae.

Habitat and distribution: Single to scattered in moss-carpets on steppe-like sandy soil in open grassland with some trees of Pinus, and in sand dunes. So far known from one locality in Denmark and four localities in southernmost Sweden.

Notes - Psathyrella sabuletorum is an ordinarily looking species characterized by small basidiomata, scanty veil, and habitat on sandy soil. The gill edge is often dominated by small clavate to obpyriform cells close to cap margin and the spore size varies significantly. It differs from Psathyrella obtusata in being smaller, in having amygdaliform spores, and a habitat on sandy soil. Psathyrella flexispora also prefers growing on sand, but has darker spores with a distinct germ pore and differently arranged gill edge where acute shaped cheilocystidia dominates. In the description we include the Danish collection with longer but not broader spores.

Additional specimens: Denmark, W. Jylland, Vejers Strand W. of Oksbøl, 3.XI.1990, M. Christensen JV 90–770, (C); Sweden, Skåne, Åhus, Älleköpinge, 31.X.2011, L. Örstadius 310–11; Horna Bränneri, 24.X.1998, L. Örstadius 187–98; 1.XI.2011, L. Örstadius 321–11; Löderup, NO Löderup, 2.XI.2011, L. Örstadius 330–11, 331–11.

Psathyrella scanica Örstadius & E. Larss., sp. nov. [MB 811493] – Holotype: Sweden, Skåne, Trolle-Ljungby, Lyckeboda, 29.X.2009, L. Örstadius 183–09 (GB). Fig. 16. (S3: Fig. 15a–d.)

Fig. 16

Psathyrella scanica. L. Örstadius 183–09; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×2; Scale bars 10 μm

Etymology. The name refers to the province Skåne where the fungus was discovered.

Pileus 6 mm latus, 5 mm altus, conicus, flavido-rubidus, striatus, hygrophanus, in sicco pallescens; velum non visum. Lamellae ascendentes, adnatae, distantes, ventricosae, rufo-brunneae; acie alba fimbriata. Stipes 38 × 0.8 mm, cylindricus, albidus, fibrillosus. Odor nullus. Sporae 8.5–9.5 × 4.5–5.5 μm, anguste ovoideae, oblongae, amygdaliformes, luteolo-rubrae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 30–60 × 10–16 μm, lageniformia, saepe subcapitata vel subclavata, numerosa. Cheilocystidia 25–50 × 8–16 μm, pleurocystidiis similia, numerosa, cellulis clavatis immixta. Fibulae adsunt. Habitat: Ad terram, in pascuis.

Cap 6 mm wide and 5 mm high, conical, when fresh yellowish red (Mu. 5YR 4/6), striate, hygrophanous, pallescent on drying, veil not seen. Gills ascendant, adnate, distant, L = 12, ventricose, reddish brown (Mu. 5YR 4/4), with white fimbriate edge. Stem 38 × 0.8 mm, cylindrical, whitish, with some scattered fibrillose veil remnants. Smell none. Taste not recorded.

Spores 8.5–9.5 × 4.5–5.5 μm, av. 9 × 5 μm, Qav. = 1.8, narrowly ovoid, oblong, in profile sometimes amygdaliform, in water yellowish red (Mu. 5YR 4/6), with distinct germ pore. Basidia 4-spored, 17–23 × 8–10 μm. Pleurocystidia 30–60 × 10–16 μm, lageniform, often with subcapitate or subclavate apex, numerous, pale. Cheilocystidia of two types: A: 25–50 × 8–16 μm, similar to pleurocystidia, numerous, B: small, clavate, very scattered. Scalp cap half-way from margin: pileipellis with subglobose to ellipsoid 20–32 μm wide cells, pale; pileitrama with strongly pigmented hyphae. Hymenophoral trama moderately pigmented. Clamps present on stem base mycelium.

Habitat and distribution: Solitary among moss on sandy calcareous soil in a cow-grazed pasture with Juniperus communis. Known from the type locality.

Notes - Psathyrella scanica can be recognized by small basidiomata, conical cap, scanty veil, lageniform cystidia with capitate to clavate upper part, and habitat on sandy calcareous soil. P. perpusilla differs in having 2-spored basidia, lageniform cystidia, often with acute to subacute apex, broad, non-amygdaliform spores, and a distinct veil.

Psathyrella siccophila Örstadius & E. Larss., sp. nov. [MB 811494] – Holotype: Sweden, Skåne, Åhus, Ripa, Älleköpingegård, 24.X.2006, L. Örstadius 417–06 (GB). Fig. 17. (S3: Fig. 16a–d.)

Fig. 17

Psathyrella siccophila. L. Örstadius 417–06; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. The epithet refers to the preference for dry places.

Pileus 8–14 mm latus, primo semiglobatus, deinde convexus, fulvo-cinnamomeus, usque ad 2/3 diametro pellucide striatus, hygrophanus, in sicco pallescens; velum fibrillosum. Lamellae distantes, adnatae, pallide rufo-brunneae. Stipes 20–30 × 1–2 mm, pallide brunneus, apice pulverulento, deorsum fibrillosus. Odor indistinctus; sapor mitis. Sporae 5.5–7 × 4.5–5 μm, ellipsoideae vel late ellipsoideae, pallidae; poro germinativo destitutae. Basidia 4-sporigera. Pleurocystidia 35–65 × 9–18 μm, anguste utriformia, lageniformia, numerosa. Cheilocystidia 25–40 × 8–14 μm, numerosa, cellulis clavatis immixta. Fibulae adsunt. Habitat in aridis, ad terram arenosam.

Cap 8–14 mm broad, at first semiglobate, then conico-convex, expanded convex with a low umbo, yellowish red (Mu. 5YR 5/6, 4/6, 6/6) with the darkest colours at centre, striate two thirds from margin, hygrophanous, pallescent on drying; veil as bundles of fibres close to margin, when young appendiculate. Gills distant, L = 12–16, adnate, light reddish brown (Mu. 5YR 6/4); edge concolorous. Stem 20–30 × 1–2 mm, pale brown, upper part sometimes with a fibrillose annulate zone of veil remnants, downwards fibrillose, evanescent, pulverulent at apex. Smell indistinct. Taste mild.

Spores 5.5–7 × 4.5–5 μm, av. 6–6.4 × 4.7–4.9 μm, Qav. = 1.3, ellipsoid to broadly ellipsoid, in profile flattened on adaxial side, sometimes amygdaliform, in water reddish yellow (Mu. 7.5YR 7/8-6/8); germ pore absent. Basidia 4-spored, 20–32 × 8–9 μm. Pleurocystidia 35–65 × 9–18 μm, narrowly utriform to lageniform, with pale yellow walls below apex, numerous. Cheilocystidia of two types: A: 25–50 × 8–15 μm, similar to pleurocystidia, numerous, B: small, clavate, scattered. Scalp cap half-way from margin: pileipellis with subglobose to ellipsoid, 20–40 μm wide, pale yellow brown cells; pileitrama with strongly pigmented hyphae. Clamps seen on mycelium hyphae.

Habitat and distribution: In small groups in open, dry, calcareous grassland with patches of sand steppe, grazed by horses and cows, together with species like Geastrum schmidelii and Poronia punctata. Known from two localities in the southernmost province of Skåne, Sweden.

Notes - The species is distinguished by small basidiomata, scanty veil, broad spores, absence of a germ pore, narrowly utriform cystidia, and habitat on dry, calcareous grassland. Psathyrella stridvallii, also in the present paper, has narrower, darker spores, and slightly larger basidiomata.

Additional specimens: Sweden, Skåne, Everöd, 500 meters east of Nybo farm, 4.XI.2010, L. Örstadius 98–10.

Psathyrella stridvallii Örstadius & E. Larss., sp. nov. [MB 811495] – Holotype: Sweden, Skåne, Kristianstad, Egna hem, 22.VIII.1998, L. Örstadius 104–98 (GB). Figs. 2b, 18. (S3: Fig. 17a–d.)

Fig. 18

Psathyrella stridvallii. L. Örstadius 104–98; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. In honour of the late Leif Stridvall and his wife Anita, who sent hundreds of specimens of Psathyrella on loan, contributing to a better understanding of the genus.

Pileus 10–30 mm latus, conico-convexus, demum expanso-planus, initio spadiceo-brunneus, striatus, hygrophanus, in sicco pallescens; velum cito evanidum. Lamellae modice confertae, anguste adnatae, fusco-brunneae; acie dilute alba. Stipes 30–40 × 2–3 mm, albidus, interdum brunnescens. Odor atque sapor indistincti. Sporae 5.5–7 × 3.5–4.5 μm, ovoideae, ellipsoideae, oblongae, saepe amygdaliformes, in aqua observatae aurantiacae, poro germinativo destitutae vel indistinctae. Basidia 4-sporigera. Pleurocystidia 25–50 × 10–16 μm, utriformia, clavata, numerosa. Cheilocystidia 25–40 × 8–14 μm, pleurocystidiis similia, numerosa, cellulis clavatis immixta. Cellulae veli 2–6 μm latae. Fibulae adsunt. Habitat: Ad terram in silva frondosa vel in horti.

Cap 10–30 mm broad, conico-convex, then expanded plane, often wavy, when young dark reddish brown (Mu. 5YR 3/4), striate at margin or halfway to centre, hygrophanous, drying from margin to brown grey, sometimes with ochraceous tinges, micaceous; veil rudimentary as scattered fibrils close to margin in young stages. Gills medium spaced, L = 30–33, narrowly adnate, when young light brown (Mu. 7.5YR 6/4), when mature reddish brown (Mu. 5YR 5/4); edge faintly white. Stem 30–40 × 2–3 mm, fistulose, whitish, brown discolouring when handled, pulverulent at apex, glabrous or with some fibrils downwards. Smell faint. Taste mild. Spore print reddish black (Mu. 10R 2.5/1).

Spores 5.5–7 × 3.5–4.5 μm, av. 6.2–6.4 × 3.9–4 μm, ovoid, ellipsoid, oblong, in profile often amygdaliform, in water yellowish red (Mu. 5YR 5/8); Qav. = 1.6; germ pore absent or indistinct. Basidia 4-spored, 18–22 × 7–8 μm. Pleurocystidia 25–50 × 10–16 μm, utriform, clavate, numerous, sometimes with yellow pigmented walls below apex. Cheilocystidia of two types: A: 25–40 × 8–14 μm, similar to pleurocystidia, numerous, scattered towards cap margin, B: small, clavate, scattered, numerous towards cap margin, sometimes difficult to separate from the first type. Pileipellis a hymeniderm of 20–55 μm wide cells; pileitrama of coarsely to moderately incrusted hyphae. Veil cells 2–6 μm broad. Clamps present.

Habitat and distribution: Solitary to subgregarious on soil in a rich deciduous forest and on a manured lawn. Known from two localities in Sweden.

Notes - The species is recognized by small basidiomata, rudimentary veil, small spores, and utriform cystidia. Psathyrella piluliformis differs in being larger, having a membranaceous veil between cap margin and stem, and a lignicolous habitat. Moreover, the spores are slightly smaller, irregular in outline, sometimes phaseoliform, and paler. P. siccophila with about the same spore length and cystidia shape can be separated by broader and paler spores. The three species are not closely related genetically.

Additional specimens: Sweden, Skåne, Nosaby, Eknabben, 29.V.1991, L. Örstadius 130–91.

Psathyrella sublatispora Örstadius, S.-Å. Hanson & E. Larss., sp. nov. [MB 811496] – Holotype: Sweden, Valleberga, Löderup – Kåseberga, 19.X.1997, S.-Å. Hanson & L. Örstadius 190–97 (GB). Fig. 19. (S3: Fig. 18a–d.)

Fig. 19

Psathyrella sublatispora. L. Örstadius 190–97; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. The name refers to the relatively broad spores.

Pileus 10–25 mm latus, convexus, interdum umbonatus vel undulatus, castaneus, striatus, in sicco pallescens; velum fibrillosum. Lamellae modice confertae, adnexae, brunneae; acie alba fimbriata. Stipes 40–70 × 1.5–2 mm, subradicatus usque ad 15 mm, pallide brunneus, apice pulverulentus, deorsum fibrillosus. Odor atque sapor indistincti. Sporae 9.5–11 × 5.5–7 μm, oblongae, ellipsoideae, raro ovoideae, interdum subamygdaliformes, rubidae, poro germinativo distinctae. Basidia 4-sporigera. Pleurocystidia 40–50 × 10–16 μm, conica, lageniformia, dispersa. Cheilocystidia 25–40 × 8–13 μm, pleurocystidiis similia, numerosa, cellulis clavatis immixta. Fibulae adsunt. Habitat: In silva frondosa vel in arvo sicco.

Cap 10–25 mm broad, convex, sometimes with umbo or wavy, dark reddish brown (Mu. 5YR 3/4) with darker umbo, striate almost to centre, pallescent on drying; veil as dispersed fibres halfway to centre. Gills medium spaced, L = 20–24, adnexed, brown, with white fimbriate edge. Stem 40–70 × 1.5–2 mm, with a 8–15 mm long pseudorhiza, pale brown, fibrillose lower part, pulverulent at apex. Smell and taste not distinctive.

Spores 9.5–11 × 5.5–7 μm, av. 10.1–10.7 × 6–6.4 μm, Qav. = 1.7, oblong, ellipsoid, rarely ovoid, with obtuse base and apex, in profile sometimes subamygdaliform, in water red (Mu. 2.5YR 4/8-5/8), with distinct germ pore. Basidia 4-spored, 18–24 × 9–10 μm. Basidia or other cells in hymenium sometimes partly brown intracellular pigmented, sometimes surrounded by pseudoparaphyses like coprinoid species. Pleurocystidia 40–50 × 10–16 μm, conical to lageniform, scattered, pale. Cheilocystidia of two types: A: 25–40 × 8–13 μm, similar to pleurocystidia, numerous or rather numerous, B: 10–15 μm wide, clavate, scattered to numerous, increasing in frequency towards cap margin. Scalp cap half-way from margin: pileipellis with subglobose to ellipsoid 10–40 μm wide cells, pale; pileitrama with strongly pigmented hyphae. Hymenophoral trama moderately pigmented. Clamps seen on stem hyphae.

Habitat and distribution: Solitary to gregarious, on the margin of an old potato field on sandy soil with Senecio vulgaris, Polygonum aviculare, Chenopodium album and Artemisia vulgaris, also among grass on a path through a grove. Found on two localities in Sweden.

Notes - The species is recognized by the relatively broad spores Qav. = 1.7, pointed cystidia, pseudorhiza, and a preference for growing on nitrophilous soil. Psathyrella orbitarum and P. orbicularis differ in having more elongate spores, reflected in the mean-values for the two species: Qav. = 1.8–2.1. Moreover, P. orbitarum lacks a pseudorhiza and P. orbicularis has less frequent clavate cells on gill edge near cap margin. The two species are molecularly well separated from P. sublatispora.

Additional specimens: Sweden, Skåne, Kristianstad, Näsby park, 22.X.2000, L. Örstadius 264–00 (GB).

Psathyrella vesterholtii Örstadius & E. Larss., sp. nov. [MB 811497] – Holotype: Sweden, Västerbotten, Saxnäs, 26.VII.2010, Laessoe & Petersen JHP-10.086 (GB). Figs. 2f, 20. (S3: Fig. 19a–d and 20).

Fig. 20

Psathyrella vesterholtii. J.H. Petersen JHP-10.086; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Etymology. In honour of the late Jan Vesterholt, who first discovered the species.

Pileus 8–25 mm latus, campanulatus, conico-convexus, deinde expanso-convexus, rufo-brunneus vel brunneus, striatus, hygrophanus, velum disperse fibrillosum. Lamellae modice confertae, angustae, adnatae, brunneae; acie alba. Stipes 30–80 × 1–4 mm, sursum annulato-membranaceus, deorsum fibrillosus. Sporae 9–11 × 5–6 μm, ovoideae, oblongae, subfusiformes, amygdaliformes, rubidae vel luteolo-rubrae, poro germinativo subdistinctae. Basidia 4-sporigera. Pleurocystidia 45–85 × 9–22 μm, lageniformia, obtuse fusiformia, utriformia, subcylindrica, luteola, numerosa. Cheilocystidia 30–60 × 10–18 μm, pleurocystidiis similia, modice numerosa, cellulis clavatis immixta. Cellulae veli 30–150 × 5–40 μm. Fibulae adsunt. In sylva inter muscos.

Cap 8–25 mm broad, campanulate, conico-convexus, then expanded convex, reddish brown to brown, striate, hygrophanous; veil remnants as dispersed fibrils. Gills medium spaced, L = c. 25–35, narrow, adnate, brown with white edge. Stem 30–80 × 1–4 mm, upper part with a membranaceous annulus, downwards fibrillose. Smell and taste unknown.

Spores 9–11 × 5–6 μm, av. 9.8–10.4 × 5.2–5.7 μm, Qav. = 1.8–1.9, ovoid, oblong, subfusiform, in profile amygdaliform or with a slight suprahilar depression, rarely subphaseoliform, in water red to yellowish red (Mu. 2.5–5YR 5/8), with rather distinct germ pore. Basidia 4-spored, 18–22 × 9–10 μm. Pleurocystidia 45–85 × 9–22 μm, lageniform, obtusely fusiform, utriform, subcylindrical, with thickened yellow walls, numerous. Cheilocystidia of two types: A: 30–60 × 10–18 μm, similar to pleurocystidia, rather numerous, B: small, clavate, numerous. Scalp cap halfway from margin: pileipellis of subglobose to ellipsoid 20–60 μm wide cells. Pileitrama made up of rather strongly pigmented hyphae. Veil cells 30–150 × 5–40 μm, ventricose, cylindrical. Clamps present.

Habitat and distribution: Solitary or in small groups in moss carpets (compare below) in coniferous forests. Known from four localities in the Faroes and Sweden, all north of latitude 62° N but south of the arctic circle.

Notes - The species is recognized by a membraneous annulus, large yellow cystidia, and habitat among mosses. T. Hallingbäck, expert on bryophytes, kindly tried to determine the mosses on a photo of the holotype collection: Hylocomium sp. (probably H. splendens), Dicranum (probably D. majus) and Lophozia (probably L. ventricosa). The genetically closely related Psathyrella sphagnicola differs from all four collections of P. vesterholtii in having smaller spores and pleurocystidia. P. fibrillosa is similar by habitat and yellow-pigmented cystidia but can be separated by smaller spores, more acute cystidia, and lack of a membraneous annulus.

Additional specimens: The Faroes, Eysturoy, Selatrad, 22.VIII.1992, J. Vesterholt F3507, (C); Streymoy, Torshavn, Vidalundin, 31.VII.1988, J. Vesterholt 514, (C). Sweden, Jämtland, Kall, Lindvallen, 12.VIII.1996, S. Jacobsson 96004, (GB).

New recognized genera, species, and combinations

Kauffmania Örstadius & E. Larss., gen. nov. [MB811613] – Type: Kauffmania larga (Kauffman) Örstadius & E. Larss.

Diagnostic characters. – Basidiomata large, veil present, often connected to wood. Cap dark reddish brown, moist striate, hygrophanous, pallescent on drying. Gills crowded, adnate to adnexed, pale, becoming reddish brown, with white fimbriate edge. Stem rather firm. Spores medium-sized, pale, with indistinct to absent germ pore. Basidia or other cells in hymenium sometimes brown intracellular pigmented. Hymenial cystidia present. Pileipellis a hymeniderm sometimes with transition to a paraderm. Clamps present.

Etymology. – The genus name refers to the American mycologist Calvin Henry Kauffman who described the type species.

Note. – A monotypic genus recognized by large sized basidiomata, scanty veil, and pale spores.

Kauffmania larga (Kauffman) Örstadius & E. Larss., comb. nov. [MB811614] ≡ Psilocybe larga Kauffman, Publications Mich. geol. biol. Surv., Biol. Ser. 5 26: 279. 1918 – Holotype: U.S.A., Michigan, Washtenaw, Dexter Rd., 6.VI.1910, C. H. Kauffman (MICH!). Figs. 3e, 21.

Fig. 21

Kauffmania larga. C. H. Kauffman holotype; Pl Pleurocystidia; Ch Cheilocystidia; Basidioma ×0,5; Scale bars 10 μm

Selected description: Smith, Mem. N.Y. bot. Gdn 24: 125–126, 1972.

Cap 30–140 mm, at first hemispherical, then expanded convex to plane, sometimes umbonate, when young dark reddish brown to dark brown (Mu. 7.5YR 4/4), at maturity cinnamon, dark yellow brown to yellowish red (Mu. 5YR 5/6), sometimes glossy, when moist striate up to half-way from margin, hygrophanous, drying pale buff or cream, veil as white fibrils at margin, connecting to the stem in young fruitbodies. Gills adnate to adnexed, ventricose, crowded, L = 30–40, at first beige, becoming reddish brown to brown (Mu. 7.5YR 4/4), with white flocculose edge under a lens. Stem 40–110 × 4–15 mm, enlarged below, hollow, rather firm, whitish, pulverulent at apex, silky fibrillose below. Smell faint, sometimes like tomato leaves. Taste mild.

Spores 7.5–10 × 4.5–6.5 μm, av. 8–9.6 × 5–5.8 μm, Qav. = 1.5–1.7, ellipsoid, oblong to ovoid, sometimes slightly irregular in outline, in profile sometimes phaseoliform, reddish yellow (Mu. 5YR 6/8), germ pore indistinct to absent. Basidia 18–36 × 8–10 μm, 4-spored. Basidia or other cells in hymenium sometimes in part with brown intracellular pigmentation. Pleurocystidia 40–90 × 10–24 μm, mostly utriform, sometimes obtusely fusiform, lageniform to clavate, rarely forked or rostrate, numerous, walls yellow. Cheilocystidia of two types: A: 35–90 × 10–22 μm, similar to pleurocystidia, numerous to scattered, B: small, clavate, scattered to numerous. Pileipellis a hymeniderm sometimes with transition to a paraderm of 20–60 μm wide cells; pileitrama of pale to moderately pigmented hyphae. Veil cells 20–150 × 2–10 μm, cylindrical. Clamps present.

Habitat and distribution: Gregarious to scattered in deciduous or coniferous forests on rotten wood, wood chips, sticks or twigs, or directly on soil, in dry or moist places, on acid- or base-rich soil; summer to autumn; so far recorded from Denmark, Finland, Germany, Iceland, Norway, Sweden, and North America, but probably overlooked.

Notes – Kauffmania larga is recognized by its large size, scanty veil, and pale spores. The cystidia are large, mostly utriform and numerous. Psathyrella rostellata differs in having a well-developed veil in shape of large patches at cap margin, also reflecting in 4–20 μm wide veil cells. Moreover, the cystidia are lageniform to fusiform, at apex with a rostrum which often is bent or forked.

Selected specimens examined: Denmark: NE. Jylland. Høstemark Skov, S. of Mou, 1.X.1994, J. Vesterholt & S. Hansen (C, JV94-763). Finland: Etelä-Häme. Tammela, Mustiala, 1.VIII.1881, P. A. Karsten 4996 (H). Germany: Brandenburg. Um Buchwäldchen, 28.IX.2002, L. Örstadius 16–02. Iceland: S.-Múl.: Hallormsstaður, Ljósárkinn, 20.VI.1986, H. Hallgrímsson (AMNH, FA-13976). Norway: Akershus. Ås, Nordbyveien v/Dylterud, 2.VIII.1977, K. Østmoe 96/77 (O, as Psathyrella sp.). Sweden: Västergötland, Habo, N. of Sjöbol, 22.10.1999, E. Grundel LÖ197-99. USA: Michigan. Washtenaw, Dexter Rd., 6.VI.1910, C. H. Kauffman (MICH, holotype of Psilocybe larga).

Cystoagaricus

Murrill (1945) described the new species Nolanea strobilomyces Murrill. According to Singer (1947) the species was not nolanoid. He found it necessary to describe a new genus and transfer the species, hence Cystoagaricus strobilomyces (Murrill) Singer. The shape and dark colour of the spores combined with structure of gill edge and pileipellis made it impossible for Singer to identify the species in genera like Inocybe, Psathyrella or Agaricus. Singer found it more proper to place Cystoagaricus in Leucocoprinaceae than in Coprinaceae, the latter family corresponding to our days Psathyrellaceae. Four species belong to Cystoagaricus according to Index Fungorum. Horak (1980), however, recognized only one species, C. strobilomyces. He examined ten collections, among them the type, and gave a detailed description.

The phylogenetic position of Cystoagaricus strobilomyces in the present paper together with C. silvestris and allies is not surprising as the species have in common a brown squamulose cap surface, rather small, subrectangular, mitriform, or phaseoliform shaped spores, and utriform cystidia.

Some species described by Smith (1972) should in the near future be examined closer to see if they belong to Cystoagaricus.

Cystoagaricus Singer emend. Örstadius & E. Larss. – Type: Cystoagaricus strobilomyces (Murrill) Singer, Mycologia 39(1): 86, 1947. Fig. 22.

Fig. 22

Cystoagaricus. Spore shapes from five species; a. C. strobilomyces; b. C. squarrosiceps; c. C. olivaceogriseus; d. C. silvestris; e. C. hirtosquamulosus; Scale bars 10 μm

Basidiomata small to medium sized, non-deliquescent, lignicolous. Cap grey, brown, dark brown, sometimes olivaceous, fibrillose, squamulose, spiny, or squarrose, striate or not; veil present. Lamellae often crowded, adnate to adnexed, pale, becoming dark brown. Stem fibrillose, scaly, or with an annulus. Context sometimes blackening. Spores small to medium-sized, often irregular, subtriangular, subrectangular, or mitriform. Cheilo- and pleurocystidia present, often utriform or clavate.

Cystoagaricus strobilomyces (Murrill) Singer, Mycologia 39(1): 86, 1947. Holotype: USA, Florida, Murrill, F-16520 (FLAS).

Selected descriptions: Horak, Sydowia 33:64–70, 1980; Singer, Mycologia 39:85, 1947.

Collection examined: New Caledonia: Paita, Mt. Mou, 1200 m, 22.II.1977, Horak (ZT, 77/35). Fig. 22a.

Cystoagaricus squarrosiceps (Singer) Örstadius & E. Larss., comb. nov. [IF551032] ≡ Psathyrella squarrosiceps Singer, Nova Hedwigia 29:53. 1978 – Holotype: Ecuador, R. Singer B7276 (F!). Fig. 22b.

Selected description: Singer (l.c.).

Additional collection examined: Ecuador, Napo, Anangu, Rio Napo, 20.6–5.7.1983, T. Laessoe 44834 (C).

Cystoagaricus olivaceogriseus (A.H. Sm.) Örstadius & E. Larss., comb. nov. [IF551033] ≡ Psathyrella olivaceogrisea A.H. Sm., Mem NY bot Gdn 24:47. 1972 – Holotype: USA, Alaska, Wells-Kempton 8/15/63-5 (MICH!). Fig. 22c.

Selected description: Smith (l.c.).

Cystoagaricus silvestris (Gillet) Örstadius & E. Larss., comb. nov. [IF551034] ≡ Hypholoma silvestre Gillet, Hyménomycètes: 568. 1878 – Holotype (France, missing?). Fig. 3b.

Selected descriptions: Kits van Waveren, Persoonia, Suppl Vol 2:121, 1985 (as P. populina); Romagnesi, Bull Soc mycol Fr 98:62, 1982.

Selected collections examined: Finland: Ahvenanmaa. Jomala, Ramsholm, 27.IX.1980, M. Korhonen & R. Tuomikoski (H). Sweden: Skåne. Benestad, Örups almskog, 18.IX.2008, I. Månsson & L. Örstadius 163–08. Fig. 22d.

Cystoagaricus hirtosquamulosus (Peck) Örstadius & E. Larss., comb. nov. [IF551031] ≡ Agaricus hirtosquamulosus Peck, Bull. Buffalo Soc. Nat. Sci. 1: 53. 1873 – Holotype: USA, New York, C.H. Peck (NYS 1479!). Fig. 22e.

Selected descriptions: Musumeci, Boll Gr Micol G Bres 49:9–57, 2006. Smith, Mem NY bot Gdn 24:44, 1972.

Additional collections examined: Estonia: Heimtali, S. of Viljandi, 26.VIII.1989, J. Vesterholt 89–459 (C). Finland, Ahvenanmaa, Jomala, Ramsholm, 27.IX.1980, M. Korhonen & R. Tuomikoski (H).

Typhrasa Örstadius & E. Larss., gen. nov. [MB 811499] – Type: Typhrasa gossypina (Bull.) Örstadius & E. Larss.

Diagnostic characters. – Basidiomata medium-sized, with well-developed veil, growing on wood or on soil. Cap brown, striate or not, hygrophanous. Gills crowded, adnate to adnexed, pale, becoming brown, with white edge. Spores small to medium-sized, with colours from red to reddish yellow. Hymenial cystidia large, rostrate, and with intracellular oily drops or globules. Pileipellis with a hymeniderm or a paraderm. Clamps present.

Etymology. – The genus name is an anagram of Psathyra an illegitimate genus earlier containing many psathyrelloid species.

Typhrasa gossypina (Bull.: Fr.) Örstadius & E. Larss., comb. nov. [MB811616] ≡ Agaricus gossypinus Bull., Herb France 9:pl 425 Fig. 2, 1789 – Lectotype designated here: Bulliard, Herbier de la France 9:plate 425, Fig. 2, 1789. Figs. 3a, 23.

Fig. 23

Typhrasa gossypina. R.K. Schumacher 024; Pl Pleurocystidia; Ch Cheilocystidia; Basidioma ×1; Scale bars 10 μm

Selected descriptions: Arnolds, Fungi non delineati 26: 63, 2003; Kits van Waveren, Persoonia, Suppl. Vol. 2:251, 1985; Kühner & Romagnesi, Fl Analyt Champ Super:362, 1953.

Cap 10–70 mm, at first conical to campanulate, then convex, finally expanded plane with umbo, when young dark brown to dark reddish brown, striate at margin, hygrophanous, drying pale ochraceous brown, veil as fibres or flocci near the margin, appendiculate at margin. Gills broadly adnate, sometimes with tooth, medium spaced, L = 30–38, when young pale brown, becoming dark brown with white fimbriate edge. Stem 25–100 × 3–10 mm, equal or thickened towards base, whitish, pulverulent at apex, with fibrils and flocci from veil remnants below, sometimes with a fugacious, hanging, striate ring. Smell none or “etwas säuerlich-kakaoartig” and taste “mild” (R. K. Schumacher pers. comm.).

Spores 7–9 × 5–6 × 4.5–5.5 μm, av. 7.6–8.5 × 5–5.8 × 4.7–5.3 μm, Qav. = 1.4–1.6 and 1.5–1.7, ovoid, subfusiform, ellipsoid, oblong, broadly ellipsoid, in profile often amygdaloid or with suprahilar depression, sometimes subphaseoliform, yellowish red (Mu. 5YR 5/8), with distinct germ pore. Basidia 4-spored, 18–30 × 9–11 μm. Pleurocystidia 35–80 × 10–18 μm, fusiform to conical, often with a long rostrum, rarely clavate, with one, rarely two large internal oily drops, pale or rarely yellow pigmented below apex, numerous. Cheilocystidia of two types: A: similar in size, shape, and frequency to pleurocystidia, B: small, clavate, scattered to numerous; gill edge rarely faintly pigmented. Pileipellis a hymeniderm sometimes a transition to a paraderm of 15–55 μm wide cells; pileitrama made up of pale pigmented hyphae with scattered incrustations; an intracellular pigmented layer sometimes present between pileipellis and pileitrama. Veil cells 25–120 × 4–10(−20) μm, cylindrical to ventricose. Clamps present.

Habitat and distribution: Solitary, gregarious or caespitose in deciduous or coniferous forests with Betula, Fagus, Quercus, Picea or Pinus, on logs, twigs, wood chips and fireplaces; rarely reported from many countries in Europe and North America.

Notes. – Typhrasa gossypina is recognized by large rostrate cystidia with intracellular oily drop and rather wide spores. It lacks outstanding macroscopical features. The cap is usually 10–30 mm broad but Orton (1960) measured it up to 70 mm for his Psathyrella xanthocystis. T. gossypina is separated from P. fibrillosa by the intracellular oily drop and in having pale, long rostrate cystidia. It differs from T. nanispora in having larger spores.

Selected collections examined: Denmark: Sjaelland. Nørreskov, 24.IX.2011, T, Laessøe (C, TL2011-389713). England: Wiltshire, VII.1941, unknown collector (L). Finland: Sompion Lappi. Sodankylä, Pullinpuoti, Ammattikoulu, 24.VIII.1992, I. Kytövuori 92–910 (H). Germany: Brandenburg. E. Fürstenwalde, “Beerenbusch”, 13.IX.2003, R. K. Schumacher 024. The Netherlands: Drente. Gees, 28.I.1975, K. Booy (L). Norway: Oslo, Oslofjord, Gressholmen, 7.X.2013, A. Molia & T. Laessøe, AM241s-2013 (O). Portugal: Beira Litoral. Esmoriz, 1.I.1990, D. Olofsson LÖ 71–90. Sweden: Uppland. Värmdö, 3.XII.2000, H.-G. Toresson LÖ 3–01. Wales, Lake Vyrnwy, 10.IX.1979 (L, coll. Kits van Waveren).

Additional collections examined: USA: New York. Suffolk, Port Jefferson, Aug. 1904, G. F. Atkinson (NYS 983, holotype of Hypholoma delineatum); Washington. Pierce Co, Lower Tahoma, Mt. Rainer Nat’l Park, 9.VIII.1948, Smith 29994 (MICH, as Psathyrella canadensis).

Typhrasa nanispora Örstadius, Hauskn. & E. Larss., sp. nov. [MB 811500] – Holotype: Austria, Wien, Lainzer Tiergarten, Wienerblick, 6.VII.1998, T. Barta (GB!). Fig. 24. (S3: Figs. 20a–d.)

Fig. 24

Typhrasa nanispora. T. Barta 980706; Pl Pleurocystidia; Ch Cheilocystidia; Basidioma ×0,5; Scale bars 10 μm

Etymology. The epithet refers to the small spores.

Pileus 35–55 mm latus, convexus, flavo-brunneus, ferrugineus, carneus tinctus, hygrophanus sed non striatus, velum gossypino-squamulosum, ad marginem appendiculatum. Lamellae anguste adnatae, confertae, griseofuscae, acie albida. Stipes 45–55 × 8–13 mm, ad basim paulum incrassatus, subradicatus, pallide brunneus, fibrilloso-flocculosus brunneus vel olivaceo-brunneus obtectus. Sporae 5–6 × 3–4 μm, ovoideae, ellipsoideae, oblongae, phaseoliformes, sub microscopium pallidae, poro germinativo instructae. Basidia 4-sporigera. Pleurocystidia 40–75 × 12–18 μm, copiosa, fusiformia, clavata, rostrata, globus intracellularis praesens similis Typhrasa gossypina. Cheilocystidia 35–70 × 10–17 μm, similia pleurocystidiis, numerosa. Cellulae veli 30–100 × 4–14 μm. Fibulae adsunt. Habitat: Ad terram vel ad lignum subterreum.

Cap 35–55 mm broad, expanded convex with broad obtuse umbo, at centre yellow brown, rust brown, tan (Kornerup & Wanscher 6E8-6E6), towards margin distinctly paler, goldblond (5-6C4), with tinge of flesh colour, hygrophanous but not striate when collected, smooth to slightly rugose; veil as large, white cottony patches scattered on entire surface (not visible on the photo), partly appendiculate at margin. Gills adnexed, crowded, narrow, grey brown with white edge. Stem 45–55 × 8–13 mm, thickened towards base, sometimes almost bulbous and with a pseudorhiza (attached to buried wood?), pale brown, beige white at apex, slightly darker lower part, fistulose, covered with brown to olive brown fibrils and flocci. Flesh whitish, without smell. Spores 5–6 × 3–4 μm, mean-value 5.4 × 3.4 μm, Qav. = 1.6, ovoid, ellipsoid, oblong, in profile often phaseoliform, reddish yellow (Mu. 7.5YR 7/6-7/8); germ pore rather distinct. Basidia 4-spored, 14–27 × 5.5–6.5 μm. Pleurocystidia 40–75 × 12–18 μm, fusiform, clavate, at apex often with a rostrum 2–10 μm in length, with intracellular globule like in Typhrasa gossypina, abundant, with slightly yellow walls. Cheilocystidia 35–70 × 10–17 μm, similar to pleurocystidia, numerous. Pileipellis a paraderm of subglobose, clavate, or irregular shaped 15–30 μm wide cells. Pileitrama of moderately yellow brown pigmented hyphae incrusted or not; gill trama rather pale. Veil cells 30–100 × 4–14 μm. Clamps seen on hyphae of stem and veil.

Habitat and distribution: On bare soil, maybe attached to buried wood, in a forest dominated by deciduous trees. So far only known from the type locality in Austria.

Notes - Typhrasa nanispora is recognized by rather large basidiomata, well-developed veil, small and pale spores, and abundant fusiform, rostrate cystidia equipped with an intracellular globule. The closely related Typhrasa gossypina also has cystidia with globules but differs in having larger and darker spores. Psathyrella laevissima, P. maculata, P. obscurotristis, P. piluliformis, all with spores less than 6 μm in length, differ in having smaller, less frequent cystidia lacking intracellular globules. In addition, the basidiomata of P. laevissima and P. obscurotristis are smaller. P. maculata has a white soon black discolouring veil.

Homophron (Britzelm.) Örstadius & E. Larss., gen. stat. nov. [MB811630] ≡ Agaricus subgenus Homophron Britzelm., Ber naturhist Ver Augsburg 27:174, 1883 ≡ Homophron (Britzelm.) W. B. Cooke, Gen Homobas 45:100, 1953, (nom. inval., no basionym cited) – Type: Agaricus spadiceus Schaeff., Fung. Bavariae 4:27, 1774, selected by Singer, Lilloa 22:468, 1951 ≡ Psilocybe spadicea P. Kumm., Führ Pilzk (Zerbst):71, 1871.

Description: Basidiomata small to large, fragile to rather rigid, non-deliquescent, lignicolous. Pileus brown, striate, hygrophanous. Lamellae close to crowded. Stem central. Veil absent. Spores small to medium sized, moderately pigmented to hyaline, with indistinct to absent germ pore. Basidia 4-spored. Pleurocystidia often thick-walled and with crystals or incrustations at apex. Pileipellis a hymeniderm or transition to a paraderm. Clamps present.

Notes – The combination of absence of veil and presence of thick-walled cystidia with crystals is unique in Psathyrellaceae.

Homophron spadiceum (P. Kumm.) Örstadius & E. Larss. comb. nov. [MB811631] ≡ Agaricus spadiceus Schaeff., Fung Bavariae 4:27, 1774 (nom. inval., non Agaricus spadiceus Scop. 1772. Psilocybe spadicea P. Kumm. in Führ Pilzk (Zerbst): 71,1871. – Lectotype (designated by Örstadius in Windahlia 24:21. 2001): Schaeffer, Fung Bavariae 1, pl. 60, Figs. 4 and 6, 1762 – Epitype (designated by Örstadius in Windahlia 24:21. 2001): Germany, Bavaria, M. Enderle (GB!, isoepitype M!, GenBank no. DQ389729). Figs. 3c, d, 25.

Fig. 25

Homophron spadiceum. M. Enderle epitype; Pl Pleurocystidia; Ch Cheilocystidia; H Hairs from cap margin; Basidioma ×1; Scale bars 10 μm

Selected descriptions: Enderle, Beitr Kenntn Pilze Mitteleur 5:58–59, 1989; Kits van Waveren, Persoonia, Suppl Vol 2:161–164, 1985; Kühner & Romagnesi, Fl Analyt Champ Super:369, 1953; Ludwig, Pilzkompendium Band 2:644, 2007; Örstadius, Windahlia 24:19–24, 2001.

Cap 25–120 mm, convex, with inflexed margin, becoming plane, sometimes wavy, at first dark reddish brown, red brown (Mu. 2.5YR 3/4-4/4, 5YR 4/4), sometimes striate at margin, faintly hygrophanous, at maturity or when drying fading to pale ochraceous brown, flesh-coloured or pink (Mu. 5YR 5/8-8/4, 7.5YR 8/4), at margin covered with minute hairs (lens); veil absent. Gills adnate, very crowded, L = 60–82, when young very pale brown (Mu. 10YR 8/3), then brown with red tinges (Mu. 5YR 5/4, 7.5YR 5/4-5/6) and with a slightly flocculose edge. Stem 25–100 × 3–12 mm, rigid, equal or slightly thickened towards base, pale brown, flocculose at apex or half-way down, fibrillose below. Context rather firm. Taste mild and smell pleasant. Spore print dark reddish brown (Mu. 5YR 3/3).

Spores 6.5–11 × 4–5.5 μm, av. 7.2–10.3 × 4.1–5.3 μm, Qav. = 1.7–1.9, ovoid, oblong, or subcylindrical, sometimes irregular in outline, in profile sometimes phaseoliform, rarely amygdaloid, thin-walled, very pale brown (Mu. 10YR 8/4), germ pore absent or rarely indistinct. Basidia 4-spored, 16–36 × 6–8 μm. Pleurocystidia 30–80 × 10–24 μm, lageniform, fusiform to conical, often rostrate with apex covered with crystals, wall up to 3 μm thick near apex, yellow or not, abundant, rarely rather scattered. Cheilocystidia of two types: A: 40–85 × 8–22 μm, similar to pleurocystidia, scattered to abundant, B: 30–90 × 6–28 μm, clavate, utriform or cylindrical, scattered or rarely numerous. Caulocystidia present at apex and similar to shape of cheilocystidia. Hairs at cap margin originating from globose cells of pileipellis, septate or not; terminal cells of the hairs 40–180 × 6–18 μm, versiform, cylindrical, clavate, lageniform etc., rarely similar to pleurocystidia. Pileipellis a hymeniderm with transition to a paraderm of 12–40 μm wide cells; pileitrama of moderately pigmented hyphae, rarely incrusted; hymenophoral trama of membranal pigmented hyphae. Clamps present on stem hyphae.

Habitat and distribution: Usually caespitose around deciduous trees and conifers, both stumps and at the base of living trees, e.g., Abies pinsapo, Acacia xanthophloea, Aesculus, Alnus, Bambusa, Betula, Castanea, Fagus, Morus alba, Picea, Pinus, Populus, Quercus faginea, and Ulmus; known from many countries in and outside Europe.

Notes – The species is distinguished by the absence of veil, hairy cap margin, very pale spores, and caespitose growth. Moreover, the rostrate cystidia are often thick-walled and provided with crystals. H. cernuum differs in having paler basidiomata, smaller and darker spores, and obtuse cystidia.

Selected collections examined: Belgium: Antwerpen. Retie, Prinsenpark, 11.XI.1993, Volders 93222 (BR, holotype of Psathyrella imleriana). Finland: Ahvenanmaa. Jomala, Prästgården Ramsholmen, 24.IX.1994, J. Vauras 9754 F (TURA). Germany: Bayern. Leipheim-Riedheim, 11.XI.1998, M. Enderle (GB, epitype of Psathyrella spadicea; M, isoepitype). The Netherlands: Overijssel. Delden, 20.X.1975 (L, coll. Kits van Waveren, as Psathyrella variata). Sweden: Skåne. Kristianstad, S:a Lingenäset, 11.IX.2003, L. Örstadius 75–03. USA: Idaho. Bonner Co., Priest Lake, 6.X.1964, A. H. Smith 71092 (MICH, holotype of Psathyrella variata). New York. North Elba, Aug., C. H. Peck (NYS 2690, holotype of Agaricus saccharinophilus).

Homophron cernuum (Vahl: Fr.) Örstadius & E. Larss., comb. nov. [MB811632] ≡ Agaricus cernuus Vahl in Fl Danic 6(17):9, pl. 1008, Fig. 1, 1790. ≡ Agaricus cernuus Vahl: Fr., Syst mycol (Lundae) 1:298, 1821. – Holotype missing (C). – Lectotype (designated here): Agaricus cernuus Vahl, Fl Danic 6(17):9, pl. 1008, Fig. 1, 1790. – Epitype (designated here): Denmark, Sjaelland, Risø, Boertmann 8.XI. 2013 (GB!, GenBank no. KJ939632)

Note: The illustration Vahl refers to when describing the species is ambiguous and therefore an epitype was selected. The ITS sequence of the type specimens of P. crenulata A.H. Sm (MICH!, KC992957) was shown to be identical with the ITS of Homophron cernuum. The two species are also congruent in morphology, so we regard P. crenulata as a later synonym.

Selected descriptions: Gröger, Boletus 1: 1, 1984; Kits van Waveren, Persoonia, Suppl Vol 2:76, 1985; Kühner & Romagnesi, Fl Analyt Champ Super:368, 1953; Ludwig, Pilzkompendium Band 2:642, 2007.

Selected collections examined: Denmark, Sjaelland, Risø, 8.XI.2013, D. Boertmann (GB, epitype, ATLAS DB2013-652266). Finland, Varsinais-Suomi. Dragsfjärd, Örö, Örö island, 14.X.1999, J. Vauras 15664 (TURA 8633). Germany, Bayern. Goaslweide, 12.X.2002, P. Karasch (herb. Karasch G/109-02). Norway, Oppland. S. Land hd, Odnes hotell, 24.IX.1977, J. Stordal 18841 (O, as Psathyrella sp.). Sweden, Dalarna. Falun, Önsbacken, 28.X.1977, R. Morander 2843 (S).

Homophron camptopodum (Sacc.) Örstadius & E. Larss., comb. nov. [MB811633] ≡ Agaricus camptopus Peck, Ann Rep NY State Mus 31:35. 1879 (nom. illegit., non Agaricus camptopus Berk., 1850). Psilocybe camptopoda Sacc., Syll Fung 5:1057. 1887; Psathyrella camptopoda (Sacc.) A.H. Sm., Mycologia 42:127. 1950. – Holotype: USA, New York, Sept, Peck (NYS!).

Selected description: Smith, Mem. N.Y. bot. Gdn 24: 245, 1972.

Collections examined: Russia, Kamchatka, Avacha River Valley, N.E. of Razdolny, 31.VII.1997, T. Laessoe (IB 1997/956). Kamchatka, Esso, Bystraya River Valley, 8 km. N.E. of Esso, 9.VIII.1997, U. Peintner (IB 1997/957). USA, Idaho, Bonner Co, Priest Lake, 11.X.1956, A.H. Smith 54450 (MICH). New York, Catskill Mountains, Sept., C.H. Peck (NYS 617, holotype of Agaricus camptopus). New York, Savannah, October, C.H. Peck (NYS 3290, holotype of Psilocybe unicolor). Washington, Jefferson Co, 30.VI.1939, A.H. Smith 14719 (MICH).

Coprinopsis canoceps (Kauffman) Örstadius & E. Larss., comb. nov. [IF551035] ≡ Hypholoma canoceps Kauffman, Pap Mich Acad Sci 5:132. 1926 – Holotype: USA, Oregon, Oct 1922, Kauffman (MICH!).

Coprinopsis cineraria (Har. Takah.) Örstadius & E. Larss., comb. nov. [IF551036] ≡ Psathyrella cineraria Har. Takah., Mycoscience 41(1):18. 2000 – Holotype: Japan, Takahashi (CBM FB-24142!, GenBank no. KC992962).

Coprinopsis melanthina (Fr.) Örstadius & E. Larss., comb. nov. [IF551037] ≡ Agaricus melanthinus (“melantinus”) Fr., Öfvers K Vetensk Akad Förh 8:49. 1852, “1851” – Holotype lacking.

Note: The species was described by Fries from a Swedish collection. Since then no further records have been reported in Sweden and therefore we await to typify the species.

Coprinopsis submicrospora (Heykoop & G. Moreno) Örstadius & E. Larss., comb. nov. [IF551038] ≡ Psathyrella submicrospora Heykoop & G. Moreno, Mycotaxon 83:426. 2002 – Holotype: Spain, Heykoop & Villarreal (AH 27055!, GenBank no. KC992959).

Coprinopsis uliginicola (McKnight & A.H. Sm.) Örstadius & E. Larss., comb. nov. [IF551038] ≡ Psathyrella uliginicola McKnight & A.H. Sm., Mem NY bot Gdn 24:100. 1972 – Holotype: USA, Wyoming, Smith 34903 (MICH!, GenBank no. KC992960).

Neotypes selected for species described by Fries

Parasola conopilus (Fr.) Örstadius & E. Larss., Mycol. Res. 112(10):1180, 2008 ≡ Agaricus conopilus Fr., Syst mycol (Lundae) 1:504, 1821 – Neotype (designated here) [MBT200684]: Sweden, Skåne, Ivetofta, Krogstorp, 2002-10-22, L. Örstadius 186–02 (GB!, GenBank no. DQ389725).

Psathyrella candolleana (Fr.) Maire apud Maire, Mém. Soc. Sci. Nat. Maroc. 45:112, 1937 ≡ Agaricus candolleanus Fr., Observ. mycol. (Havniae) 2:182, 1818 – Neotype (designated here) [MBT200683]: Sweden, Bohuslän, Lysekil, Bro, SO Kibbestens torvemyr, 1973-08-17, L. & A. Stridvall 73/030 (GB!, GenBank no. KM030175)

Psathyrella caput-medusae (Fr.) Konrad & Maubl., Encyclop. Mycol. 14: 127, 1949 [1948] ≡ Agaricus caput-medusae Fr., Epicr. syst. mycol. (Upsaliae):216, 1838 [1836–1838] – Neotype (designated here) [MBT200681]: Sweden, Västergötland, Trollhättan, Lextorp, 1993-08-30, L. & A. Stridvall 93/062 (GB!, GenBank no. KM030174)

Psathyrella fatua (Fr.) Konrad & Maubl., Encyclop. Mycol. 14(1):125, 1949 [1948] ≡ Agaricus stipatus y. fatuus (Fr.) Fr., Syst. mycol. (Lundae):296 (1821). – Neotype (designated here) [MBT 200685]: Sweden, Skåne, Österslöv, Lövhall, 1997-10-11, L. Örstadius 132–97 (GB!, GenBank no. DQ389681).

Psathyrella noli-tangere (Fr.) A. Pearson & Dennis, Trans. Br. mycol. Soc. 31(3–4): 184, 1948 [1947]. ≡ Agaricus noli-tangere Fr., Epicr. syst. mycol. (Upsaliae):234, 1838 [1836–1838] – Neotype (designated here) [MBT200686]: Sweden, Skåne, V. Vram, Stackedala, 2003-09-14, L. Örstadius 83–03 (GB!, GenBank no. DQ389713).

Psathyrella pennata (Fr.) A. Pearson & Dennis, Trans. Br. mycol. Soc. 31(3–4):184, 1948 [1947]. ≡ Agaricus pennatus Fr., Syst. mycol. (Lundae) 1:297, 1821 – Neotype (designated here) [MBT200687]: Sweden, Skåne, Kiaby, Kjugekull, 1984-10-01, L. Örstadius 216–84 (GB!, GenBank no. KJ939633).

Psathyrella prona (Fr.) Gillet, Hyménomycètes (Alençon): 618, 1878 ≡ Agaricus pronus Fr., Epicr. syst. mycol. (Upsaliae):239, 1838 [1836–1838]. – Neotype (designated here) [MBT200688]: Sweden, Skåne, Trolle-Ljungby, Fårabäck, 2000-10-17, L. Örstadius 237–00 (GB!, GenBank no. KJ939634).

Species new to Europe and/or to the Nordic countries

Psathyrella hololanigera (G.F. Atk.) A.H. Sm., Mem NY bot Gdn 24:130. 1972 ≡ Hypholoma hololanigerum G.F. Atk., Annls mycol 7:371. 1909. Fig. 26. (S3: Fig. 21a–d.)

Fig. 26

Psathyrella hololanigera. G.F. Atkinson 15192; Pl Pleurocystidia; Ch Cheilocystidia; Scale bars 10 μm

Cap 18–30 mm broad, up to 10 mm high, flat convex, sometimes with low umbo, fresh brown, yellowish brown, near the margin paler, and here soon drying to ochraceous, brownish-ochre, hygrophanous, striate; veil strongly devoloped as squamules or flocci, appendiculate at margin. Gills distant, adnate, pale brown, becoming moderately dark brown, with concolorous edge. Stem up to 80 × 3 mm, white to whitish, with fibrillose remnants from the white veil. Context without smell.

Spores 7.5–9.5 × 4–5 μm, av. 8–8.7 × 4.3–4.5 μm, Q = 1.9, oblong with acute to obtuse base, in profile flattened on adaxial side, amygdaliform or with a distinct suprahilar depression, phaseoliform, with oil drop, in water yellowish red (Mu. 5YR 5/8-6/8), with distinct germ pore. Basidia 4-spored, 17–28 × 8–10 μm. Pleurocystidia 35–60 × 11–20 μm, narrowly utriform, subcylindrical, lageniform, pale, numerous. Cheilocystidia of two types: A: 25–40 × 8–11 μm, similar to pleurocystidia, scattered near cap margin, then increasing and becoming numerous, B: small, clavate to sphaeropedunculate, numerous close to cap margin, then decreasing and becoming scattered. Scalp cap half-way from margin: pileipellis with 20–30 μm wide, pale, ellipsoid to subglobose cells; pileitrama with strongly pigmented hyphae. Hymenophoral trama with rather pale hyphae. Veil cells on cap 30–125 × 5–22 μm. Clamps seen on veil and stem hyphae.

Habitat and distribution: Solitary or in a small group on leaf litter in pine forest on sandy soil with Quercus ilex, Rhuscus, Phillyria etc., in the USA on strongly decayed wood in woods. Known from single collections in Italy and North America.

Specimens examined. Italy, Emilia-Romagna, Ravenna, Pineta di Classe, 9.XI.2007, A. Hausknecht (WU). USA, New York, Ithaca, 9.VII.1903, G.F. Atkinson 15192 (CUP!, holotype of Hypholoma hololanigerum).

Additional specimen examined. The Netherlands, Drenthe, Anloo, Plot Q65, 10.XI.1987, P.-J. Keizer 87324 (WAG-W!, holotype of Psathyrella rhombispora P.-J. Keizer).

Notes - The species can be recognized by the copious veil, the spores with a distinct suprahilar depression and the cystidia shape. The examined type agrees satisfactory with the European find. Psathyrella kitsiana differs in having smaller spores, 6.5–8 × 3.5–5 μm, av. 7.2–7.8 × 3.9–4.8 μm and moreover the germ pore is lacking or indistinct. The two species differ molecularly. Psathyrella rhombispora (Keizer 1993) is difficult to separate from the two mentioned species. The spores of the examined type lacked a germ pore and measured 7.8–9 × 4.2–5 μm, mean-value 8.3 × 4.6 μm. The collector observed no velar remains on the cap but some flocci on the stem. P. rhombispora should be sequenced to get information of its phylogenetic position.

Psathyrella parva A.H. Sm., Mem NY Bot Gdn 24:433. 1972. Fig. 27. (S3: Fig. 22b–d.)

Fig. 27

Psathyrella parva. Contu 071227; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Cap 2–15 mm broad, when young semiglobose, then expanded convex, moist chestnut-brown, date-brown, dark brown, subfulvous, dingy tan, then cinnamon-buff, hygrophanous, faintly striate, drying to buff or pinkish buff; surface covered by white flocci or fibrils from veil remants, soon disappearing. Gills adnate to adnexed, distant L = 12–15, ventricose, at first ochraceous or pale brown, becoming avellaneous to greyish-brown with paler edges. Stem 10–35 × 0.3–1.5 mm, equal, delicate, fragile, white overall, pruinose above, fibrillose below. No smell or taste present. Spore print brown.

Spores 6–8 × 4–5 μm, av. 6.2–7.4 × 3.9–4.6 μm, Qav. = 1.4–1.7, ellipsoid, ovoid, with obtuse poles, in profile rarely subamygdaliform or subphaseoliform, in water yellowish red (Mu. 5YR 5/8-6/8); germ pore indistinct to absent. Basidia 4-spored, 12–23 × 6–9 μm. Pleurocystidia 25–60 × 7–13 μm, narrowly lageniform, conical, fusiform, with acute to obtuse apex, rarely bent or forked, with pale to slightly yellow walls, scattered to numerous. Cheilocystidia of two types: A: 25–55 × 5–14 μm, similar to pleurocystidia, numerous to scattered, B: small, clavate, scattered, absent to rather numerous. Scalp cap halfway from margin: pileipellis with subglobose to ellipsoid 15–50 μm wide cells, pale; pileitrama with incrusted hyphae. Veil cells 15–90 × 4–16 μm, pale, clamped.

Habitat and distribution: Solitary to gregarious on sand in wasteland near scrub oak, in open calcareous grassland or on acid sand in a coastal grassland near the sea; autumn to nearly winter; known from Denmark, Italy, Norway, Sweden, and the USA.

Notes - Psathyrella parva is recognized by small basidiomata, a rather scanty veil, small spores, indistinct germ pore, small often acute cystidia, and a habitat on sandy soil. Also P. seymourensis grows on sand but can be separated by slightly larger basidiomata, slightly larger spores, and a distinct germ pore. More material is needed to separate the two species more clearly. P. umbrina differs in having larger basidiomata and paler spores.

Specimens examined. Denmark, E. Jylland, Vorsø, Horsens Fjord, 1.XI.1982, T. Laessoe TL-Vorsø-854, (C); Italy, Sardinia, Olbia, Pittulongu, 27.XII. 2007, M. Contu, (GB); Norway, Telemark, Bamble, Langøya, 10.X.2012, A. Molia & T. Laessøe AM-330 g-2012, (O 245716); Bamble, Langøya, 15.X.2013, A. Molia & T. Laessøe AM-263a-2013, (O 22035); Bamble, Eikstrand, 13.X.2012, A. Molia & T. Laessøe AM-336 g-2012, (O 245581); Sweden, Medelpad, Stöde, Vigge, 12.IX.1995, E. Malm LÖ81-95, (GB); Skåne, Åhus, Älleköpingegård, 14.X.1995, L. Örstadius 211–95; USA, Michigan, Washtenaw, Crooked Lake, Waterloo State Recreation Area, 20.X.1971, A.H. Smith 81026, (MICH! holotype).

Psathyrella tenera Peck, Rep NY St Mus nat Hist 47:144. 1894. Fig. 28. (S3: Fig. 23b–d.)

Fig. 28

Psathyrella tenera. L. Örstadius 81–83; Pl Pleurocystidia; Ch Cheilocystidia; Basidiomata ×1; Scale bars 10 μm

Cap 5–15(−20) mm broad, at first semiglobose to paraboloid, then expanded conico-convex or convex, dark reddish brown (Mu. 2.5YR 3/4-2.5/4), dark brown, when moist striate at margin, hygrophanous, drying to reddish brown and later to ochraceous grey, cream or whitish; veil when young covering entire surface with dispersed flocci, evanescent. Gills broadly adnate to narrowly adnate with tooth, distant, L = 13–14, when young pale ochraceous grey, grey brown, becoming dark brown, with white fimbriate edge, faintly to distinctly red underlined. Stem 30–70 × 0.7–2 mm, cylindrical, slightly thickened towards base or subbulbous, sometimes curved, pale brown to ochraceous, towards base brown, covered with white veil remnants, pulverulent at apex. Taste not recorded. Smell faint.

Spores 12–17 × 6–7.5 μm, av. 13–14.7 × 6.3–7.1 μm, Qav. 1.9–2.2, subcylindrical, oblong, narrowly limoniform, in profile with faint to distinct suprahilar depression, amygdaliform, in water red to dark red (Mu. 2.5YR 4/6-4/8-3/6), with distinct to very distinct germ pore. Basidia 4-spored, 18–34 × 10–14 μm. Pleurocystidia 30–65 × 10–16 μm, lageniform to conical with often narrow and pointed upper part, sometimes flexuous, scattered, sometimes rather numerous, pale. Cheilocystidia of two types: A: 25–50 × 9–17 μm, similar to pleurocystidia, numerous to scattered, B: small, clavate, numerous especially close to cap margin. Scalp cap halfway from margin: pileipellis made up of subglobose, ellipsoid to clavate cells 15–50 μm wide, pale to yellow. Pileitrama of heavily or sometimes moderately encrusted hyphae; gill trama strong to pale pigmented. Veil cells 15–60 × 4–14 μm. Clamps seen at stem and veil hyphae.

Habitat and distribution: Psathyrella tenera grows single to subcaespitose on wet or moist ground in deciduous forest or open grassland in calcareous areas, in fens or close to rivulets, directly on earth or attached to decaying leaves, woody debris and rotten straw. So far known from Austria, Norway, Sweden, and North America.

Notes - In the original description nothing is said about a veil on the cap but other characters agree satisfactory with the examined European material. In the type material the spores were seen with suprahilar depression and a small papillate apex. Other features of the species are the small basidiomata, copious veil, large spores, and a moist habitat. It comes close to Psathyrella potteri from which it differs in having a darker, striate cap with well-developed veil. Moreover, P. potteri prefers to grow in dry, nutrient rich places. P. prona can be separated by 2-spored basidia, a scanty veil, and dry habitat.

Specimens examined. Austria, Wien, Mauerbach, 11.IX.1990, E. Ludwig; Norway, Oppland, Lunner, Søndre Oppdalen øvre, 14.VI.1978, T. Brandrud 18–78, (O, 171816); 25.VI.1978, T. Brandrud 30–78 (O, 171815); Svalbard, Barentsburg, 15.VIII.1988, G. Gulden 100/88, (O, 74539); Sweden, Skåne, Baldringe, Lyckås, 1.X.1989, L. Örstadius 382–89, (GB); 25.V.2011, L. Örstadius 24–11; Ivetofta, Gyetorp, 12.VI.2000, A. Ryberg LÖ18-00; Kristianstad, Björket, 5.VII.1992, L. Örstadius 143–92; Maglehem, Blåherremölla, 2.VIII.1990, L. Örstadius 158–90; Ravlunda, Haväng, 10.VII.1983, L. Örstadius 81–83, (GB); USA, New York, Jefferson Co., Pierrepont Manor, June Peck, (NYS3146!, holotype).