Mycosphere

Occurrence of coprophilous Agaricales in Italy, new records, and comparisons with their European and extraeuropean distribution

Doveri F*

Via Baciocchi 9, I-57126-Livorno [email protected]

Doveri F 2010 – Occurrence of coprophilous Agaricales in Italy, new records, and comparisons with their European and extraeuropean distribution Mycosphere 1(2), 103–140.

This work is the successor to a recent monograph on coprophilous ascomycetes and basidiomycetes from Italy. All Italian identifications of coprophilous Agaricales, which the author has personally studied over an 18 year period, are listed and categorized depending on the dung source. All collections were subjected to the same procedure and incubated in damp chambers and an estimate of occurrence of fungal species on various dung types is made. A second collection of Coprinus doverii is described and discussed, while the southern most finding of Panaeolus alcis is listed. An additional collection of Psilocybe subcoprophila, a species previously reported from Italy, is described and illustrated with colour photomicrographs. The morphological features of each species is briefly described, and substrate preferences compared with those reported from previous data.

Key words – Coprinus doverii – damp chambers – fimicolous basidiomycetes – frequency – natural state – Panaeolus alcis – Psilocybe subcoprophila – survey.

Article Information Received 25 March 2010 Accepted 21 May 2010 Published online 19 July 2010 *Corresponding author: Francesco Doveri – e-mail –[email protected]

Introduction have recently been made and despite a The commencement of our systematic relatively slow increase in the numbers of studies on the dung fungi of Italy started in coprophilous basidiomycetes known from Italy 1992 resulting in Doveri (2004) and Doveri et and the inability to use field records for al. (2005) reporting and describing 83 copro- statistical analysis, I have obtained useful philous basidiomycetes. Since these publica- information from abundant cultured material tions more species have been collected and have been able to calculate the frequency throughout Italy and faecal material has been of occurrence of the commonest species on received from colleagues. The increase in several types of dung. species found is lower than expected, possibly I therefore consider it timely to update owing to their differing nutritional require- the list of coprophilous basidiomycetes from ments; coprophilous basidiomycetes are Italy and provide details on their ecology. noticeably fewer than ascomycetes. Unlike ascomycetes, and with the exception of several Materials and Methods Coprinus s.l. species, coprophilous basidio- Samples of dung were collected inter- mycetes hardly grow or do not develop at all in mittently from January 1992 to December 2009 damp chamber cultures. However, new and incubated in non-sterilised damp chambers collections of Coprinus doverii L. Nagy, within a few days, or dried and subsequently Panaeolus alcis M.M. Moser, and Psilocybe cultured within 3 months. Other samples were subcoprophila (Britzelm.) Sacc. from dung not incubated, but simply listed as substrates of

103 species collected in the field by the author or Leaving out 29 records from unidentified colleagues. herbivores, most findings (96%) were made on The design of the damp chambers bovine (54%) and equine (42%) dung, the followed that suggested by Richardson & remaining 4% on dung of other herbivores, Watling (1997) and Richardson (2001), slightly with a marked preponderance of Coprinus s.l. modified by Doveri (2004), utilising both the (47% of records). The preference for bovine lower and the upper third of a 0.5 or 1.5 l and equine dung is typical for all genera, (depending on the size of dung to be contained) although a reverse ratio can be noticed in mineral water plastic bottles. The bases were Bolbitius Fr. (67% of findings on equine, 28% filled with sheets of filter or blotting paper on bovine dung) and Psathyrella (Fr.) Quel. wetted with tap water, and the dung placed on (75% on equine), but findings of the latter are them was in turn wetted by tap water. The scarce at present. Another 415 dung samples of upper third of a bottle functioned as a lid, and 36 animal species were cultured in damp its cap was replaced with a cotton wool plug. chambers. No growth of Agaricales was The incubating samples were placed under observed on dung of badger (Meles meles, 2 natural light at room temperature (18°–25°) and samples), beech marten (Martes faina, 3), were observed from day to day for 5 weeks dormouse (Glis glis, 3), duck (Anas sp., 2), fox with the unaided eye and at × 7–45 (Vulpes vulpes, 7), gecko (Tarentola maurita- magnification using a stereomicroscope. The nica, 1), hedgehog [Erinaceus europaeus (12)], macroscopic features of species appearing on insect [various (9)], lizard [Lacerta sp. (3)], incubated dung were described, and the micro- mouse [Mus musculus (2)], polecat [Mustela scopic characteristics were studied from fresh putorius (5)], porcupine [Hystrix cristata (1)], material with a binocular light microscope, rat [Rattus rattus (1)], snail [Helix sp. (3)], using water, Congo red and Melzer’s reagent as squirrel [Sciurus sp. (3)], toad [Bufo sp. (2)], mounting media. The macroscopic features of weasel [Putorius nivalis (1)] and wolf [Canis species collected from dung in the field were lupus (1)]. reported by the respective collectors, while the The remaining 353 samples provided 237 microscopic characteristics were studied by the collections of 30 Agaricales species (Table 2). author, usually from dried material. Spore size Samples of chamois (Rupicapra rupi- was measured in water and calculated on a capra), marmot (Marmota marmota), marten minimum of 20 mature samples, excluding any (Martes martes), pig (Sus scrofa domesticus), ornamentations from the measurements. The and tortoise (Testudo sp.) dung with their desiccation of small Agaricales, the majority of respective records, have been left out of Coprinus s.l. included, was performed in a few statistics owing to the small number (five or minutes with an artificial light, whereas that of less) of incubations. No Agaricales developed larger species was done with an electric on tortoise dung apart from a few collections of desiccator. All collections have been preserved Coprinus s.l. in the author’s personal herbarium (CLSM) as Samples of the remaining animals—bird dried material or, exceptionally, as slides. (various), cattle (Bos taurus), donkey (Equus asinus), horse (Equus caballus), deer (Cervus Results elaphus), fallow deer (Dama dama), roe deer I recorded 522 collections of 86 Agari- (Capreolus capreolus), goat (Capra hircus), cales species from 700 dung samples in Italy – hare (Lepus sp.), rabbit (Oryctolagus cunicu- 285 records of 82 species came from 285 lus), rock goat (Capra ibex ibex), sheep (Ovis samples detected and identified in the field, aries), and wild pig (Sus scrofa)—have been listed as substrate sources, but not incubated. It included to estimate the frequency of is difficult to use records from the field for a occurrence of Agaricales species on different statistical survey but the samples provide a dung types (Table 3). view of substrate preferences. Thus, data is Coprophilous species of Agrocybe Fayod, limited to findings of each Agaricales genus Conocybe Fayod, Lepista (Fr.) W.G. Sm., and species on different dung types (Table 1). Leucocoprinus Pat., and Volvariella Speg. Did

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Table 1. Records from Italy of coprophilous Agaricales in the natural state.

equine cattle donkey horse mule deer fallow deer roe rabbit wild unidentified deer pig herbivore Agrocybe molesta 1 Agrocybe pediades s. Watling 4 1 1 Agrocybe praecox 1 Agrocybe subpediades s. Watling 2 Agrocybe temulenta s. Watling 1 1 Total Agrocybe 7 0 4 0 0 0 0 0 0 1 Bolbitius coprophilus 4 Bolbitius vitellinus var. titubans 1 Bolbitius vitellinus var. variicolor 8 Bolbitius vitellinus var. vitellinus 4 1 Total Bolbitius 5 1 12 0 0 0 0 0 0 0 Conocybe alboradicans 2 Conocybe antipus 2 Conocybe aurea 1 Conocybe brunneidisca 5 Conocybe cettoiana 1 Conocybe coprophila 6 Conocybe fuscimarginata 4 Conocybe gigasperma 1 Conocybe pubescens 2 Conocybe rickenii 4 1 2 Conocybe siennophylla 4 1 Conocybe siliginea 2 Conocybe singeriana 1 Total Conocybe 28 1 10 0 0 0 0 0 0 0 Coprinellus bisporus 2 1 1 3 Coprinellus brevisetulosus 3 Coprinopsis cinerea 4 1 1 5

105 Table 1. (Continued). Records from Italy of coprophilous Agaricales in the natural state.

equine cattle donkey horse mule deer fallow deer roe rabbit wild unidentified deer pig herbivore Coprinellus congregatus 3 2 1 1 Coprinopsis cothurnata 4 1 Coprinellus curtus 2 “Coprinus” doverii 1 “Coprinus” ephemeroides 1 3 Coprinellus ephemerus 5 2 1 Coprinellus flocculosus 1 1 Coprinellus heptemerus 5 1 1 Coprinellus heterosetulosus 2 Coprinopsis luteocephala 1 Coprinopsis macrocephala 1 1 Coprinellus marculentus 1 2 Parasola misera 2 Coprinopsis nivea 10 1 4 “Coprinus” patouillardii 2 3 Coprinellus pellucidus 1 Coprinopsis poliomallus 1 Coprinopsis pseudocortinata 1 Coprinopsis pseudonivea var. pseudo- 2 nivea Coprinopsis pseudoradiata 1 1 Coprinopsis radiata 1 8 1 Coprinellus sassii 1 Parasola schroeteri 2 Coprinus spadiceisporus 2 Coprinopsis stercorea 1 1 Coprinus sterquilinus 2 Coprinopsis tuberosa 1 2

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Table 1 (Continued). Records from Italy of coprophilous Agaricales in the natural state.

equine cattle donkey horse mule deer fallow deer roe rabbit wild unidentified deer pig herbivore Coprinopsis utrifera 1 Coprinopsis xenobia 4 Total Coprinus s.l. 54 2 43 0 1 2 1 4 1 13 Lepista sordida 1

Leucocoprinus cretaceus 4

Panaeolus acuminatus 4 Panaeolus alcis 1 Panaeolus antillarum 5 Panaeolus cinctulus 2 1 1 Panaeolus fimicola 1 Panaeolus guttulatus 1 Panaeolus papilionaceus 2 3 1 Panaeolus retirugis 2 1 Panaeolus semiovatus 3 1 1 Panaeolus sphinctrinus 6 3 1 Panaeolus subfirmus 2 Total Panaeolus 25 0 9 3 0 0 0 0 0 5 Psathyrella hirta 1 1 Psathyrella lacrymabunda 1 Psathyrella prona var. prona f. prona 1 Total Psathyrella 1 0 3 0 0 0 0 0 0 0 Psilocybe coprophila 2 3 1 1 5 Psilocybe crobulus 1 Psilocybe cyanescens 1 Psilocybe inquilina 1 Psilocybe liniformans 1

107 Table 1 (Continued). Records from Italy of coprophilous Agaricales in the natural state.

equine cattle donkey horse mule deer fallow deer roe rabbit wild unidentified deer pig herbivore Psilocybe subcoprophila 1 Total Psilocybe 7 0 4 1 0 0 0 1 1 6 Stropharia dorsipora 1 6 1 Stropharia luteonitens 1 Stropharia semiglobata 9 3 2 Total Stropharia 10 0 9 0 0 0 0 0 0 4 Volvariella gloiocephala 1

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Table 2. Records from Italy of coprophilous Agaricales in damp chamber cultures (n° of dung samples in brackets).

equine bird cattle chamois donkey horse deer fallow roe goat hare marmot marten pig rabbit rock sheep tortoise wild (27) (54) (4) (6) (56) deer deer (16) (18) (5) (2) (5) (18) goat (51) (2) pig (24) (10) (37) (8) (10) Bolbitius 1 coprophilus

Coprinellus 1 brevisetulosus Coprinopsis 1 1 1 candidolanata Coprinopsis 3 1 1 cinerea Coprinellus 1 congregatus Coprinellus 5 1 2 3 3 curtus “Coprinus” 1 ephemeroides Coprinellus 2 2 1 2 1 ephemerus Coprinopsis 1 1 1 1 2 1 1 3 filamentifera Coprinellus 4 1 1 1 3 2 7 2 1 1 2 9 heptemerus Coprinellus 2 14 1 heterosetulosus Parasola misera 6 5 3 1 1 2 5 Coprinopsis 1 1 1 nivea “Coprinus” 4 1 13 patouillardii Coprinellus 2 1 1 1 2 pellucidus Coprinopsis 1 poliomallus

109 Table 2 (Continued). Records from Italy of coprophilous Agaricales in damp chamber cultures (n° of dung samples in brackets).

equine bird cattle chamois donkey horse deer fallow roe goat hare marmot marten pig rabbit rock sheep tortoise wild (27) (54) (4) (6) (56) deer deer (16) (18) (5) (2) (5) (18) goat (51) (2) pig (24) (10) (37) (8) (10) Coprinopsis 1 1 pseudocortinata Coprinopsis 1 1 1 pseudoradiata Coprinopsis 5 4 17 2 6 radiata Coprinopsis 8 2 2 7 3 9 1 stercorea Coprinus 1 sterquilinus Coprinopsis 1 tuberosa Coprinopsis 1 1 1 1 1 3 utrifera Coprinopsis 1 vermiculifera Total 3 43 1 5 62 10 8 14 16 5 1 0 1 9 8 42 1 1 Coprinus s.l.

Panaeolus 1 papilionaceus

Psathyrella 1 1 coprinoides

Psilocybe 1 coprophila

Stropharia 1 dorsipora Stropharia 1 semiglobata Total Stropharia 1 1

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Table 3. Frequency (%) of Agaricales on different dung types (n° of samples in brackets).

bird cattle equine deer fallow deer roe goat hare rabbit rock sheep wild pig (27) (54) (62) (24) (10) deer (16) (18) (18) goat (8) (51) (10) (37) Bolbitius coprophilus 2

Coprinellus brevisetulosus 2 Coprinopsis candidolanata 4 6 2 Coprinopsis cinerea 11 2 12 Coprinellus congregatus 2 Coprinellus curtus 8 3 12 37 17 “Coprinus” ephemeroides 2 Coprinellus ephemerus 4 3 3 4 Coprinopsis filamentifera 2 2 10 3 12 12 6 Coprinellus heptemerus 7 2 4 30 5 44 11 6 25 18 Coprinellus heterosetulosus 4 23 2 Parasola misera 11 8 12 10 6 25 10 Coprinopsis nivea 2 2 12 “Coprinus” patouillardii 7 23 Coprinellus pellucidus 4 4 3 6 4 Coprinopsis poliomallus 2 Coprinopsis pseudocortinata 2 2 Coprinopsis pseudonivea var. 2 pseudonivea Coprinopsis pseudoradiata 2 4 2 Coprinopsis radiata 9 34 11 12 Coprinopsis stercorea 15 8 20 19 19 18 10 Coprinus sterquilinus 2 Coprinopsis tuberosa 6 Coprinopsis utrifera 2 4 10 3 6 6 Coprinopsis vermiculifera 12 Total 11 50 80 25 60 30 56 22 33 75 61 10 Coprinus s.l.

111 Table 3 (Continued). Frequency (%) of Agaricales on different dung types (n° of samples in brackets)

bird cattle equine deer fallow deer roe goat hare rabbit rock sheep wild pig (27) (54) (62) (24) (10) deer (16) (18) (18) goat (8) (51) (10) (37) Panaeolus papilionaceus 2

Psathyrella coprinoides 3

Psilocybe coprophila 2

Stropharia dorsipora 2 Stropharia semiglobata 2 Total Stropharia 2 2

112 Mycosphere not grow in the non-sterilised damp chambers to 0,3 mm broad, at first white, then blackish. (unlike in a natural state), and species of Stipe 0.2–0.6 × 10–25 mm, slender, whitish Bolbitius, Panaeolus (Fr.) Quél., Psathyrella, also when old, pilose all over.” Spores (5.2–) Psilocybe (Fr.) P. Kumm., and Stropharia (Fr.) 5.8–7.3 × 4.2–4.7 × 3.7 µm, mitriform in Quél. rarely developed. Their low frequency of frontal view (Q = 1.20–1.75; Q = 1.46), occurrence (1.4%), calculated based on all subamygdaliform in side view, knotted with incubated samples, slightly rises (to 1.7%) low, roundish warts, reddish brown, with a well when samples of dung types not developing developed apiculum and an eccentric, about 1 any Agaricales species (see above) are µm wide, germ pore. Basidia 4-spored, 15–20 excluded, but rises (to 9%) when only bovine × 6–7.5 µm, dimorphic, spheropedunculate or and equine dung are considered. This increase claviform. “Cheilocystidia very sparse, globose confirms their preference for cattle and horse to balloon–shaped, 10 – 14 μm diam.; Pleuro- dung as in the natural state (99% of records). cystidia absent; Pileocystidia short, lageniform The occurrence of Coprinus s.l. on incubated with a tapering neck and a narrow, more or samples is much higher (55% to 65%) than less acute tip, rarely somewhat encrusted, 35– other genera, and higher (86%) on dung of 65 × 6–13 μm; Sclerocystidia absent, but some domestic (cattle, equine, goat, pig, rabbit, pileocystidia with slightly thickened walls are sheep) rather than of wild animals (chamois, observable; Veil on pileus made up of 11–25 deer, fallow deer, roe deer, hare, marmot, rock μm wide, globose elements, often covered with goat, wild pig, = 41%). coarse, strongly refringent crystals; Pileipellis Twenty four Coprinus species were a hymeniderm.” Caulocutis with the outermost recorded from incubated dung, 32 from the hyphae encrusted at intervals, 2–4 µm diam., field, and Coprinopsis candidolanata (Doveri supporting numerous, sometimes crowded, & Uljé) Keirle et al., C. filamentifera (Kühner) often encrusted (particularly at their bases), Redhead et al., and C. vermiculifera (Joss. ex lageniform caulocystidia, 25–75 × 13–21 µm, Dennis) Redhead et al. were observed in damp with a bulbous base, and a tapering, excep- chambers only. The most frequent species tionally cylindrical neck, 4–8 µm diam. at its (each recorded more than 30 times) were base, 2.5–4.5 µm upwards. Clamp-connections Coprinellus heptemerus (M. Lange & A.H. Sm.) observed in the caulocutis. Vilgalys et al., Coprinopsis radiata (Bolton) *In inverted commas and italics, Nagy’s Redhead et al., and C. stercorea (Fr.) Redhead description from Hungary (pers. comm., and et al., together representing 43% of all records Mycotaxon 98, 2006). In normal type Doveri’s from cultures in their group, each with a ca. description from Italy (see also under “Discus- 10% occurrence. sion”). Material examined – Italy, Livorno, Taxonomy Livorno city, 0 m a.s.l., one specimen on dung of an unidentified animal, F. Doveri, 16 Apr Coprinus doverii L. Nagy, Mycotaxon 98: 148, 2004, 283.1-Livorno, CLSM 016.04. 2006. Figs 1–4 *“Pileus up to 1.5 × 2 mm when still Panaeolus alcis M.M. Moser, Mycologia 76: closed, mainly broadly elliptical to obtusely 551, 1984 (as “alcidis”). Figs 5–14 conical, hemispherical to campanulate later, *Cap 6 mm high, 3 mm diam., cylindric- flattened when fully mature, up to 5 (7) mm conical, grey, neither striate nor umbonate. diam., dark olive-brown in primordial stages, Cuticle smooth, not hygrophanous, lacking any with brown olive–brown centre and pale veil remnant on the margin. Stem filiform, ochraceous margin later, uniformly dark somewhat paler and much longer than cap ochraceous with faint olive tint when old; height, lacking a ring. Spores 18–20 × 10–11 surface strongly translucently striate when µm, ellipsoidal to narrowly ellipsoidal in young, becoming radially sulcate, shortly frontal view (Q = 1.71–1.90; Q = 1.80), pilose. Veil macroscopically not observable, exceptionally slightly angular, narrowly ellip- see below under microscopical features. soidal to subamygdaliform in side view, dark Lamellae free, crowded, strongly ventricose, up maroon to dark brown, opaque or sometimes

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Figs 1–4 – Original drawing of Coprinus doverii microscopical features from Italy. 1 Caulocutis. 2 Spores. 3 Basidia. 4 Detail of hymenium in frontal view. Bars = 10 µm. semitransparent, smooth, thick-walled, apicu- crusted cells, 15–30 × 14–20 µm, interspaced late, with a prominent, central to slightly with scarce, deformed pileocystidia. Caulocutis eccentric (in side view) germ pore, more than 2 of parallel, somewhat encrusted hyphae, the µm diam. Basidia 30–35 × 12–15 µm, clavate outermost 1.5–3 µm diam., supporting many with a slight middle constriction, usually 4- cystidia, very close to cheilocystidia but spored, rarely 2-spored, with thorn-shaped usually larger. Clamp-connections observed on sterigmata, short-stipitate or sessile. Gill-trama caulocutis. regular, with hyphae 5–15 µm diam. *Unfortunately notes on macroscopic Cheilocystidia abundant (gill edge sterile to features are from the field, and are scarce but subfertile), 19–31 × 4–9 µm, polymorphous enough to identify this species when combined (subcylindric, lageniform, utriform), often with with the microscopic characteristics. an enlarged base, with a slightly to strongly Material examined – Italy, Sardinia wavy, rarely subcapitate neck, 2–3 µm diam. island, Ogliastra, Gairo Taquisara-Leperccei, Pleurocystidia absent. Epicutis hymenidermal, 950 m a.s.l., three specimens on dung (possibly several layers of clavate or globose, short- cattle), L. Arras, 2 Dec 2006, 531.3-Ussàssai, stalked, quite thick-walled, somewhat en- CLSM 009.06.

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Figs 5–7 – Italian collection of Panaeolus alcis. 5, 6, 7 Spores from different specimens. Bars 5, 6 = 15 µm, 7 = 20 µm.

Psilocybe subcoprophila (Britzelm.) Sacc., striate, evenly brown to reddish brown or Syll. Fung. 11: 72, 1895. Figs 15–29 somewhat darker at the disc, with a paler ≡ Agaricus subcoprophilus Britzelm., margin due to scarce veil remnants, expallant Hymenomyc. Südbayern 8: 9, 1891. in dry conditions and becoming yellowish ≡ Geophila coprophila var. subcopro- cream with a pale brown disc. Gills quite dense, phila (Britzelm.) Kühner & Romagn., Fl. Anal. interspaced with lamellulae (L/l = 1/3), broadly Champ. Sup.: 338, 1953 (inval. publ.). adnate to slightly decurrent, broad, thin, ≡ Deconica subcoprophila (Britzelm.) E. ochreous at first, greyish later, greyish brown Horak, Darwiniana 14: 363, 1967. with purple shades at maturity, with a paler, Cap 5-20 mm diam., subglobose in the furfuraceous edge. Stem 20–30 × 2–3 mm, early stages, becoming convex, finally convex- usually straight, cylindrical, hollow, somewhat plane, sometimes with a small papilla. Cuticle enlarging at the apex and base, lacking an smooth, slightly viscous, hygrophanous, in annulus, the same colour or slightly darker than moist conditions more or less translucently cap, indistinctly striate, white furfuraceous in

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Figs 8–14 – Italian collection of Panaeolus alcis. 8, 9 Basidia. 10, 11 Caulocistidia. 12, 13 Cheilocystidia. 14 Epicutis. Bars 8 = 15 µm, 9, 10, 11, 12 = 20 µm, 13 = 10 µm; 14 = 30 µm. the upper third. Context pale brown, inodorous. cylindrical or cylindric-claviform, often with a Spores (12.7–) 15.5×19.8 (22.6) × (7.2–) 9.1– slight median constriction, short-stalked. Sub- 10.7 µm, narrowly ellipsoidal to subcylindrical hymenium of small polygonal cells. Gill trama in frontal view (Q = 1.71–1.91; Q = 1.82), regular, with slightly encrusted hyphae, 5–12 slightly flattened at one side in lateral view, µm diam. Gill edge sterile. Cheilocystidia 27– thick-walled, smooth, pale greyish brown in 49 × 8.3–11.7 µm, predominantly lageni-form, water, containing numerous vacuoles, with a sometimes subutriform or fusiform, with an central, slightly flattened germ pore, about2 µm usually long and flexuous neck, 4–6 µm diam., diam. Basidia 28–35 × 12–13 µm, 4-spored, and a roundish or clavate or even (sub)

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Figs 15–18 – Italian collection of Psilocybe subcoprophila. 15, 16 Fruitbodies in the natural state. 17 Ixocutis. 18 Spores. Bars 15 = 5 µm, 16 = 20 µm, 17 = 50 µm, 18 = 15 µm. capitate apex. Pleurocystidia absent. Epicutis coprophilous. They are distinguishable from of elongated, indistinct hyphae, embedded in a other Strophariaceae on dung especially by hyaline gelatinous material (ixocutis). Subcutis their hymeniform pileipellis, from Conocybe of interwoven and branched, encrusted, spp. by usually more fleshy, collybioid or yellowish ochreous hyphae, 6–12 µm diam. tricholomatoid fruitbodies, and absence of Caulocutis with yellowish-ochreous, more or lecythiform cheilocystidia. less parallel, quite thick-walled, encrusted Agrocybe pediades (Fr.) Fayod s. outermost hyphae, 2–3 µm diam., supporting, Watling; A. subpediades (Murrill) Watling s. numerous, often crowded caulocystidia, 32–80 Watling; A. temulenta (Fr.) Singer s. Watling: I × 12–20 µm, similar to cheilocystidia. Clamp- separately described (Doveri 2004) A. pediades connections present, widespread. s. Watling and A. subpediades s. Watling Material examined – Italy, Teramo, San (1982), which also now are listed as Biagio-Rocca Santa Maria, 1200 m a.s.l., about independent taxa in our tables (see above). ten specimens on wild pig dung, B. de Ruvo, 5 These species, characterised by comparatively Jun 2007, 339.3-Teramo, CLSM 010.07. small fruitbodies, absence of pleurocystidia and coarse veil remnants on the cap edge, are very Discussion similar to each other and described by Nauta I have recorded from Italy one species of (in Noordeloos et al. 2005) as an aggregate all those Agaricales genera (Agrocybe, taxon under A. pediades. Nauta (2004; in Bolbitius, Conocybe, Coprinus s.l., Panaeolus, Noordeloos et al. 2005) also described A. Psathyrella, Psilocybe, Stropharia) commonly pediades var. fimicola (Speg.) Nauta, different regarded as potentially coprophilous (Wicklow, from A. pediades var. pediades in occurring on 1992), and some others, like Lepista, cattle and horse dung, patent coarse veil Leucocoprinus, Volvariella, that occasionally remnants on the cap edge, also when mature, occur on dung. They need to be briefly and a late areolate cap centre. Nauta (2004) discussed: chose a Consiglio’s (1999a) colour photo from Agrocybe, placed in Strophariaceae cattle dung in Italy (sub nomine A. subpediades) Singer & A.H. Sm. after molecular studies by as a significant picture of the var. fimicola. Moncalvo et al. (2002), includes about 100 Nauta (2004) also described the new species A. species, a few of which are regarded as ochracea Nauta, which possibly is the same

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Figs 19–22 – Italian collection of Psilocybe subcoprophila (Congo red). 19 Subcutis. 20–22 Basidia. Bars 19 = 10 µm, 20, 22 = 15 µm, 21 = 30 µm. as A. temulenta s. Watling (an ambiguous name Agrocybe molesta (Lasch) Singer: it has in her opinion), and differs from A. pediades already been expounded (Cacialli et al. 1997) var. pediades particularly in having why this name is kept, although it was regarded pleurocystidia, well differentiated from by Nauta (in Noordeloos et al. 2005) as a later cheilocystidia. synonym of A. dura (Bolton) Singer. Like A. Dealing with these species as a whole, I praecox, but unlike A. pediades s.l., it has large have recorded them from Italy, always in the fruitbodies and very coarse veil remnants, often field, particularly from cattle (78% of all in the shape of an annulus. Habitat: in rich soils, records), but also from horse dung. A species but no records known by us, except our own, belonging to the pediades-group, possibly A. from dung or manure. fimicola, was described by Contu (2006) in Agrocybe praecox (Pers.) Fayod: with Italy on unspecified excrements. Other findings less stout fruitbodies than A. molesta, a more worldwide (Watling 1992, De Meulder 2001) membranous and persistent annulus, a mealy confirm the preference for cattle and horse smell, and never a coarsely cracked cap. Like dung. the latter, it has never been recorded from dung.

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Figs 23–29 – Italian collection of Psilocybe subcoprophila (Congo red). 23, 24 Cheilocystidia. 25, 27, 29 Caulocystidia. 26 Caulocutis. 28 Detail of caulocutis. Bars 23 = 50 µm, 24, 25, 27, 29 = 20 µm, 26, 28 = 40 µm.

Nauta (in Noordeloos et al. 2005) mentions, Bolbitius coprophilus (Peck) Hongo: however, also compost-heaps as substratum. refer to Cacialli et al. (1995) and Doveri (2004) Bolbitius, in Bolbitiaceae Singer, for comment about the synonymy between B. includes several coprophilous species, which coprophilus, B. demangei (Quél.) Sacc. & D. sometimes can be mistaken for some Conocybe Sacc. and B. incarnatus Hongo. In subgen. spp. on dung, although they are recognisable Bolbitius (species with a hymeniderm of non- by more or less deliquescent fruitbodies with a articulate claviform cells), it is microscopically glutinous cap, usually free gills, a constantly very similar to B. vitellinus s.lat., but with coprinoid hymenium, and polymorphous, not somewhat stouter fruitbodies and clear pinkish lecythiform, cheilocystidia. shades on the cap. A widespread species

119 worldwide, in Italy always recorded from horse and Hausknecht (2009) regard Conocybe and dung (Narducci & Petrucci 1994, Cacialli et al. Pholiotina as distinct, the former characterised 1995). Most records are from equine dung by absence of a partial veil, constantly (Quélet 1901, Hübsch 1985, Täglich 1991, lecythiform cheilocystidia and a hymenophoral Hausknecht & Krisai-Greilhuber 2003), but trama with a thin mesostratum. also from cattle (Watling 1995, Enderle 1996) Conocybe alboradicans Arnolds: in sect. and elephant (Thomas et al, 2001, Manimohan Conocybe (pruinose stem with strongly et al. 2007). dominant caulocystidia), it is distinguishable Bolbitius vitellinus (Pers.) Fr. var. by its rooting stem, large, ellipsoidal spores, vitellinus and B. vitellinus var. titubans (Bull.) and 2-spored basidia. Recorded in Italy from Bon & Courtec.: described by Doveri 2004 as horse dung in the natural state, it is also independent taxa at variety rank, and reported elsewhere from manured pastures distinguished from each other by slight habit (Arnolds in Noordeloos et al. 2005), rarely and colour differences, they were regarded by from pure or mixed dung (Hausknecht 1996, Arnolds (2003a in Noordeloos et al. 2005) and 2009). Hausknecht & Vesterholt (in Knudsen & Conocybe antipus (Lasch) Fayod: with a Vesterholt 2008) as synonyms under the single rooting stem in sect. Conocybe, it differs from taxon B. titubans (Bull.) Fr. They grow C. alboradicans in having 4-spored basidia and particularly on dung, manured soil, and rotten smaller, angular to submitriform spores. Two hay. Our Italian fimicolous records are records from cattle dung in Italy. Collections especially from cattle dung. Widespread all from unspecified dung types are also over the world, with most records of fimicolous mentioned (Svrček 1959, Hausknecht 1996, collections from equine (Persoon 1801, 2009, Hausknecht et al. 2005, Arnolds in Josserand 1936, Dennis 1970, Pegler 1983) and Noordeloos et al. 2005). bovine (Jamoni 1994, 2001, De Meulder 2001, Conocybe aurea (Jul. Schäff.) Hongo: in Welt & Heine 2007) dung, but also from sect. Conocybe, recognisable by its vivid egg elephant (Pegler 1977) and water vole yellow pileic cuticle. A single collection in (Sengupta & Laessøe 2004). Italy from old cattle dung. Widespread Bolbitius vitellinus var. variicolor (G.F. throughout Europe, but exceptionally recorded Atk.) Krieglst.: differs from var. vitellinus and from old dung (Bon 1992, Hausknecht & var. titubans in having a ridged cap with olive Vesterholt in Knudsen & Vesterholt 2008). tinges. Described by Arnolds (2003a, in Conocybe brunneidisca (Murrill) Noordeloos et al. 2005) as a synonym of B. Hauskn.: previously described by Doveri (2004) titubans var. olivaceus (Gillet) Arnolds, and by under C. lenticulispora Watling, which Hausknecht & Vesterholt (in Knudsen & Hausknecht (2009) has stated to be a later Vesterholt 2008) as B. variicolor G.F. Atk. synonym. In section Pilosellae Kühner ex Recorded from Italy always on horse dung in Singer (a pubescent stem with variable, often the natural state. Although mentioned world- hairy caulocystidia, sometimes with additional, wide as facultatively coprophilous, with an sparse lecythiform), it is distinguishable by its habitat similar to B. titubans var. titubans, there habitat and large, lentiform, and slightly is a single record (Jamoni 1997) from cattle. angular spores. All collections from cattle dung Conocybe, in Bolbitiaceae, propor- in Italy. Widespread throughout Europe, and tionally includes fewer coprophilous species most fimicolous records are from horse dung than Bolbitius, from which they can usually be (Watling 1980, 1982, Enderle 1993, Hausk- recognised in the field by their duller and less necht 1995, Arnolds in Noordeloos et al. 2005). delicate fruitbodies. Usually lecythiform Conocybe cettoiana Hauskn. & Enderle: cheilocystidia and an exceptionally coprinoid in sect. Mixtae Kühner ex Singer (lecythiform hymenium help to recognise Conocybe. Our caulocystidia mixed with polymorphous ones), concept of Conocybe is wide and follows stands out for often having fasciculate Watling & Gregory (1981) and Watling (1982), fruitbodies with rooting stems. Recorded from also including Galerella Earle and Pholiotina cattle dung (Cetto 1994, Doveri 2004, Hausk- Fayod, but Arnolds (in Noordeloos et al. 2005) necht et al. 2005) and horse dung (Hausknecht

120 Mycosphere

2003), and manured meadows grazed by sheep dung, there are many others from equine and (Arnolds in Noordeloos et al. 2005). bovine dung (Gillet 1877, Kühner 1949, Singer Conocybe coprophila (Kühner) Kühner: & Digilio 1952, Singer 1953, Pegler 1977, described by Hausknecht (2009) under Pholio- Watling & Taylor 1987, Watling 1988, 1992, tina coprophila (Kühner) Singer, belongs to Bon 1992, Enderle 1993, Fernández Sasia 2001, subgen. Piliferae (Kühner ex Singer) Watling Arnolds in Noordeloos et al. 2005). (veil absent, cheilocystidia polymorphous, Conocybe rickenii (Jul. Schäff.) Kühner: pileocystidia well developed), and is dis- in sect. Pilosellae, is distinguished by its tinguishable by its habitat, a non-striate, viscid habitat, a cap with olive shades, 2-spored cap, comparatively large spores, and long basidia, and large, ellipsoidal spores. A filiform pileocystidia. All records from Italy on common species throughout Europe. All cattle dung. There are numerous records, some records from Italy are on bovine and equine from unspecified dung (Singer 1950, Watling dung. Many records from Europe on 1982, 1985, Hausknecht et al. 2009), most unspecified dung, but findings, especially on from cattle and horse (Kühner 1926, 1935, cattle and horse dung are known (Watling 1988, Singer 1953, Moreno 1976, Moser 1978, Bon Bon 1992, Hausknecht & Passauer 1997, 1992, Jamoni 1992, Contu 1997, Consiglio Enderle & Hübner 1999, De Meulder 2001). 1999a, Arnolds in Noordeloos et al. 2005, Conocybe siennophylla (Berk. & Broome) Hausknecht 2007, 2009, Welt & Heine 2007, Singer: in section Pilosellae, with a small, Hausknecht & Vesterholt in Knudsen & striate, conical cap, and small, ellipsoidal Vesterholt 2008), with a preponderance of spores. It is facultatively coprophilous. It is cattle (60%), rarely from sheep (Hausknecht & rarer in Europe generally than in Italy, where it Vesterholt in Knudsen & Vesterholt 2008). is recorded particularly from cattle dung. There Conocybe fuscimarginata (Murrill) are also records from unspecified dung, some Singer: in sect. Pilosellae, with a hemispheric from cattle and horse (Hausknecht 2005). to conic-convex, non-striate cap and ellipsoidal Conocybe siliginea (Fr.) Kühner: close to spores, it differs from the very similar C. C. rickenii, but with paler, smaller, and more rickenii, particularly in having 4-spored basidia. delicate fruitbodies, and it never grows on pure Widespread worldwide and also growing on dung, preferring rich soils. In Italy it was found very rich soil, in Italy fimicolous collections on a manured soil. were always recorded from cattle dung in the Conocybe singeriana Hauskn.: in sect. natural state. Most records of fimicolous Pilosellae, with large fruitbodies, a long, collections are from cattle dung (Watling 1988, bulbous stem, 4-spored basidia, and large, Enderle 1993, Consiglio 1999a, Karasch 2002, ellipsoidal spores. Mainly recorded from dung, Fernández Sasia 2008, Suárez & Suárez-Gracia in Italy from cattle, elsewhere in Europe from 2009), some from horse (Enderle 1991a, cattle and horse (Hausknecht & Krisai- Sammler 1998, Hausknecht 2009), one from Greilhuber 1997, 2003, Hausknecht 1998, 2005, goat (Kaşik et al. 2004). Enderle & Hübner 1999, Arnolds in Conocybe gigasperma Enderle & Noordeloos et al. 2005, Hausknecht et al. 2005). Hauskn.: in sect. Pilosellae is distinguished by Coprinus s.l. is the commonest its thimble-like cap, 2-spored basidia, and very Agaricales genus on dung (Bell 1983, large spores. It is very rare, and does not grow Richardson 2001), confirmed by surveys in on pure dung, but has always been found in Italy. It is a heterogeneous assemblage rich soils, especially when strongly manured (Reijnders 1979) of non-monophyletic species (Hausknecht & Enderle 1992). (Hopple & Vilgalys 1994, 1999, Johnson 1999) Conocybe pubescens (Gillet) Kühner: in few of which are retained in Coprinus s.str. by sect. Mixtae, it differs from C. cettoiana in Redhead et al. (2001), and the remaining lacking fasciculate fruitbodies and having transferred in Coprinellus P. Karst., bulbous, not rooting stems and larger spores. A Coprinopsis P. Karst. and Parasola Redhead et common species worldwide, mainly on pure al. dung. Our two Italian collections are from Coprinellus bisporus (J.E. Lange) horse dung. Besides records from unspecified Vilgalys et al.: similar to C. sassii in having 2-

121 spored basidia, but with smaller spores, and Never grown in our cultures, but recorded from without pleuro- and sclerocystidia. It usually the field particularly on cattle dung. Recorded grows fasciculate. Rarely recorded from Italy, from The Netherlands (Uljé & Noordeloos in the natural state only. There are no recent 1993, Uljé in Noordeloos et al. 2005) and records other than that of Jamoni (2007) from Scandinavia (Vesterholt in Knudsen & Italy on unidentified dung. Vesterholt 2008) on cattle and horse dung. Coprinellus brevisetulosus (Arnolds) Always recorded in association with cattle Redhead et al. (= Coprinus stellatus Buller in dung from Hawaiian Islands (Keirle et al. Bisby et al.): our experience allows us to 2004). identify it also in the field, as it is hardly Coprinellus curtus (Kalchbr.) Vilgalys et pubescent (very small setulae on cap and stem), al.: recognisable for its habitat, ochreous and to distinguish it from C. pellucidus (P. yellowish sphaerocysts, and subcapitate pileo- Karst.) Redhead et al., which has quite smaller and caulocystidia. It was stated to be present in fruitbodies and a more variable habitat. Four Italy on horse dung only (Doveri, 2004), but records only (three in the field) from Italy, all now its substrate preference appears to be from cattle dung. Also recent records from wider, in culture at least. It was reported from France (Cheype 2008) and Spain (Rubio & Great Britain (Orton & Watling 1979) and the Miranda 2008), both on cattle dung in the Netherlands (Uljé & Bas 1991, Uljé in natural state. Noordeloos et al. 2005) on horse dung only. Coprinopsis candidolanata (Doveri & Most records are from equine dung (Lange Uljé) Keirle et al.: close to C. pseudoradiata, 1915, Buller 1920, Rea 1922, Nathorst- but also with diverticulate, narrower hyphae of Windahl 1961, Hausknecht & Krisai- the veil, and ellipsoidal to ovoid spores. Very Greilhuber 2003, Richardson 2008a, Vesterholt rare worldwide. Recorded only four times, in Knudsen & Vesterholt 2008), but some are from Italy and the Netherlands (Uljé et al., from deer (Keirle et al. 2004) and sheep 2000), respectively on fallow deer and sheep (Richardson 2008b). dung, from Germany (Melzer, 2009) on alpaca, Coprinus doverii Nagy: This was and from the Hawaiian Islands (Keirle et al., originally discovered by Doveri (unpublished) 2004) on goat. and confirmed as distinct by Uljé (pers. Coprinopsis cinerea (Schaeff.) Redhead comm.), but the material was inadequate for a et al.: similar to C. candidolanata, but with proper description. Subsequently Nagy, somewhat larger spores and the filamentous studying Uljé’s material, recognised that it was veil of only broad hyphae in chains. Common the same as a species found by him from in Italy in the natural state (66% of records Hungary, from deer dung, and described as from cattle dung), less common in our cultures Coprinus doverii (Nagy 2006; pers. comm.). (three records out of five from bird). Common According to the systematics suggested in Europe and widespread worldwide on dung by Redhead et al. (2001) Coprinus doverii is heaps or manured straw (Orton & Watling undoubtedly a Coprinellus species, although it 1979, Uljé in Noordeloos et al. 2005). There has not been transferred to this genus. In are two records from cattle dung (Vila & Coprinellus, only C. angulatus (Peck) Redhead Rocabruna 1996, Schafer 2001). et al. has similar, but larger spores. The latter, Coprinellus congregatus (Bull.) P. Karst.: besides, is carbonicolous, has pleurocystidia very similar to C. ephemerus (Bull.) Redhead and larger fruitbodies (Nagy 2006). et al., in comparison with which it tends to be Coprinus ephemeroides (Bull.) Fr.: in caespitose, in the field, rather than scattered or comparison with C. patouillardii it has an gregarious. It preferably grows in the natural annulate stem and always lacks lageniform state in Italy, on cattle and horse dung. Recent cheilocystidia. Recorded from Italy in the records are from Belgium (Wuilbaut 2006) and natural state, once from our cultures, from Spain on cattle dung (Rubio & Miranda 2008). cattle and horse dung. Mentioned as quite Coprinopsis cothurnata (Godey) Red- common all over Europe, particularly on head et al.: close to C. nivea, but easily equine dung (Uljé & Noordeloos 1993, distinguishable by having hexagonal spores. Richardson 2008a,b, Vesterholt in Knudsen &

122 Mycosphere

Vesterholt 2008). Sporadic records from cattle (Uljé & Bas 1991), whereas a greater (Rubio & Miranda 2008), mouflon (Richardson variability is recorded from Great Britain 2008b), and elk (Vesterholt in Knudsen & (Orton & Watling 1979) and Scandinavia Vesterholt 2008) dung. (Vesterholt in Knudsen & Vesterholt 2008) and Coprinellus ephemerus (Bull.) Redhead Castro (2006) has recorded it from cattle from et al.: it can be distinguished from C. Spain. congregatus not only by its scarce tendency to Coprinellus flocculosus (DC.) Vilgalys et be caespitose, but also by the presence of al.: owing to its floccose-scaly veil of clamp-connections and somewhat larger spores ellipsoidal or subglobose, sometimes thick- (Uljè in Noordeloos et al. 2005). Quite frequent walled and pigmented hyphae, it belongs to in Italy, where it prefers cattle dung in the Coprinus s.l. subsect. Domestici Singer, and is natural state, but with a wider substratum regarded as a facultatively coprophilous choice for fruiting in culture. These data only species, and the few records known by us are partly match those from France (Josserand from straw mixed with dung (Orton & Watling 1933, Locquin 1947, Kühner & Romagnesi 1979) or manured soil (Lanconelli 1997). 1953) and the Netherlands (Uljé & Bas 1991, Coprinopsis luteocephala (Watling) Uljé in Noordeloos et al. 2005), where it Redhead et al.: is the latest Coprinus s.l. prefers cattle dung. described by Doveri et al. (2005) from Italy. Coprinopsis filamentifera (Kühner) The lemon yellow veil, and its preference for Redhead et al.: well recognisable in equine dung (Watling 1972, Orton & Watling Coprinopsis particularly by its subrectangular 1979, Uljé & Noordeloos 1997, Doveri et al. spores. Recorded from Italy in culture only, 2005), distinguishes C. luteocephala from C. where it quite frequently grows (ca. 5% of total xenobia, which prefers bovine dung (Orton Coprinus s.l. records) on a variety of dungs, 1976, Bender 1991, Cacialli et al. 1995, 1996, and there are numerous records from Europe Uljé in Noordeloos et al. 2005). (Kühner & Romagnesi 1957, Moreno & Coprinopsis macrocephala (Berk.) Calonge 1975, Orton & Watling 1979, Cacialli Redhead et al.: see also under C. radiata. et al. 1999, Doveri 2004, Richardson 2005, Commoner on straw and manure rather than in 2008a, b, Uljé in Noordeloos et al. 2005, Nagy pure dung. Our few records in Italy, from the 2007, Vesterholt in Knudsen & Vesterholt field only, are from bovine and equine dung. 2008). Coprinellus marculentus (Britzelm.) Coprinellus heptemerus: a very Redhead et al.: the whole of a cap with purple distinctive species with its minute fruitbodies, shades, hexagonal spores, and pileocystidia velar sphaerocysts, strongly tapering, pointed with a swollen, but not capitate tip, charac- pileocystidia, and ellipsoidal spores with an terises this taxon. Few records from Italy on eccentric germ pore. Very common in Italy, cattle and horse dung in the natural state. Other both in the field, where it seems to prefer cattle records known by us are from equine dung dung, and in culture (15% of all Coprinus s.l. (Smith 1948, Richardson 2008c). records, with ca. 10% occurrence on all dung Parasola misera (P. Karst.) Redhead et samples), where its preferences are more al.: the small, glabrous, orange caps on dung variable. allow this species to be readily identified in the Coprinellus heterosetulosus (Locq. ex field. It is frequent in our cultures (10% of Watling) Vilgalys et al.: growth on dung, cap Coprinus s.l. records) on dung of several with reddish shades, and the contemporaneous herbivores. It appears to be one of the presence of lepto- and scleropileocystidia help commonest Coprinus spp. on various dung its identification. The sclerocystidia, smaller types, as confirmed by numerous records from than leptocystidia and variable in number, can Europe and Africa (Richardson 2004a,b, 2005, easily be missed. It grows with the highest 2008a,b). frequency on equine dung (82% of our records Coprinopsis nivea (Pers.) Redhead et al.: from cultures). A confirmation of the another coprophilous species easily recog- preference for this substratum type comes from nisable by its snow white, pruinose veil and France (Locquin 1947) and the Netherlands large, limoniform spores. Rarely developed in

123 our cultures, but fairly common in the natural in Coprinus s.l. Rare in Italy on cattle and state in Italy, particularly on cattle dung (66% horse dung. Possibly rare also in Europe, of its records from the field). Recorded as a judging by the few published records (Locquin species particularly frequent on cattle and horse 1947, Uljé & Noordeloos 1993, Cacialli et al. dung (Orton & Watling 1979, Uljé & 1999). A recent collection from Spain (Rubio Noordeloos 1993, Vesterholt in Knudsen & & Miranda, 2008) from roe deer dung. Vesterholt 2008), and from horse dung by Coprinopsis pseudonivea (Bender & Uljé) Miranda & Rubio (2000) in Spain and Redhead et al. var. pseudonivea: close to Poumarat (2008b) in France. C. nivea, but with smaller spores and patent Coprinus patouillardii Quél.: it also pinkish shades on the veil. Coprinus pseudo- includes C. cordisporus Gibbs, the latter niveus var. tenuicystidiatus Chalange (not considered a later synonym (see Doveri 2004). recombined yet in Coprinopsis) differs from It is identifiable by its finely granulose veil and var. pseudonivea in having smaller fruitbodies, angular to cordate spores with an apical papilla. velar sphaerocysts, and pleurocystidia. The Common in Italy, where almost all records type variety has been recorded only three times come from cattle and horse dung, more in Italy, always from cattle dung. Other records frequently from the latter. Common also in also from Europe on cattle dung (Vila & other European countries, where it is recorded Rocabruna 1996, Hausknecht et al. 1999, from several dung types (Richardson 2004a, Vesterholt in Knudsen & Vesterholt 2008). 2005, 2008a, Uljé in Noordeloos et al. 2005). Coprinopsis pseudoradiata (Watling) Coprinellus pellucidus: see also under C. Redhead et al.: among Coprinopsis spp. with a brevisetulosus. Not very common in Italy, fibrous-floccose veil of broad, more or less where it grows on different dung types. parallel hyphae, it can be recognised by its Reported to be common on cattle dung from small fruitbodies and small, ellipsoidal to Great Britain (Orton & Watling 1979), the oblong spores. Few collections from Italy, Netherlands (Uljé & Bas 1991, Uljé in recorded from different dung types. Other Noordeloos et al. 2005) and Scandinavia records in Europe from various types (Vesterholt in Knudsen & Vesterholt 2008), it (Richardson 2004a, 2008a, b, Rubio & is also recorded from various dung types Miranda 2008, Vesterholt in Knudsen & elsewhere in Europe (Beyer 2004, Coué 2004, Vesterholt 2008). Richardson 2004a, 2008a). Outside Europe, it Coprinopsis radiata (Bolton) Redhead et appears frequent on cattle and horse dung al.: differs from C. pseudoradiata usually in (Malençon & Bertault 1970, Keirle et al. 2004, having larger fruitbodies and spores, and from Richardson 2008c). C. macrocephala in somewhat narrower spores Very common in Italy, both in the natural state Coprinopsis poliomallus (Romagn.) Doveri et (80% of records from horse dung) and in our al.: recognisable on dung by its minute cultures, where 62% of records are from equine, fruitbodies with a pruinose (mainly formed of with 34% of occurrence on this dung type. It is sphaerocysts), greyish veil, and small, ellip- also common on horse dung in Europe (Orton soidal spores. Very rare in Italy on cattle and & Watling 1979, Miranda & Rubio 2000, horse dung. Most records from Europe on Richardson 2008b) and elsewhere (Keirle et al. cattle dung (Romagnesi 1945, Kühner & 2004, Richardson 2004b). Romagnesi 1953, Moreno & Barrasa 1977, Coprinellus sassii (M. Lange & A.H. Sm.) Orton & Watling 1979, Enderle & Bender 1990, Redhead et al.: very close to C. heterosetulosus Uljé & Noordeloos 1993, Uljé in Noordeloos et with the presence of sclerocystidia, but al. 2005, Vesterholt in Knudsen & Vesterholt distinguishable by its 2-spored basidia and 2008), and only one record from sheep (Coué larger spores. It is rare both in Italy and et al. 2004). elsewhere, with few collections reported, all Coprinopsis pseudocortinata (Locq. ex from horse dung (Sass 1929, Lange & Smith Doveri et al.) Doveri et al.: close to C. 1953, van de Bogart 1975, Doveri et al. 2005). poliomallus, but with a white veil and even Parasola schroeteri (P. Karst.) Redhead smaller spores. Perhaps the smallest fruitbodies et al.: differs from P. misera in having larger,

124 Mycosphere cream-yellowish fruitbodies, larger spores, and it differs from C. stercorea in having larger presence of pleurocystidia. Rare in Italy, where spores and lacking a disagreeable smell. Unlike it has been recorded from cattle dung in the the latter, it is very rare in Italy, known from field (Cacialli et al. 1995). Recorded from only one recent record on dung of an Europe on cattle and horse dung (Karsten 1879, unidentified herbivore (Jamoni 2007). There Orton 1972, Orton & Watling 1979), and it has are three recent records from Europe (Miranda also been reported as terrestrial (Uljé & Bender & Rubio 2000, Roux 2006, Melzer 2009) on 1997, Consiglio 2005). cattle, unspecified, and alpaca dung, Coprinus spadiceisporus Bogart: in respectivey. Orton & Watling (1979) mention Coprinus s.str., it is close to C. sterquilinus it particularly on horse dung, like our record from which it differs in having smaller from the field. fruitbodies and smaller, submitriform to Coprinopsis utrifera (Watling) Redhead rhomboid spores. It is a North American et al.: close to C. poliomallus and C. species (van de Bogart 1976), which grows on pseudocortinata, but with somewhat larger cervine dung. Few records from Italy, all from fruitbodies, subcylindrical spores, and a veil the same place on fallow deer dung in the formed both of sphaerocysts and diverticulate natural state (Uljé et al. 1998). There is a single hyphae. In Europe few records from sheep non-fimicolous collection from Sardinia (Contu (Orton & Watling 1979), horse (Uljé in 2007) and one other record from Europe, from Noordeloos et al. 2005), and cattle (Richardson Spain on rabbit dung (Tabarés & Rocabruna 2008a). Our Italian findings extend the 2002). substratum to rabbit and cervine dung. Coprinopsis stercorea (Scop. ex Fr.) Coprinopsis vermiculifera (Dennis) Redhead et al.: with its habitat on dung and a Redhead et al.: in comparison with C. mealy-pruinose veil of diverticulate sphaero- filamentifera it has somewhat larger spores and cysts, it is similar to C. tuberosa, but it has a ascending, very thick-walled, and pigmented faecal smell and smaller spores. Common in end hyphae of the veil. In Italy recorded only our Italian cultures (14% of all Coprinus s.l. once from rock goat dung in culture. Less rare records, with a ca. 9% frequency of occurrence in Europe and particularly recorded from sheep on all samples), where it develops on various (Orton & Watling 1979, Richardson 2005, dung types. Widespread in Europe and one of 2008a, b) or dung of various herbivores the commonest Coprinus s.l. in culture world- (Moreno 1976, Enderle et al. 1986, Vesterholt wide (Richardson, 2004b), on dung of several in Knudsen & Vesterholt 2008). herbivores (Keirle et al. 2004, Richardson Coprinopsis xenobia (P.D. Orton) 2004a,b, 2005, 2008a,b,c). Additional, quite Redhead et al.: see under C. luteocephala. recent records from Spain (Moreno-Arroyo et Lepista (Fr.) W.G. Sm. is recognisable in al. 2005, Siquier & Salom 2005b, Moreno et al. Tricholomataceae R. Heim ex Pouzar espe- 2008), Austria (Dämon 2005), and Germany cially by its often tricholomatoid fruitbodies (Melzer 2009). with crowded, easily separable gills, non- Coprinus sterquilinus (Fr.) Fr.: in amyloid, ornamented, strongly cyanophilous Coprinus s.str. (species with an annulate stem spores, presence of clamp-connections, and and an adherent veil of filamentous hyphae), it absence of veil and cystidia. Often in rich soils usually has, unlike C. spadiceisporus, a but exceptionally fimicolous. L. sordida has a pseudovolva, and grows on equine dung, as striate, strongly hygrophanous pileic cuticle, reported from Italy (Cacialli et al. 1995) and violet gills and stem, and a faint fruity smell. other European (Orton & Watling 1979, Justo There are a few records from specified dung, in et al. 2005, Uljé in Noordeloos et al. 2005) or Italy from cattle (Consiglio & Contu 2003). extraeuropean (Keirle et al. 2004, Richardson Leucocoprinus Pat. in Agaricaceae 2004b) localities. Rarely found on excrements Chevall. has coprinoid fruitbodies with pale of different herbivores (Orton & Watling 1979, gills, a strongly striate cap and a well Babos 2004). developed veil, pale, dextrinoid and meta- Coprinopsis tuberosa (Quél.) Doveri et chromatic spores, and presence of pseudo- al.: often with a sclerotium and a rooting stem, paraphyses. L. cretaceus (Bull.) Locq. has a

125 large, white, furfuraceous-floccose cap and Vesterholt 2008), in Canada and Scandinavia. comparatively large spores with a germ pore. It But recently it has spread southwards, up to has often been recorded from rich soils or France and Spain on horse dung (Poumarat compost heaps but, besides a record from horse 2008c), and Italy (Hausknecht & Krisai- (Doveri 2004), it is not known directly from Greilhuber 2009) on unidentified dung. Ours in pure dung or manures. Sardinia is the southernmost finding so far Panaeolus, in Bolbitiaceae after known. molecular studies by Moncalvo et al. (2002) Panaeolus antillarum (Fr.) Dennis: in and in agreement with Petersen & Knudsen (in subgen. Anellaria (P. Karst.) Ew. Gerhardt, Knudsen & Vesterholt 2008), differs from the with its fleshy and pale fruitbodies, a viscid other genera of this family in having darker and non-hygrophanus cap, and a constant spores in mass. It includes non-deliquescent, presence of sulphidia, it differs from often hygrophanous species with a cellular P. semiovatus particularly in having an hemi- pileipellis, spotted gills, and spores non- spheric-campanulate cap and lacking a veil. All discolouring in H2SO4. Most have a direct or our Italian collections from cattle dung. indirect connection with dung. Recorded from cattle (Dennis 1961, Guzmán & Panaeolus acuminatus (Schaeff.) Quél.: Pèrez-Patraca 1972, Guzmán & Johnson 1974, distinguishable by its dark, slender, and non- Yokoyama 1979, Pegler 1983, Young 1989, fasciculate fruitbodies lacking a veil, with a Grgurinovic 1997, Reid & Ecker 1999), conic-paraboloid, hygrophanous cap and a very frequently from horse (Yokoyama 1979, 1984, long, exannulate stem, microscopically for Pegler 1983, Young 1989, Stijve & Meijer lacking sulphidia and having limoniform or 1993, Alves & Cavalcanti 1996, Grgurinovic mitriform, flattened spores. All our fimicolous 1997, Consiglio 1999a, Cortés & Montón 2002, findings are directly on cattle dung. Also re- Hausknecht & Krisai-Greilhuber 2003, Mir & corded from manured or rich soils (Gerhardt in Melis 2008), sometimes from elephant dung Knudsen & Vesterholt 2008). Most fimicolous (Pegler 1977, Natarajan & Raman 1983, collections from cattle dung (Maire 1937, Manimohan et al. 2007). Jamoni 1994, Pegler 1977, Ortega et al. 1997, Panaeolus cinctulus (Bolton) Sacc.: it De Meulder 2001, Hausknecht & Krisai- lacks veil, weeping gills, and ornamented Greilhuber 2009), fewer from equine (Yoko- spores, i.e. is placed in sect. Laevispora yama 1979, Pegler 1983, Hausknecht & Krisai- (Gerhardt 1996), where it is distinguished in Greilhuber 2009) and elephant (Vrinda et al. having often fasciculate fruitbodies, broad, 1999, Manimohan et al. 2007). even capitate cheilocystidia, and no sulphidia. Panaeolus alcis M.M. Moser: Moser Facultatively fimicolous, it has been recorded (1984), in the protologue, compared P. alcis in Italy and elsewhere both from cattle with P. sphinctrinus var. minor (Fr.) Singer, (Saccardo 1916, Christiansen 1941, Enderle with fruit-bodies similar in size, shape and 1982, Stijve & Meijer 1993) and horse dung colour. The latter, however, has patent veil (Parker-Rhodes 1951, Calonge & Menezes de remnants on the cap edge, clearly angular Sequeira 2003). One record from elephant spores in frontal view (in these respects it dung (Natarajan & Raman 1983). belongs to sect. Panaeolus ss. Gerhardt 1996), Panaeolus fimicola (Fr.) Gillet: in sect. is widespread and usually grows on horse Laevispora, it differs from P. acuminatus in (Singer 1960) or cattle (Guzmán & Pérez- having less slender fruitbodies, a greyish- Patraca 1972), occasionally on moose (Moser brown pileic cuticle, and much less flattened 1984) dung, while P. alcis lacks veil remnants, spores. Fimicolous records are not so numerous both on the stem and cap edge (it belongs to as those from rich soils. In Italy there is only sect. Laevispora Ew. Gerhardt), and has one fimicolous collection, from an unidentified ellipsoidal, exceptionally slightly angular, and herbivore. Elsewhere there are records from more tapered (Q = 1.80 versus 1.50, personal unspecified dung (Gillet 1877, Morgan 1907a, data) spores. Besides it grows on moose, rarely Christiansen 1941, Guzmán & Pèrez-Patraca on roe-deer and reindeer dung (Moser 1984, 1972, Pegler 1977, Stamets 1996, Ludwig Noordeloos 1998, Gerhardt in Knudsen & 2001), cattle (Britzelmayr 1883, De Meulder

126 Mycosphere

2001, Richardson 2008a) and horse dung Dennis 1961, Miller 1968, Bon 1970, Ola’h (Hongo 1959, Hausknecht & Krisai-Greilhuber 1970, Calonge 1971, Guzmán & Pèrez-Patraca 2009). 1972, Pilát 1972, Watling & Gregory 1980, Panaeolus guttulatus Bres.: easily Bon & Marchand 1987, Robich 1992, Stamets recognisable by its weeping gills, secreting 1996, Ortega et al. 1997, Jamoni 2001, cheilocystidia, and small spores. Our Italian Gerhardt in Knudsen & Vesterholt 2008, collections are the only fimicolous ones known, Poumarat 2008a, Hausknecht & Krisai- but it has been reported on manured soil Greilhuber 2009) in the natural state. (Gerhardt 1996). Panaeolus subfirmus P. Karst.: in sect. Panaeolus papilionaceus (Bull.) Quél, P. Laevispora, distinguishable by its pale buff cap retirugis (Fr.) Gillet, P. sphinctrinus (Fr.) Quél.: and large, opaque, subhexagonal, lenticular all described (Gerhardt 1996 in Knudsen & spores. It is a particularly northern European Vesterholt 2008), together with P. campanu- species, but recorded in Italy from horse dung latus (Bull.) Quél., as synonyms under the prior (2004), and elsewhere from heavily manured Panaeolus papilionaceus, which so is con- soils or cattle (Noordeloos 1998), horse and ceived as a species with a variable cap in sheep dung (Ludwig 2001). colour and shape. It can be recognised by its Psathyrella, in Psathyrellaceae Vilgalys abundant veil on the cap margin (sect. et al., includes species with a habit similar to Panaeolus), particularly in the young speci- many Coprinus s.l. in the same family, and to mens, and growth on dung or manures, some Panaeolus spp. in Bolbitiaceae but, sometimes on strongly manured soils. This unlike the former, it lacks deliquescent gills aggregate taxon has been recorded several and a regular coprinoid hymenium (basidia times from Italy from cattle and horse dung in surrounded by brachycystidia), unlike the latter, the natural state, once from horse dung in usually it has unmottled gills and ellipsoidal culture (the only Panaeolus sp. developing in spores that discolour in H2SO4. Usually our damp chambers). Widely recorded with terricolous or graminicolous, but some species levels of occurrence very close to ours on are fimicolous. bovine (Gillet 1877, Maire 1937, Christiansen Psathyrella coprinoides Delannoy et al.: 1941, Singer 1960, Dennis 1961, Ola’h 1970, distinguishable on dung by its small fruitbodies, Guzmán & Pèrez-Patraca 1972, Guzmán & a granulose-pubescent cap with pileocystidia Johnson 1974, Blanco & Moreno 1986, and veil sphaerocysts (sect. Cystopsathyra Moreno et al. 1990, Treu 1996, Ortega et al. (Singer) Kits van Wav.) and very small spores. 1997, De Meulder 2001, Jamoni 2001, Together with Psathyrella minima Peck, Poumarat 2008b, Richardson 2008a, Hausk- P. berolinensis Ew. Gerhardt, P. granulosa necht & Krisai-Greilhuber 2009) and equine Arnolds, and Coprinus parvulus P.-J. Keiser & dung (Gillet 1877, Romagnesi 1937, Chris- Uljé, it also has been synonymised under tiansen 1941, Guzmán 1961, Miller 1968, Psathyrella tenuicula (P. Karst.) Örstadius & Guzmán & Pèrez-Patraca 1972, Young 1989, Huhtinen s.l. (Larsson & Örstadius 2008). A Enderle 1991a,b, Siquier & Lillo 1994, Treu rare species, twice recorded in Italy from our 1996, Grgurinovic 1997, Siquier & Salom cultures of marten and roe deer dung, but never 2005a, Poumarat 2008a, b, Hausknecht & from the field, rare also elsewhere, and Krisai-Greilhuber 2009). Isolated records from recorded from cervine (Peck 1888, Delannoy et sheep, guanaco (Horak 1979), and rabbit dung al. 2002), cattle (Arnolds 2003b), horse, and (Arnolds 1982). wild boar (Larsson & Örstadius 2008) dung, Panaeolus semiovatus (With.) S. Lundell: always in the natural state. in subgen. Anellaria, distinguished by its Psathyrella hirta Peck: in subgen. semiovate cap and patent veil remnants on its Psathyrella, sect. Atomatae Romagn. ex Singer margin or in the shape of an annulus. (small and delicate fruitbodies, not rooting Worldwide in occurrence, recorded in Italy and stems, gills with pinkish shades, comparatively elsewhere, especially from bovine or equine large and smooth spores, and broad, dung (Spegazzini 1881, 1925, Maire 1937, spheropedunculate and often 2-spored basidia), Christiansen 1941, Lacaze 1953, Hongo 1959, it is regarded as a constantly fimicolous species,

127 which is recognisable, in the early stages at and annulus. According to Noordeloos (2009), least, by its fibrillose veil strongly developed following phylogenetic studies (Moncalvo et al. both on the cap and stem. Widespread in 2000) and a proposal to conserve the name Europe and recorded from cattle (Kits van Psilocybe (Redhead et al. 2007), the halluci- Waveren 1972, 1985, Heykoop & Esteve- nogenic (bluing) species must be kept in Raventós 1994, Tassi 1997, De Meulder 2001, Psilocybe, whereas the non-hallucinogenic Vašutová 2006, Larsson & Örstadius 2008) and ones must be accomodated in Deconica (W.G. horse dung (Kits van Waveren 1972, 1985, Sm.) P. Karst. Perez Losantos & Bascones Carretero 1981, Psilocybe coprophila (Bull.) P. Kumm.: Enderle & Christan 1992, Tassi 1997, Larsson in sect. Merdariae (Fr.) Singer (coprophilous, & Örstadius 2008, Siquier & Salom 2008) in not bluing, with a central stem and com- the natural state, but also in culture (Hugueney paratively large, thick-walled spores), it is 1965, Esteve-Raventós & Barrasa 1989), similar to and macroscopically mistakable for unusually from roe deer dung (Derbsch & P. subcoprophila, in the same section, and P. Schmitt 1987). liniformans, in sect. Semilanceatae Guzmán. It Psathyrella lacrymabunda (Bull.) M.M. differs from the former in having hexagonal Moser: in subgen. Lacrymaria (Pat.) Singer & rather than ellipsoidal spores, from the latter in A.H. Sm. (fleshy fruitbodies with an abundant lacking a gelatinous gill edge and not bluing by cortiniform veil and an appendiculate cap handling. Common in Italy, particularly on margin, weeping gills, and verrucose spores), it cattle and horse dung in the natural state, and is distinguishable by its ochreous brown cap once isolated in culture. One of the commonest and a long stem. According to phylogenetic Psilocybe spp. from all continents, particularly analyses (Hopple & Vilgalys 1999, Larsson & from cattle (Bresadola 1931, Christiansen 1941, Örstadius 2008, Padamsee et al. 2008) Hongo 1965, Guzmán et al. 1977, Pegler 1977, P. lacrymabunda and allied species should be Watling & Gregory 1980, Stamets 1996, De described under the distinct genus Lacrymaria Meulder 2001, Calonge & Menezes de Pat. Exceptionally fimicolous; I have studied Sequeira 2003, Gularte Cortez & Coelho 2004) numerous Italian collections, but only one from and horse dung (Christiansen 1941, Dennis horse manure; reported elsewhere from rich 1961, Guzmán et al. 1977, Bigelow 1978, nitrogenous (Ortega Díaz & Gálan Márquez Yokoyama 1979, 1987, Stamets 1996, Granito 1981) or manured soils (Esteve-Raventós & & Lunghini 2004, Gularte Cortez & Coelho Barrasa 1989). 2004, Siquier & Salom 2005a) but, as supposed Psathyrella prona (Fr.) Gillet: some by Guzmán (1983), also growing on other dung forms and varieties are known of this types, e.g. hare (Hongo 1959, Richardson facultatively fimicolous species, which can be 2008a), sheep (Watling & Gregory 1987, distinguished from P. hirta (both in sect. Johnston & Buchanan 1995), elephant (Thomas Atomatae) particularly by its hardly developed & Manimohan 2002, Manimohan et al. 2007), veil and larger spores. I have revised several and marmot (Jamoni 1990). collections from Italy, only one from pure dung Psilocybe crobulus (Fr.) Singer: in sect. (horse). Reported on rich soils or rotten wood, Psilocybe (not bluing fruitbodies with a central rarely recorded from manure (Saccardo 1816) stem and comparatively small, usually flattened or dung (Larsson & Örstadius 2008, Örstadius spores), it is distinguishable by a well & Knudsen in Knudsen & Vesterholt 2008). developed veil, particularly at the cap margin, a There are also records on cattle (Esteve- viscid cuticle, and thin-walled spores. Usually Raventós & Barrasa 1989) and horse dung growing on vegetable debris, it has (Kits van Waveren 1985). occasionally been recorded on unspecified Psilocybe in Strophariaceae includes dung (Guzmán 1983, Watling & Gregory 1987, species with a hyphal pileipellis (cutis or Ludwig 2001, Vesterholt in Knudsen & ixocutis) and purple or purple-brown spores in Vesterholt 2008) or sawdust mixed with dung mass, which can be distinguished from (Noordeloos in Kuyper et al. 1999). Besides coprophilous Stropharia spp. by constantly our Italian records there are other records from lacking chrysocystidia and a glutinous stem

128 Mycosphere cattle dung (Breitenbach & Kränzlin 1995, water vole (Sengupta & Laessøe 2004), and elk Consiglio 1999b). (Vesterholt in Knudsen & Vesterholt 2008). Psilocybe cyanescens Wakef.: in sect. Psylocybe semilanceata (Fr.) P. Kumm.: Cyanescentes Guzmán (bluing species with a type species of sect. Semilanceatae, it stands convex cap and thick-walled, ellipsoidal to out for having a hardly expanding, typical mitriform, flattened spores), it is dis- campanulate to conic-paraboloid, umbonate tinguishable from P. semilanceata by stouter cap, almost free, not gelatinised gills, and not fruitbodies with a different cap shape, more strongly bluing tissues. It is a worldwide flattened spores, and presence of pleurocystidia. temperate species, especially recorded from It usually grows on rotten wood and among grazed meadows and fields, exceptionally from leaves on nitrogenous soils. Apart from our manure (Morgan 1907b), cattle (Doveri 2004), Italian records there appear to be no other or unspecified dung (Castro 2005). coprophilous collections. Psilocybe subcoprophila (Britzelm.) Psilocybe inquilina (Fr.) Bres.: similar to Sacc.: very close to and indistinguishable in the P. crobulus, from which it differs in having a field from P. coprophila, but with ellipsoidal, less developed veil and slightly larger spores. somewhat larger spores. It also resembles P. Occasionally fimicolous, it has been described angustispora A.H. Sm., a North American on horse dung (Høiland 1978, Ludwig 2001), fimicolous species both on domestic and wild rabbit (Vila & Llimona 1998), unspecified animal dung (Guzmán 1983), but it differs in dung (Moreno-Arroyo et al. 2005), and having wider cheilocystidia, and larger, meadows grazed by cattle and sheep (Watling especially broader spores (Guzmán 1983, & Gregory 1987). Guzmán & Trappe 2005). Widespread but not Psilocybe liniformans Guzmán & Bas: in so common as P. coprophila, with a single sect. Semilanceatae Guzmán (bluing, umbonate collection from our studies from wild pig dung, species with ellipsoidal, hardly flattened spores, and another Italian record from horse (Granito and no pleurocystidia), it is macroscopically & Lunghini 2004). Elsewhere especially closer to P. coprophila and P. subcoprophila observed on horse (Svrček 1959, Orton 1969, than to P. semilanceata (the latter in the same Høiland 1978, Noordeloos 1998, Guzmán & section). It is distinguished by a strongly Trappe 2005, Poumarat 2008b), but also gelatinised gill edge. Recorded only from horse recorded from sheep (Orton 1969, Watling & dung (Guzmán & Bas 1977, Guzmán 1983, Gregory 1987, Ludwig 2001), cattle (Horak Stamets 1996, Noordeloos 1998, in Kuyper 1979, Poumarat 2008b), hare (Noordeloos 1999, Esteve-Raventós et al. 2001, Fernández- 1998), and from dung of unspecified domestic Sasia 2001, Guzmán et al. 2006, Siquier & herbivores (Guzmán 1983, Noordeloos in Salom 2008, Vesterholt in Knudsen & Kuyper et al. 1999, Vesterholt in Knudsen & Vesterholt 2008). Vesterholt 2008). Psilocybe merdaria (Fr.) Ricken: like Stropharia, in Strophariaceae, includes P. coprophila, it has hexagonal, but slightly usually annulate species with a viscid cap and smaller spores, and differs also in having a acanthocytes in the basal mycelium, sharing a paler, usually non-striate cap, and patent veil hyphal pileipellis with Psilocybe, from which remnants, often in the shape of an annulus. they particularly differ in having pleuro- Twice recorded on cattle dung from Italy, and chrysocystidia. Most coprophilous species are widespread elsewhere, especially on bovine accomodated in sect. Stercophila (Romagn.) dung (Pegler 1977, Guzmán 1978, Jamoni Singer, which further differs from Psilocybe in 1993, Ortega et al. 1997, De Meulder 2001, having glutinous veil, cap, lower portion of Ludwig 2001, Pirlot 2003), and also from other stem, and annulus. Stropharia has also been domestic herbivores (Pilát 1972, Bigelow 1978, regarded as a later synonym of Psilocybe Watling & Gregory 1987, Noordeloos 1998, in (Noordeloos in Kuyper et al., 1999; Kirk et al., Kuyper et al. 1999, Richardson 2005, 2008b, 2008). All three species observed by us in Italy Vesterholt in Knudsen & Vesterholt 2008), belong to sect. Stercophila. occasionally from hare (Richardson 2005), Stropharia dorsipora Esteve-Rav. &

129 Barrasa: except for its always well developed sheep (Arnolds 1982, Watling & Gregory 1987, annulus, it is practically indistinguishable in Richardson 2005, Ryman in Knudsen & the field from S. semiglobata, from which it Vesterholt 2008), elk (Kytövuori 1999, Ryman also differs in having spores with an eccentric in Knudsen & Vesterholt 2008), other cervid germ pore, and a variable presence of (Batsch 1783) and hare dung (Richardson cheilochrysocystidia and caulochrysocystidia. 2008a). Usually recorded by us from horse dung, once Volvariella, in Pluteaceae Kotl. & in culture, rarely from cattle, it appears to be Pouzar (free gills, an inverse hymenophoral quite common in Europe and possibly over- trama, smooth spores without a germ pore), is looked. Besides some records from unspecified characterised by a volvate stem, pink gills, a dung, there are records from horse and cattle hyphal pileipellis, and pink spores in mass. dung (Esteve-Raventós & Barrasa 1995, They usually grow on soil, wood and compost, Kytövuori 1999, Noordeloos in Kuyper et al. and are rarely fimicolous. Volvariella gloioce- 1999, Hausknecht & Krisai-Greilhuber 2003, phala (DC.) Boekhout & Enderle: recognisable Gularte Cortez & Silveira 2008, Rubio & by having robust fruitbodies with a viscid, dirty Miranda 2008, Ryman in Knudsen & white to grey cap, a raphanoid smell, and Vesterholt 2008). comparatively large spores. Widespread in Stropharia luteonitens (Vahl) Quél.: the temperate zones. It usually grows on straw, only species in sect. Stercophila without sawdust, humus, manured soils. Besides our pleurochrysocystidia, easily recognisable also Italian record from horse dung, it has also been by its slender fruitbodies, a conic-campanulate reported from dung-hills (Massee 1893, Rea and papillate-umbonate cap with yellowish 1922). shades, large spores, and 2-spored basidia. Although it can grow in pastures and rich soils Acknowledgements (Baş Sermenli & Işiloğlu 2006), it has been The author is particularly indebted to recorded from dung-rich soils (Clemençon & Mike Richardson for critically revising the text. Roffler 2003), sometimes from unspecified He also thanks Luigi Arras, Bruno de Ruvo, dung, and from horse or cattle dung and the “Mycological Forum A.M.B. Gruppo (Spegazzini 1899, Cleland & Cheel 1918, di Muggia e del Carso” (http://www.amb Hongo 1959, Granito & Lunghini 2004). muggia.it/forum/) for providing him with a Stropharia semiglobata (Batsch) Quél.: share of the material subject of this study. unlike S. dorsipora, it has a rudimentary annulus, spores with a central germ pore, and a References constant presence of cheilo-, pleuro- and caulo- Alves MH, Cavalcanti MA. 1996 – Coprina- chrysocystidia. A common, widespread copro- ceae en el campus de la Universidad philous species worldwide, it has been reported Federal de Pernambuco (Recife. Pe, from herbivore dung in general (Noordeloos in Brasil). Boletin Micológico 11, 33–40. Kuyper et al. 1999), but most Italian records Arnolds E. 1982 – Ecology and coenology of come from cattle and horse dung in the field, macrofungi in grasslands and moist and one record also from cattle dung in culture. heathlands in Drenthe, the Netherlands. There is a noticeable preference for cattle and Bibliotheca Mycologica 90. J. Cramer. horse dung (Spegazzini 1899, Buller 1922, Vaduz. Spegazzini 1925, Christiansen 1941, Romag- Arnolds E. 2003a – Notulae ad Floram nesi 1937, Hongo 1959, Dennis 1961, Calonge Agaricinam Neerlandicam – XXXIX. 1968, Bon 1970, Pilát 1972, Watling & Bolbitius. Persoonia 18, 201–214. Gregory 1980, Yokoyama 1984, Jamoni 1994, Arnolds E. 2003b – Rare and interesting Esteve-Raventós & Barrasa 1995, Mayoral & species of Psathyrella. In: Fungi non Angel 1995, Stamets 1996, Ortega et al. 1997, Delineati XXVI (ed. M Candusso). Kytövuori 1999, De Meulder 2001, Cortez & Edizioni Candusso, Alassio, Italia. Coelho 2004, Welt & Heine 2007, Gularte Babos M. 2004 – Macromycètes des dunes de Cortez & Silveira 2008, Ryman in Knudsen & l’île Szentendre (Danube, Hongrie). Vesterholt 2008). There are other records from

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