Fourteen coprophilous species of Psathyrella identified in the Nordic countries using morphology and nuclear rDNA sequence data

Ellen Larsson; Leif Örstadius

Novel species of fungi described in the present study include the following from Australia: Neoseptorioides eucalypti gen. & sp. nov. from Eucalyptus radiata leaves, Phytophthora gondwanensis from soil, Diaporthe tulliensis from rotted stem ends of Theobroma cacao fruit, Diaporthe vawdreyi from fruit rot of Psidium guajava, Magnaporthiopsis agrostidis from rotted roots of Agrostis stolonifera and Semifissispora natalis from Eucalyptus leaf litter. Furthermore, Neopestalotiopsis egyptiaca is described from Mangifera indica leaves (Egypt), Roussoella mexicana from Coffea arabica leaves (Mexico), Calonectria monticola from soil (Thailand), Hygrocybe jackmanii from littoral sand dunes (Canada), Lindgomyces madisonensis from submerged decorticated wood (USA), Neofabraea brasiliensis from Malus domestica (Brazil), Geastrum diosiae from litter (Argentina), Ganoderma wiiroense on angiosperms (Ghana), Arthrinium gutiae from the gut of a grasshopper (India), Pyrenochaeta telephoni from the screen of a mobile phone (India) and Xenoleptographium phialoconidium gen. & sp. nov. on exposed xylem tissues of Gmelina arborea (Indonesia). Several novelties are introduced from Spain, namely Psathyrella complutensis on loamy soil, Chlorophyllum lusitanicum on nitrified grasslands (incl. Chlorophyllum arizonicum comb. nov.), Aspergillus citocrescens from cave sediment and Lotinia verna gen. & sp. nov. from muddy soil. Novel foliicolous taxa from South Africa include Phyllosticta carissicola from Carissa macrocarpa, Pseudopyricularia hagahagae from Cyperaceae and Zeloasperisporium searsiae from Searsia chirindensis. Furthermore, Neophaeococcomyces is introduced as a novel genus, with two new combinations, N. aloes and N. catenatus. Several foliicolous novelties are recorded from La Réunion, France, namely Ochroconis pandanicola from Pandanus utilis, Neosulcatispora agaves gen. & sp. nov. from Agave vera-cruz, Pilidium eucalyptorum from Eucalyptus robusta, Strelitziana syzygii from Syzygium jambos (incl. Strelitzianaceae fam. nov.) and Pseudobeltrania ocoteae from Ocotea obtusata (Beltraniaceae emend.). Morphological and culture characteristics along with ITS DNA barcodes are provided for all taxa.

mycological research 112 (2008) 1165–1185 journal homepage: www.elsevier.com/locate/mycres Fourteen coprophilous species of Psathyrella identified in the Nordic countries using morphology and nuclear rDNA sequence data Ellen LARSSONa,*, Leif ÖRSTADIUSb a Deptartment of Plant and Environmental Sciences, University of Gothenburg, Box 461, SE-405 30 Göteborg, Sweden Lyckans väg 39A, SE 291 43 Kristianstad, Sweden b article info abstract Article history: Psathyrella species growing on dung or occasionally on dung in the Nordic countries were Received 31 October 2007 studied using morphological characters and nu-rDNA sequence data and type collections Received in revised form were examined when available. Fourteen species capable of growing on dung were identi- 19 March 2008 fied. Descriptions are given of all dung-inhabiting species and to a lesser extent of the spe- Accepted 1 April 2008 cies occasionally growing on dung. Three new species are described: Psathyrella fimiseda, Corresponding Editor: P. merdicola, and P. scatophila. P. stercoraria is described as a new species in order to validate David L. Hawksworth the name. A key to the coprophilous species in Europe including the species described by Peck & Smith from North America is provided. The phylogenetic analyses recovered four Keywords: major supported clades within Psathyrellaceae corresponding to Parasola, Coprinopsis, Lacry- Agaricales maria/Spadiceae pro parte, and Psathyrella. The status of Coprinellus was ambiguous. The Basidiomycota current morphology-based infrageneric classification of Psathyrella was not supported by Molecular systematics the phylogenetic analyses and a coprophilous habit has apparently evolved on multiple Phylogeny occasions. Three new combinations are proposed: Parasola conopilus, Coprinopsis marcescibi- Taxonomy lis, and Coprinopsis pannucioides. ª 2008 The British Mycological Society. Published by Elsevier Ltd. All rights reserved. Introduction The genus Psathyrella was traditionally placed in the family Coprinaceae. Molecular phylogenetic studies based on nurDNA sequences have considerably altered the understanding of the relationships within coprinoid fungi (Hopple & Vilgalys 1999; Park et al. 1999; Moncalvo et al. 2000, 2002). These studies demonstrate that Coprinus is non-monophyletic. Although the majority of species form a large clade (/psathyrellaceace in Moncalvo et al. 2002), the type species, C. comatus, belongs to the lepiotoid fungi in Agaricaceae (Vellinga 2004). A new classification recognizes Psathyrellaceae (Redhead et al. 2001a) as a separate family. Psathyrellaceae comprises six genera in Europe, viz. Psathyrella (Fr.) Quél. 1872, Lacrymaria Pat. 1887, Coprinellus P. Karst 1879, Parasola Redhead, Vilgalys & Hopple 2001, Coprinopsis P. Karst. 1881, and the small genus Macrometrula Donk & Singer 1948. However, some limits and subdivision of genera are still unresolved, especially around Psathyrella, Coprinopsis, and Coprinellus. Several recent molecular studies on these genera have indicated that Psathyrella is non-monophyletic (Keirle et al. 2004; Walther et al. 2005; Matheny et al. 2007; Padamsee et al. 2008). The species in Psathyrella have a saprotrophic habit and most species occur on soil or on wood. However, a number of them grow on dung and a few occur on old bonfires or in swamps. The ecological specialization to dung of various animals, such as cow, horse, * Corresponding author. Tel.: þ46 31 7862662. E-mail address: ellen.larsson@dpes.gu.se 0953-7562/$ – see front matter ª 2008 The British Mycological Society. Published by Elsevier Ltd. All rights reserved. doi:10.1016/j.mycres.2008.04.003 1166 and wild boar, is widespread among species in Coprinopsis, Coprinellus, and Psathyrella. Psathyrella was first introduced as a tribus, within Agaricus (Fries 1838) and a few years later reclassified as a subgenus (Fries 1849). Fries included ten species, but only four of them are currently recognized in Psathyrella. The sole species for which Fries mentions dung as a substrate is Agaricus subtilis (Fries 1821), a name adopted by few later authors. Kirchner & Eichler (1894) mentioned it from Germany and also Massee (1902) used the name and gave the spore measurement 12– 16 5–8 mm. Romagnesi (1937) reported Drosophila subtilis as ‘assez rare’ from France, but did not include it in his later works. The interpretation of the name is unclear, but the description recalls a tiny Psathyrella growing on dung in moist forest (‘in fimo locis udis silvaticis’). Kühner & Romagnesi (1953) described two coprophilous species in Drosophila (syn. Psathyrella), viz. D. coprobia (syn. P. hirta) and D. stercoraria, both referred to subgenus Psathyrella, but within the subgenus placed in sections Atomatae and Pronae, respectively. Smith (1972) reported five species as exclusively growing on dung in North America (P. galericolor, P. minima, P. argentata, P. equina, and P. sphaerocystis) and nine species associated with dung or manured, fertilized soil (P. pseudolimicola, P. pratensis, P. odorata, P. nezpercii, P. potteri, P. pruinosipes, P. prona, P. hirta, and P. conopilus). With the exception of P. sphaerocystis and P. conopilus, all species were classified in the subgenus Psathyrella sect. Atomatae, a section containing coprophilous species and species growing on well-fertilized soils. Psathyrella conopilus was placed in subgenus Psathyrella section Subatratae, a sect. distinguished by the absence of pleurocystidia. P. sphaerocystis was placed in subgenus Cystopsathyra on the strength of the powdery veil with cells mostly in the form of sphaerocysts (Smith 1972). In the systematic arrangement of Psathyrella by Romagnesi (1982) the taxa Atomatae and Pronae were united to sect. Atomatae. P. cobrobia was placed in subgenus Psathyrella sect. Bipelles together with the non-coprophilous P. bipellis (syn. P. odorata). Kits van Waveren (1985) listed five dung-inhabiting species from France, the UK, and The Netherlands, viz. P. waverenii, P. hirta, P. stercoraria, P. coprophila, and P. sphaerocystis. In addition, P. prona var. prona f. cana, normally a non-coprophilous taxon, was recorded from horse dung. In the systematic arrangement P. sphaerocystis was placed in subgenus Psathyra section Cystopsathyra, whereas all other species, including P. prona var. prona f. cana, were placed in subgenus Psathyrella section Atomatae. In literature from the Nordic countries (e.g. Hansen & Knudsen 1992) P. coprophila and P. hirta are often the only species mentioned as dung-inhabiting. Lange (1936) described Psathyra gordonii f. minor with pale cap colour, growing on grass, with a spore size of 10.5–12.5 5.5–6 mm and obtusely lageniform cystidia, characters reminiscent of Psathyrella marcescibilis. Besides growing on grass, Lange also remarked in brackets ‘with remnants of horse-dung’, a habitat not known for P. marcescibilis. Lange (1939) also reported Psathyra cobrobia (syn. Psathyrella hirta) from dung. A species of uncertain identity is Psathyrella polaris collected by H. G. Simmons on dung of muskox (Bovis maschatis) in northern Canada during the Second Norwegian Arctic Expedition on the Fram 1898–1902 (Rostrup 1906). Neither the type nor any other collections of the species have been located so E. Larsson, L. Örstadius far. Based upon the short original decription, it is not clear if the species belongs in Psathyrella. Freire & Losa (1977) introduced Psathyrella ascarioides based on a single basidiome collected on horse dung in Spain. The cap is described as lemon yellow, viscid, and 10.5 mm broad. Other characters are the viscid upper part of the stem, the spore size 18 9.5 mm, and the absence of pleurocystidia. Some of these characters deviate from the concept of Psathyrella and the species may belong to another genus. According to EsteveRaventós & Barrasa (1989), it could be the same as Psilocybe luteonitens (syn. Stropharia luteonitens). In current classifications of Psathyrella (e.g. Kits van Waveren 1985), all coprophilous species, except P. sphaerocystis and P. conopilus, are placed in sect. Atomatae. However, when carefully studied the dung-inhabiting species show considerable variation both in macro- (veil characteristics) and micromorphology. These observations suggested to us that dunggrowth might not reflect common ancestry and that section Atomatae could be non-monophyletic. The principal aims of the present study were to identify the number of coprophilous species occuring in the Nordic region (Denmark, the Faroe Islands, Finland, Iceland, Norway, and Sweden), to identify and describe new taxa, to infer the phylogenetic relationships of coprophilous species within Psathyrella, and contribute to a more fully resolved phylogeny of Psathyrella and related taxa. Materials and methods Morphological studies 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. On dung or on heaps of manure basidiomata sometimes grow in smaller or larger groups. The features of each basidiome were then carefully compared to prevent a mixture of species being collected, and the specimens were put into separate boxes. 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 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 Coprophilous species of Psathyrella 1167 Table 1 – Data of specimens sequenced in this study Species Psathyrella sp. P. albofloccosa P. berolinense P. berolinense P. calcarea P. candolleana P. cernua P. clivensis P. conopilus P. corrugis P. dicrani P. effibulata P. fatua P. fibrillosa P. fimiseda P. hirta P. kitsiana P. larga P. larga P. longicauda P. longicauda P. lutensis P. marcescibilis P. merdicola P. microrhiza P. mucrocystis P. noli-tangere P. obtusata P. odorata P. odorata P. olympiana P. orbicularis P. orbitarum P. panaeoloides P. pannucioides P. pennata P. pertinax P. piluliformis P. potteri P. prona P. pseudocasca P. pseudogracilis P. purpureobadia P. purpureobadia P. pygmaea P. romagnesii P. romagnesii P. romagnesii P. rostellata P. saponacea P. scatophila P. senex P. spadicea P. spadiceogrisea P. sphaerocystis P. spintrigeroides P. squamosa P. stercoraria P. stercoraria P. stercoraria P. tenuicula P. tenuicula P. typhae Coll. ID./Origin Ecology, substrate GenBank no. LÖ382-89/Sweden Sivertsen 65-89 (TROM)/Norway LÖ37-04/Sweden LÖ148-91/Sweden LÖ211-03/Sweden LÖ38-00/Sweden LÖ134-98/Sweden LÖ182-03/Sweden LÖ186-02/Sweden LÖ171-01/Sweden LÖ270-04/Sweden LÖ37-96 type/Sweden LÖ132-97/Sweden LÖ138-00/Sweden LÖ56-96 type/Sweden LÖ142-00/Sweden LÖ217-85 type/Sweden LÖ223-90/Sweden LAS97-054/Sweden LÖ201-02/Sweden Kytövuori 94-009 (H)/Finland LÖ98-03/Sweden LÖ31-03/Sweden LÖ45-02 type/Sweden LÖ185-02/Sweden LÖ103-98/Sweden LÖ83-03/Sweden LÖ88-01/Sweden LÖ207-96/Sweden LÖ50-04/Sweden LÖ32-02/Sweden LÖ211-04/Sweden LÖ257-90/Sweden LÖ44-03/Sweden LÖ143-03/Sweden LÖ206-03/Sweden LÖ259-91/Sweden LÖ162-02/Germany LÖ271-01/Sweden LÖ91-99/Sweden LÖ17-04/Sweden LÖ172-02/Sweden LÖ23-94/Sweden 9956 (L) type/The Netherlands LÖ97-04/Sweden LÖ213-96/Sweden LÖ85-98/Sweden LÖ267-04/Sweden LÖ 228-85 type/Sweden LÖ204-96/Sweden LÖ64-95 type/Sweden LÖ115-02/Germany Enderle epitype/Germany LÖ92-01/Sweden LÖ126-99/Sweden LÖ122-86/Sweden LÖ104-95/Sweden LAS80-94/Sweden Kytövuori Virrat 1991 (H)/Finland LÖ460-05 type/Sweden LÖ58-03/Sweden Brown (K, 49734)/England LÖ21-04/Sweden In a moist, rich forest On grass remnants Wild boar dung Wild boar dung On dry, calcareous soil In a rich deciduous forest Caespitose on a stump On the great alvar On buried wood Attached to buried wood On dry, sandy soil In a field margin On nitrophilous soil Among leaves of Fagus Cow dung Cow dung On or near a stump In a rich deciduous forest Well decayed stump In a copse Between compost heaps On soil attached to sticks In a park with Urtica dioica Cow dung In a rich deciduous forest On a stump Among leaves in a forest On moist soil On soil in a pasture In a rich deciduous forest On soil attached to wood On dry, sandy soil In a rich deciduous forest In mud of a fen On clayey soil, Fagus forest On burnt soil On mossy twigs of Picea On stump of Fagus On nitrophilous soil In a rich deciduous forest In a rich deciduous forest In a shrubbery Gregarious on cow dung In a grazed grassland On wood of Salix Horse dung Horse dung Horse dung Deciduous wood Horse dung Horse dung In a park, among leaves Caespitose on a tree base In a deciduous forest Horse dung On a stump In a gravel at a roadside Horse dung In a pasture Cow dung On soil rich in humus Horse dung On remnants of Typha DQ389667 DQ389708 DQ389704 DQ289705 DQ389671 DQ389720 DQ389726 DQ389683 DQ389725 DQ389674 DQ389698 DQ389672 DQ389681 DQ389686 DQ389690 DQ389702 DQ389689 DQ389694 DQ389695 DQ389676 DQ389677 DQ389685 DQ389728 DQ389688 DQ389684 DQ389700 DQ389713 DQ389711 DQ389679 DQ389680 DQ389722 DQ389692 DQ389673 DQ389719 DQ389727 DQ389710 DQ389701 DQ389699 DQ389665 DQ389666 DQ389691 DQ389675 DQ389678 EU126026 DQ389718 DQ389714 DQ389716 DQ389715 DQ389693 DQ389717 DQ389703 DQ389712 DQ389729 DQ389682 DQ389709 DQ389696 DQ389687 DQ389668 DQ389670 DQ389669 DQ389706 DQ389707 DQ389721 (continued on next page) 1168 E. Larsson, L. Örstadius Table 1 – (continued) Species P. umbrina Agrocybe pusiola Conocybe siliginea Coprinus cordisporus Galerina marginata Lacrymaria lacrymabunda Psilocybe semilanceata Coll. ID./Origin LÖ235-04/Sweden LÖ 304-05/Sweden LÖ93-04/Sweden LÖ41-01/Sweden RM3225/Sweden EL70-03/Sweden Holst84/Sweden 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 Beck Zenith V microscope equipped with phase contrast. For each collection ten to 20 mature spores were measured in water at 1250 magnification. Unusually large or small spores were not considered. Other morphological 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 half-way from the margin by cutting tangential to the pileus, 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. Drawings were made with the aid of a drawing tube attached to the microscope. Material studied Collections are deposited in the herbarium at Plant and Environmental Sciences, Göteborg University (GB) if not otherwise indicated. Numerous collections from Nordic herbaria have been examined. Many types and other collections from Europe and North America were studied as loans, among these the Charles Peck collections from NYS and collections from AMNH, B, C, E, G, GB, H, K, L, LD, MICH, NYS, O, S, TROM, TURA, UPS, and WBS. Material was also received as gifts or loans from several private herbaria. Data on sequenced specimens are provided in Table 1. Additional specimens of the four new species, Psathyrella fimiseda, P. merdicola, P. scatophila, and P. stercoraria are listed under each species description. Additional specimens studied are provided as a supplementary data file. Taxon sampling Sequences of the complete ITS region and 1200 bp of the 50 end of nu-rDNA from 65 specimens of Psathyrella were generated for the study. The specimens represent almost half of the species of Psathyrella that occur in North Europe. Specimens were Ecology, substrate In a rich deciduous forest On calcareous soil In a pasture Cow dung On wood In a pasture In grass GenBank no. DQ389697 DQ389732 DQ389731 DQ389723 AF195590 DQ389724 EU029945 selected to represent a broad spectrum of morphological characters and infrageneric classification groups but with an emphasis on coprophilous species. Availability of type material, the state of the material, and the possibility to make a complete description of the collection was also considered. Type collections were sequenced only when deemed unharmful and permission received. Fifteen LSU sequences of Coprinopsis, Coprinellus, Parasola, and Psathyrella were taken from GenBank (AF261489, AF041488, AF041489, AF041503, AF041507, AF041510, AF041517, AF041518, AF041519, AF041527, AF041525, AF041515, AF041523, AF041520, AF041511). Based on results of earlier molecular phylogenetic studies of Agaricales (Moncalvo et al. 2002; Redhead et al. 2001a) Agrocybe pusiola, Conocybe siliginea, Psilocybe semilanceata, and Galerina marginata were selected as out-group. DNA extraction, PCR, and sequencing Sequence data were obtained from herbarium specimens (Table 1). Total DNA was isolated using DNeasy plant mini kit (QIAGEN, Valencia), following the manufacturers recommendations. PCR reactions were carried out using Ready-To-GoÔ PCR beads (Amersham Biosciences, Uppsala). Primers used to amplify the complete ITS region and the 50 end of the LSU region were ITS1F (Gardes & Bruns 1993) and LR21, LR0R and LR7 (Hopple & Vilgalys 1999). Amplified products were purified using Qiaquick spin columns (QIAGEN, Hilden). Primers used for sequencing were ITS1, ITS3, ITS4 (White et al. 1990), Ctb6 (http://plantbio.berkeley.edu/wbruns/), Lr5 and LR3R (Hopple & Vilgalys 1999). Fifty to 75 ng of PCR products were used in each sequencing reaction using DTCS Quick Start Kit (Beckman Coulter, Fullerton). Sequences were obtained using CEQ 8000 DNA analysis system (Beckman Coulter). Phylogenetic analyses Sequences were edited and assembled using Sequencher 3.1 (Gene Codes, Ann Arbor). Sequences were aligned automatically using the software MAFFT (Katoh et al. 2002) and adjusted manually using the data editor in PAUP (Swofford 2003). Sequences have been deposited in GenBank and accession numbers are listed in Table 1. Heuristic searches for most parsimonious trees were performed using PAUP. 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 K random-addition sequence replicates and tree bisection–reconnection (TBR) branch swapping were performed. Relative robustness of Coprophilous species of Psathyrella clades was assessed by the BS method using 1 K heuristic search replicates with 100 random taxon addition sequence replicate, TBR swapping, saving 100 trees in each replicate. Bayesian analysis of phylogeny was performed using MrBayes 3.0B4 (Ronquist & Huelsenbeck 2003). MrModelTest 2.2 (Nylander 2004) was used to estimate separate best-fit models of evolution for ITS1, 5.8S, ITS2, and LSU. A heterogeneous Bayesian inference was set up with model parameters estimated separately for each partition. Eight Metropoliscoupled (MCMCMC) chains with a temperature of 0.2 were initiated; these were run for 10 M generations with tree and parameter sampling every 5 K generations (2 K trees). The initial burn-in was set to 50 % (1 K trees). A 50 % majority-rule consensus cladogram was computed from the remaining trees; the proportions of this tree correspond to Bayesian PPs (BPP). Results The aligned dataset had 1753 characters. After exclusion of ambiguous regions 1427 characters remained for the analysis. Of these 925 were constant, 136 were variable and parsimony uninformative, and 366 were parsimony informative. The MP analysis yielded 5645 equally most parsimonious trees (length ¼ 1884, CI ¼ 0.3896, RI ¼ 0.6320) one of which is presented in Fig 1. BS values above 50 % are indicated above branches. The BS analysis recovered Psathyrellaceae as monophyletic with 99 % support. Four major clades received support corresponding to Parasola including P. conopilus (99 %), Coprinopsis including P. marcescibilis and P. pannucioides (50 %), L. lacrymabunda, P. cernua, P. spadicea and P. camptopoda (100 %), Psathyrella including Coprinellus and Coprinus cordisporus (73 %), and P. larga that occurs on a single branch but cluster with the Psathyrella clade with 74 % BS support. The structure within the large Psathyrella–Coprinellus–Coprinus cordisporus clade was not resolved and many nodes were unsupported. Species belonging to Coprinellus did not form a monophyletic clade. For the separate regions, MrModelTest suggested GTRþG (ITS1), K80 (5.8S), and GTRþIþG (ITS2 and LSU), as optimal models; this information was employed in MrBayes. Chain convergence was attained well ahead of the initial burn-in threshold, which was not modified any further. The 50 % majority-rule consensus phylogram from the Bayesian anlysis is presented in Fig 2. BPP are indicated on branches. Also in this analysis Psathyrellaceae is recovered as monophyletic with a BPP value of 1. The four major clades recovered in the MP analysis, are present also in the Bayesian tree, all of them with BPP values 1. Like before P. larga occurs on a single branch but clusters with the Psathyrella clade (BPP 1). The Bayesian tree is better resolved with more supported nodes. The topology within the Psathyrellaceae clade differs in some respects from the BS tree topology. Coprinellus and Coprinus cordisporus form a single monophyletic clade (1), albeit still nested within the Psathyrella clade, while in the parsimony analysis the same sequences are separated in two clades (Fig 1). Other minor differences in topology within the Psathyrella clade affect mainly unsupported clade constellations. Subclades supported in the BS analysis were recovered with support also in the Bayesian analysis. 1169 Through the morphological investigation of Nordic material 14 coprophilous Psathyrella species were identified. Sequence data confirm that all of them are distinct species. Five species grow exclusively on dung, viz. P. purpureobadia, P. sphaerocystis, and the three new species P. fimiseda, P. merdicola, and P. scatophila. Five species occur mostly on dung but also on manured soil and in nutrient rich grassland, viz. P. hirta, P. romagnesii, P. saponacea, P. tenuicula, and P. stercoraria. Four species grow occasionally on dung, viz. P. conopilus, P. microrhiza, P. potteri, and P. prona. Coprophilous species are marked with bold type in the phylogenetic trees (Figs 1 and 2). Results of the phylogenetic analyses and the present morphology-based infrageneric classification of Psathyrella are not in congruence. No distinct pattern of substrate preferences was revealed and the coprophilous species are spread rather evenly throughout the Psathyrella clade. Substrate records are listed in Table 2. The average spore size was calculated for all coprophilous species. The results are provided in Table 3 where species are arranged according to decreasing spore size. Psathyrella tenuicula in a broad sense was subject to intense morphological studies but unfortunately remains an unresolved issue. Four specimens of P. tenuicula s.lat. were sequenced, one from humus rich soil, one from horse dung, and two from dung of wild boar. We identified two ITS sequence types. The sequence differences between the types are five substitutions in the ITS1 and one substitution and one insertion/deletion event in the ITS2 region. Molecular data indicate a differentiation that may correlate with substrate preferences but further data are needed to disentangle the species complex. Taxonomy Species exclusively or mainly growing on dung Psathyrella fimiseda Örstadius & E. Larss., sp. nov. (Figs 3A, 4A–D) MycoBank no.: 511717 Etym.: fimus- dung, sedes- inhabiting, living. Pileus 3–6 mm latus, convexus, modice fuscus, humido striatus paene usque ad medium, hygrophanus, in sicco fulvescens, primo omnino flocculosus vel fibrillosus. Lamellae adnatae, distantes. Stipes 10–20 0.5–1 mm, albus, apice pruinosus, deorsum flocculosus. Sporae 8–9 4.5–5.5 mm, ovoideae, oblongae, ellipsoideae, amygdaliformes, in aqua observatae castaneae; poro germinativo distincto. Basidia 2- vel 4-sporifera. Pleurocystidia 25–45(–60) 8–14 mm, conica, lageniformia, spansa, pallida. Cheilocystidia 25–40 8–14 mm, numerosa. Fibulae absentes. In stercore vaccino. Typus: Sweden: Skåne: Ivö, N. of Flötö, 2 Aug 1996, L. Örstadius 56–96 (GB–holotypus). Additional specimen: Sweden: Skåne: Ivö, N. of Flötö, 17 Aug 2006, L. Örstadius 194-06 (GB). Cap 3–6 mm broad, convex, fulvous, yellowish red (Mu. 5YR 5/6), striate almost to centre, hygrophanous, drying to pale brown; veil when young with flocci to cap centre. Gills broadly adnate, distant, L ¼ 11–13, rather pale brown, with white fimbriate edge. Stem 10–20 0.5–1 mm, whitish, pulverulent at apex, flocculose below. Smell not distinctive. Taste not recorded. Spores 1170 E. Larsson, L. Örstadius 74 P.potteri LÖ127-01 Psathyrella sp. LÖ382-89 P. prona LÖ91-99 97 P. stercoraria LAS80-94 84 P. stercoraria Kytövuori Virrat 1991 P. stercoraria LÖ460-05 Type 82 P. calcarea LÖ211-03 P. orbitarum LÖ257-90 84 P. corrugis LÖ171-01 P. pseudogracilis LÖ172-02 P. effibulata LÖ37-96 Type 80 P. purpureobadia LÖ23-94 P. purpureobadia Arnolds 99-56 Type P. odorata LÖ50-04 P. odorata LÖ207-96 P. longicauda LÖ201-02 P. longicauda Kytövuori 94-009 P. lutensis LÖ98-03 P. microrhiza LÖ185-02 97 P. fatua LÖ132-97 P. spadiceogrisea LÖ92-01 P. clivensis LÖ182-03 P. obtusata LÖ88-01 P. orbicularis LÖ211-04 P. fimiseda LÖ56-96 Type P. hirta LÖ142-00 P. spintrigeroides LÖ122-86 P. rostellata LÖ128-85 Type 66 P. fibrillosa LÖ138-00 P. pennata LÖ206-03 P. dicrani LÖ270-04 P. kitsiana LÖ217-85 Type P. pseudocasca LÖ17-04 P. umbrina LÖ235-04 57 P. squamosa LÖ104-95 P. merdicola LÖ45-02 Type P. scatophila LÖ64-95 Type P. tenuicula LÖ37-04 P. tenuicula LÖ148-91 99 73 P. tenuicula LÖ58-03 P. tenuicula Brown (K, 49734) P. albofloccosa Sivertsen 65-89 71 P. sphaerocystis LÖ126-99 P. senex LÖ115-02 P. noli-tangere LÖ83-03 97 P. romagnesii LÖ213-96 100 P. romagnesii LÖ85-98 P. romagnesii LÖ267-04 P. piluliformis LÖ162-02 99 P. pertinax LÖ259-91 P. mucrocystis LÖ103-98 55 P. saponacea LÖ204-96 P. panaeoloides LÖ44-03 P. pygmaea LÖ97-04 79 P. olympiana LÖ32-02 100 Coprinus cordisporus LÖ41-01 Coprinus cordisporus AF041511 57 80 C. heterosetulosus AF041520 C. bisporus AF041523 97 P. candolleana LÖ38-00 candolleana P. typhae LÖ21-04 C. flocculosus AF041515 98 C. disseminatus AF041525 COPRINELLUS C. curtus Af041527 100 P. larga LÖ223-90 larga P. larga LAS97-054 Lacrymaria lacrymabunda EL70-03 LACRYMARIA 100 84 P. cernua LÖ134-98 P. spadicea Enderle Epitype Spadiceae P. camptopoda AF261489 P. pannucioides LÖ143-03 C. friesii AF041503 C. cothurnata AF041507 C. latispora AF041510 50 P. marcescibilis LÖ31-03 C. lagopides AF041488 C. macrocephala AF041489 Pa. nudiceps AF041517 Pa. megasperma AF041518 99 Pa. auricoma AF041519 P. conopilus LÖ186-02 Psi. semilanceata Holst84 Agrocybe pusiola LÖ304-05 G.marginata RM3225 C. siliginea LÖ93-04 99 PSATHYRELLA cordisporus COPRINELLUS COPRINOPSIS PARASOLA 10 changes Fig 1 – One of the 5645 most parsimonious trees from the MP analysis presented as a phylogram. BS values are indicated above branches. Coprophilous species are marked in bold. Major supported clades have been named and marked with scale bars. 8–9 4.5–5.5 mm, av. 8.5 5–5.1 mm, ovoid, oblong, ellipsoid, in profile flattened on adaxial side, sometimes amygdaliform, in water dark reddish brown (Mu. 2.5YR 3/4), opaque; germ pore distinct. Basidia 2- or 4-spored, 18–28 8–9 mm. Pleurocystidia 25–45(–60) 8–14 mm, obtusely conical, lageniform, rarely forked, scattered, pale. Cheilocystidia of two types: (A) similar to shape of pleurocystidia, 25–40 8–14 mm, numerous; (B) clavate, small, scattered. Cap scalp halfway from margin: pileipellis made Coprophilous species of Psathyrella 1171 P. spintrigeroides LÖ127-86 P. fibrillosa LÖ138-00 P. rostellataL Ö228-85 Type P. pennata LÖ206-03 P. dicrani LÖ207-04 P. kitsiana LÖ217-85 Type P. orbicularis LÖ211-04 P. fimiseda LÖ56-96 Type 0.9 P. hirta LÖ142-00 P. squamosa LÖ104-95 1.0 P. merdicola LÖ45-02 Type P. pseudocasca LÖ17-04 P. umbrina LÖ235-04 P. scatophila LÖ64-95 Type 1.0 P. tenuicula LÖ37-04 0.95 P. tenuicula LÖ148-91 1.0 P. tenuicula LÖ58-03 1.0 P. tenuicula Brown K49734 P. albofloccosa Sivertsen 65-89 1.0 P. sphaerocystis LÖ126-99 1.0 P. romagnesii LÖ213-96 1.0 P. romagnesii LÖ85-98 P. romagnesii LÖ267-04 1.0 P. noli-tangere LÖ83-03 P. senex LÖ115-02 1 1.0 PSATHYRELLA 2 3 C. floccosus AF041515 C. curtus AF041527 C. disseminatus AF041525 1.0 C. heterosetulosus AF041520 COPRINELLUS 1.0 C. bisporus AF041523 1.0 Coprinus cordisporus LÖ41-01 cordisporus Coprinus cordisporus AF041511 P. saponacea LÖ204-96 1.0 P. panaeoloides LÖ44-03 P. pygmea LÖ97-04 1.0 4 P. olympiana LÖ32-02 P. piluliformis LÖ162-02 0.94 1.0 P. pertinax LÖ259-91 5 P. mucrocystis LÖ103-98 P. obtusata LÖ88-01 P. spadiceogrisea LÖ92-01 1.0 P. clivensis LÖ182-03 6 P. fatua LÖ132-97 P. potteri LÖ171-01 Psathyrella sp. LÖ382-89 P prona LÖ91-99 1.0 P. stercoraria LAS80-94 7 P. stercoraria Kytövuori Virrat 1991 PSATHYRELLA P. stercoraria LÖ460-05 Type 1.0 0.93 P. calcarea LÖ211-03 P. orbitarum LÖ257-90 P. purpureobadia LÖ23-94 1.0 P. purpureobadia Arnolds 99-56 Type 8 P. effibulata LÖ37-96 Type P. corrugis LÖ171-01 1.0 9 P. pseudogracilis LÖ172-02 1.0 P. odorata LÖ50-04 P. odorata LÖ207-96 0.99 P. longicauda LÖ201-02 P. longicauda Kytövuori 94-009 1.0 P. lutensis LÖ98-03 P. microrhiza LÖ185-02 P. candolleana LÖ38-00 1.0 10 P. typhae LÖ21-04 P. larga LÖ223-90 1.0 11 P. larga LAS97-054 0.87 C. corthunata AF041507 0.99 C. latispora AF041510 1.0 P. pannucioides LÖ143-03 COPRINOPSIS P. marcescibilis LÖ31-03 1.0 C. macrocephala AF041489 1.0 C. friesii AF041503 0.9 C. lagopides AF041488 1.0 P. spadicea Enderle Epitype 0.96 P. camptopoda AF261489 Spadiceae 1.0 P. cernua LÖ134-98 Lacrymaria lacrymabunda EL7-03 LACRYMARIA Pa. nudiceps AF041517 1.0 Pa. megasperma AF041518 PARASOLA 1.0 Pa. auricoma AF041519 P. conopilus LÖ186-02 Agrocybe pusiola LÖ304-05 Galerina marginata RM3225 Psilocybe semilanceata Holst84 Conocybe siliginea LÖ93-04 1.0 0.1 Fig 2 – The 50 % majority rule consensus phylogram from the Bayesian analysis. BPPs have been indicated on branches. Minor supported clades discussed in the text have been numbered 1–11. 1172 E. Larsson, L. Örstadius Table 2 – Recorded substrates of examined specimens of Psathyrella Psathyrella species conopilus fimiseda hirta merdicola microrhiza potteri prona purpureobadia romagnesii saponacea scatophila sphaerocystis stercoraria tenuicula s. lat. Cow dung Horse dung Deer dung Wild boar dung Elk dung Badger dung Unspecified dung Not on dung 63 2 27 6 15 13 1 4 5 1 26 2 1 2 1 12 30 3 2 1 1 4 1 1 1 1 4 1 1 8 1 1 3 213 69 76 1 6 1 2 The figures show the number of examined specimens. The name tenuicula s. lat. represents P. tenuicula, P. minima, P. berolinensis, P. coprinoides, and P. granulosa. up of clavate to subglobose cells 10–35 mm wide, pale yellow brown; pileitrama of rather heavily pigmented hyphae. Hymenophoral trama of moderately pigmented hyphae. Veil 20–120 4–20 mm. Clamps absent. Habitat: Growing solitary in small groups on cow dung in a nonfertilised pasture with Crataegus sp., Rosa sp., on calcareous soil. Distribution: So far only known from the type locality in Sweden. Psathyrella fimiseda is recognized by small basidiomata, absence of clamps, and a well-developed veil. Clamps were carefully looked for on stem hyphae without success but some pseudoclamps were seen. P. sphaerocystis differs by smaller cystidia, a veil of sphaerocysts, and numerous true clamps. Psathyrella hirta Peck, Rep. N. Y. State Mus. nat. Hist. 50: 107 (1897). Table 3 – Average spore size of species, based on studied collections, arranged after size Epithets conopilus prona potteri stercoraria saponacea microrhiza hirta romagnesii purpureoabadia scatophila fimiseda sphaerocystis galericolor merdicola tenuicula s. lat. Spore length (mm) Spore width (mm) 13.5–17 12.5–17 12.5–16 11–14.5 10.5–14.5 10–14 10–13 10–13 9–11.5 8.5–10.5 8–9 7.5–9 8–8.5 6.5–8 6–8.5 6.5–9 6.5–8 6.5–8.5 6–7 6–8 5.5–7 5.5–7 5–6.5 5–6 4.5–5.5 4.5–5.5 5–5.5 4.5–5 3.5–4.5 3–4 Agaricus pennatus var. fimicola G. E. Bernard, Champ. Rochelle: 136, pl. 36, fig. 2 (1882). Psathyra semivestita var. coprobia J. E. Lange, Dansk Bot. Ark. 9: 7 (1936); nom. inval. (Art. 36.1). Psathyra coprobia J. E. Lange, Fl. Agaric. Danic. 5: VII (1940). Psathyrella coprobia (J. E. Lange) A. H. Sm., Contr. Univ. Michigan Herb. 5: 44 (1941). Drosophila coprobia (J. E. Lange) Kühner & Romagn., Fl. Analyt. Champ. Supér: 355 (1953); comb. inval. (Art. 33.2). Psathyra semivestita sensu Kauffman, Agaric. Mich.: 271 (1918). Selected illustrations: Bon, Mushr. Toads. Br. N. Western Europe: 267 (1987); Doveri, Fungi Fimicoli Ital.: pl. 88 (2004); Enderle & Christan, Z. Mykol. 58: 80 opposite (1992); Engel, Pilzf. Nordwestoberfrankens 6, pl. 9:026 (1982); Gerhardt, Gr. Pilzf.: 369 above (1997); J. E. Lange, Fl. Agaric. Danic. 4, pl. 152 F as Psathyra coprobia (1939); M. Lange, Nordens Svampar: 164 (1990). Selected descriptions: Enderle & Christan, Z. Mykol. 58: 73–75 (1992); Kits van Waveren, Persoonia, Suppl. 2: 96–98 (1985); Kühner & Romagnesi, Fl. Analyt. Champ. Super.: 355 as Drosophila coprobia (1953). Cap 5–30 mm broad, at first semiglobose and dark reddish brown (Mu. 2.5YR 3/4), then expanded convex and brown to yellow brown (Mu. 10YR 3/6), hygrophanous, striate when moist, drying to pale ochraceous or pale brown (Mu. 10YR 7/ 3); veil well developed as flocci to centre, evanescent. Gills subdistant, L ¼ 14–26, broadly adnate, grey brown, brown (Mu. 7.5YR 5/4), with white fimbriate edge. Stem 20–45 1–2 mm, enlarged below, pale brown, at apex pulverulent, lower part with flocci. Smell not distinctive. Spore print almost black (Mu. 7.5YR N 2/). Spores 10–13 5.5–7 mm, av. 10.3–12.1 5.9– 6.6 mm, Qav. ¼ 1.6–1.9, oblong to narrowly ovoid, rarely ellipsoid, in profile sometimes amygdaliform or with a slight suprahilar depression, in water dark red (Mu. 2.5YR 3/6–4/6, 10R 3/6); germ pore distinct. Basidia 4-spored, 20–30 9–11 mm. Pleurocystidia 30–60 10–16 mm, lageniform, fusiform, at apex sometimes acute, rarely subcapitate or bent, scattered to numerous, hyaline. Cheilocystidia of two Coprophilous species of Psathyrella 1173 Fig 3 – Photos of the four newly described species: (A) Psathyrella fimiseda LÖ194-06, (B) P. merdicola LÖ45-02 (holotype), (C) P. scatophila LÖ64-95 (holotype) and (D) P. stercoraria LÖ460-05 (holotype). types: A. similar to shape of pleurocystidia, 25–50 7–14 mm, numerous; B. clavate, small, numerous close to cap margin and scattered for the rest. Pileipellis a hymeniderm of 15–60 mm wide cells sometimes pigmented at base; pileitrama made up of pigmented hyphae sometimes strongly incrusted. Veil cells 40–250 4–32 mm, cylindrical to fusiform. Clamps present on stem hyphae. Habitat: Solitary, scattered or gregarious on dung of horse and cow, ‘auf Rehlosung’ according to Derbsch & Schmitt (1987), also ‘on manure or fertilized soil’ (Smith 1972), all records from the Nordic countries are from pure dung except the one collected by I. Kytövuori (H) that grew on ‘open hay field grazed by cattle’. Distribution: Known from Denmark, the Faroe Islands, Finland, Norway, and Sweden; the Baltic States (Urbonas et al. 1986); the Czech Republic (Vašutová 2006), France, and the Netherlands (Kits van Waveren 1985); Germany (Derbsch & Schmitt 1987); Spain (Heykoop & Esteve-Raventós 1994); Switzerland (Breitenbach & Kränzlin 1995); North America (Smith 1972). Psathyrella hirta is recognised by a copious veil, broadly adnate gills, spore size, and cystidia shape. P. scatophila has slightly smaller spores and cystidia with a more acute apex. P. purpureoabadia has smaller spores, narrowly utriform cystidia, and no clamps. P. saponacea differs by larger spores, a scanty veil, and utriform cystidia. Psathyrella merdicola Örstadius & E. Larss., sp. nov. (Figs 3B, 4F–H) MycoBank no.: 511718 Etym.: merda- dung, cola- dwelling. Pileus 5–15(–20) mm latus, primo semiglobatus, deinde convexus postremo planus, primo castaneus, deinde fuscus, hygrophanus, leviter striatus; velum squamuloso-floccosus paene usque ad medium, glabrescens. Lamellae subdistantes, adnatae vel anguste adnatae; acie interdum albae. Stipes 15–35 0.5–2 mm, apice pruinosus, deorsum floccosus. Sporae 6.5–8 3.5–4.5 mm, oblongae, interdum amygdaliformes, raro subphaseoliformes; poro germinativo distincto. Basidia 4-sporifera. Pleurocystidia 25–50 8–15 mm, conica, lageniformia, subutriformia, numerosa vel modice numerosa. Cheilocystidia 20–40 6–13 mm, spansa vel modice numerosa. Cellulae veli 20–180 6–40 mm. Fibulae adsunt. In stercore vaccino. Typus: Sweden: Skåne: Kristianstad, Näsby fält, Lerduvedungen, 27 June 2002, L. Örstadius 45-02 (GB–holotypus). Additional specimens examined: Sweden: Skåne: Baldringe, Skogshejdan, 18 Aug 2006, L. Örstadius 201-06; Kiaby, Kjugekull, 2 July 1991, L. Örstadius 161-91; Kristianstad, Näsby fält, Lerduvedungen, 14 Aug. 2001, L. Örstadius 59-01, 60-01, 61-01. Cap 5–15(–20) mm broad, at first semiglobose, then expanded convex and finally plane, umbonate or not, when young dark reddish brown (Mu. 2.5YR 2.5/4-2.5/2), becoming strong brown (Mu. 7.5YR 4/6), slightly paler towards margin, 1174 E. Larsson, L. Örstadius Fig 4 – Micromorphology of the newly described species. Psathyrella fimiseda (LÖ56-96 holotype), (A) spores, (B) pleurocystidia, (C) cheilocystidia, (D) basidiomata; P. merdicola (LÖ45-02 holotype), (E) spores, (F) pleurocystidia, (G) cheilocystidia, (H) basidiomata; P. scatophila (LÖ64-95 holotype). (I) spores, (J) pleurocystidia, (K) cheilocystidia, (L) basidiomata; P. stercoraria (LÖ460-05 holotype), (M) spores, (N) pleurocystidia, (O) cheilocystidia, (P) basidiomata. Bars [ 10 mm (microscopic features), [ 10 mm (basidiomata). hygrophanous, faintly striate on the margin, fading to pale yellow brown; veil with white flocci on entire cap, appendiculate at margin, glabrescent. Gills subdistant, L ¼ 15–24, adnate to adnexed, yellowish red (Mu. 5YR 5/6); edge sometimes minutely white fimbriate. Stem 15–35 0.5–2 mm, often bent, very pale brown, pulverulent at apex, flocculose below from veil remnants. Smell not distinctive. Taste mild. Spore print dusky red (Mu. 2.5YR 3/2). Spores 6.5–8 3.5–4.5 mm, av. 7–7.3 3.9–4 mm, Qav. ¼ 1.7–1.8, oblong, narrowly ovoid, in profile sometimes narrowly amygdaliform, rarely subphaseoliform, in water red to yellowish red (Mu. 2.5YR 4/8, 5YR 5/6); germ pore distinct to rather distinct. Basidia 4-spored, Coprophilous species of Psathyrella 14–22 7–8 mm. Pleurocystidia 25–50 8–15 mm, conical, lageniform, subutriform, sometimes rostrate or swollen at apex, numerous to rather numerous, faintly yellow below apex. Cheilocystidia of two types: A. similar to shape of pleurocystidia, 20–40 6–13 mm, scattered to rather numerous; B. clavate, small, most frequent towards cap margin. Pileipellis most similar to a hymeniderm with clavate, spheropedunculate, ellipsoid to subglobose cells 15–35 mm wide; pileitrama of moderately to strongly pigmented hyphae. Veil cells 20–180 6–40 mm, inflated. Clamps present. Habitat: Growing on moderately old cow dung in open pastures or in copses with Quercus robur, Larix decidua, Rosa sp., and Crataegus sp. 1175 rather dry, sandy soil in extensively grazed grassland on river-dunes, near solitary Crataegus shrub’ (Arnolds 2003), probably overlooked. Distribution: Known from Finland, the Netherlands, and Sweden. This species is recognised by a purple tinged cap, copious veil, usually narrowly utriform cystidia, absence of clamps, and a coprophilous habitat. Psathyrella hirta and P. scatophila also have a copious veil but differ by having clamps and acute cystidia. P. saponacea is distinguished by larger spores and a scanty veil. The type collection of P. pratensis A. H. Sm. was examined (MICH). The species differs by a paler cap, scanty veil, and lageniform to fusiform cystidia. Moreover, clamps were rare on stem hyphae. Distribution: Known from three localities in Sweden. Psathyrella merdicola can be recognised by its small spores and a copious veil, microscopically marked by large cells. P. tenuicula differs by a rudimentary veil and a pubescent cap and stem surface. P. fimiseda and P. sphaerocystis have slightly larger spores, whereas P. squamosa differs by having paler more pronounced amygdaliform spores, numerous cheilocystidia, and a non-coprophilous habitat. Psathyrella purpureobadia Arnolds, Fungi non delineati 26: 8 (2003). Selected illustration: Arnolds, Fungi non delineati 26: pl. 7C (2003). Selected description: Arnolds, Fungi non delineati 26: 8–11 (2003). Cap 3–15 mm broad, semiglobose, expanded convex, at first dusky red, dark reddish brown with purple tinges (Mu. 10R 3/4-2.5/2, 2.5YR 3/2-3/4-3/6-4/6), at mature stage fading to rust brown, reddish yellow, hazel brown (Mu. 5YR 6/8), hygrophanous, when moist striate up to 2/3 from margin, drying to reddish yellow, pale brown (Mu. 5YR 6/8); veil when young with flocci on entire cap, appendiculate at margin, evanescent. Gills distant, L ¼ 10–16, adnate to adnexed, ventricose, reddish grey or brown (Mu. 5YR 5/2-5/3-5/4), edge white fimbriate. Stem 10–25 0.5– 1.5 mm, pale brown, darker towards base, at apex pulverulent, downwards with fibrils or floccules. Smell faint or absent. Taste not recorded. Spores 9–11.5 5–6 mm, av. 9.8–10.8 5.4–5.9 mm, Qav. ¼ 1.8–1.9, oblong, rarely ovoid, obovoid, limoniform, subfusiform, in profile sometimes with a suprahilar depression, rarely amygdaliform, in water dark red (Mu. 2.5YR 3/4-3/6-4/8), almost opaque, with distinct germ pore. Basidia 4-spored, 18–30 9– 11 mm. Pleurocystidia 30–65 8–18 mm, narrowly utriform to lageniform, rarely with flexuous walls or forked apex, pale, scattered to rather numerous. Cheilocystidia of two types: A. similar to shape of pleurocystidia, 25–65 6–20 mm, numerous; B. clavate, small, scattered to numerous. Cap scalp halfway from margin: pileipellis consists of often subglobose 15–40 mm wide cells; pileitrama of strongly incrusted hyphae. Veil cells 10–80 5–26 mm, inflated to cylindrical, hyaline to yellow brown. Clamps absent. Habitat: Growing single or in small groups on cow and horse dung in pastures and margin of forests, ‘on calcareous, Psathyrella romagnesii Kits van Wav., Persoonia 7: 44 (1972), non P. romagnesii Bon (1988). Psathyrella waverenii Arnolds, Bibliotheca Mycol. 90: 439 (1983). Selected illustrations: Arnolds, Bibliotheca Mycol. 90: pl. 8d (1983); Arnolds, Fungi non delineati 26: 41, pl. 7B (2003). Selected descriptions: Arnolds, Bibliotheca Mycol. 90: 436– 437 (1983); Kits van Waveren, Persoonia, Suppl. 2: 94–95 (1985). Cap 2–10 mm broad, when young semiglobose to conicoconvex and dark reddish brown, reddish brown (Mu. 5YR 4/64/4-3/3-3/4), then becoming convex and dark brown to grey– brown (Mu. 7.5YR 4/2, 4/4, 10YR 4/3, 6/3, 6/2), hygrophanous, striate up to centre, drying to pale ochraceous brown, pale brown or grey; veil at margin as fibrils or rarely as flocci, fugacious. Gills distant, L ¼ 6–13, broadly adnate, brown (Mu. 7.5YR 5/4-5/2-4/2). Stem 10–40 0.3–1.5 mm, cylindrical, pale brown, upper part pulverulent, lower part with fibrils or flocci from veil remnants. Smell not distinctive. Spores 10–13 5– 6.5 mm, av. 10.6–11.7 5.3–6.2 mm, Qav. ¼ 1.7–2, oblong, ovoid, subcylindrical, rarely limoniform, sometimes irregular in outline, in profile sometimes narrowly amygdaliform or with a suprahilar depression, rarely subphaseoliform, in water red (Mu. 2.5YR 3/6-4/6-4/8); germ pore distinct, central or rarely eccentric. Basidia 2–4-spored, 16–28 9–11 mm. Pleurocystidia 25– 60 8–16 mm, narrowly utriform, lageniform, sometimes widened or subcapitate at apex, scattered to rather numerous, pale. Cheilocystidia of two types: A. similar to shape of pleurocystidia, 20–50 6–14 mm, scattered to numerous; B. clavate, small, scattered to numerous. Cap scalp halfway from margin: pileipellis with subglobose to ellipsoid 10–50 mm wide cells, pale yellow brown; pileitrama with rather coarsely incrusted hyphae. Veil cells 15–100 4–16 mm. Clamps seen on stem hyphae. Habitat: Single or scattered in pastures on rotten straw, on mixtures of straw and dung or directly on dung of horse more seldom of cow, once on a burnt spot. Distribution: So far known from Denmark, Iceland, the Netherlands, Norway, and Sweden. The description is based on examination of collections from Denmark, Iceland, The Netherlands, Norway, and 1176 Sweden, including the type collections of Psathyrella romagnesii (L) and P. waverenii (WBS). Three collections of P. romagnesii were sequenced and found to have identical ITS regions. In one of them (L. Örstadius 213-96) one basidioma had 4-spored basidia with normal sized and shaped spores while another basidioma had 2-spored basidia with both small and normal sized spores, which moreover sometimes had an irregular shape. Kits van Waveren (1985) reported disturbed spore production in P. tephrophylla. P. waverenii, only known from the type locality, was described with 2-spored basidia and growing ‘on old cow dung’ (Arnolds 1983). ‘Exceptionally’ (Arnolds 1983) or ‘very seldom’ (Kits van Waveren 1985) 3-spored basidia were demonstrated. The characters separating P. waverenii from P. romagnesi are small and P. waverenii is, therefore, reduced to synonymy with P. romagnesii. P. purpureoabadia differs by a well-developed veil, a purple tinged cap, and by lacking clamps. P. hirta has slightly wider spores and a copious veil as reflected by the larger cells, and often larger basidiomata. Psathyrella saponacea F. H. Møller, Fungi of the Faeröes 1: 179 (1945). Psathyrella coprophila Watling, in Watling & Jurand, Notes Roy. Bot. Gard. Edinb. 31: 146 (1971). Psathyrella fimetaria Watling, apud Watling & Jurand, Notes Roy. Bot. Gard. Edinb. 31: 143 (1971). Psathyrella equina A. H. Sm., Mem. New York Bot. Gard. 24: 269 (1972). Selected descriptions: Hausknecht & Rücker, Z. Mykol. 55: 105– 110 (1989); Kits van Waveren, Persoonia, Suppl. 2: 99–101 (1985). E. Larsson, L. Örstadius Distribution: Known from a few countries in Europe, i.e. Austria (Hausknecht & Rücker 1989), Denmark, the Faeroes, Finland, Great Britain, Germany, Norway, Spain (EsteveRaventos & Barrasa 1989), Sweden, and North America. The species is recognized by the rudimentary veil, large spores, and utriform cystidia. Psathyrella hirta differs by slightly smaller spores, pointed cystidia, and a copious veil. P. microrhiza is distinguished by the red gill edge, a well-developed veil, and often pointed cystidia. With the exception of larger basidiomata P. equina agrees with P. saponacea. Contrary to Smith (1972) pleurocystidia were found in the type collection (MICH). We examined the type material of P. coprophila and P. fimetaria (E). As no morphological differences against P. saponacea could be discerned both species are reduced to synonomy. The cystidia and basidia are collapsed in the type material of P. saponacea (C) but in the original description Møller (1945) illustrates narrowly utriform cystidia. We measured the spores of the type to 12–13.5 6.5–8 mm. These spores deviate from other species on dung in some respects: (1) mean-value of the size; (2) shape sometimes subhexagonal or irregular; (3) dark pigmented; (4) a difference of width in front view and profile; (5) germ pore sometimes eccentric. We noticed a smell in fresh collections of P. saponacea but found it difficult to describe it. Møller characterized the smell as ‘unpleasant, distinctly soap-like (like that of Tricholoma saponaceum)’. Hausknecht & Rücker (1989) described the smell of P. coprophila as ‘leicht unangenehm’. Psathyrella scatophila Örstadius & E. Larss., sp. nov. (Figs 3C, 4I–L) MycoBank no.: 511719 Etym.: Gk. Skatos - dirt, Gk. phila - loving. Cap 3–25 mm broad, at first semiglobose, hardly expanding or becoming convex, dark reddish brown (Mu. 2.5YR 3/2-2.5/4), dark brown (Mu. 7.5YR 3/4), later fading to yellow brown, hygrophanous, when moist striate up to 2/3 from margin, drying to pale brown with ochraceous centre; veil rudimentary as scattered fibrils near the margin. Gills adnate, sometimes with tooth, subdistant, L ¼ 16–23, pale brown, becoming brown (Mu. 7.5YR 5/4); edges white flocculose. Stem 15–55 1– 2.5 mm, pale brown, pulverulent 1⁄4 from apex, fibrillose lower part. Smell unpleasant, of soap, or not distinctive. Taste not recorded. Spores 10.5–14.5 6–8 mm, av. 11.7–13.8 6.4–7.3 mm, Qav. ¼ 1.7–1.9, oblong, subcylindrical, sometimes irregular in outline, subhexagonal, in profile flattened on adaxial side, sometimes with a slight suprahilar depression, in water dark red (Mu. 2.5YR-10R 3/6), opaque; germ pore indistinct to distinct, central to eccentric. Basidia 4-spored, rarely mixed with 2-spored, 18–30 10–13 mm. Pleurocystidia 25–60 10– 16 mm, narrowly utriform, sublageniform, scattered, hyaline. Cheilocystidia of two types: A. similar to shape of pleurocystidia, 20–45 8–16 mm, numerous; B. clavate, small, scattered, rarely numerous. Pileipellis a paraderm sometimes with transition to a hymeniderm of subglobose, ellipsoid, oblong to clavate 15–50 mm wide cells; pileitrama made up of weakly to strongly incrusted hyphae. Veil cells 10–50 2–8 mm. Clamps present. Additional specimens: Finland: Etelä-Häme: Lankaa, Hitonhauta Protection Area, 8 Sep 2002, T. Talvitie (TUR 158155); PohjoisHäme: Toivakka, 11 Sep 2004, I. Kytövuori (H). d Sweden: Gotland: Gerum, Russpark, 24 Sep 2004, L. Örstadius 173-04. Halland: Vallda Sandö, 11 Sep 2006, S. Jacobsson 426-06. Medelpad: Borgsjö, 300 m E. of the church, 30 Aug 1993, L. Örstadius 143-93; Skåne: Kristianstad. Näsby fält, Lerduvedungen, 6 Aug 2000, L. Örstadius 94-00. Södermanland: Österhaninge, Tyresta NP, 21 Sep 1997, H.-G. Toresson (herb. Toresson). Habitat: Solitary to gregarious in pastures on horse dung, once on cow dung, or on manured soil, so far not known from dung of other animals. Cap 5–20 mm broad, at first semiglobose, becoming expanded convex, reddish brown, reddish yellow (Mu. 5YR 4/3, 5/8, 6/8), when mature fading to brown, hygrophanous, striate Pileus 5–20 mm latus, primo semiglobatus, deinde convexus, brunneus, hygrophanus, striatus, in sicco pallide brunneus; velum squamuloso-floccosus paene usque ad medium. Lamellae adnatae vel anguste adnatae, distantes vel subconfertae, rufo-griseae. Stipes 20–60 1–2.5 mm, pallidus, apice pruinosus, deorsum squamulosus. Sporae 8.5–10.5 4.5–5.5 mm, oblongae, subcylindricae, ovoideae, interdum amygdaliformes, in aqua observatae ferrugineae; poro germinativo distincto. Basidia 4-sporifera. Pleurocystidia 30–60 7–16 mm, anguste fusiformia, lageniformia, numerosa. Cheilocystidia 20–45 7–16 mm. Cellulae veli 30– 300 5–35 mm. Fibulae adsunt. Coprobia. Typus: Sweden: Medelpad: Borgsjö, 10 Sep 1995, L. Örstadius 64–95 (GB–holotypus; UPS–isotypus). Coprophilous species of Psathyrella up to half-way from margin, drying to pale brown; veil welldeveloped, surface covered by floccules to centre, at margin appendiculate, evanescent. Gills adnate to adnexed, distant to subclose, L ¼ 11–25, pale brown, becoming reddish grey (Mu. 5YR 5/2) with flocculose edge sub lente. Stem 20–60 1– 2.5 mm, whitish to pale brown, sometimes slightly darker towards base, pulverulent at apex, downwards small scales from veil remnants. Smell not distinctive. Taste not recorded. Spores 8.5–10.5 4.5–5.5 mm, av. 9.2–10.3 4.7–5.5 mm, Qav. ¼ 1.8–2, oblong, subcylindrical, sometimes narrowly ovoid, in profile sometimes narrowly amygdaliform, in water red (Mu. 2.5YR 4/6–4/8); germ pore distinct. Basidia 4-spored, 16–24 8–10 mm. Pleurocystidia 30–60 7–16 mm, narrowly fusiform, lageniform, sometimes flexuous or with yellow thickened wall, most often upper part pointed and rarely at apex bent or provided with crystals or incrustations, numerous. Cheilocystidia of two types: A. similar to shape and frequency of pleurocystidia, 20–45 7–16 mm; B. clavate, small, scattered; edge rarely pigmented. Cap scalp half-way from margin: pileipellis made up of subglobose, clavate to ellipsoid cells 10–50 mm wide; pileitrama of moderately to pale pigmented, rarely incrusted hyphae. Veil cells 30–300 5–35 mm, cylindrical to fusiform. Clamps present, often also pseudoclamps. Habitat: Solitary, gregarious or subcaespitose on dung of badger, horse, cow, and elk. Distribution: So far known from seven localities in Finland and Sweden but probably overlooked. The species is recognized by spore size and a copious veil. Psathyrella hirta differs in having larger spores (10–13 5.5– 7 mm, av. 10.3–12.1 5.9–6.6 mm), and in addition it has slightly darker cap and spores. However, more material must be studied in order to understand the variation of all characters. P. fimiseda has smaller basidiomata and smaller spores. P. purpureoabadia differs by its purple tinged cap, utriform cystidia, and absence of clamps. Psathyrella sphaerocystis P. D. Orton, Notes Roy. Bot. Gard. Edinb. 26: 57 (1964). Selected icon: Arnolds, Fungi non delineati 26: 41, pl. 7E (2003). Selected descriptions: Arnolds, Fungi non delineati 26: 21–23 (2003); Enderle, Z. Mykol. 64: 229–230 (1998); Kits van Waveren, Persoonia, Suppl. 2: 113–115 (1985); Orton, Notes Roy. Bot. Gard. Edinburgh 26: 57–58 (1964). Cap 4–10 mm broad, campanulate, conico-convex or convex, seemingly not expanded, pale ochraceous-buff or clayochraceous then darkening to clay-buff, faintly striate when moist, hygrophanous, entirely granular mealy-scaly or furfuraceous, becoming matt and atomate when veil is washed off, edge of cap joined to stem by pale buff or ochraceous cortinate veil when young. Gills rather broadly adnate with tooth, whitish or pale clay then pale clay-umber or with slight olivaceous-clay tinge, finally clay-umber, distant, L ¼ 14–18, edge minutely white denticulate at first. Stem 15–24 0.5–1.5 mm, equal or slightly swollen at base, pale creamy yellowish or tinged ochraceous below, often hyaline at apex or in upper 1177 part, sometimes becoming deeper ochraceous-cream with age, at first pale ochraceous-buff furfuraceous or floccose-scaly from veil, later more scattered adpressed fibrillose-scaly, apex white pulverulent when fresh, hollow. Flesh ochraceous in cap drying paler, creamy-whitish or pale yellowish in stem, sometimes darker at base. Spores 7.5–9 5–5.5 4–5 mm, av. 8.2 5– 5.4 4.5–4.6 mm, Qav. ¼ 1.5–1.7 (breadth) and 1.8 (witdth), ovoid, oblong, ellipsoid, in profile sometimes amygdaliform but never pronounced, rarely subphaseoliform, with a distinct germ pore, in water dark red (Mu. 2.5YR 4/6-3/6), opaque. Basidia 4-spored, 14–22 6–8 mm. Pleurocystidia 20–35 8–12 mm, narrowly lageniform to subutriform, hyaline, yellow, or sometimes strongly orange pigmented below apex, scattered to rather numerous. Cheilocystidia of two types: A. similar to shape of pleurocystidia, 20–30 6–12 mm, scattered to numerous; B. clavate, small, scattered. Veil cells (sphaerocysts) from cap surface, single or in chains, 10–35 mm broad, subglobose, ellipsoid, subcylindrical, often brown pigmented sometimes incrusted. Cells of pileipellis hardly possible to distinguish from the veil cells. Pileitrama with strongly incrusted hyphae. Clamps found at stem hyphae. Habitat: Single to gregarious, ‘on old horse dung’ (Orton 1964), ‘on old cow dung (Scottish Highland cattle) in heathland on dry, acidic sandy soil’ (Arnolds 2003), in the Swedish material on horse dung in open grassland on dry, sandy soil. Distribution: Known from the type locality in Scotland, Germany (Enderle 1998), the Netherlands (Arnolds 2003), Sweden, and North America (Smith 1972). The species is recognized by small basidiomata, often subglobose veil cells, small cystidia, and small spores. Psathyrella tenuicula can be distinguished by its pubescent cap and stem surface. P. merdicola differs by smaller spores and a fibrillose veil, and P. fimiseda differs by slightly larger cystidia and absence of clamps. P. albofloccosa, another species with veil of sphaerocysts, is so far not found on dung. It has slightly larger spores and cystidia than P. sphaerocystis and is known from Norway and Spain. Psathyrella stercoraria Örstadius & E. Larss., sp. nov. (Figs 3D, 4M–P) MycoBank no.: 511720 Etym.: stercus – dung, growing on dung. Drosophila stercoraria Kühner & Joss., Fl. Analyt. Champ. Super.: 356 (1953); nom. inval. (Art. 36.1). Psathyrella stercoraria (Kühner & Joss.) Park.-Rhodes, Trans. Br. mycol. Soc. 37: 335 & 339 (1954); nom. inval. (Art. 33.9 & 36.1). Drosophila stercoraria Kühner & Joss., in Kühner and Romagnesi, Mém. Hors. Série 2. Suppl. Bull. Soc. Naturalistes d’Oyonnax 10-11: 4, 57, 59 (1957); nom. inval. (Art. 34.2). Psathyrella stercoraria Kühner & Joss., in Kühner and Romagnesi, Mém. Hors. Série 2. Suppl. Bull. Soc. Naturalistes d’Oyonnax 10-11: 6 (1957); nom. inval. (Art. 34.2). Psathyrella stercoraria (Kühner & Joss.) M.M. Moser, Kleine Krypt.-Fl., 2 b/2, 3. Aufl.: 215 (1967); comb. inval. (Art. 33.4 & 37.1). Psathyrella stercoraria (Kühner & Joss.) M.M. Moser, in Kits van Waveren, Persoonia 7: 48 (1972); comb. inval. (Art. 33.4 & 37.1). 1178 Psathyrella stercoraria (Kühner & Joss.) Arnolds, Bibliotheca Mycol. 90: 439 (1983); comb. inval. (Art. 33.4 & 37.1). Pileus 4–10(–13) mm latus, semiglobatus vel campanulatus, deinde convexus, madefactus brunneus, leviter striatus, hygrophanus, in sicco pallescens interdum roseus; velum fibrillosofloccosum ad dimidium ad medium, glabrescens. Lamellae adnatae, distantes, acie interdum roseae. Stipes 15–40 0.5– 1.5 mm, apice pruinosus, deorsum fibrillosus. Sporae 11–14.5 6–7 mm, oblongae, subcylindricae, subfusiformes, interdum amygdaliformes; poro germinativo distincto. Basidia 2- vel 4-sporifera. Pleurocystidia 35–65 8–18 mm, lageniformia, ventricosa-rostrata, interdum conica, anguste utriformia, dispersa vel numerosa. Cheilocystidia 20–80 6–28 mm. Cellulae veli 10–40 3–7 mm. Fibulae adsunt. Saepe coprophila. Typus: Sweden: Skåne: Äsphult, Liarum, 7 Oct 2005, L. Örstadius 460-05 (GB–holotypus; UPS-isotypus). Additional specimens: Finland: Pohjois-Häme: Virrat, Rajaniemi, 1991, I. Kytövuori (H). d France: Rhône: Environs de Lyon, Francheville le Haut, 8 June 1947, R. Kühner (G). d Sweden: Skåne: Äsphult, Liarum, 10 Oct 2005, L. Örstadius 471-05, 472-05, 473-05. Västergötland: Norra Björke, Hunneberg, Håstensklev, 20 July 1980, L. Stridvall LAS 94-80 (herb. Stridvall). Cap 4–10(–13) mm broad, at first semiglobose, campanulate, then conico-convex, finally expanded convex, sometimes shining, when young dark reddish brown (Mu. 5YR 3/3), faintly striate up to 2/3 from margin, hygrophanous, drying to pale brown, pale ochraceous, sometimes with pink tinge (Methuen 6B3); veil when young as dispersed fibrils or flocci half-way from margin, soon evanescent. Gills adnate with small tooth, distant, L ¼ 12–16, when young pale brown, becoming brown (Mu. 7.5YR 5/4); edge paler and sometimes red pigmented. Stem 15–40 0.5–1.5 mm, slightly enlarged or with a small bulb at base, not with a pseudorhiza, pale brown, pulverulent at apex, below with dispersed fibrils. Taste mild, not farinaceous. Smell not distinctive. Spores 11–14.5 6–7 mm, av. 12.2–12.5 6.4–6.6 mm, Qav. ¼ 1.9, oblong, subcylindrical, subfusiform, narrowly ovoid, in profile sometimes narrowly amygdaliform or with a slight suprahilar depression, in water dark red to red (Mu. 2.5YR 3/6-4/6-5/6); germ pore distinct. Basidia 4-spored, sometimes mixed with 2-spored, 22–32 10–12 mm. Pleurocystidia 35–65 8–18 mm, polymorph, lageniform, ventricose–rostrate, sometimes conical, clavate, narrowly utriform, scattered to numerous. Cheilocystidia 20– 80 6–28 mm, similar to shape and frequency of the pleurocystidia. Pileipellis a hymeniderm of clavate to ellipsoid 10–40 mm wide cells; pileitrama, hymenophoral trama, and subhymenium strongly pigmented. Veil cells 10–40 3–7 mm. Clamps present on hyphae of stem and veil. Habitat: Gregarious or single on horse and cow dung, once in a pasture grazed by horses (the Finnish record). Our Liarum material grew on cow dung in a pasture rich of herbs, e.g., Scorzonera humilis, Polygala vulgaris, Arnica montana, Filipendula vulgaris, Orchis mascula, Listera ovata, and Dactylorhiza maculata. Distribution: Rarely recorded from Finland, France (Kühner & Romagnesi 1957, reprint 1977) as Drosophila (Psathyrella) stercoraria, Germany (Krieglsteiner 1991), Great Britain (Watling & Jurand 1971), Italy (Cetto 1989), the Netherlands (Kits van Waveren 1985) all as P. stercoraria, and Sweden. E. Larsson, L. Örstadius The species is recognized by the small basidiomata that sometimes have a red pigmented gill edge, spore size, the often conspicuously acute cystidia, and by habitat. Psathyrella hirta is distinguished by a copious veil and slightly smaller spores. P. romagnesii and P. saponacea differ in having obtuse cystidia. The binomial Drosophila stercoraria Kühner & Joss. (Kühner & Romagnesi 1953) was not validly published as it neither had a latin description nor a diagnosis as required by Art. 36.1 of the Code. On page 4 in Compléments à la Flore Analytique VII (Kühner & Romagnesi 1957) a latin description of D. stercoraria is given. However, on page 6 the authors add ‘Psathyrella stercoraria Kühner-Josserand ¼ Drosophila stercoraria K.-J.’ thereby simultaneously supplying an alternative name by the same authors. Neither name is valid because simultaneous publication of alternative names by the same author(s) is forbidden after 1953 (Art. 34.2). The proposed combination in Psathyrella by Moser (1967) is also invalid because the basionym was invalid and the requirements for valid publication of a new name were not fulfilled. Later acceptance of Moser’s (1967) combination together with a direct citation of the invalid basionym and subsequent latin description by Kits van Waveren (1972) and the new combination proposed by Arnolds (1983) are also invalid because the basionym was invalid and all requirements for valid publication of a new name were not met. Specifically, after 1958 (Art. 37.1), designation of a type was required, and no such designation was made in 1967, 1972 or 1983. Because no alternative name is available, the species is here described as new but under the same name. Thus the latin binomial is retained and only the author citation changed. We examined the material of D. stercoraria (Kühner 08/06/ 1947) preserved G and the morphology agrees with the collections from the Nordic countries. The macroscopical description above is for the greater part taken from the two Swedish records. The microscopical description is based on examination of the material from Finland and Sweden. Gills from the French material had a red pigmented edge while gill edges from the Liarum material were only partly red pigmented. In the material from Håstensklev and Finland no red pigmentation on gill edges was detected. Psathyrella tenuicula (P. Karst.) Örstadius & Huhtinen, Oesterr. Z. Pilzk. 5: 146 (1996). Psathyra tenuicula P. Karst., Bidr. Känn. Finl. Natur Folk 32: 511 (1879). Cap 2–15 mm broad, at first semiglobose, then expanded convex, brown, yellow brown with grey tinge, ochraceous buff (Mu. 5YR-7.5YR 5/6), pubescent (sub lente), striate when moist, hygrophanous, pallescent on drying, when young a granular rudimentary veil present. Gills distant, L ¼ 10–18, narrowly adnate, ascendant, when young grey yellow, becoming ochraceous brown. Stem 10–60 0.2–1.5 mm, whitish, pale yellow, when moist entirely pubescent (sub lente). Smell not distinctive. Taste mild. Spores 6.5–8 3–4 mm, av. 6.7–7.9 5–7 mm. Pseudoparaphyses absent or rarely arranged in the specific pattern occurring in most coprinoid species (compare below). Pleurocystidia 20–40 6–10 mm, narrowly lageniform to conical, scattered to numerous, pale. Cheilocystidia of two types: A. similar to pleurocystidia, 15–30 5–10 mm, numerous or sometimes scattered; Coprophilous species of Psathyrella B. clavate, small, scattered. Pileipellis similar to a hymeniderm with subglobose, ellipsoid to clavate, 5–50 mm wide cells, mixed with pileocystidia similar to shape of the pleurocystidia. Pileitrama made up of moderately to pale pigmented hyphae. Caulocystidia 25–50 6–10 mm, similar to shape of the pleurocystidia or widened at base. Veil cells 15–40 mm wide, subglobose to ellipsoid, often brown pigmented, smooth or delicately warty. Clamps only seen in mycelium of stem base. Habitat: Solitary, gregarious to subcaespitose, on dung of wild boar, deer or their allies (Peck 1888; Smith 1972; Delannoy et al. 2002; Doveri 2004), marten (Doveri 2004), cow (Arnolds 2003), and horse, on soil rich in humus, on decaying plant remains, with Alnus, Corylus, Fagus, Picea, and Pinus, dry to moist. Distribution: Known from the former Czechoslovakia, Finland, France, Germany, Great Britain, Italy, the Netherlands, Sweden, and North America. Psathyrella tenuicula is here treated in a broad sense including P. minima (Peck 1888), P. berolinense (Gerhardt 1978), P. coprinoides (Delannoy et al. 2002), P. granulosa (Arnolds 2003), and Coprinus parvulus (Uljé & Keizer 2003). The latter species was included in Coprinus because the authors found basidia to be surrounded by four or five pseudoparaphyses and because pleurocystidia were absent. In our experience presence or absence of pseudoparaphyses in Psathyrella is an unreliable character (cf. also Smith 1972: 15–16). Species normally lacking pseudoparaphyses can in single collections have these pavement-like cells that characteristically surround the basidia. A similar variability exists when it comes to pleurocystidia. In the specimens we examined only scattered pleurocystidia were found. This corresponds with reports in the literature, e.g. Smith (1972; ‘rare’), Delannoy et al. (2002; ‘peu abondantes’), and Arnolds (2003; ‘scattered’). Doveri (2004) found no pleurocystidia at all. It should be noted that the description of C. parvulus is based on a single collection. P. tenuicula s. lat. is distinguished by having small basidiomata, pubescent cap and stem, a rudimentary veil, and small cystidia. It is closely related to P. sphaerocystis that differs by larger spores and a non-pubescent cap and stem. Species occasionally growing on dung Psathyrella conopilus (Fr.) Ulbr., Krypt.-Fl. Anf., edn. 3: 289 (1928). Selected icons: Breitenbach & Kränzlin, Pilze Schweiz 4, pl. 326 (1995); Cetto, Gr. Pilzf. 3: pl. 890 (1979) as P. subatrata; Courtecuisse & Duhem, Champ. Fr.: pl. 790 (1994); Dähncke, 1200 Pilze: 566, 567 (1993); Enderle, Pilzf. Ulmer Raumes: 404 (2004); Gerhardt, Gr. Pilzf.: 371 below (1997); Laessøe et al., Bonniers Stora Svampbok: 151 (1997); Laessøe & Petersen, MycoKey 1.0 CD ROM (2003); J. E. Lange, Fl. Agaric. Danic. 4: 99, pl. 155 D as Psathyra conopilea & pl. 155 E as P. conopilea var. subatrata (1939); M. Lange, Nordens Svampar: 165 (1990); Moser & Jülich, Farbatl. Basidiomyc. III. Psathyrella 1 (1993); Ricken, Blätterpilze: pl. 68, fig. 3 as P. subatrata (1913); Vesterholt, Danmarks Svampe: 300 above (2004). 1179 Selected descriptions: Enderle, Beitr. Kenntn. Pilze Mitteleur. 8: 87–90, 1992; Kits van Waveren, Persoonia, Suppl. 2: 104–107 (1985); Krisai, Libri Botanici 6: 104 (1992); Kühner & Romagnesi, Fl. Analyt. Champ. Super.: 354 as Drosophila subatrata (1953); Pegler, Kew Bull., Addit. Ser. 6: 422 (1977); Smith, Mem. New York Bot. Gard. 24: 273 (1972). Psathyrella conopilus is distinguished by the conical cap with brown setae, the often tall stem, large spores, and absence of pleurocystidia. It also differs from the other dung-inhabiting species by lacking a veil. Psathyrella microrhiza (Lasch) Konrad & Maubl., Encycl. mycol. 14: 123 (1949) [‘‘1948’’]. Selected illustrations: Courtecuisse & Duhem, Champ. Fr.: pl. 786 (1994); Enderle, Beitr. Kenntn. Pilze Mitteleur. 3: 256 (1987); Enderle, Pilzfl. Ulmer Raumes: 411 (2004); Kasparek, Tintling 20: 38 (2000); Lange, Fl. Agaric. Danic.: 100, pl. 154 A as Psathyra squamifera (1939); Ricken, Blätterpilze: pl. 67, fig. 4 as Psathyra semivestita (1910-15); Vesterholt, Danmarks Svampe: 298 below (2004). Selected descriptions: Enderle, Beitr. Kenntn. Pilze Mitteleur. 3: 252–253 (1987); Kits van Waveren, Persoonia, Suppl. 2: 59–63 (1985); Orton, Trans. Brit. Mycol. Soc. 43: 368 as Psathyrella badiovestita (1960); Smith, Mem. New York Bot. Gard. 24: 198–199 as Psathyrella squamifera (1972). Psathyrella microrhiza is recognized by a dark reddish brown cap, rather well-developed veil, an often unbroken red pigmentation of the gill edge, a pseudorhiza, and by many of the cheilocystidia having the same morphology as pleurocystidia. Psathyrella stercoraria differs by smaller basidiomata, scantier veil, lack of a pseudorhiza, and cystidia with a narrow and more acute neck. Psathyrella hirta also lacks a pseudorhiza and also has no red pigmentation on the gill edge. The pleurocystidia of P. hirta hardly ever exceed 60 mm in length, whereas those of P. microrhiza often do. The stem length of P. hirta is 20–45 mm compared with 35–150 mm of P. microrhiza. Psathyrella potteri A. H. Sm., Mem. New York Bot. Gard. 24: 265 (1972). Psathyra subatomata J. E. Lange, Dansk Bot. Ark. 9(1): 16 (1936), not nom. inval. (Art. 36.1); ex J. E. Lange, Fl. Agar. Dan. 5: VII (1940). Psathyrella argentata A. H. Sm., Mem. New York Bot. Gard. 24: 268 (1972). Psathyrella prona f. cana Kits van Wav., Persoonia 7: 37 (1972). Drosophila atomata sensu Kühner & Romagnesi, Fl. Analyt. Champ. Super.: 355 (1953). Drosophila albidula sensu Kühner & Romagnesi, Fl. Analyt. Champ. Super.: 356 (1953). Selected icons: Kasparek, Tintling 35: 8 (2003) as Psathyrella cf. hirta; Lange, Fl. Agaric. Danic. 4: 102, pl. 153 C (1939) as Psathyra subatomata; Enderle & Zuccherelli, Boll. Gruppo Micol. G. Bres. 3– 6: 130 (1993) as Psathyrella prona. Selected descriptions: Lange, Fl. Agaric. Danic. 4: 102 (1939) as Psathyra subatomata; Romagnesi, Bull. Soc. Mycol. France 91: 207–210 (1975) as Drosophila atomata. 1180 Cap 5–25 mm broad, at first obtusely conical to campanulate, becoming convex, often uneven or rough, when young reddish brown to yellowish brown (Mu. 5YR 5/4, 10YR 5/4), often grey tinged, when mature fading from margin to very pale brown (Mu. 10YR 8/4-7/4), ochraceous-grey or buff, hygrophanous, not or rarely faintly striate at margin, drying to white grey without pink tinges; veil present as fibres close to margin. Gills broadly adnate, subdistant, L ¼ 10–22, mature greyish brown, grey, reddish grey; edge exceptionally red pigmented close to cap margin. Stem 20–60 0.5–2 mm, whitish, pulverulent at apex, fibrillose lower part. Smell not distinctive. Taste mild. Spore print reddish black. Spores 12– 16 6.5–8.5 mm, av. 13–15.3 6.7–7.8 mm, oblong, subcylindrical, subovoid, rarely sublimoniform, in profile sometimes narrowly amygdaliform or rarely with a slight suprahilar depression, in water dark red (Mu. 2.5YR 4/6-3/6, 10R 4/8-3/6), opaque; germ pore distinct. Basidia 4-spored, rarely mixed with 2-spored, 20– 34 11–14 mm. Pleurocystidia 35–80(–140) 10–20 mm, narrowly lageniform–fusiform, sometimes forked or flexuous, scattered, hyaline. Cheilocystidia of two types: A. similar to shape of the pleurocystidia, 25–65(–80) 8–16 mm, scattered to numerous; B. clavate, small, numerous, increasing in frequency towards cap margin. Veil cells 20–100 2–8 mm. Clamps present. Habitat: Single to subcaespitose in grasslands, in fields, on manured soil, on rotten straw or mixtures of dung and straw, sometimes on raw dung; out of 79 collections four of them grew on horse dung, five on cow dung, and one on unspecified dung, also on poor soil, with plants like Filipendula vulgaris, Rumex thyrsiflorus, Stellaria graminea, Symphytum sp., and Urtica dioica, ‘sur paille pourrie et débris de pulpe de betterave’ (Romagnesi 1975). Distribution: Known from Denmark, Finland, France, Germany, the Netherlands, Norway, Sweden, and North America. Exact distribution unknown due to confusion with species like Psathyrella prona. The species is recognised by a rather pale and non-striate cap, large dark spores, acute cystidia, and by growing on manured soil. P. prona differs by a dark brown striate cap that often becomes pink upon drying, gills that often have a red edge when mature, and spores that are papillate at apex or have a suprahilar depression. Often 2-spored basidia dominate in P. prona. Psathyrella prona (Fr.) Gillet, Hyménomycètes: 618 (1878). Selected illustrations: Bresadola, Iconogr. Mycol. (Milan): Tab. 890 (1931); Enderle, Beitr. Kenntn. Pilze Mitteleur. 8: 96 (1992); Enderle, Pilzf. Ulmer Raumes: 414 as P. prona, 415 as P. prona var. prona f. picta (2004); Fries, Ic. Sel. Hymenomyc.: Tab. 139, f. 3 (1879); Lange, Fl. Agaric. Danic. 4: 101, Tab. 155 C, C1 as Psathyra prona (1939); Ricken, Blätterpilze: 265, Taf. 68 (1915). Selected description: Kühner & Romagnesi, Fl. Analyt. Champ. Super.: 367 as Drosophila prona (1953). Cap 5–30 mm broad, conical to campanulate, becoming convex, sometimes umbonate, when young dark reddish brown (Mu. 5YR 3/4-4/4), when mature becoming brown, strong brown (7.5YR 4/6), sometimes shining, striate almost E. Larsson, L. Örstadius to centre, hygrophanous, drying to ochraceous grey or cream sometimes with pink tones; veil rudimentary as scattered fibres close to margin, evanescent. Gills adnate, subdistant, L ¼ 14–22, mature reddish grey to reddish brown, with a white fimbriate edge often red pigmented; close to cap margin the red pigmentation typically turns off from gill margin and disappears diffusely and then the red pigment is replaced by a white broad non-pigmented zone reaching to cap margin (visible sub lente). Stem 30–80 1–3 mm, cylindrical, slightly thickened towards base, at apex pale brown, lower part rather dark brown, at apex pulverulent, downwards fibrillose from veil remnants. Smell not distinctive. Taste mild. Spore print reddish black.Spores 12.5–17(–20) 6.5–8 mm, av. 13.1–15.9 6.4– 7.6 mm, subcylindrical, limoniform, oblong, narrowly ovoid, subhexagonal, in profile sometimes narrowly amygdaliform or with a suprahilar depression, in water red (Mu. 2.5YR 4/83/6); germ pore distinct. Basidia often (1-) 2-spored, rarely 4-spored dominate, 18–30 9–13 mm. Pleurocystidia 30–70 9– 16 mm, narrowly lageniform, fusiform, often pointed at apex, rarely forked, sometimes flexuous, scattered to numerous, hyaline. Cheilocystidia of two types: A. similar to shape and frequency of pleurocystidia, 25–50 8–15 mm; B. clavate, small, numerous becoming extremely abundant in the nonpigmented zone close to cap margin, here no seldom a layer of about ten cells. Veil cells 30–80 2–10 mm. Clamps present. Habitat: Solitary, gregarious to subcaespitose, on base-rich soil, on rotten wood, on sticks or twigs, in grass on paths, on manured soil, twice on dung, and once on a fire place. Distribution: Known from all Nordic countries, many countries in the rest of Europe, and North America, often confused with related taxa, probably overlooked. Psathyrella prona is recognised by a striate, dark brown cap often drying to pinkish tones and with a red gill edge. It is usually found with 2-spored basidia and spores with a suprahilar depression or a papilla at apex. The closely related Psathyrella potteri differs by the non-striate paler cap lacking pink tinges when drying. The gill edge is only exceptionally red pigmented. Basidia are 4-spored and the spores are rarely slightly papillate or have a suprahilar depression. New combinations Parasola conopilus (Fr.) Örstadius & E. Larss., comb. nov. MycoBank no.: 511721 Basionym: Agaricus conopilus Fr., Syst. mycol. 1: 504 (1821). Coprinopsis marcescibilis (Britzelm.) Örstadius & E. Larss., comb. nov. MycoBank no.: 511722 Basionym: Agaricus marcescibilis Britzelm., Bot. Centrlbl. 54: 69 (1893) Coprinopsis pannucioides (J. E. Lange) Örstadius & E. Larss., comb. nov. MycoBank no.: 511723 Basionym: Hypholoma pannucioides J. E. Lange, Fl. Agar. Dan . 5: iv (1940). Coprophilous species of Psathyrella Discussion Some taxonomic characters previously not considered significant for the classification of coprinoid fungi gained importance when correlated with molecular evidence (Redhead et al. 2001a). One example is Psathyrella conopilus that clusters with the species in Parasola with strong support (99 % and BPP 1). Psathyrella conopilus shares with Parasola the large, dark spores with an eccentric germ pore, the utriform cystidia, and the absence of a veil. In addition, both Psathyrella conopilus and 1181 Parasola auricoma have setae. Except for Parasola auricoma, the basidiomata of Parasola are not or only weakly deliquescent (Orton & Watling 1979 as stirps Auricomus and stirps Hemerobius within Coprinus) and P. conopilus shares that character. Walther et al. (2005) and Padamsee et al. (2008) recovered the same sister-group relationship of P. conopilus and Parasola as we found. Walther et al. (2005) point to the setae of the pileipellis as a morphological support for the topology, whereas Padamsee et al. (2008) suggest that P. conopilus could represent a separate genus. As arguments for that conclusion they stress presence of thin-walled pileocystidia and absence of A key to the coprophilous psathyrelloid species occurring in Europe Species described by C. H. Peck and A. H. Smith from North America have been considered and Psathyrella galericolor (Smith 1972) known from North America on porcupine dung has been included. P. polaris on musk-ox dung is not included because of the absence of data. 1 Spores on average > 10.5 mm long ...................................................................................................................................................2 Spores on average < 10.5 mm long .................................................................................................................................................16 2(1) Spores on average 13 mm long ......................................................................................................................................................3 Spores on average < 13 mm long ......................................................................................................................................................8 3(2) Hymenial cystidia utriform to lageniform with obtuse apex . ....................................................................................................4 Hymenial cystidia lageniform to conical with acute apex ..........................................................................................................6 4(3) Pleurocystidia and veil absent. Setae on cap present .................................................................................................. conopilus Pleurocystidia and veil present. Setae on cap absent ..................................................................................................................5 5(4) Gill edge not red pigmented. Veil when fresh as fibrils . ............................................................................................saponacea Gill edge often red pigmented. Veil when fresh as flocci .......................................................................................... microrhiza 6(3) Veil when fresh as flocci. Spores on average 10.5–13.5 mm long . ............................................................................. microrhiza Veil when fresh as fibrils. Spores on average 13–15 mm long . ....................................................................................................7 7(6) Cap when moist dark brown and striate. Gill edge often red pigmented . ....................................................................... prona Cap when moist ochraceous-grey to buff, rare fainly striate at margin. Gill edge exceptionally red pigmented .....potteri 8(2) Pleurocystidia with acute apex . ......................................................................................................................................................9 Pleurocystidia with obtuse apex . ..................................................................................................................................................12 9(8) Cap 5–30(–40) mm broad. Veil when fresh as flocci and fibrils at least half-way from cap margin to centre . ..................10 Cap 2–10 mm broad. Veil when fresh as fibrils or flocci at cap margin . .................................................................................11 10(9) Gill edge not red pigmented. Stem 20–45 mm long, without pseudorhiza . ....................................................................... hirta Gill edge often red pigmented. Stem 35–150 mm long, often with pseudorhiza . .................................................. microrhiza 11(9) L ¼ 12–16. Spores on average > 12 mm long ...................................................................................................................stercoraria L ¼ 6–13. Spores on average < 12 mm long ....................................................................................................................romagnesii 12(8) Veil when fresh as flocci and fibrils half-way from cap margin or to centre .........................................................................13 Veil when fresh as fibrils rarely as flocci at cap margin . ..........................................................................................................15 13(12) Clamps absent. L ¼ 10–16 . ....................................................................................................................................... purpureobadia Clamps present. L ¼ 14–36 . ............................................................................................................................................................14 14(13) Gill edge not red pigmented. Stem 20–45 mm long, without pseudorhiza . ....................................................................... hirta Gill edge often red pigmented. Stem 35–150 mm long, often with pseudorhiza . .................................................. microrhiza 15(12) Spores on average 11.7–13.8 6.4–7.3 mm. L ¼ 16–23 . ..................................................................................................saponacea Spores on average 10.6–11.7 5.3–6.2 mm. L ¼ 6–13 . ...................................................................................................romagnesii 16(1) Spores on average > 9 mm long ......................................................................................................................................................17 Spores on average < 9 mm long ......................................................................................................................................................19 1182 E. Larsson, L. Örstadius 17(16) Cystidia narrowly utriform to lageniform. Clamps absent . ............................................................................. purpureoabadia Cystidia narrowly lageniform to conical. Clamps present . .......................................................................................................18 18(17) Spores on average 5.9–6.6 mm wide. Cystidia moderately acute . ........................................................................................ hirta Spores on average 4.7–5.5 mm wide. Cystidia distinctly acute . .................................................................................. scatophila 19(16) Spores 4.5–5.5 mm wide . .................................................................................................................................................................20 Spores 3–4.5 mm wide . ....................................................................................................................................................................22 20(19) On porcupine dung from North America. Spores pale . .............................................................................................. galericolor Not on porcupine dung from North America. Spores dark .......................................................................................................21 21(20) Cap 3–6 mm, fulvous, with a veil of fibrils. Clamps absent. Pleurocystidia 25–45(–60) mm long .............................. fimiseda Cap 4–10 mm, pale ochraceous–buff to clay–buff, with a veil of sphaerocysts. Clamps present. Pleurocystidia 20–35 mm long . .................................................................................................................................. sphaerocystis 22(19) Veil floccose, consisting of hyphae, cells 20–180 6–40 mm ........................................................................................ merdicola Veil of sphaerocysts, cells 10–40 mm wide . ..................................................................................................................... tenuicula pleurocystidia in P. conopilus. However, neither we nor Kits van Waveren (1985) or Smith (1972) found that type of pileocystidia in P. conopilus. That would make the absence of pleurocystidia the main argument for placing P. conopilus in a separate genus. In our experience the presence/absence of pleurocystidia is not always a reliable character, which is why we find it more logical to include P. conopilus in Parasola. Another discrepancy concerning the species concept of P. conopilus is that P. subatrata is distinguished by Padamsee et al. (2008) as a separate species placed in the Coprinopsis clade, whereas in our opinion P. subatrata is a synonym of P. conopilus. Psathyrella marcescibilis and P. pannucioides cluster in the phylogenetic trees with species in the Coprinopsis clade. The clade receives a low BS support (50 %) but a BPP value of 1. This is in congruence with the results of Walther et al. (2005) and Padamsee et al. (2008). Like in Coprinopsis, the pileipellis of P. marcescibilis and P. pannucioides is a cutis of short to elongated cells and not the typical hymeniderm or paraderm seen in most Psathyrella species. We propose that P. marcescibilis and P. pannucioides are moved to Coprinopsis. Species belonging to Coprinellus did not form a monophyletic clade in our analyses. In other recent analyses based either on ITS (Keirle et al. 2004) or LSU data (Padamsee et al. 2008) Coprinellus is monophyletic. In Keirle et al. (2004), like in our study, Coprinus cordisporus is a sister taxon to Coprinellus, whereas in Padamsee et al. (2008) it occurs on a single branch nested within the (Psathyrella) cystidiosa clade. We conclude that the position of C. cordisporus and the delimitation of Coprinellus is still an unresolved issue. Lacrymaria lacrymabunda clusters with P. cernua, P. spadicea, and P. camptopoda in a strongly supported clade (BS 100 % and BPP 1). These species are apparently closely related and clearly separated from the main clade of Psathyrella. Lacrymaria has by some authors (e.g. Smith 1972) been treated as a subgenus of Psathyrella and by others (e.g. Watling 1979) as a genus of its own. Our phylogenetic analyses and those presented by Hopple & Vilgalys (1999) and Padamsee et al. (2008) confirm that Lacrymaria should be treated as a distinct genus. However, the three Psathyrella species deviate in morphology from the genus concept of Lacrymaria. In the infrageneric classification of Psathyrella by Smith (1972) P. cernua, P. spadicea, and P. camptopoda are placed in subgenus Homophron. These species are united by the lack of a veil, smooth spores, and muricate cystidia with crystals. In contrast Lacrymaria has a strongly developed cortina, ornamented spores with very dark verrucae or rarely reduced to insignificant verruculae (Watling 1979), and cystidia that do not have abundant crystalline incrustations. Because of the discrepancy in morphological characters and our limited sampling in Lacrymaria, we do not presently want to make any nomenclatural changes. However, we believe that the Spadiceae clade may represent yet another genus within Psathyrellacae. The infrageneric classification of Psathyrella has shifted throughout the years. Smith (1972) and Kits van Waveren (1985) place the majority of coprophilous species together with non-coprophilous species in Atomatae, a section typified by P. prona. We included in our phylogenetic analyses about half of the Psathyrella species occurring in the Nordic countries. The results indicate that neither the coprophilous species placed in Atomatae nor section Atomatae form monophyletic clades. Instead the coprophilous species are distributed rather evenly over the Psathyrella clade (Figs 1 and 2). The phylogenetic trees, even if based on a limited selection of Psathyrella species, also indicate that the present infrageneric classification is composed of non-monophyletic taxa. A revised infrageneric classification of Psathyrella is an issue for future studies with more species included and data from additional genes added. The confusion of species limits and concepts is another issue of great concern in the effort to clarify the infragenetic classification of Psathyrella. The problem of naming species is clearly demonstrated when our results are compared with other phylogenetic studies of Psathyrellaceae. We agree with Padamsee et al. (2008) that increased ITS sequencing is necessary to clarify species limits and names for taxa occurring in both Europe and North America. For the time being we only make minor comments on some of the infrageneric clades that were recovered. These numbered clades are marked in Fig 2. Clade 1: Psathyrella spintrigeroides, P. fibrillosa (syn P. artemisiae s. auct.), P. rostellata, P. pennata (66 %, BPP 1). In the infrageneric classification by Kits van Waveren (1985) P. spintrigeroides is placed in section Spadiceae, whereas P. fibrillosa and P. pennata are placed in Pennatae. All four species have a copious veil and acute cystidia with pigmented walls. Clade 2: Psathyrella albofloccosa and P. sphaerocystis form one supported group (71 %, BPP 1), and P. tenuicula s. lat. another Coprophilous species of Psathyrella (99 %, BPP 1), while the two groups combined is not supported. The species are morphologically united by the presence of subglobose veil cells, small basidiomata, small spores, and small pleurocystidia. Kits van Waveren (1985) introduced section Cystopsathyra for species with subglobose veil cells. Clade 3: Psathyrella senex (syn. P. fusca sensu Kits van Waveren, P. ocellata, P. fulvescens var. brevicystis), P. noli-tangere, P. romagnesii (97 %, BPP 1). We find no morphological similarities that unite the species and they are placed in different sections by Kits van Waveren. Extended taxon sampling may change the topology of the clade. Our placement of P. noli-tangere differs from Padamsee et al. (2008) where it is nested in clade G, (/spadicea) among species lacking veil. Our concept of P. nolitangere follows Kits van Waveren (1985) and Nathorst-Windahl (1961) and material collected by these authors was examined (L, GB). The species is characterized by large utriform cystidia that are numerous on the gill edge, moderately developed veil, phaseoliform spores, and a moist habitat. The descriptions by Romagnesi (1975) and Smith (1972) also report a veil but differ regarding spore-form that is described as non-phaseoliform. Clade 4: Psathyrella pygmea, P. olympiana (79 %, BPP 1). The two species have muricate and thick-walled pleurocystidia and are placed in section Spadiceae by Kits van Waveren (1985). Clade 5: Psathyrella piluliformis, P. pertinax (syn. P. chondroderma), P. mucrocystis (99 %, BPP 1). These three species have large basidiomata, small pale spores, close gills, and a lignicolous habitat. They are all placed in section Hydrophilae by Kits van Waveren (1985). Clade 6: Psathyrella spadiceogrisea, P. clivensis, P. fatua (97 %, BPP 1). All three species have utriform pleurocystidia and a gill edge dominated by clavate cheilocystidia. They are placed in subsection Spadiceogriseae by Kits van Waveren (1985). Clade 7: Psathyrella potteri, Psathyrella sp., P. prona, P. stercoraria, P. calcarea, P. orbitarum (82 %, BPP 1). The species have basidiomata that are rather small in size and with a poorly developed veil. The spores are moderately large to large and moderately dark to dark. Some species can have a red pigmented gill edge. The species are sometimes nitrophilous and hardly lignicolous. Clade 8: Psathyrella purpureobadia, P. effibulata (80 %, BPP 1). Both species lack clamps and occur on nitrophilous soil or on dung. Clade 9: Psathyrella corrugis, P. pseudogracilis (84 %, BPP 1). These species are characterized by a scanty veil, pale caps, and large spores. They are placed in section Psathyrella by Kits van Waveren (1985). Psathyrella gracilis, here named P. corrugis, is conserved as type species of the genus Psathyrella (Redhead et al. 2001b; McNeill et al. 2005). Throughout the years the names corrugis and gracilis have been used either to separate two species or to separate infraspecific taxa of one species (e.g. Psathyrella gracilis f. corrugis). We use the name Psathyrella corrugis (Pers.: Fr.) Konrad & Maubl. instead of Psathyrella gracilis (Fr.) Quél. as we consider the latter to be a younger synonym (Hirsch 1984; Enderle 1987). This name is also better defined as a neotype from Persoon’s herbarium has been selected (Kits van Waveren 1971) whereas a type for P. gracilis is currently lacking. In Padamsee et al. (2008) sequences labelled Psathyrella gracilis, P. corrugis, and P. microrhiza occur repeatedly in 1183 different places in clade V /psathyrella s. str. indicating the concepts of the species vary. Our concept of P. corrugis (syn. P. gracilis) and P. microrhiza follows Kits van Waveren (1985) with the addition that red pigmented pieces of the gill edge alternate with short non-pigmented areas in P. corrugis, whereas such non-pigmented areas are rare or lacking in P. microrhiza. Clade 10: Psathyrella candolleana and P. typhae cluster with high support values (97 %, BPP 1). In morphology the two species are united by the lack of pleurocystidia and the presence of utriform cheilocystidia. In the parsimony analyses the clade clusters with a portion of Coprinellus (Fig 1), however without support. In the Bayesian tree clade 10 comes out as a separate group between P. larga and the majority of the included Psathyrella species (Fig 2). The same two species are in Padamsee et al. (2008) nested among other Psathyrella species in a supported clade that comes out as a sister clade to Coprinellus. Clade 11: Psathyrella larga takes a basal position in the Psathyrella clade in both phylogenetic analyses (Figs 1 and 2). Because of the unexpected position in the trees two specimens were sequenced. The species may represent a clade of its own but may also eventually, when more species will be included, emerge as a section within Psathyrella. In morphology P. larga is similar to P. rostellata but lacks outstanding characters. The basidiomata are rather large, the veil is scanty, and the cystidia are large. Morphological characters indicate that it belongs in Psathyrella. The genus Cystoagaricus was first shown by Vellinga (2004) to belong in Psathyrellaceae and in Padamsee et al. (2008) it is placed in a sister clade to P. larga, however without support. Characters as irregular-shaped spores and a cap with pigmented fibrils, spines, or squamules are reminescent of some members of section Pseudostropharia in Psathyrella. Acknowledgements We express our gratitude to Nils Lundqvist, Svengunnar Ryman, Jacques Melot, and Scott Redhead for advice particularly with Latin and nomenclatural issues. Henning Knudsen and Karl-Henrik Larsson for critical comments and suggestions of improvements of the manuscript. Lennart Holm for help with the Latin diagnoses. We are grateful to the curators of the following herbaria: AMNH, B, C, E, G, GB, H, K, L, LD, MICH, NYS, O, S, TROM, TURA, UPS, and WBS. Thanks are also due to individuals who supplied us with the loan of collections or valuable information: Eef Arnolds, Katriina Bendiksen, Kerstin Bergelin, Manfred Enderle, Guðrı́ður Gyða Eyjólfsdóttir, Guillaume Eyssartier, Gro Gulden, Sven-Åke Hanson, Anton Hausknecht, Seppo Huhtinen, Stig Jacobsson, Ilkka Kytövuori, Thomas Laessøe, Per Lassen, Erhard Ludwig, Geir Mathiassen, the late Henri Romagnesi, Arne Ryberg, Karl Soop, Leif & Anita Stridvall, Sigmund Sivertsen, Sigvard Svensson, Tor Tønsberg, Hans-Göran Toresson, and Jukka Vauras. Financial support was received from the Swedish species initiative project, ArtDatabanken, SLU, Uppsala to EL (grant dha 146/05). 1184 Supplementary material Supplementary data associated with this article can be found, in the online version, at doi:10.1016/j.mycres.2008.04.003 references Arnolds E, 1983 [‘1982’]. Ecology and coenology of macrofungi in grasslands and moist heathlands in Drenthe, The Netherlands, Vol. 2. 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Fourteen coprophilous species of Psathyrella identified in the Nordic countries using morphology and nuclear rDNA sequence data

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