Fungi Fimicoli Italici - Mycosphere-online journal

Mycosphere 

Additions to “Fungi Fimicoli Italici”: An update on the occurrence of 

coprophilous Basidiomycetes and Ascomycetes in Italy with new records and 

descriptions. 

Doveri F* 

Via Baciocchi 9, I–57126–Livorno. 

Doveri F. 2011 – Addition to “Fungi Fimicoli Italici”: An update on the occurrence of coprophilous 

Basidiomycetes and Ascomycetes in Italy with new records and descriptions. Mycosphere 2(4), 

331–427. 

“Fungi Fimicoli Italici”, the first monograph on coprophilous fungi from Italy, is regarded as the 

starting point of a survey on Basidiomycetes and Ascomycetes obligatorily or facultatively growing 

on any kind of dung. All fimicolous species recorded from Italy and described in that work or in 

subsequent author's papers are listed and their dung sources are mentioned. The occurrence on dung 

of different genera and species is reported in separate Tables, and their frequency on different types 

of excrements is discussed and compared with other parts of the world. An update on the families 

Gymnoascaceae, Microascaceae and Sporormiaceae, and keys to genera of Gymnoascaceae and 

Microascaceae, and to Italian species of Gymnoascus s.l. and Sporormiella with 8-celled ascospores 

are provided. Chaetomidium fimeti, C. ancistrocladum, C. murorum, Gymnoascus dankaliensis, G. 

littoralis, G. ruber, Hypocopra equorum, Iodophanus difformis, Kernia cauquensis, Lophotrichus 

bartlettii, Orbicula parietina, Pithoascus intermedius, and Sporormiella affinis are first described 

from Italy. 

Key words – damp chambers – fimicolous fungi – frequency – keys – natural state – survey 

Article Information 

Received 7 March 2011 

Accepted 18 April 2011 

Published online 14 September 2011 

*Corresponding author: Francesco Doveri – e-mail – f.doveri@sysnet.it 

Introduction 

“Fungi Fimicoli Italici” (Doveri 2004a) is 

the first monograph on Basidiomycetes and 

Ascomycetes living on faecal material in Italy, 

and one of few recent monographs worldwide 

on coprophilous fungi. I have stressed before 

(Doveri 2004a, 2010b) that this work has more 

than a regional significance and can be helpful 

to studies of these fungi in any part of the 

world, which so very easily spread through cattle 

freight. The finding, after my monograph, of 

several taxa new to Italy, developed either in 

the naural state or in damp chamber cultures, 

induced me to update the work with the introduction 

of new keys, descriptions of species, 

and an account on their ecology. 

The present work must be regarded as 

complementary to the paper recently published 

in this journal (Doveri 2010b). 

Methods 

Dung samples of the following animals 

were collected intermittently from January 

1992 to December 2010 and usually incubated 

in non-sterile damp chambers. 

The design of the damp chambers, the 

examination of the incubating dung samples, 

and the description of macroscopic features of 

species appearing on incubated dung or detected 

in the field, follows those suggested by 

Doveri (2004a, 2010b). 

331

The microscopic characteristics of species 

from damp chambers were studied on fresh 

material, those of species collected from the 

field were usually studied on dried material, 

using water, Congo red, Melzer's reagent, lactic 

cotton blue (or methyl blue), and Indian ink as 

mounting media. 

All Basidiomycetes and most Ascomycetes 

were preserved as dried material or, 

exceptionally, as slides in the author's personal 

herbarium (CLSM) or rarely in herbarium of 

Pisa Botanical Garden (PI). Chaetomium murorum, 

Cleistothelebolus nipigonensis, and Rodentomyces 

reticulatus were also preserved as 

living cultures (PI) 

Results 

In my monograph on coprophilous fungi 

from Italy (Doveri 2004a) I described 81 taxa 

332 

of Basidiomycota and 214 of Ascomycota 

growing both in the natural state or in damp 

chamber cultures. Since then my research has 

continued resulting in many publications 

(Doveri 2004b, 2005, 2006, 2007a,b, 2008a,b,c, 

2010a,b, Doveri et al. 2005, 2010a,b, Doveri & 

Coué 2008a) reporting 43 additional species (5 

Basidiomycota; 38 Ascomycota) new to Italy, 

13 of which are described in this work, and one 

genus (Rodentomyces Doveri et al.), four 

species (Podospora alexandri Doveri, Sporormiella 

hololasia Doveri, Thecotheus neoapiculatus 

Doveri & Coué, Coprinellus mitrinodulisporus 

Doveri & Sarrocco), one variety 

(Tripterosporella heterospora var. octaspora 

Doveri) and one form (Thecotheus formosanus 

f. collariatus Doveri & Coué) new to science 

(Table 1). 

badger (Meles meles) goose (Anser sp.) porcupine (Hystrix cristata) 

beech marten (Martes faina) hare (Lepus sp.) rabbit (Oryctolagus cuniculus) 

bird (various, except for Anser sp.) hedgehog (Erinaceus europaeus) rat (Rattus rattus) 

cattle (Bos taurus) horse (Equus caballus) rock goat (Capra ibex ibex) 

chamois (Rupicapra rupicapra) insect (various) roe deer (Capreolus capreolus) 

deer (Cervus elaphus) lizard (Lacerta sp.) sheep (Ovis aries) 

dog (Canis familiaris) marmot (Marmota marmota) snail (Helix sp.) 

donkey (Equus asinus) marten (Martes martes) squirrel (Sciurus sp.) 

dormouse (Glis glis) mouflon (Ovis musimon) toad (Bufo sp.) 

fallow deer (Dama dama) mouse (Mus musculus) tortoise (Testudo sp.) 

ferret (Mustela furo) mule (Equus mulus) weasel (Putorius nivalis) 

fox (Vulpes vulpes) ostrich (Struthio camelus) wild pig (Sus scrofa) 

gecko (Tarentola mauritanica) pig (Sus scrofa domesticus) wolf (Canis lupus) 

goat (Capra hircus) polecat (Putorius foetidus) 

I have recorded, at present, 544 collections 

of 92 Basidiomycota species from 722 

dung samples in Italy. 

303 records of 88 species came from 303 

samples detected and identified in the field, 

listed as substrate sources, but not cultured. All 

but two (Sebacina epigaea; Sphaerobolus 

stellatus) are Agaricales, whose occurrence in 

Italy has recently been reported (Doveri 2010 

b). The overall picture is unchanged, so I limit 

myself to summarise the results (Tables 2–3) 

and refer to that work for details. 

I have recorded, at present, 2312 collections 

of 256 Ascomycota species from Italy. 

573 records of 156 species came from samples 

detected and identified in the field. Leaving out 

29 records from unidentified animals, most 

findings (96%) were made on bovine (27%), 

cervine (23%), equine (18%), leporine (13%), 

ovine (12%), and caprine (3%) dung, the 

remaining 4% on dung of other animals, with a 

slight preponderance of pyrenomycetes s.l. 

(53%) on discomycetes (45%), and with a 

marked preponderance of Sporormiella (19%), 

followed by Ascobolus Pers. (8%), Lasiobolus 

Sacc. (8%), Sordaria Ces. & De Not. (8%), 

Cheilymenia Boud. (7%), Podospora Ces., 

(7%), and Schizothecium Corda (7%). The 

preference for bovine dung is marked (72%) in 

Cheilymenia, slighter in Ascobolus (29%) and 

Schizothecium (20%), while a preference for 

cervine dung is manifest in Lasiobolus (38%), 

Sporormiella (30%) and Podospora (29%), for 

leporine dung (36%) in Sordaria. 

Another 432 dung samples of 42 animal 

species were cultured in damp chambers and

Table 1 List of coprophilous Basidiomycota and Ascomycota from Italy. 

BASIDIOMYCOTA 

Agrocybe molesta (Lasch) 

Singer 

Agrocybe pediades (Fr.) 

Fayod s. Watling 

Agrocybe praecox (Pers.) 

Fayod 

Agrocybe subpediades 

(Murrill) Watling s. 

Watling 

Agrocybe temulenta (Fr.) 

Singer s. Watling 

Bovista aestivalis (Bonord.) 

Demoulin 

Bolbitius coprophilus 

(Peck) Hongo 

Bolbitius vitellinus var. 

titubans (Bull.) Bon & 

Courtec. 

Bolbitius vitellinus var. 

variicolor (G.F. Atk.) 

Krieglst. 

Bolbitius vitellinus (Pers.) 

Fr. var. vitellinus 

Conocybe alboradicans 

Arnolds 

Conocybe antipus (Lasch) 

Fayod 

Conocybe aurea (Jul. 

Schäff.) Hongo 

Conocybe brunneidisca 

(Murrill) Hauskn. 

Conocybe cettoiana 

Hauskn. & Enderle 

Conocybe coprophila 

(Kühner) Kühner 

Conocybe fuscimarginata 

(Murrill) Singer 

Conocybe gigasperma 

Enderle & Hauskn. 

Conocybe pubescens 

(Gillet) Kühner 

Conocybe rickenii (Jul. 

Schäff.) Kühner 

Conocybe siennophylla 

(Berk. & Broome) Singer 

Conocybe siliginea (Fr.) 

Kühner 

Conocybe singeriana 

Hauskn. 

Stropharia dorsipora 

Esteve–Rav. & Barrasa 

Stropharia luteonitens 

(Vahl) Quél. 

Stropharia semiglobata 

(Batsch) Quél. 

Volvariella gloiocephala 

(DC.) Boekhout & Enderle 

ASCOMYCOTA 

Aphanoascus fulvescens 

(Cooke) Apinis 

Arnium arizonense 

(Griffiths) N. Lundq. & 

J.C. Krug 

Arnium caballinum N. 

Lundq. 

Arnium cervinum N. 

Lundq. 

Coprotus sexdecimsporus 

(H. Crouan & P. Crouan) 

Kimbr. & Korf 

Coprotus 

subcylindrosporus J. 

Moravec 


Saccobolus saccoboloides 

(Seaver) Brumm. 

Saccobolus succineus 

Brumm. 

Delitschia didyma Auersw. Saccobolus truncatus 

Velen. 

Delitschia gigaspora Cain Saccobolus verrucisporus 

Brumm. 

Delitschia gigaspora Cain 

var. ceciliae Doveri 

Delitschia intonsa Luck– 

Allen 

Delitschia leptospora 

Oudem. 

Delitschia marchalii Berl. 

& Voglino 

Arnium imitans N. Lundq. Delitschia patagonica 

Speg. 

Arnium inaequilaterale 

(Cain) N. Lundq. & J.C. 

Krug 

Arnium septosporum N. 

Lundq. 

Delitschia vulgaris 

Griffiths 

Delitschia winteri Plowr. 

ex G. Winter 

Arnium sudermanniae N. Enterocarpus grenotii 

Lundq. 

Locq.-Lin. 

Ascobolus albidus H. Fimetariella microsperma 

Crouan & P. Crouan J.C. Krug 

Ascobolus brassicae H. Gymnoascus dankaliensis 

Crouan & P. Crouan (Castell. ex J.F.H. Beyma) 

Arx 

Ascobolus carletoni Boud. Gymnoascus devroeyi (G.F. 

Orr) Arx 

Ascobolus costantinii Gymnoascus littoralis (G.F. 

Rolland 

Orr) Arx 

Ascobolus crenulatus P. 

Karst. 

Saccobolus versicolor (P. 

Karst.) P. Karst. 

Schizothecium aloides 

(Fuckel) N. Lundq. 

Schizothecium conicum 


Schizothecium dakotense 

(Griffiths) N. Lundq. 

Schizothecium glutinans 

(Cain) N. Lundq. 

Schizothecium inaequale 

(Cain) N. Lundq. 

Schizothecium 

miniglutinans (Mirza & 

Cain) N. Lundq. 

Schizothecium pilosum 

(Mouton) N. Lundq. 

Schizothecium simile (E.C. 

Hansen) N. Lundq. 

Schizothecium 

squamulosum (H. Crouan 

& P. Crouan) N. Lundq. 

Schizothecium tetrasporum 

(G. Winter) N. Lundq. 

Schizothecium vesticola 

(Berk. & Broome) N. 

Lundq. 

Gymnoascus reessii Baran. Schizothecium 

vratislaviense (Alf. 

Schmidt) Doveri & Coué 

Ascobolus elegans J. Klein Gymnoascus ruber Tiegh. Scutellinia crinita (Bull) 

Lambotte 

Ascobolus furfuraceus Pers. Hypocopra antarctica Selinia pulchra (G. Winter) 

(Speg.) Furuya & Udagawa Sacc. 

Ascobolus hawaiiensis Hypocopra brefeldii Zopf Sordaria fimicola 

Brumm. 

(Roberge) Ces. & De Not. 

Ascobolus immersus Pers. Hypocopra equorum Sordaria humana (Fuckel) 

ex Pers. 

(Fuckel) G. Winter G. Winter 

Ascobolus lineolatus Hypocopra aff. festucacea Sordaria lappae Potebnia 

Brumm. 

J.C. Krug & Cain 

Ascobolus mancus (Rehm) Hypocopra leporina (Niessl Sordaria macrospora 

Brumm. 

ex Rehm) Doveri 

Auersw. 

333

Table 1 (Continued) List of coprophilous Basidiomycota and Ascomycota from Italy. 

Coprinellus bisporus (J.E. Ascobolus michaudii Boud. Hypocopra lojkaeana Sordaria superba De Not. 

Lange) Vilgalys et al. 

(Rehm) Doveri 

Coprinellus brevisetulosus Ascobolus aff. pseudocainii Hypocopra merdaria (Fr.) “Sordaria” minima Speg. 

(Arnolds) Redhead et al. Prokhorov 

J. Kickx f. 

& Sacc. 

Coprinellus congregatus Ascobolus reticulatus Iodophanus carneus (Pers.) Sphaeronaemella fimicola 

(Bull.) P. Karst. 

Brumm. 

Korf 

Marchal 

Coprinellus curtus Ascobolus 

Iodophanus difformis (P. Sporormia fimetaria (De 

(Kalchbr.) Vilgalys et al. roseopurpurascens Rehm Karst.) Kimbr. et al. Not.) De Not. 

Coprinellus doverii (Nagy) Ascobolus stictoideus Speg. Kernia cauquensis Sporormiella affinis (Sacc. 

Nagy 

Calviello 

et al.) S.I. Ahmed & Cain 

Coprinellus ephemerus Ascodesmis microscopica Kernia nitida (Sacc.) Sporormiella antarctica 

(Bull.) Redhead et al. (H. Crouan & P. Crouan) 

Seaver 

Nieuwl. 

(Speg.) S.I. Ahmed & Cain 

Coprinellus flocculosus Ascodesmis nana Brumm. Lasiobolus ciliatus (JC. Sporormiella australis 

(DC.) Vilgalys et al. 

Schmidt) Boud. 


Coprinellus heptemerus Ascodesmis nigricans Lasiobolus cuniculi Velen. Sporormiella capybarae 

(M. Lange & A.H. Sm.) 

Vilgalys et al. 

Tiegh. 


Coprinellus heterosetulosus Ascozonus woolhopensis Lasiobolus diversisporus Sporormiella corynespora 

(Locq. ex Watling) 

Vilgalys et al. 

(Renny) Boud. 

(Fuckel) Sacc. 

(Niessl) S.I. Ahmed & Cain 

Coprinellus marculentus Bombardioidea stercoris Lasiobolus intermedius J.L. Sporormiella cylindrospora 

(Britzelm.) Redhead et al. (DC.) N. Lundq. 

Bezerra & Kimbr. 

S.I. Ahmed & Cain 

Coprinellus pellucidus (P. Cercophora anisura N. Lasiobolus macrotrichus Sporormiella cymatomera 

Karst.) Redhead et al. Lundq. 

Rea 


Coprinellus sassii (M. Cercophora coprophila Lasiobolus microsporus Sporormiella dodecamera 

Lange & A.H. Sm.) 

Redhead et al. 

(Fr.) N. Lundq. 

J.L. Bezerra & Kimbr. S.I. Ahmed & Cain 

Coprinopsis candidolanata Cercophora gossypina N. Lasiobolus monascus Sporormiella dubia S.I. 

(Doveri & Uljé) Keirle et 

al. 

Lundq. 

Kimbr. 

Ahmed & Cain 

Coprinopsis cinerea Cercophora mirabilis Lasiobolus ruber (Quél.) Sporormiella gigantea 

(Schaeff.) Redhead et al. Fuckel 

Sacc. 

(E.C. Hansen) Doveri 

Coprinopsis cothurnata Cercophora septentrionalis Lophotrichus bartlettii Sporormiella grandispora 

(Godey) Redhead et al. N. Lundq. 

(Massee & E.S. Salmon) 

Malloch & Cain 


Coprinopsis filamentifera Chaetomidium 

Melanospora brevirostris Sporormiella heptamera 

(Kühner) Redhead et al. cephalothecoides (Malloch (Fuckel) Höhn. 

(Auersw.) S.I. Ahmed & 

& Benny) Arx 

Cain 

Coprinopsis luteocephala Chaetomidium fimeti Melanospora zamiae Corda Sporormiella hololasia 

(Watling) Redhead et al. (Fuckel) Sacc. 

Doveri 

Coprinopsis macrocephala Chaetomidium 

Neocosmospora vasinfecta Sporormiella intermedia 

(Berk.) Redhead et al. megasporum Doveri et al. E.F. Sm. var. vasinfecta (Auersw.) S.I. Ahmed & 

Cain 

Coprinopsis nivea (Pers.) Chaetomium 

Orbicula parietina Sporormiella kansensis 


ancistrocladum Udagawa (Schrad.) S. Hughes (Griffiths) S.I. Ahmed & 

& Cain 

Cain 

Coprinopsis poliomallus Chaetomium bostrychodes Persiciospora moreaui P.F. Sporormiella lageniformis 

(Romagn.) Doveri et al. Zopf 

Cannon & D. Hawksw. (Fuckel) S.I. Ahmed & 

Cain 

Coprinopsis 

Chaetomium carinthiacum Peziza fimeti (Fuckel) E.C. Sporormiella lasiocarpa 

pseudocortinata (Locq. ex 

Doveri et al.) Doveri et al. 

Sörgel 

Hansen 

Lorenzo 

Coprinopsis pseudonivea Chaetomium crispatum Peziza merdae Donadini Sporormiella leporina 

(Bender & Uljé) Redhead 

et al. var. pseudonivea 

(Fuckel) Fuckel 


Coprinopsis pseudoradiata Chaetomium cuniculorum Peziza perdicina (Velen.) Sporormiella 

(Watling) Redhead et al. Fuckel 

Svrček 

longisporopsis S.I. Ahmed 

& Cain 

334


Coprinopsis radiata 

(Bolton) Redhead et al. 

Coprinopsis stercorea 

(Scop. ex Fr.) Redhead et 

al. 

Coprinopsis tuberosa 

(Quél.) Doveri et al. 

Coprinopsis utrifera 

(Watling) Redhead et al. 

Coprinopsis vermiculifera 

(Dennis) Redhead et al. 

Coprinopsis xenobia (P.D. 

Orton) Redhead et al. 

Coprinus ephemeroides 

(Bull.) Fr. 

Coprinus patouillardii 

Quél. 

Coprinus spadiceisporus 

Bogart 

Coprinus sterquilinus (Fr.) 

Fr. 

Crucibulum laeve (Huds.) 

Kambly 

Cyathus stercoreus 

(Schwein.) De Toni 

Leucocoprinus cretaceus 

(Bull.) Locq. 

Lepista sordida 

(Schumach.) Singer 

Panaeolus acuminatus 

(Schaeff.) Quél. 

Chaetomium elatum Kunze: 

Fr 

Chaetomium funicola 

Cooke 

Chaetomium fusisporum G. 

Sm. 

Chaetomium gangligerum 

L.M. Ames 

Chaetomium globosum 

Kunze: Fr 

Chaetomium homopilatum 

Omvik 

Chaetomium medusarum 

J.A. Mey. & Lanneau 

Chaetomium mollicellum 

L.M. Ames 

Chaetomium murorum 

Corda 

Chaetomium oblatum 

Dreyfuss & Arx 

Chaetomium robustum 

L.M. Ames 

Chaetomium semen–citrulli 

Sergeeva 

Chaetomium spinosum 

Chivers 

Chaetomium subaffine 

Sergeeva 

Chaetomium 

trigonosporum (Marchal) 

Chivers 

Chaetomium variostiolatum 

A. Carter 

Chalazion erinaceum 

Panaeolus alcis M.M. 

Moser 

Panaeolus antillarum (Fr.) 

Dennis 

Doveri et al. 

Panaeolus cinctulus Cheilymenia 

(Bolton) Sacc. 

aurantiacorubra K.S. 

Thind & S.C. Kaushal 

Panaeolus fimicola (Fr.) Cheilymenia coprinaria 

Gillet 

(Cooke) Boud. 

Panaeolus guttulatus Bres. Cheilymenia dennisii J. 

Moravec 

Panaeolus papilionaceus Cheilymenia fraudans (P. 

(Bull.) Quél. 

Karst.) Boud. 

Panaeolus retirugis (Fr.) Cheilymenia granulata 

Gillet 

(Bull.) J. Moravec 

Panaeolus sphinctrinus Cheilymenia insignis (H. 

(Fr.) Quél. 

Crouan & P. Crouan) Boud. 

Panaeolus semiovatus 

(With.) S. Lundell 

Cheilymenia pulcherrima 

(H. Crouan & P. Crouan) 

Boud. 


Peziza vesiculosa Bull. Sporormiella megalospora 


Pithoascus intermedius 

(C.W. Emmons & B.O. 

Dodge) Arx 

Pleospora herbarum (Pers.) 

Rabenh. 

Podospora alexandri 

Doveri 

Podospora anserina (Ces. 

ex Rabenh.) Niessl 

Podospora australis 

(Speg.) Niessl 

Podospora bifida N. 

Lundq. 

Podospora communis 

(Speg.) Niessl 

Podospora curvicolla (G. 

Winter) Niessl 

Podospora dasypogon N. 

Lundq. 

Podospora decipiens (G. 

Winter ex Fuckel) Niessl 

Podospora excentrica N. 

Lundq. 

Podospora fimiseda (Ces. 

& De Not.) Niessl 

Podospora gigantea Mirza 

& Cain 

Podospora globosa 

(Massee & E.S. Salmon) 

Cain 

Podospora granulostriata 

N. Lundq. 

Podospora intestinacea N. 

Lundq. 

Podospora myriaspora (H. 

Crouan & P. Crouan) 

Niessl 

Podospora pleiospora (G. 


Podospora pyriformis (A. 

Bayer) Cain 

Podospora setosa (G. 


Poronia erici Lohmeyer & 

Benkert 

Poronia punctata (L.) 

Rabenh. 

Preussia fleischhakii 

(Auersw.) Cain 

Cain 

Sporormiella minima 


Cain 

Sporormiella minimoides 


Sporormiella minipascua 


Sporormiella octomera 


Cain 

Sporormiella octonalis S.I. 

Ahmed & Cain 

Sporormiella pascua 


Sporormiella pilosa (Cain) 


Sporormiella pulchella 

(E.C. Hansen) S.I. Ahmed 

& Cain 

Sporormiella teretispora 


Sporormiella trogospora 


Sporormiella vexans 


Cain 

Strattonia insignis (E.C. 

Hansen) N. Lundq. 

Thecotheus cinereus (H. 


Chenant. 

Thecotheus crustaceus 

(Starbäck) Aas & N. 

Lundq. 

Thecotheus formosanus Y.– 

Z. Wang 

Thecotheus formosanus f. 

collariatus Doveri & Coué 

Thecotheus holmskjoldii 

(E.C. Hansen) Chenant. 

Thecotheus lundqvistii Aas 

Thecotheus neoapiculatus 

Doveri & Coué 

Thecotheus pelletieri (H. 

Crouan & P. Crouan) Boud. 

Thecotheus strangulatus 

(Velen.) Aas & N. Lundq. 

Thelebolus caninus 

(Auersw.) Jeng & J.C. 

Krug 

Thelebolus crustaceus 

(Fuckel) Kimbr. 

335


Panaeolus subfirmus P. 

Karst. 

Parasola misera (P. Karst.) 


Parasola schroeteri (P. 

Karst.) Redhead et al. 

Psathyrella coprinoides 

Delannoy et al. 

336 

Cheilymenia rubra (W. 

Phillips) Boud. 

Cheilymenia stercorea 

(Pers.) Boud. 

Cheilymenia theleboloides 

(Alb. & Schwein.) Boud. 

Cleistothelebolus 

nipigonensis Malloch & 

Cain 

Psathyrella hirta Peck Coniochaeta leucoplaca 

(Sacc.) Cain 

Psathyrella lacrymabunda Coniochaeta scatigena 

(Bull.) M.M. Moser (Berk. & Broome) Cain 

Psathyrella prona (Fr.) 

Gillet var. prona f. prona 

Psilocybe coprophila 

(Bull.) P. Kumm. 

Psilocybe crobulus (Fr.) 

Singer 

Psilocybe cyanescens 

Wakef. 

Psilocybe inquilina (Fr.) 

Bres. 

Psilocybe liniformans 

Guzmán & Bas 

Psilocybe merdaria (Fr.) 

Ricken 

Psilocybe subcoprophila 

(Britzelm.) Sacc. 

Psylocybe semilanceata 

(Fr.) P. Kumm. 

Sebacina epigaea (Berk. & 

Broome) Neuhoff 

Sphaerobolus stellatus 

Tode : Pers. 

Coniochaeta vagans 

(Carestia & De Not.) N. 

Lundq. 

Copromyces bisporus N. 

Lundq. 

Coprotus aurora (H. 


Kimbr. et al. 

Coprotus disculus Kimbr. 

et al. 

Coprotus glaucellus 

(Rehm) Kimbr. 

Coprotus granuliformis (H. 


Kimbr. 

Coprotus lacteus (Cooke & 

W. Phillips in Cooke) 

Kimbr. et al. 

Coprotus leucopocillum 

Kimbr. et al. 

Coprotus aff. luteus Kimbr. 

et al. 

Coprotus niveus (Fuckel) 

Kimbr. et al. 

Coprotus aff. ochraceus 

(H. Crouan & P. Crouan) J. 

Moravec 

provided 1739 collections of 203 Ascomycota 

species. Leaving out 7 collections from unidentified 

animals, most records (81%) came from 

equine (20%), bovine (17%), ovine (15%), cervine 

(10%), leporine (10%), and caprine (9%) 

dung, with a preponderance of pyrenomycetes 

s.l. (59%) on discomycetes (40%) and a high 

percentage (14%) of Podospora, followed by 

Ascobolus (10%), Saccobolus Boud. (9%), 

Schizothecium (9%), Chaetomium Kunze (8%), 

Sporormiella (8%), and Sordaria (7%). 

Preussia funiculata Thelebolus dubius var. 

(Preuss) Fuckel 

lagopi (Rea) Doveri 

Preussia isomera Cain Thelebolus microsporus 

(Berk. & Broome) Kimbr. 

Preussia terricola Cain Thelebolus polysporus (P. 

Karst.) Y. Otani & 

Kanzawa 

Preussia typharum (Sacc.) Thelebolus stercoreus Tode 

Cain 

Pseudombrophila bulbifera 

(E.J. Durand) Brumm. 

Pseudombrophila 

fuscolilacina (Grélet) 

Brumm. 

Pseudombrophila merdaria 

(Fr.) Brumm. 

Pseudombrophila minuta 

Brumm. 


theioleuca Rolland 

Trichobolus octosporus 

J.C. Krug 

Trichobolus sphaerosporus 

Kimbr. 

Trichobolus zukalii 

(Heimerl) Kimbr. 

Trichodelitschia minuta 


Trichodelitschia munkii N. 

Lundq. 

Rodentomyces reticulatus Trichophaea gregaria 

Doveri et al. 

(Rehm) Boud. 

Saccobolus beckii Heimerl Tripterosporella 

heterospora var. octaspora 

Doveri 

Saccobolus caesariatus Tripterosporella pakistani 

Renny 

(Mirza) Malloch & Cain 

Saccobolus citrinus Boud. 

& Torrend 

Saccobolus depauperatus 

(Berk. & Broome) E.C. 

Hansen 

Saccobolus dilutellus 

(Fuckel) Sacc. 

Saccobolus glaber (Pers.) 

Lambotte 

Saccobolus minimus Velen. 

Xanthothecium peruvianum 

(Cain) Arx & Samson 

Zopfiella erostrata 

(Griffiths) Udagawa & 

Furuya 

Zopfiella longicaudata 

(Cain) Arx 

Zygopleurage zygospora 

(Speg.) Boedijn 

In order to make the reading easier, I 

have gathered groups of species belonging to 

different genera in separate Tables, containing 

records from the field and the frequency in 

damp chamber cultures, and I have organised 

them following the systematics adopted by 

Kirk et al. (2008): 

Dothideomycetes – Pleosporomycetidae – 

Pleosporales – Delitschiaceae 

Delitschia Auersw (Table 4)

The genus Delitschia as a whole has a 

low frequency of occurrence on total dung 

samples in culture (1.1%), and a significant 

substrate preference (14%) for cattle dung. D. 

winteri preferably grows on cattle dung and, 

after Lundqvist (1972), it can be called 

“species with a high substrate preference” but 

also “with a wide tolerance” as it can inhabit a 

variety of dung types. Like mine from Italy, 

most records elsewhere (Phillips & Plowright 

1874, Winter 1874, Spegazzini 1878,Marchal 

1883, Griffiths 1901, Massee & Salmon 1901, 

Griffiths & Seaver 1910, Munk 1948, 1957, 

Lundqvist 1960, Breton 1965, Tóth 1965, 

1967, García–Zorron 1973, Wicklow et al. 

1980, Dennis 1981, Barrasa 1985, Barrasa & 

Checa 1990, Moyne & Petit 2006, Welt & 

Heine 2006a, Richardson 2008a,b) are from 

cattle dung (34%), many from horse (25%), 

rabbit (20%), and sheep (14%). 


Pleosporales – Phaeotrichaceae 

Trichodelitschia Munk (Table 5) 

Leaving out pig dung (samples too scarce 

to estimate the frequency of occurrence), T. 

minuta, the commonest Trichodelitschia species 

in Italy, shows a slight preference for leporine 

dung, compared with cervine and leporine, 

and an increased preference when also records 

from the field are considered. It is a common 

species worldwide, and there are many records 

from Europe, mostly (56%) from leporine dung 

(Marchal 1883, Lundqvist 1960, Tóth 1963, 

Barrasa 1985, Valldosera 1991, Eriksson 1992, 

Richardson 2005, Heine & Welt 2008). 


Pleosporales – Pleosporaceae 

Pleospora Rabenh ex Ces. & De Not. (Table 6) 


Pleosporales – Sporormiaceae 

Preussia Fuckel (Table 7) 

Refer to the taxonomic part of this work, 

under Sporormiella affinis, for discussion on 

taxonomy of Preussia and Sporormiella. In 

Italy the genus Preussia is less frequent (0.9%) 

on dung than Sporormiella (6.6%), and Preussia 

spp. appear to have a wide substrate tole- 


rance, with the exception of P. isomera, which 

has always been found on horse dung. Unlike 

mine, the few world records of P. isomera 

known by me are usually from rabbit dung 

(Cain 1961, Mukerji 1970, Angel & Wicklow 

1975). 

Sporormia De Not. (Table 8) 

Sporormia fimetaria is a rare but widespread 

species, preferably growing on bovine 

(44% of records worldwide) (De Notaris 1849, 

Pirotta 1878, Spegazzini 1878, Marchal 1883, 

Griffiths 1901, Massee & Salmon 1901, Angel 

& Wicklow 1975, Eriksson 1992, Chang & 

Wang 2009) and ovine (23%) dung (Stratton 

1921, Narendra 1973, Richardson 1998). It 

shows, however, a wide substrate tolerance, as 

it has been recorded also from goat (Ahmed & 

Cain 1972, Ahmad & Sultana 1973, Udagawa 

& Sugiyama 1982) and a variety of dungs 

(Khan & Cain 1979, Dissing 1992, Richardson 

2004a, 2008b) 

Sporormiella Ellis & Everh. (Table 9) 

Sporormiella is one of the most commonly 

encountered genera on dung (Bell 2005) and 

the most recorded by me from the field (19% 

of all records), particularly from cervine (30%) 

and leporine (22%) dung. It frequently occurs 

(8% of all samples) also in damp chamber cultures, 

and its frequency is much higher when 

only caprine and leporine dungs are considered. 

It shows, however, a wide tolerance for a 

variety of dungs. 

S. intermedia and S. minima are two of 

the most common coprophilous pyrenomycetes 

s.l. in Italy, but less frequent than Sordaria 

fimicola, Podospora decipiens and Chaetomium 

bostrychodes. Both have a wide substrate 

tolerance, but S. intermedia preferably grows 

on caprine (26%) and leporine (26%) dung in 

culture, S. minima on ovine (25%) and caprine 

(23%) dung. My data concerning S. intermedia 

are in agreement with Richardson (2001a), who 

found it to be most frequent on leporine dung. 

I have found more than 300 records worldwide 

of S. minima, confirming its wide substrate 

tolerance, but also a preference (23%) for 

bovine dung (Griffiths 1901, Lundqvist 1960, 

Tóth 1965, Parker 1979, Lundqvist 1997, De 

Meulder 2000b, Welt & Heine 2006b, 

Richardson 2008c, Chang & Wang 2009). 

337

Table 2 Records from Italy of coprophilous Basidiomycota in the natural state. 

equine cervine 

cattle donkey horse mule deer fallow deer roe deer chamois rabbit wild pig unidentified herbivore 

Bovista aestivalis 1 

Agrocybe molesta 1 

Agrocybe pediades s. Watling 4 1 1 

Agrocybe praecox 1 

Agrocybe subpediades s. Watling 2 1 

Agrocybe temulenta s. Watling 1 1 

Total Agrocybe 7 5 1 

Bolbitius coprophilus 5 

Bolbitius vitellinus var. titubans 1 

Bolbitius vitellinus var. variicolor 8 

Bolbitius vitellinus var. vitellinus 4 1 

Total Bolbitius 5 1 13 

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 

Coprinellus bisporus 2 1 1 3 

Coprinellus brevisetulosus 3 

Coprinopsis cinerea 4 1 2 5 

Coprinellus congregatus 3 2 1 1 

Coprinopsis cothurnata 4 1 

Coprinellus curtus 2 

Coprinellus doverii 1 

338

Table 2 (Continued) Records from Italy of coprophilous Basidiomycota in the natural state. 




“Coprinus” ephemeroides 1 3 

Coprinellus ephemerus 6 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 

Coprinellus mitrinodulisporus 1 

Coprinopsis nivea 10 1 4 

“Coprinus” patouillardii 2 4 

Coprinellus pellucidus 1 

Coprinopsis poliomallus 1 

Coprinopsis pseudocortinata 1 

Coprinopsis pseudonivea 

2 

var. pseudonivea 

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 

Coprinopsis utrifera 1 

Coprinopsis xenobia 4 

Total Coprinus s.l. 55 2 45 1 2 1 1 4 1 13 

Crucibulum laeve 1 

Cyathus stercoreus 1 2 

Lepista sordida 1 

Leucocoprinus cretaceus 4 

Panaeolus acuminatus 4 

339

Table 2 (Continued) Records from Italy of coprophilous Basidiomycota in the natural state. 



Panaeolus alcis 1 

Panaeolus antillarum 5 

Panaeolus cinctulus 2 3 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 11 3 5 

Psathyrella hirta 1 1 

Psathyrella lacrymabunda 1 

Psathyrella prona var. 

1 

prona f. prona 

Total Psathyrella 1 3 

Psilocybe coprophila 2 3 1 2 5 

Psilocybe crobulus 1 

Psilocybe cyanescens 1 

Psilocybe inquilina 1 

Psilocybe liniformans 1 

Psilocybe merdaria 2 

Psilocybe semilanceata 1 

Psilocybe subcoprophila 1 

Total Psilocybe 7 4 1 2 1 6 

Sebacina epigaea 1 

Sphaerobolus stellatus 1 1 

Stropharia dorsipora 1 6 1 

Stropharia luteonitens 1 

Stropharia semiglobata 9 3 2 

Total Stropharia 10 9 4 

Volvariella gloiocephala 1 

340


Table 3 Records from Italy and frequency (%) of coprophilous Basidiomycota on different dung types in damp chamber cultures (n° of dung samples 

in square brackets) * . 

Bolbitius 

coprophilus 

Coprinellus 

brevisetulosus 

Coprinopsis 

candidolanata 

Coprinopsis 

cinerea 

Coprinellus 

congregatus 

Coprinellus 

curtus 

“Coprinus” 

ephemeroides 

Coprinellus 

ephemerus 

Coprinopsis 

filamentifera 

Coprinellus 

heptemerus 

Coprinellus 

heterosetulosus 

Parasola 

misera 

Coprinopsis 

nivea 

“Coprinus” 

patouillardii 

Coprinellus 

pellucidus 

Coprinopsis 

poliomallus 

Coprinopsis 

pseudocortinata 

Coprinopsis 

pseudocortinata 

bird 

[27] 

3 

(11%) 

cattle 

[55] 

1 

(2%) 

1 

(2%) 

1 

(2%) 

1 

(2%) 

2 

(4%) 

1 

(2%) 

4 

(7%) 

2 

(4%) 

6 

(11%) 

1 

(2%) 

4 

(7%) 

2 

(4%) 

1 

(2%) 

1 

(2%) 

1 

(2%) 

donkey 

[6] 

1 

(17%) 

equine cervine caprine leporine ovine 

horse 

[63] 

1 

(2%) 

6 

(9%) 

2 

(3%) 

1 

(2%) 

1 

(2%) 

14 

(22%) 

5 

(8%) 

1 

(2%) 

14 

(22%) 

1 

(2%) 

1 

(2%) 

Deer 

[24] 

1 

(4%) 

1 

(4%) 

3 

(12%) 

1 

(4%) 

fallow 

deer 

[10] 

1 

(10%) 

3 

(30%) 

1 

(10%) 

roe 

deer 

[37] 

1 

(3%) 

1 

(3%) 

1 

(3%) 

2 

(5%) 

1 

(3%) 

goat 

[16] 

1 

(6%) 

2 

(12%) 

2 

(12%) 

7 

(44%) 

1 

(6%) 

chamois 

[6] 

1 

(17%) 

1 

(17%) 

rock 

goat 

[8] 

1 

(12%) 

1 

(12%) 

2 

(25%) 

2 

(25%) 

1 

(12%) 

hare 

[20] 

3 

(15%) 

2 

(10%) 

rabbit 

[19] 

3 

(16%) 

1 

(5%) 

1 

(5%) 

marmot 

[5] 

1 

(20%) 

marten 

[2] 

pig 

[5] 

1 

(20%) 

sheep 

[51] 

1 

(2%) 

2 

(4%) 

3 

(6%) 

9 

(18%) 

1 

(2%) 

5 

(10%) 

2 

(4%) 

Mouflon 

[1] 

tortoise 

[2] 

1 

341 

wild 

pig 

[10]

Table 3 (Continued) Records from Italy and frequency (%) of coprophilous Basidiomycota on different dung types in damp chamber cultures (n° of 

dung samples in square brackets) * . 

Coprinopsis 

pseudonivea 

var. 

pseudonivea 

Coprinopsis 

pseudoradiata 

Coprinopsis 

radiata 

Coprinopsis 

stercorea 

Coprinus 

sterquilinus 

Coprinopsis 

tuberosa 

Coprinopsis 

utrifera 

Coprinopsis 

vermiculifera 

Total ** 

Coprinus s.l. 

Panaeolus 

papilionaceus 

Psathyrella 

coprinoides 

Psilocybe 

coprophila 

Stropharia 

dorsipora 

Stropharia 

semiglobata 

Total ** 

Stropharia 

342 

bird 

[27] 

cattle 

[55] 

1 

(2%) 

donkey 

[6] 

equine cervine caprine leporine ovine 

horse 

[63] 

Deer 

[24] 

fallow 

deer 

[10] 

roe 

deer 

[37] 

goat 

[16] 

chamois 

[6] 

rock 

goat 

[8] 

1 1 

1 

(2%) (4%) 

(2%) 

5 4 17 2 

6 1 

(9%) (67%) 

(10%) 

(12%) 

8 

2 2 7 3 

9 

1 

(14%) 

(8%) (20%) (19%) (19%) 

(18%) 

(10%) 

1 

(2%) 

1 

1 1 1 

1 

(5%) 

1 

3 

(2%) 

(10%) (3%) 

(5%) 

(6%) 

(4%) 1 

(12%) 

11% 49% 83% 73% 25% 60% 30% 56% 33% 75% 20% 31% 20% 20% 61% 10% 

77% 32% 57% 26% 

1 

(2%) 

1 

(3%) 

1 

1 

(2%) 

1 

(2%) 

1 

(2%) 

2% 

2% 

*The frequency (%) is not reported when samples of a dung type are less than five. °Records from dung in the natural state, in normal type. Records from dung in damp chambers 

and frequency (%), in bold type. **The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of 

that genus and the number of dung samples [ ]. The total occurrence of a genus does not result from a simple addition of occurrences of all species belonging to it, because the genus 

as whole must be regarded as occurring only once in one sample even if two species, or more, occur in that sample. 

hare 

[20] 

rabbit 

[19] 

marmot 

[5] 

marten 

[2] 

pig 

[5] 

sheep 

[51] 

Mouflon 

[1] 

tortoise 

[2] 

wild 

pig 

[10]


Table 4 Records from Italy of coprophilous Delitschia on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

cervine caprine 

cattle horse deer fallow deer roe deer hare goat rock goat pig sheep 

[55] [63] [24] [10] [37] [20] [16] [8] [5] [51] 

D. didyma 1 

D. gigaspora 1 (12%) 

D. gigaspora var. ceciliae 2 (4%) 

D. intonsa 3 (5%) 2 (4%) 1 (3%) 1 

D. leptospora 1 1 (3%) 1 (5%) 

D. marchalii 1 (5%) 1 (6%) 1 (12%) 

D. patagonica 1 (2%) 

D. vulgaris 1 (2%) 

D. winteri 1+4 (7%) 1+1 (2%) 1 (10%) 1 1 (2%) 

Total Delitschia in natural state 1 1 1 1 2 

Total Delitschia in damp chambers** 9 

5 

1 2 2 

1 2 

1 

(10%) (3%) 

(6%) (8%) 

(14%) (6%) 

(3%) 

(10%) 

(12%) 

(2%) 

*The frequency (%) is not reported when samples of a dung type are less than five. °Records from dung in the natural state, in normal type. Records from dung in damp chambers 

and frequency (%), in bold type. **The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of 



343

Table 5 Records from Italy of coprophilous Trichodelitschia on different dung types (n° of dung 

samples in damp chamber cultures, in square brackets)°. 

cervine leporine 

deer roe deer hare rabbit pig sheep 

[24] [37] [20] [19] [5] [51] 

T. minuta 2+1 (4%) 1 (3%) 3+1 (5%) 2+1 (5%) 1 (20%) 2 (4%) 

T. munkii 1 (2%) 

Total Trichodelitschia in natural state 2 3 2 

Total 

Trichodelitschia in damp chambers** 

1 1 

(4%) (3%) 

1 1 

(5%) (5%) 

1 3 

(3%) 

(5%) 

(20%) (6%) 

°Records from dung in the natural state, in normal type. Records from dung in damp chambers and frequency (%), in 

bold type. **The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio 

between the total number of occurrences of that genus and the number of dung samples [ ]. The total occurrence of a 

genus does not result from a simple addition of occurrences of all species belonging to it, because the genus as whole 

must be regarded as occurring only once in one sample even if two species, or more, occur in that sample. 

Table 6 Records from Italy of coprophilous Pleospora on different dung types (n° of dung samples 

in damp chamber cultures, in square brackets)°. 

sheep 

[51] 

P. herbarum 1 

° Records from dung in the natural state, in normal type. Records from dung in damp chambers, in bold type. 

Table 7 Records from Italy of coprophilous Preussia on different dung types (n° of dung samples 


344 

cattle 

[55] 

donkey 

[6] 


horse 

[63] 

deer 

[24] 

fallow 

deer [10] 

roe 

deer 

[37] 

rabbit 

[19] 

P. fleischhakii 1 (2%) 1 2 (3%) 

P. funiculata 1 (3%) 1(2%) 1+1 

(10%) 

P. isomera 2+3 (5%) 

P. terricola 2 (4%) 

P. typharum 1 (2%) 2 (8%) 1 (10%) 1 (5%) 

Total Preussia 

in natural state 

1 2 1 

Total Preussia 4 

5 2 1 1 1 1 1 

in Damp 

(6%) (8%) (10%) (3%) 

chambers** (7%) (6%) 

(6%) 

(5%) (2%) (10%) 

° Records from dung in the natural state, in normal type. Record s from dung in damp chambers and frequency (%), in 

bold type. ** The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio 

between the total number of occurrences of that genus and the number of dung samples [ ]. The total occurrence of a 

genus does not result from a simple addition of occurrences of all species belonging to it, because the genus as whole 

must be regarded as occurring only once in one sample even if two species, or more, occur in that sample. 

Table 8 Records from Italy of coprophilous Sporormia on different dung types (n° of dung samples 


deer 

marmot 

[24] 

[5] 

S. fimetaria 2 (8%) 1 (20%) 

° Records from dung in the natural state, in normal type. Records from dung in damp chambers and frequency (%), in 

bold type. 

sheep 

[51] 

wild 

pig 

[10]


Table 9 Records from Italy of coprophilous Sporormiella on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

beechmarten 

[3] 

bird 

[27] 

cattle 

[55] 

donkey 

[6] 

equine cervine leporine caprine 

horse 

[63] 

deer 

[24] 

fallow 

deer 

[10] 

roe 

deer 

[37] 

fox 

[7] 

hare 

[20] 

S. affinis 1 

S. antarctica 1 

S. australis 1+1 

4+2 

6+3 1 3 1 1 

1 1+1 

1 

(4%) 

(8%) 

(8%) (14%) 

(6%) 

(2%) 

S. capybarae 1 1 

S. 

1 

1 

corynespora 

(3%) 

(5%) 

S. 

3 1+1 

1 

1 1 

cylindrospora 

(4%) 

(3%) 

S. 

1 

cymatomera 

(3%) 

S. 

1 

dodecamera 

(3%) 

S. dubia 1 1 

S. gigantea 1 

S. 

2 

1 1 1 1 1 

grandispora 

(4%) 

(4%) 

S. heptamera 1 

(3%) 

1 1 

S. hololasia 1 

(5%) 

S. intermedia 1+5 

2+2 4+1 1+2 4+4 2 3+5 3+5 4 3 1+1 1 1 

4+6 

1 

(9%) 

(3%) (4%) (20%) (11%) (28%) (25%) (26%) (25%) (50%) (12%) (8%) (20%) 

(12%) 

S. kansensis 1 

S. 

2 

1 1 

lageniformis 

(3%) 

(3%) (14%) 

S. lasiocarpa 2 

(10%) 

S. leporina 1 1+2 

1+2 

3 1 1 

2+1 

(2%) (8%) 

(5%) 

(17%) 

S. 

longisporopsis 

1(2%) 1 1(5%) 

S. 

2+2 

1+1 2+2 

1+1 

1 

1(20%) 2 

megalospora 

(4%) 

(2%) (8%) 

(3%) 

(5%) 

S. minima 1 7+9 2 2+6 

1+2 2+3 

2 2+2 2+4 

3 

1 

2+13 

1+1 

(16%) (33%) (9%) 

(20%) (8%) 

(10%) (10%) (25%) 

(37%) 

(20%) 

(25%) 

S. minimoides 1 1+1 

(5%) 

S. minipascua 1 

rabbit 

[19] 

goat 

[16] 

chamois 

[6] 

rock 

goat 

[8] 

hedgehog 

[12] 

marmot 

[5] 

pig 

[5] 

sheep 

[51] 

wild 

pig 

[10] 

unidentified 

animal 

[2] 

345

Table 9 (Continued) Records from Italy of coprophilous Sporormiella on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

346 

beech– 

marten 

[3] 

bird 

[27] 

cattle 

[55] 

donkey 

[6] 


horse 

[63] 

deer 

[24] 

fallow 

deer 

[10] 

roe 

deer 

[37] 

S. octomera 1 1 

S. octonalis 1 1 1 

fox 

[7] 

hare 

[20] 

1 

(20%) 

S. pascua 1 

(2%) 

1 

S. pilosa 1 

1 

(2%) 

(14%) 

S. pulchella 1 

S. teretispora 1 1 

S. trogospora 1 

S. vexans 1 

(3%) 

1 1 

Total 

Sporormiella 

natural state 

1 18 1 8 15 3 15 14 10 2 2 4 10 1 6 

Total 

1 1 19 2 14 9 4 20 5 7 14 9 4 4 1 2 2 20 

3 

Sporormiella 

(4%) (29%) (33%) (17%) (25%) (40%) (38%) (29%) (25%) (63%) (44%) (67%) (37%) (8%) (40%) (20%) (37%) 

in damp 

chambers** 

(19%) (31%) (44%) (47%) 

*The frequency (%) is not reported when samples of a dung type are less than five. 

°Records from dung in the natural state, in normal type. 

Records from dung in damp chambers and frequency (%), in bold type. 

**The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of that genus and the number of dung 

samples[ ]. The total occurrence of a genus does not result from a simple addition of occurrences of all species belonging to it, because the genus as whole must be regarded as 

occurring only once in one sample even if two species, or more, occur in that sample. 

rabbit 

[19] 

goat 

[16] 

chamois 

[6] 

rock 

goat 

[8] 

Hedgehog 

[12] 

marmot 

[5] 

pig 

[5] 

sheep 

[51] 

wild 

pig 

[10] 

unidentified 

animal 

[2]


Table 10 Records from Italy of coprophilous Gymnoascus on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

badger bat bird cattle horse roe deer dormouse hedgehog pig sheep 

[2] [1] [27] [55] [63] [37] 

[4] 

[12] 

[5] [51] 

G. dankaliensis 

1 1 

1 

2 

(2%) (2%) 

(20%) (4%) 

G. devroeyi 1 1 

1 1 

(3%) 

(8%) 

G. littoralis 1 

G. reessii 1 

(4%) 

G. ruber 1 

(2%) 

Total Gymnoascus in damp 

1 1 1 2 1 

1 

1 1 

1 

2 

chambers** 

(4%) (4%) (2%) (3%) 

(8%) (20%) (4%) 

*The frequency (%) is not reported when samples of a dung type are less than five. 

°Records from dung in the natural state, in normal type. Records from dung in damp chambers and frequency (%), in bold type. 

**The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of that genus and the number of dung 

samples [ ]. The total occurrence of a genus does not result from a simple addition of occurrences of all species belonging to it, because the genus as whole must be regarded as 


347

Eurotiomycetes – Eurotiomycetidae –Onygenales 

– Gymnoascaceae 

Gymnoascus Baran. (Table 10) 

Refer to the taxonomic part of this work 

for discussion about the facultative coprophily 

of Gymnoascaceae. In Italy, like elsewhere, 

species of Gymnoascus grow on a variety of 

dungs, where they are able to decompose substances 

(cellulose) unutilised by other fungi. 

Eurotiomycetes – Eurotiomycetidae – Onygenales 

– Onygenaceae 

Aphanoascus Zukal. (Table 11) 

Table 11 Records from Italy of coprophilous 

Aphanoascus on different dung types (n° of 

dung samples in damp chamber cultures, in 

square brackets)*°. 

348 

beechmarten 

[3] 

deer 

[24] 

cervine 

fallow 

deer 

[10] 

roe deer 

[37] 

goat 

[16] 

A. fulvescens 1 1 (4%) 1 (10%) 1 (3%) 2 (12%) 

(4%) 

*The frequency (%) is not reported when samples of a 

dung type are less than five. ° Records from dung in the 

natural state, in normal type. Records from dung in damp 

chambers and frequency (%), in bold type. 

Xanthothecium Arx & Samson (Table 12) 


Xanthothecium on different dung types (n° of 


square brackets)°. 

cattle sheep 

[55] [51] 

X. peruvianum 1 (2%) 1 (2%) 

° Records from dung in the natural state, in normal type. 

Records from dung in damp chambers and frequency 

(%), in bold type. 

Like the Gymnoascaceae, the Onygenaceae 

are facultatively coprophilous. Their keratinolytic 

capability allows them to inhabit a 

variety of dungs, particularly carnivore dung, 

which contains large amounts of keratinic remnants 

(hairs, nails, etc.). Xanthothecium is an 

atypical Onygenacea as it is not keratinophilic 

(Currah 1985), but Xanthothecium peruvianum 

has frequently been isolated from carnivore 

dung (Stolk 1955, Cain 1957). 

The keratinolytic Aphanoascus has often 

been isolated from carnivore dung and bird of 

prey pellets (Cain 1957, Currah 1985, Cano & 

Guarro 1990, Valldosera 1991, Valldosera & 

Guarro 1992, Doveri 2006). 

Leotiomycetes – Thelebolales – Thelebolaceae 

Ascozonus (Renny) E.C. Hansen. (Table 13) 


Ascozonus on different dung types (n° of dung 

samples in damp chamber cultures, in square 

brackets)*°. 

unidentified animal 

[2] 

A. woolhopensis 1 

* The frequency (%) is not reported when samples of a 

dung type are less than five. ° Records from dung in the 

natural state, in normal type. Records from dung in damp 

chambers, in bold type. 

The psychrophilic genus Ascozonus is 

absolutely coprophilous (Eckblad 1968), and 

most records worldwide come from leporine 

dung (75%). Also A. woolhopensis preferably 

develops on leporine dung (58% of records 

found by us) (Otani & Kanzawa 1970, Dennis 

1981, Derbsch & Schmitt 1987, Prokhorov, 

1989a, Valldosera & Guarro 1992, Schavey 

1999, Coste & Rey 2000, Richardson 2008b). 

Cleistothelebolus Malloch & Cain (Table 14) 


Cleistothelebolus on different dung types (n° of 



fox [7] 

C. nipigonensis 1 (14%) 




Mine is the second finding worldwide of 

C. nipigonensis. I have made a morphological, 

cultural and molecular study, and the work is in 

preparation.

Coprotus Korf & Kimbr. (Table 15) 

The absolutely coprophilous genus Coprotus 

has a wide substrate tolerance (Kim 

brough et al. 1972), but decidedly prefers cattle 

and horse dung in Italy, where C. disculus and 

C. sexdecimsporus are the most frequent. The 

former does not have a particular substrate 

preference, while the latter preferably grows on 

horse and cattle dung. There are numerous 

records worldwide confirming the wide substrate 

tolerance of C. disculus and C. sexedcimsporus, 

but the former has a slight 

preference for cattle dung (Barrasa 1985, 

Valldosera & Guarro 1985, Korf & Zhuang 

1991a, Korf & Dirig 2009), the latter a high 

preference (52% of records) for horse (Cooke 

1864, Fuckel 1866, Phillips 1893, Bednarczyk 

1974, Aas 1978, Engel & Hanff 1985, Wang 

1995, Ramos et al. 2008) and cattle (Crouan & 

Crouan 1858, Massee 1895, Prokhorov 1989a, 

Abdullah & Alutbi 1994, De Meulder 2000) 

dung. 

Thelebolus Tode (Table 16) 

The psychrophilic genus Thelebolus is 

widespread worldwide, and the commonest 

species, T. stercoreus s. str., has a wide substrate 

tolerance and no preference for particular 

types of dung (de Hoog et al. 2005). 

Refer to de Hoog et al. (2005) for the synonymy 

between T. stercoreus and other Thelebolus 

spp. 

Pezizomycetes – Pezizomycetidae – Pezizales 

– Ascobolaceae 

Ascobolus Pers (Table 17) 

In Italy the genus Ascobolus is one of the 

commonest on dung, with 45 (8%) records 

from the field and 174 (10%) from damp chambers. 

Its frequency of occurrence on total samples 

is quite high (8%) and the highest frequencies 

are on ovine, caprine and marmot dung. 

The cosmopolitan A. immersus is the 

most commonly recorded species from dung in 

Italy. There are 27 finding in the natural state 

and 98 in damp chambers (frequency of occurrence 

on total samples = 6%). Its preference for 

ovine dung is considerable (frequency = 61%). 

I have found 234 records worldwide, partly 

confirming my data, as bovine dung is the most 

mentioned (30% of all records) (Crouan & 


Crouan 1857, Coemans 1862, Denison 1963, 

Ahmed et al. 1971, Barrasa 1985, Derbsch & 

Schmitt 1987, Abdullah & Alutbi 1993, Wang 

1993, 1999, De Meulder 2000a, Delgado Avila 

et al. 2001a, Calonge & Menezes de Sequeira 

2003, García Alonso, 2007, Cheype 2008), 

followed by ovine (23%) (Berkeley & Broome 

1865, Guarro Artigas 1983, Engel & Hanff 

1984, Jahn 1997, Richardson 2005, 2008a, De 

Meulder 2007a) and equine (19%) (Green 

1931, Marasas et al. 1966, Dissing & Raitviir 

1973, Coste & Rey 1996, Van Vooren & 

Gaignou 2002). 

I have recorded the rare A. carletoni only 

once from tetraonid dung, like most records 

elsewhere (Boudier 1912, van Brummelen 

1967, Richardson 1972, 1998, 2007). A. 

michaudii is quite frequent on dung in Italy, 

particularly in damp chambers, where it has a 

slight preference for ovine (12%) and equine 

(10%) dung. Unlike mine, most records worldwide 

(46%) are from cattle dung (Persoon 

1796, Rattan & El-Buni 1980, Aguirre-Acosta 

& Ulloa 1982, Abdullah & Alutbi 1993, Wang 

1993, 1999, Richardson 1998). 

Saccobolus Boud (Table 18) 

In Italy the occurrence on dung of the 

genus Saccobolus, both in the natural and 

artificial state, is very similar to Ascobolus, but 

Saccobolus shows a high preference for equine 

and bovine dung, with a fairly good tolerance 

for other types. A lower tolerance is possessed 

by S. verrucisporus which has been recorded 

almost exclusively (91%) from cervine dung, 

with a 23% frequency on this type in culture. 

Records from deer dung (van Brummelen 

1967, Larsen 1970, Häffner 1986, Richardson 

2001b) are prevalent (28%) elsewhere, but not 

with the same frequency as in Italy. 

S. depauperatus is very common in Italy, 

where it absolutely prefers equine dung (41% 

of records from damp chambers). I have found 

numerous records worldwide proving its preference 

for equine dung (26%) (Seaver 1928, 

Guarro Artigas 1983, Ranalli & Mercuri 1995a, 

Jahn 1997, Spooner & Butterfill 1999), but also 

for ovine (24%) (Raitviir & Prokhorov 1988, 

Valldosera 1991, Richardson 2004b, 2005, De 

Meulder 2007a), and a wide tolerance for other 

types of dung. 

349

Table 15 Records from Italy of coprophilous Coprotus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 


cattle horse deer fallow roe deer fox rabbit goat rock sheep wild pig unidentified 

[55] [63] [24] deer [10] [37] [7] [19] [16] goat [8] [51] [10] animal [2] 

C. aurora 2 1 

C. disculus 1+1 

1+1 

1 

1+1 1 1 

1 

(2%) 

(4%) 

(3%) 

(5%) (6%) (12%) 

(10%) 

C. glaucellus 1 2+1 

(3%) 

C. granuliformis 2+4 

1+1 

(7%) 

(2%) 

C. lacteus 1 1 

1 

(3%) 

(2%) 

C. leucopocillum 1 (2%) 1+1 

(2%) 

C. aff. luteus 2 1 

(4%) (2%) 

C. niveus 1 (2%) 

C. aff. ochraceus 1 

C. sexdecimsporus 1+10 15 

1 4 1 1 

1 4 1 

(18%) (24%) 

(10%) (11%) (14%) (5%) 

(12%) (8%) (10%) 

C. subcylindrosporus 1 1 1 (3%) 

Total Coprotus in 6 2 2 2 2 1 

natural state 

Total Coprotus in damp 14 17 1 1 8 1 2 1 2 7 ( 2 

chambers** 

(4%) (10%) (22%) (14%) (5%) (6%) (12%) 14%) 

(25%) (25%) 

(14%) (8%) 

(20%) 

* The frequency (%) is not reported when samples of a dung type are less than five. 

° Records from dung in the natural state, in normal type. Records from dung in damp chambers and frequency (%), in bold type. 

** The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of that genus and the number of dung 



350


Table 16 Records from Italy of coprophilous Thelebolus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

bird 

[27] 

cattle 

[55] 

horse 

[63] 

deer 

[24] 

cervine leporine caprine 

roe deer 

[37] 

T. caninus 1 

(3%) 

T. crustaceus 1 1 

hare 

[20] 

rabbit 

[19] 

goat 

[16] 

rock goat 

[8] 

porcupine 

[1] 

sheep 

[51] 

wild 

pig 

[10] 

unidentified 

animal 

[2] 

1 

T.dubius var. lagopi 

1 

1 

(5%) 

1 1 

(2%) 

(3%) 

(5%) 

T. microsporus 1 1+2 

1 

2 

2 

(4%) (4%) 

(3%) 

(25%) 

(4%) 

T. polysporus 1 1 1+1 

1 1+1 

1+3 1 1 

(3%) 

(6%) 

(6%) (10%) 

T. stercoreus 1 1 1 1 (4%) 4+3 2 1+1 

2 

1 

(4%) (2%) (2%) 

(8%) (10%) (5%) 

(4%) 

Total 

Thelebolus in natural state 

3 1 5 2 1 1 1 

Total 

2 4 1 1 7 3 2 1 2 1 7 1 1 

Thelebolus in damp 

(7%) (5%) (2%) (4%) (19%) (15%) (5%) (6%) (25%) 

(14%) (10%) 

chambers** 

(13%) (10%) (12%) 






351

Table 17 Records from Italy of coprophilous Ascobolus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

352 

bird 

[27] 

cattle 

[55] 

A. albidus 1 

(2%) 

donkey 

[6] 

equine cervine leporine caprine ovine 

horse 

[63] 

1+9 

(14%) 

deer 

[24] 

fallow 

deer 

[10] 

roe 

deer 

[37] 

duck 

[1] 

fox 

[7] 

goose 

[1] 

hare 

[20] 

rabbit 

[19] 

goat 

[16] 

1 2 

(12%) 

A. brassicae 1 1 

A. carletoni 

1 

(4%) 

A. costantinii 1 

A. crenulatus 1 

(5%) 

2 

(10%) 

1 

(8%) 

1 1 1 

(10%) 

1+1 

A.elegans 5 

(8%) 

A. furfuraceus 5+5 

1+3 1+1 

2 

1 

2 

1 1 

1 

(9%) 

(5%) (4%) 

(5%) 

(6%) 

(25%) 

(2%) 

(10%) 

A. hawaiiensis 1 3 

A. immersus 7+16 

(29%) 

3 

(50%) 

8+13 

(21%) 

1+1 

(4%) 

2 

(20%) 

2+8 

(22%) 

2 

(28%) 

4 

(20%) 

1+2 

(10%) 

1+5 

(31%) 

1 

(17%) 

5 

(62%) 

3 

(60%) 

7+30 

(59%) 

1 1 

(10%) 

1 

A. lineolatus 1 

(14%) 

A. mancus 1 1 1+2 

1 

(2%) (17%) (3%) 

(6%) 

A. michaudii 2 1 1+6 1 2 

1 1 

1 6 1 

(4%) (17%) (9%) (4%) (20%) 

(5%) (6%) 

(12%) 

A. reticulatus 1 

A. 

roseopurpurascens 

1 

A. stictoideus 1 1+1 

(2%) 

1 1 

Total Ascobolus 


13 13 3 2 1 1 1 1 7 1 2 

Total Ascobolus** 

in damp chambers 

1 

(4%) 

25 

(34%) 

5 

(67%) 

40 

(48%) 

3 

(8%) 

4 

(25%) 

10 

(24%) 

3 

(43%) 

1 5 

(25%) 

5 

(21%) 

10 

(62%) 

1 

(17%) 

7 

(87%) 

1 

(8%) 

3 

(60%) 

1 3 37 

(63%) 

2 

(63%) 

1 3 

(20%) 

1 2 

(49%) (21%) (23%) (60%) 






chamois 

[6] 

rock 

goat 

[8] 

hedgehog 

[12] 

marmot 

[5] 

marten 

[2] 

ostrich 

[1] 

rat 

[1] 

sheep 

[51] 

mouflon 

[1] 

weasel 

[1] 

wild 

pig 

[10] 

1 

wolf 

[1] 

unidentified 

animal 

[2]


Table 18 Records from Italy of coprophilous Saccobolus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

cattle 

[55] 

donkey 

[6] 


horse 

[63] 

deer 

[24] 

fallow 

deer 

[10] 

1 

(10%) 

roe 

deer 

[37] 

2+2 

(5%) 

hare 

[20] 

rabbit 

[19] 

goat [16] chamois 

[6] 

S. beckii 1 3+4 

2 

(17%) 

(33%) 

S. caesariatus 3 (5%) 1+2 

(10%) 

1 (6%) 1 (12%) 2 (4%) 

S. citrinus 2 (4%) 1 (2%) 3 

(12%) 

1 2 (5%) 2 (4%) 

S. depauperatus 3 (5%) 3 (50%) 1+25 1 (4%) 1 2+1 1+1 

1 (6%) 1 

6 

(40%) 

(10%) (3%) (5%) 

(20%) 

(12%) 

S. dilutellus 1 

S. glaber 1+5 

1 1+1 

1 

(9%) 

(20%) (2%) 

S. minimus 1 (2%) 2 1+7 1 (4%) 2 1 (3%) 2 (12%) 1 (12%) 4 (8%) 1 

(11%) 

(20%) 

(10%) 

S. saccoboloides 1 (2%) 1 (2%) 1 (3%) 1 (2%) 

S. succineus 7 

(13%) 

2 (3%) 

S. truncatus 3 (5%) 7 

(11%) 

1 (5%) 1 (6%) 3 (6%) 

S. verrucisporus 1 (2%) 3+5 

1+11 

1 (2%) 

(21%) 

(30%) 

S. versicolor 3+9 1 (17%) 2 (3%) 1 2+3 1 (6%) 3 (37%) 1 (2%) 1 

(16%) 

(5%) (16%) 

Total 

Saccobolus 

(natural state) 

5 2 2 6 1 5 1 4 1 1 

Total 

34 4 46 14 4 8 2 6 6 2 5 1 1 21 1 1 

Saccobolus** in (54%) (67%) (67%) (46%) (30%) (43%) (10%) (31%) (31%) (33%) (50%) (20%) (20%) 

(10%) 

damp chambers 

(67%) (42%) (20%) (37%) 

(33%) 

rock 

goat [8] 

marmot 

[5] 

pig 

[5] 

sheep 

[51] 

wild 

pig 

[10] 

wolf 

[1] 

unidentified 

animal 

[2] 






353

Table 19 Records from Italy of coprophilous Thecotheus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

cattle corse 

[55] [63] 

T. cinereus 1 

T. crustaceus 2 

354 

(3%) 

T. formosanus 1 

T. formosanus f. collariatus 1 

T. holmskjoldii 1+10 

(18%) 

T. lundqvistii 6 

(11%) 

T. neoapiculatus 2 

T. pelletieri 1+1 

(2%) 

deer 

[24] 


fallow deer 

[10] 

1 2 

(20%) 

1 2 

(20%) 

roe deer 

[37] 

2+2 

(5%) 

2+1 

(3%) 

4 1 

1 

(6%) (4%) 

T. strangulatus 1 

Total Thecotheus in natural state 5 2 2 4 3 

Total Thecotheus ** 

17 6 1 

4 

3 

2 

4 

1 

2 

1 

10 


(4%) (40%) (8%) 

(25%) (17%) (25%) (20%) (20%) 

(31%) (9%) 

(11%) (10%) 

(23%) 






rabbit 

[19] 

1 

(5%) 

1 

(5%) 

goat 

[16] 

3 

(19%) 

1 

(6%) 

chamois 

[6] 

1 

(17%) 

rock goat 

[8] 

2 

(25%) 

pig 

[5] 

1 

(20%) 

sheep 

[51] 

1+7 

(14%) 

3 

(6%)

Also S. minimus is quite frequent in Italy, 

where it has not a particular substrate preference. 

It grows on a variety of dung also elsewhere, 

preferably (35% of records) on sheep 

dung (Rattan & El-Buni 1980, Guarro Artigas 

1983, Valldosera 1991, Richardson 2008c). 

Thecotheus Boud (Table 19) 

The genus Thecotheus is quite frequent in 

Italy, where it has a wide substrate tolerance 

but preferably grows on bovine and caprine 

dung. T. holmskjoldii is the most frequent (46% 

of all records) and has a slight preference for 

bovine dung. 

I refer to Aas' (1992) world monograph 

for the most detailed information about the 

ecology and distribution of Thecotheus. In that 

work, the records of T. holmskjoldii from 

bovine dung are 54% out of a total of 145. 


– Ascodesmidaceae 

Ascodesmis Tiegh (Table 20) 

Brummelen (1981) monographed Ascodesmis 

and recorded about 50 collections 

worldwide, more than 50% from carnivore 

dung. Several subsequent records have been 

found by me, but a minority (20%) from 

carnivore dung (Currah 1986, Derbsch & 

Schmitt 1987, Patil & Ghadge 1987, 

Kristiansen 1994, Jahn 1997). 

Lasiobolus Sacc (Table 21) 

Lasiobolus was monographed by Bezerra 

& Kimbrough (1975), who studied numerous 

collections worldwide but did not specify their 

dung sources. 

The genus Lasiobolus is quite frequent in 

Italy, both in the natural and artificial state, and 

it has a wide substrate tolerance, with a 

preference for cervine dung in the field (38% 

of records), for caprine in damp chambers, 

followed by ovine and wild pig dung. 

L. cuniculi is the most frequent (68% of 

records from Italy) and prefers caprine dung 

(30% of frequency of occurrence), particularly 

rock goat dung (62%). Also elsewhere its 

frequeny of occurrence on caprine dung is high 

(25% of records found by us) (Moyne 2006, De 

Meulder 2007b, Richardson 2004a, 2008a), and 


equal to ovine (Barrasa 1985, Korf & Zhuang 

1991b, Richardson 2004b). 

Trichobolus (Sacc.) Kimbr. & Cain (Table 22) 

Trichobolus zukalii is one of the commonest 

discomycetes and the commonest 

Trichobolus sp. (96% of all records) in Italy. It 

has a fair substrate tolerance but a preference 

for caprine and cervine dung, particularly for 

goat. T. zukalii is less frequent elsewhere: I 

have found 30 records only, similar to mine, 

i.e. 43% from caprine, particularly goat dung 

(Korf & Zhuang 1991b, Valldosera 1991, 

Moyne 2006, Richardson 2008a,b), 33% from 

cervine dung (Heimerl 1889, Kimbrough 1966, 

Richardson 1972, 1998, Bronckers 2002, 

Moyne & Petit 2006). 


– Pezizaceae 

Iodophanus Korf (Table 23) 

Iodophanus carneus is a very common 

discomycete in Italy, second only to Ascobolus 

immersus. It has been recorded especially from 

bovine dung in the natural state, and from 

many types of dung in damp chambers, with a 

marked preference for bovine, caprine, and 

ovine dung (61% of records from these types). 

Kimbrough et al. (1969) monographed 

Iodophanus and described I. carneus as one of 

the commonest coprophilous discomycetes, 

particularly recording it from bovine dung 

(57% of records worldwide). There are more 

than 200 additional records worldwide, 

partularly from bovine (24%) (Karsten 1861, 

Coemans 1862, Spegazzini 1878, Heimerl 

1889, Denison 1963, Bednarczyk 1974, Thind 

& Kaushal 1978, Mouso & Ranalli 1989, 

Prokhorov 1989b, Wang, 1999, De Meulder 

2000a, Wang & Wang 2000, Cinto et al. 2007, 

Richardson 2008c) and ovine dung (23%) 

(Ahmed et al. 1971, Guarro Artigas 1983, 

Barrasa 1985, Abdullah & Alutbi 1993, 

Richardson 2004b, 2005, 2008b,c). 

Peziza Fr. (Table 24) 

Collections of coprophilous Peziza spp. 

are very scarce in Italy, and the overwhelming 

majority of records are from the field. Only P. 

fimeti has been recorded once from a damp 

355

Table 20 Records from Italy of coprophilous Ascodesmis on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

badger beech- bird rabbit hedgehog insect ostrich pig rat sheep toad 

[2] marten[3] [27] 

[19] 

[12] 

[9] 

[1] 

[5] [1] [51] [2] 

A. microscopica 1 

1 

1 

(8%) 

(11%) 

(2%) 

A. nana 1 

(4%) 

A. nigricans 2 1 2 

1 

1 1 1 1 

(7%) (5%) 

(20%) 

Total Ascodesmis ** 

3 

1 

1 

1 

1 

1 


(11%) (5%) 

(8%) 

(11%) 

(20%) 

(2%) 

* The frequency (%) is not reported when samples of a dung type are less than five. ° Records from dung in the natural state, in normal type. 

°Records from dung in damp chambers and frequency (%), in bold type. ** The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio 

between the total number of occurrences of that genus and the number of dung samples [ ]. The total occurrence of a genus does not result from a simple addition of occurrences of 

all species belonging to it, because the genus as whole must be regarded as occurring only once in one sample even if two species, or more, occur in that sample. 

Table 21 Records from Italy of coprophilous Lasiobolus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 


cattle donkey horse deer fallow roe deer fox hare rabbit goat chamois rock marmot marten pig porcupine rat sheep wild unidentified 

[55] [6] [63] [24] deer [10] [37] [7] [20] [19] [16] [6] goat [8] [5] [2] [5] [1] [1] [51] pig[10] animalm[2] 

L. ciliatus 4 7+2 

(3%) 

1 1 1 1 (12%) 1 

L. cuniculi 5+3 

2+7 4 2+2 3+5 

1 

1+4 

5 (62%) 1 

1 6+12 2 (20%) 2 

(5%) 

(11%) 

(20%) (13%) 

(5%) 

(25%) 

(20%) 

(23%) 

L. diversisporus 2 1 (2%) 1 (12%) 

L. intermedius 1 

1 (3%) 1+1 

(17%) 

(2%) 

L. macrotrichus 1 

(4%) 

1 (3%) 1 1 1 

L. microsporus 1 (2%) 1 (6%) 1 

(20%) 

1 1 (10%) 

L. monascus 1 

(14%) 

L. ruber 1 3 3 1 

(17%) 

Total Lasiobolus in 

natural state 

12 4 9 2 7 1 2 8 2 

Total Lasiobolus in 3 1 11 1 2 7 1 1 (5%) 5 1 7 1 1 1 1 1 13 3 

damp chambers ** (5%) (17%) (17%) (4%) (20%) (19%) (14%) (3%) (31%) (17%) (87%) (20%) 

(20%) 

(25%) (30%) 

(17%) (14%) (43%) 


° Records from dung in the natural state, in normal type. °Records from dung in damp chambers and frequency (%), in bold type. ** The frequency (%) of a genus as whole, on each 

dung type in damp chamber, results from the ratio between the total number of occurrences of that genus and the number of dung samples [ ] The total occurrence of a genus does not 

result from a simple addition of occurrences of all species belonging to it, because the genus as whole must be regarded as occurring only once in one sample even if two species, or 

more, occur in that sample. 

356


Table 22 Records from Italy of coprophilous Trichobolus on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 


cattle deer fallow roe deer hare rabbit goat chamois rock goat [8] marmot marten sheep wild pig unidentified 

[55] [24] deer [10] [37] [20] [19] [16] [6] 

[5] 

[2] [51] [10] animal [2] 

T. octosporus 1 (5%) 

T. sphaerosporus 1 

T. zukalii 1+2 1+2 1+3 (30%) 4+10 (27%) 1 (5%) 1 1+9 1 

1 1 

1 3+8 1 (10%) 1 

(4%) (8%) 

(56%) (17%) 

(20%) 

(16%) 

Total Trichobolus in 

natural state 

1 1 1 5 1 1 1 3 1 

Total Trichobolus in 2 2 3 10 1 1 9 1 1 1 8 (16%) 1 (10%) 

damp chambers ** (4%) (8%) (30%) (27%) (5%) (5%) (56%) (17%) (20%) 

(21%) (5%) (33%) 


° Records from dung in damp chambers and frequency (%), in bold type. ** The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio 

between the total number of occurrences of that genus and the number of dung samples [ ] The total occurrence of a genus does not result from a simple addition of occurrences of all 

species belonging to it, because the genus as whole must be regarded as occurring only once in one sample even if two species, or more, occur in that sample. 

Table 23 Records from Italy of coprophilous Iodophanus on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

bird 

[27] 

cattle 

[55] 

horse 

[63] 

deer 

[24] 


fallow 

deer 

[10] 

1 

(10%) 

roe 

deer 

[37] 

8 

(22%) 

goose 

[1] 

hare 

[20] 

rabbit 

[19] 

goat 

[16] 

I. carneus 3 8+24 1+8 1+2 

1 2 2 5 2 

1 1 1 1 1+17 1 2 1 

(11%) (44%) (13%) (8%) 

(10%) (10%) (31%) (33%) 

(11%) (20%) 

(33%) 

I. difformis 1(2%) 

Total 

Iodophanus 

in natural 

state 

8 1 1 1 1 

Total 

3 25 8 2 1 8 1 2 2 5 2 5 1 1 1 1 17 1 2 

Iodophanus (11%) (45%) (13%) (8%) (10%) (22%) 

(10%) (10%) (31%) (33%) (62%) (11%) (20%) 

in damp 

chambers** 

(15%) (10%) (40%) 

(33%) 

chamois 

[6] 

rock 

goat 

[8] 

5 

(62%) 

insect 

[9] 

marmot 

[5] 

mouse 

[2] 

rat 

[1] 

sheep 

[51] 

snail 

[1] 

tortoise 

[2] 

unidentified 

animal [2] 






357

Table 24 Records from Italy of coprophilous Peziza on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 


donkey 

horse 

deer fallow deer human sheep [51] unidentifiedanimal 

[6] 

[63] 

[24] 

[10] 

[0] 

[2] 

P. fimeti 3+1 (2%) 1 1 (10%) 1 

P. merdae 1 1 

P. perdicina 1 

P. vesiculosa 1 6 

Total Peziza in natural state 1 9 1 1 1 2 

Total Peziza in damp chambers 1 (2%) 

1 (10%) 

(1%) 

(3%) 






Table 25 Records from Italy of coprophilous Chalazion on different dung types (n° of dung samples in damp chamber cultures, in square brackets)°. 

Hedgehog 

[12] 

C. erinaceum 1 (8%) 

° Records from dung in damp chambers and frequency (%), in bold type. 

358

chamber. P. vesiculosa is the most frequent in 

the natural state, and absolutely prefers equine 

dung (67% of collections). It is a cosmopolitan 

species, which preferably grows (56% of 

records found by us) on equine dung or manure 

(Dodge 1914, Jülich 1968, Donadini 1979, 

Barrasa 1985, Derbsch & Schmitt 1987, 

Arroyo et al. 1990, Calonge et al. 1991, Siquier 

& Lillo 1994, Hansen et al. 2000). 


– Pyronemataceae 

Chalazion Dissing & Sivertsen (Table 25) 

Refer to Doveri et al. (1998b) for the 

original diagnosis of C. erinaceum, which is 

known so far only from the type collection. The 

other two Chalazion spp., C. sociabile Dissing 

& Sivertsen and C. helveticum Dissing, are 

terricolous (Dissing & Sivertsen 1975, Dissing 

1980, Kristiansen 1990). 

Cheilymenia Boud. (Table 26) 

The cosmopolitan genus Cheilymenia 

does not develop in damp chambers in Italy, so 

all records (40) are from the field, mostly from 

cattle (72%) and horse (17%) dung. C. granulata, 

C. stercorea, and C. theleboloides are the 

most frequent, all with preference for cattle 

dung. 

The genus was monographed by Moravec 

(2005), who confirmed "the impossibility, with 

only few exceptions, to obtain fresh apothecia 

by an incubation of dung in moist chambers". 

He studied numerous world collections of C. 

granulata (87% of records from cattle dung), 

C. stercorea (66% from cattle dung), and C. 

theleboloides, the latter with a wide range of 

substrate tolerance, but with preference for 

cattle dung (50% of fimicolous records). 

Orbicula Cooke (Table 27) 

See the taxonomic part of this work for 

discussion about Orbicula. 

Pseudombrophila Boud. (Table 28) 

Pseudombrophila is the only genus of 

Pyronemataceae to easily develop also in damp 

chamber cultures (50% of my records). It does 

not grow frequently in Italy, so it is too early to 

estimate its occurrence on different types of 

dung. The genus was monographed by van 

Brummelen (1995) who regarded it as predo- 


minantly fimicolous and, like us, recorded P. 

cervaria particularly (55%) from deer dung. 

Scutellinia (Cooke) Lambotte (Table 29) 

Scutellinia is exceptionally coprophilous 

and S. crinita has rarely been found on excrements 

(Schumacher 1990). 

Trichophaea Boud. (Table 30) 

Trichophaea spp. usually grow on bare or 

burnt soils, humus, and decaying woody debris, 

while they have rarely been recorded from 

dung (Valldosera & Guarro 1990a, Häffner & 

Christan 1991, Jamoni 1998). As far as I know, 

no collection of T. gregaria, except for mine, is 

known from dung. 

Sordariomycetes – Sordariomycetidae – 

Coniochaetales – Coniochaetaceae 

Coniochaeta (Sacc.) Cooke (Table 31) 

Coprophilous collections of Coniochaeta 

are quite infrequent in Italy (less than 1% of 

records from damp chambers), and C. leucoplaca 

is the commonest species on a variety of 

dungs, while it has been recorded elsewhere 

particularly from bovine (26% of records found 

by us) (Griffiths 1901, Khan & Krug 1994, 

Richardson 2008b) and leporine dung (23%) 

(Cain 1957, Tóth 1967, Muroi & Udagawa 

1984, Valldosera & Guarro 1984b, Valldosera 

1991, Moyne & Petit 2006). 

Sordariomycetes – Hypocreomycetidae – 

Hypocreales – Bionectriaceae 

Selinia P. Karst. (Table 32) 

The four recognised Selinia spp. have 

always been recorded from dung of various 

herbivores, and S. pulchra particularly from 

bovine (34% of records found by me worldwide) 

(Wang 1994, Ranalli & Mercuri 1995b, 

Rossman et al. 1999) and ovine (34%) dung 

(Saccardo 1878, Richardson 2001b, 2004b, 

Welt & Heine 2006a). 


Hypocreales – Nectriaceae 

Neocosmospora E.F. Sm. (Table 33) 

Neocosmospora spp. are usually isolated from 

soil or plants. N. vasinfecta var. vasinfecta is 

pathogenic on crops (Rossman et al. 

359

Table 26 Records from Italy of coprophilous Cheilymenia on different dung types (n° of dung 


cattle horse dog human unidentified animal 

[55] 

[63] [0] [0] 

[2] 

C. aurantiacorubra 2 1 1 

C. coprinaria 2 1 

C. dennisii 1 1 

C. fraudans 1 

C. granulata 6 

C. insignis 3 

C. pulcherrima 2 

C. rubra 2 1 

C. stercorea 6 1 

C. theleboloides 5 2 1 1 

Total Cheilymenia in natural state 29 7 1 1 2 


Table 27 Records from Italy of coprophilous Orbicula on different dung types (n° of dung samples 


hare 

[20] 

O. parietina 1 (5%) 


bold type. 

Table 28 Records from Italy of coprophilous Pseudombrophila on different dung types (n° of dung 

samples in damp chamber cultures, in square brackets)*°. 

cattle horse deer 

cervine 

roe deer rabbit goat marten sheep unidentified 

[55] [63] [24] [37] [19] [16] [2] [51] animal [2] 

P. bulbifera 1 

P. cervaria 1 

(4%) 

2 1 1 

P. fuscolilacina 1 

P. merdaria 3 1 (5%) 1 

P. minuta 1 (2%) 1 (6%) 1 (2%) 

P. theioleuca 2 (5%) 1 (6%) 

Total 



3 2 1 2 1 

Total 

1 (2%) 1 2 1 (5%) 2 (12%) 1 1 (2%) 


(4%) (5%) 

in damp 

chambers** 

(5%) 



bold type. 

** The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total 

number of occurrences of that genus and the number of dung samples [ ]. The total occurrence of a genus does not result 

from a simple addition of occurrences of all species belonging to it, because the genus as whole must be regarded as 


Table 29 Records from Italy of coprophilous Scutellinia on different dung types (n° of dung 



[2] 

S. crinita 1 


360


Table 30 Records from Italy of coprophilous Trichophaea on different dung types (n° of dung 



[2] 

T. gregaria 1 


Table 31 Records from Italy of coprophilous Coniochaeta on different dung types (n° of dung 


cattle horse deer 

cervine 

roe deer rabbit chamois rock goat pig 

[55] [63] [24] [37] [19] [6] 

[8] [5] 

C. leucoplaca 2+3 1 1 

1 

2 

1 

(5%) (2%) (4%) (3%) (10%) (17%) 

C. scatigena 1 

1 

(5%) 

(20%) 

C. vagans 1+2 1+1 1 1 1 

(4%) (2%) 

(12%) 

Total 

Coniochaeta 


3 1 1 1 

Total 

5 

2 1 

1 

3 

1 

1 1 

Coniochaeta in (9%) (3%) (4%) (3%) (15%) (17%) (12%) (20%) 

damp chambers** 

(3%) 


bold type. 


number of occurrences of that genus and the number of dung samples. [ ] The total occurrence of a genus does not result 



Table 32 Records from Italy of coprophilous Selinia on different dung types (n° of dung samples in 

damp chamber cultures, in square brackets)°. 

cattle 

sheep 

[55] 

[51] 

S. pulchra 1 (2%) 1 (2%) 


bold type. 

Table 33 Records from Italy of coprophilous Neocosmospora on different dung types (n° of dung 


rabbit 

[19] 

N. vasinfecta var. vasinfecta 1 (5%) 


bold type. 

Table 34 Records from Italy of coprophilous Rodentomyces on different dung types (n° of dung 


dormouse 

[4] 

R. reticulatus 1 


361

1999), and it has exceptionally isolated from 

dung (Cailleux 1971a). 

Rodentomyces Doveri, Pecchia, Sarrocco & 

Vannacci (Table 34) 

Refer to Doveri et al. (2010a) for the 

protologue of Rodentomyces gen. nov., R. 

reticulatus sp. nov. 


Melanosporales – Ceratostomataceae 

Melanospora Corda (Table 35) 

The genus Melanospora is usually 

isolated from decaying vegetables and soil, or 

detected in association with other fungi, also on 

dung, possibly behaving as a mycoparasite 

(Cannon & Hawksworth 1982). Coprophilous 

collections of Melanospora are unusual (Faurel 

& Schotter 1965, Calviello 1976, Lorenzo 

1990, Richardson 2004a, 2008b), and I have 

rarely found records of M. brevirostris (Hansen 

et al. 1998, Delgado Avila et al. 2001b, Moyne 

et Petit 2006) and M. zamiae (Seth 1968, 

Lorenzo 1992, Valldosera & Guarro 1992, 

Piontelli et al. 2006) directly from dung. 

Persiciospora P.F. Cannon & D. Hawksw. 

(Table 36) 

Mine and Eriksson's (2009) are the only 

three collections of Persiciospora known from 

dung, where it possibly behaves as a 

mycoparasite (Cannon & Hawksworth 1982). 

“Sordaria” minima (Table 37) 

“Sordaria” minima is possibly a 

Melanospora sp. (Lundqvist pers. comm.), but 

it has not been recombined yet in this genus. 

Like some Melanospora spp., this rare species 

has been found on dung in association with 

other fungi, particularly with Thelebolus (Lund 

qvist pers. comm.). Besides mine, a few other 

collections of S. minima have been recorded 

from various kinds of dung (Massee & Salmon 

1901, Larsen 1971, Richardson 1998, 2004b, 

Moyne & Petit 2006, Welt & Heine 2007). 


Microascales – Microascaceae 

Enterocarpus Locq.–Lin. (Table 38) 

362 

See the taxonomic part of this work, 

under Lophotrichus bartlettii, for discussion on 

Enterocarpus grenotii. 

Kernia Nieuwl. (Table 39) 

K. nitida is the commonest species of this 

genus, found by me in Italy on a variety of 

dungs with low frequency of occurrence in 

damp chambers (1%). Like other Kernia spp. 

and many Microascaceae with a cellulytic and 

proteolytic activity (Lumley et al. 2000), it has 

been found on a variety of substrates and 

dungs, particularly (24% of fimicolous records 

worldwide) on sheep dung (Tóth 1965, 1967, 

Udagawa 1980, Guarro Artigas 1983, 

Valldosera 1991, Richardson 2004a). 

Lophotrichus R.K. Benj. (Table 40) 


under Lophotrichus bartlettii, for discussion on 

Lophotrichus. 

Pithoascus Arx (Table 41) 


under Pithoascus intermedius, for discussion 

on Microascaceae, Pithoascus, and Microascus. 

Sphaeronaemella P. Karst. (Table 42) 

Out of five recognised species of 

Sphaeronaemella (Kirk et al. 2008), only one, 

S. fimicola, has been isolated from dung, where 

it can behave as a facultative mycoparasite 

(Cain & Weresub 1957, Weber & Webster 

1997, 1998). S. fimicola has a wide substrate 

tolerance, carnivore dung included (Marchal 

1891, Pease 1948, Cain 1957), but it prefers 

leporine (32% of several records worldwide) 

(Massee & Salmon 1902, Dennis 1981) and 

cervine (26%) (Cannon & Hawksworth 1982, 

Lundqvist 1989) dung. 


Sordariales – Chaetomiaceae 

Chaetomidium (Zopf) Sacc. (Table 43) 


under Chaetomidium fimeti, for discussion on 

the genus Chaetomidium, C. megasporum, and 

C. fimeti.


Table 35 Records from Italy of coprophilous Melanospora on different dung types (n° of dung 


badger dormouse sheep 

[2] 

[4] 

[51] 

M. brevirostris 1 

M. zamiae 1 1 (2%) 

Total Melanospora in damp chambers** 1 1 1 (2%) 

* The frequency (%) is not reported when samples of a dung type are less than five. ° Records from dung in the natural 

state, in normal type. Records from dung in damp chambers and frequency (%), in bold type. 

Table 36 Records from Italy of coprophilous Persiciospora on different dung types (n° of dung 


fox 

polecat 

[7] 

[5] 

P. moreaui 1 (14%) 1 (20%) 


bold type. 

Table 37 Records from Italy of “Sordaria” minima on different dung types (n° of dung samples in 

damp chamber cultures, in square brackets)°. 

roe deer 

rabbit 

[37] 

[19] 

S. minima 1 (3%) 1 (5%) 


bold type. 

Table 38 Records from Italy of coprophilous Enterocarpus on different dung types (n° of dung 


horse 

[63] 

E. grenotii 1 (2%) 


bold type. 

Table 39 Records from Italy of coprophilous Kernia on different dung types (n° of dung samples in 

damp chamber cultures, in square brackets)*°. 

cattle 

[55] 

horse 

[63] 

goat 

[16] 

ostrich 

[1] 

pig 

[5] 

sheep 

[51] 

unidentified 

animal 

[2] 

1 

K. cauquensis 1 

(2%) 

K. nitida 5 4 1 1 1 2+4 

(9%) (6%) (6%) 

(20%) (8%) 

Total Kernia in natural state 2 

Total Kernia in damp 

5 4 1 1 1 5 

1 

chambers** 

(9%) (6%) (6%) 

(20%) (10%) 



bold type. 


number of occurrences of that genus and the number of dung samples [ ]. The total occurrence of a genus does not result 



363


Lophotrichus on different dung types (n° of 


square brackets)*°. 

sheep 

[51] 

L. bartlettii 1 (2%) 





Pithoascus on different dung types (n° of dung 

samples in damp chamber cultures, in square 

brackets)*°. 

pig 

[5] 

P. intermedius 1 (20%) 





Sphaeronaemella on different dung types (n° of 



sheep 

[51] 

S. fimicola 2 (4%) 



(%), in bold type 

C. cephalothecoides is quite common in 

Italy, particularly on polecat and goat dung. It 

is less frequent elsewhere, and recorded from 

mouse (Malloch & Benny 1973) and sheep 

dung (Valldosera & Guarro 1992, Moyne & 

Petit 2006). 

Chaetomium Kunze (Table 44) 

The genus Chaetomium is not obligetorely 

coprophilous, and non-coprophilous 

collections show a wide substrate tolerance, 

like those from dung. It is quite frequent in my 

damp chamber cultures (8% of records), 

exceeded only by Podospora (14%), Ascobolus 

(10%), Saccobolus (9%), and Schizothecium 

(9%), and it has the highest frequency on wild 

pig and leporine dung. 

C. bostrychodes is the most frequent in 

Italy (47% of all Chaetomium records from 

364 

damp chambers), and it particularly occurs on 

wild pig, hedgehog, rabbit, and bird dung. It is 

very common elsewhere, and I have found a lot 

of records from dung, particularly from leporine 

(36%) (Marchal 1884a, Tóth 1963, Dennis 

1986, Valldosera 1991, Malaval 2004, Favre 

2008, Richardson 2008b), also from carnivore 

(9%) (Mouton 1886, Bainier 1909, Palliser 

1910, Stratton 1921, Meyer 1941). 

Also C. globosum and C. murorum are 

fairly common on a variety of dungs in Italy. 

Both are widespread worldwide and have a 

wide substrate tolerance. Coprophilous collections 

of C. globosum have been recorded also 

from carnivore dung (4% of records found by 

me) (Skolko & Groves 1953, Seth 1970), especially 

from leporine (16%) (Udagawa 1960, 

Calviello 1971, Roy 1971, Lorenzo 1993), 

cervine (15%) (Tóth 1965, 1967, Caretta & 

Piontelli 1996, Cruys & Ericson 2008, Piasai & 

Manoch 2009), and bovine (14%) (Ahmed et 

al. 1971, Jeamjitt et al. 2007) dung. 

C. murorum has been recorded from a 

variety of dungs, carnivore included (6% of all 

records worldwide) (Karsten 1888, Skolko & 

Groves 1953, Ames 1963, Hubálek 1974), but I 

have found that it has a slight preference (19% 

of records) for leporine droppings (Mirza & 

Nasir 1968, Seth 1968, Piontelli et al. 2006, 

Favre 2008). 


Sordariales – Lasiosphaeriaceae 

Arnium Nitschke ex G. Winter (Table 45) 

The genus Arnium has a slow development 

on dung in damp chambers, and its 

frequency of occurrence has possibly been 

underestimated by me, as it often grows semi- 

to almost fully immersed. Leaving out its 

frequency on pig dung (few samples incubated), 

it prefers bovine and cervine dung in 

Italy. A. arizonense is the commonest species, 

with the higest occurrence on bovine dung, 

while the less common A. cervinum has been 

isolated from cervine dung exclusively. 

A. arizonense is widespread worldwide, 

and there are many records (28%) from bovine 

dung (Lundqvist 1972, Rattan & El-Buni 1980, 

Lorenzo & Havrylenko 2001), but also many 

(27%) from ovine (Valldosera 1991, 

Richardson 2008a,b), while the seven, mostly

European, collections of A. cervinum are 

known from cervine droppings only (Lundqvist 

1972, 1974, 1989). 

Bombardioidea C. Moreau ex N. Lundq. (Table 

46) 

Lundqvist (1972) described the European 

species of Bombardioidea, and Krug & Scott 

(1994) produced a world monograph of this 

genus, which preferably grows on leporine and 

cervine dung. 

B. stercoris is widespread but uncommon, 

and mostly grows on leporine (90% of 

world records), particularly hare (67%) dung 

(Winter 1887, Cain 1957, Lundqvist 1972, 

Muroi & Udagawa 1984, Valldosera 1991, 

Lorenzo 1992, Krug & Scott 1994, Leenurm 

1998, Richardson 2005, 2008b, Moyne & Petit 

2006, Eriksson 2009). 

Cercophora Fuckel (Table 47) 

Lundqvist (1972) widely described the 

nordic species of Cercophora, and Doveri 

(2004a) provided a world key to coprophilous 

species. 

Cercophora is uncommon in Italy and 

restricted to bovine and equine dung. These 

types of dung are the most preferred also 

elsewhere (Lundqvist 1972). 

I have found C. mirabilis both in the 

natural state and in damp chambers, always on 

cattle dung, and there are records worldwide 

(78%) confirming its marked preference for 

bovine dung (Lundqvist 1972, 1981, Udagawa 

& Muroi 1979, Barrasa & Moreno 1984, 

Barrasa 1985, Wang 1994, Chang & Wang 

2005, Richardson 2008c; Eriksson 2009). 

Fimetariella N. Lundq. (Table 48) 

The fimicolous genus Fimetariella was 

monographed by Krug (1995), who also studied 

its phylogenetic relationships, and recorded 

the rare F. microsperma from North and 

Central America and Europe. The European 

collections of F. microsperma are recorded 

from the Mediterranean area (Richardson, 

Lundqvist in Krug 1995, Richardson 2008b), 

all from cattle dung, another collection from 

goat dung in the Canary Islands (Lundqvist 

1997), whereas the North and Central American 

records (Krug 1995) are from American 

moose and burro dung. 


Podospora Ces. (Table 49) 

Mirza & Cain (1969) and Lundqvist 

(1972) monographed Podospora and Doveri 

(2008b) provided a key to the species and a 

world bibliography. 

Podospora is the most common coprophilous 

genus in Italy, with 41 (7%) records 

from the field and 240 (14%) from my damp 

chambers, and with the higest frequency of 

occurrence (39%) on all incubated samples. It 

preferably grows on equine (67% frequency), 

leporine (67%) and bovine (65%) dung, and is 

represented by a wide variety of species on 

horse and cattle dung (number of records/ 

number of dung samples = >1). 

In Italy P. decipiens is one of the commonest 

coprophilous pyrenomycetes s.l. (7% of 

records from damp chambers), second only to 

Sordaria fimicola. It has a wide substrate 

tolerance, but absolutely prefers bovine dung. 

Lundqvist (1972) verified 286 findings worldwide 

of this very common and widespread 

species and stated that "it is fastidious in its 

substrate choice" as 75% of all records are 

from cow and horse dung" and "the gap to the 

third preferred matrix (5%) is large". After 

Lundqvist, I have found 81 records worldwide, 

which prove a larger substrate tolerance 

and a smaller gap, as 31% are from horse 

(Abdullah & Rattan 1978, Caillet & Moyne 

1983, Barrasa 1985, Bokhary et al. 1989, Beyer 

1999) and cattle (García-Zorrón 1973, Dennis 

1981, Khan & Krug 1989, Wang 1994), 30% 

from ovine (Lundqvist 1973, Rattan & El–Buni 

1980, Guarro Artigas 1983, Wang 2000, 

Richardson 2005, 2008a,b), 17% from leporine 

dung (Slupinski 1991, Moyne & Petit 2006). 

Adding these latest records to Lundqvist's and 

mine from Italy, I have 38% from bovine dung, 

26% from equine, 11% from ovine, and 10% 

from leporine. 

Lundqvist (1972) also verified findings 

worldwide of P. myriaspora and P. pleiospora, 

two less common species morphologically 

similar to P. decipiens, but the former with64spored 

asci, the latter with 16–32-spored asci, 

and he concluded that P. myriaspora is linked 

to cow and horse dung (64% of records), while 

365

Table 43 Records from Italy of coprophilous Chaetomidium on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

366 

beech– 

marten 

[3] 

bird 

[27] 

cattle 

[55] 

horse 

[63] 

C. cephalothecoides 2 4 2 4 

(15%) (4%) (6%) 

C. fimeti 1 

(2%) 

C. megasporum 1 

Total 

Chaetomidium in 

natural state 

1 

Total 

2 4 2 5 

Chaetomidium in 

damp chambers** 

(15%) (4%) (8%) 

fallow 

deer 

[10] 

cervine leporine 

roe deer 

[37] 

goose 

[1] 

hare 

[20] 

1 (10%) 3 (8%) 1 2 

(10%) 

1 

(10%) 

3 

(8%) 

1 2 

(10%) 

rabbit 

[19] 

3 

(16%) 

3 

(16%) 

goat 

[16] 

5 

(31%) 

5 

(31%) 

hedgehog 

[12] 

insect 

[9] 

2 (17%) 1 

(11%) 

2 

(17%) 

1 

(11%) 

lizard 

[3] 

marmot 

[5] 

pig 

[5] 

polecat 

[5] 

sheep 

[51] 

2 1 (20%) 1(20%) 2(40%) 8 

(16%) 

2 1 (20%) 1 

(20%) 

2 

(40%) 

8 

(16%) 

tortoise 

[2] 

weasel 

[1] 

1 1 

1 1 

(8%) (13%) 

* The frequency (%) is not reported when samples of a dung type are less than five. ° Records from dung in the natural state, in normal type. Records from dung in damp chambers 

and frequency (%), in bold type. ** The frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of 


as whole must be regarded as occurring only once in one sample even if two species, or more, occur in that sample.


Table 44 Records from Italy of coprophilous Chaetomium on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

badger[2] 

bat [1] 

beech– 

marten[3] 

bird[27] 

cattle[55] 

donkey[6] 


horse[63] 

deer[24] 

fallow deer [10] 

roe deer [37] 

dormouse[4] 

fox[7] 

gecko [1] 

hare[20] 

rabbit[19] 

C. 

ancistrocladum 

1 

C. 

1 2 1+8 1 

4 

2 3 1 2 1 3 1+6 1 1 4 3 1 1 1 1 9 

1 1 2 1 1 4 

bostrychodes 

(30%) (2%) 

(6%) 

(20%) (8%) 

(28%) (15%) (31%) (6%) (17%) (33%) 

(20%) 

(18%) 

(40%) 

C. 

1 

carinthiacum 

(4%) 

C. crispatum 1 

(3%) 

C. cuniculorum 1+2 3 

1 

(10%) (16%) 

(2%) 

C. elatum 1 1 

(2%) 

C. funicola 2 

1 

1 

(3%) 

(3%) 

(5%) 

C. fusisporum 1 

(2%) 

C. 

1 

1 1 

1 

1 

gangligerum 

(2%) 

(3%) 

(6%) 

(2%) 

C. globosum 1 1 1 2 

2 1+1 

1 

2 

(4%) 

(3%) 

(5%) 

(8%) 

(4%) 

C. 

1 

homopilatum 

(3%) 

C. medusarum 1 

(4%) 

C. mollicellum 1 1 

1 2 1 2 

(14%) 

(8%) 

(20%) 

C. murorum 1 1 

2 1 1 

1 

4 1 1 

(2%) 

(5%) (5%) 

(8%) 

(8%) 

C. oblatum 1 

(17%) 

C. robustum 1+1 1 3 2 

1 1 

(3%) 

(10%) 

C. 

1 1 

1 1 

semencitrulli 

(4%) (2%) 

C. spinosum 2 

(4%) 

C. subaffine 2 

1 

(4%) 

(20%) 

C. 

1 

trigonosporum 

(20%) 

goat [16] 

chamois[6] 

hedgehog[12] 

lizard[3] 

mouse [2] 

pig[5] 

polecat[5] 

porcupine[1] 

rat[1] 

sheep[51] 

mouflon[1] 

snail[1] 

squirrel[3] 

toad[2] 

tortoise[2] 

weasel[1] 

367 

wild pig[10]

Table 44 (Continued) Records from Italy of coprophilous Chaetomium on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

368 

badger[2] 

bat [1] 

beech– 

marten[3] 

bird[27] 

cattle[55] 

donkey[6] 


horse[63] 

deer[24] 

fallow deer [10] 

roe deer [37] 

dormouse[4] 

fox[7] 

gecko [1] 

hare[20] 

rabbit[19] 

C. 

2 

1 

variostiolatum 

(4%) 

(5%) 

Total 

Chaetomium in 

natural state 

1 1 1 1 2 1 

Total 

3 1 3 12 10 1 8 2 8 9 3 1 9 12 2 1 7 6 1 1 2 2 2 19 1 3 1 2 1 2 6 

Chaetomium in 

damp 

chambers** 

(37%) (14%) (17%) (11%) 

(12%) 

(20%) 

(13%) 

(19%) (43%) (35%) (58%) 

(46%) 

(12%) (17%) 

(14%) 

(42%) 

(20%) (40%) 

(31%) 

(33%) 

(60%) 

* The frequency (%) is not reported when samples of a dung type are less than five. ° Records from dung in the natural state, in normal type. Records from dung in damp chambers and frequency (%), in bold type. ** The 

frequency (%) of a genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of that genus and the number of dung samples [ ]. The total occurrence of a genus does not 

result from a simple addition of occurrences of all species belonging to it, because the genus as whole must be regarded as occurring only once in one sample even if two species, or more, occur in that sample. 

Table 45 Records from Italy of coprophilous Arnium on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

cervine 

cattle 

horse 

deer 

roe deer 

sheep 

pig 

[55] 

[63] 

[24] 

[37] 

[51] 

[5] 

A. arizonense 2+13 

1 2+8 

(24%) 

(16%) 

A. caballinum 1 2 

2 

(3%) 

(8%) 

A. cervinum 3 

2+3 

(12%) 

(8%) 

A. imitans 1+4 

4 

(17%) 

(11%) 

A. inaequilaterale 1 

1 

(3%) 

(20%) 

A. septosporum 1 (20%) 

A. sudermanniae 1 

Total Arnium in natural state 3 3 2 2 

Total Arnium in damp chambers** 13 

2 

9 

8 

8 

2 

(24%) 

(3%) 

(25%) 

(16%) 

(16%) 

(40%) 

(20%) 


° Records from dung in damp chambers and frequency (%), in bold type. 




goat [16] 

chamois[6] 

hedgehog[12] 

lizard[3] 

mouse [2] 

pig[5] 

polecat[5] 

porcupine[1] 

rat[1] 

sheep[51] 

mouflon[1] 

snail[1] 

squirrel[3] 

toad[2] 

tortoise[2] 

weasel[1] 

wild pig[10]


Table 46 Records from Italy of coprophilous Bombardioidea on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)°. 

rabbit 

[19] 

B. stercoris 1 (5%) 


Table 47 Records from Italy of coprophilous Cercophora on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

cattle 

horse 

[55] 

[63] 

C. anisura 1 

(2%) 

C. coprophila 2 

C. gossypina 1 

C. mirabilis 2+1 (2%) 

C. septentrionalis 1 (2%) 

Total Cercophora in natural state 5 

Total Cercophora in damp chambers** 1 (2%) 2 (3%) 





Table 48 Records from Italy of coprophilous Fimetariella on different dung types (n° of dung samples in damp chamber cultures, in square brackets)°. 

cattle 

[55] 

F. microsperma 1 (2%) 


369

Table 49 Records from Italy of coprophilous Podospora on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

370 

badger 

[2] 

beech– 

marten 

[3] 

bird 

[27] 

cattle 

[55] 

donkey 

[6] 


horse 

[63] 

deer 

[24] 

fallow 

deer 

[10] 

roe 

deer 

[37] 

duck 

[1] 

P. alexandri 1 

P. anserina 1 3 1 2+8 1 1 1 3 

1 

1+5 

(4%) (5%) (17%) (13%) 

(5%) (16%) 

(20%) 

(10%) 

P. australis 1 

3 

1 1+2 

1 1 2 

1 

(2%) 

(5%) 

(10%) (5%) 

(5%) (5%) (12%) 

(2%) 

P. bifida 1 

1 

(2%) 

(2%) 

P. communis 10 1 10 1 1 

1 1 

3 

(18%) (17%) (16%) 

(3%) 

(5%) (6%) 

(6%) 

P. curvicolla 1 1 1 1+2 

(10%) 

(5%) (10%) 

P. dasypogon 1 

(2%) 

P. decipiens 10+24 1 2+18 3+2 1 1+4 

4 1 1 1 2 

1 2+16 1 

(44%) 

(28%) (8%) (10%) (11%) 

(20%) (5%) 

(17%) (25%) 

(20%) (31%) 

P. excentrica 1 

(5%) 

P. fimiseda 5+4 

1+3 

1 

2 

(7%) 

(5%) 

(5%) 

(4%) 

P. gigantea 1 

P. globosa 1 

1 

(2%) 

(5%) 

P. granulostriata 1 1 

1 

(3%) 

(2%) 

P. intestinacea 1 

3 

1 

(2%) 

(5%) 

P. myriaspora 1 4 

1 

1 

1 

(17%) (6%) 

(3%) 

(6%) 

P. pleiospora 1+1 

3 1 1 

5 1+5 1 

1 

(2%) 

(5%) 

(3%) 

(25%) (26%) (6%) 

P. pyriformis 6 

1 

(11%) 

(2%) 

P. setosa 1 1 3 5 1 2+12 1 3 1+1 1 1+3 5 

1 1 3+11 1 

(11%) (9%) 

(19%) (4%) (30%) (3%) 

(16%) (31%) 

(8%) 

(21%) (10%) 

Total Podospora in 

natural state 

7 2 8 7 5 3 1 1 8 1 

Total Podospora 1 1 4 56 3 65 3 6 11 1 13 18 10 1 2 1 1 1 1 41 1 

in damp 

chambers** 

(11%) (65%) (50%) (68%) (8%) (30%) (24%) 

(45%) (89%) (37%) (17%) (25%) (8%) 

(20%) (20%) (49%) (10%) 

(67%) (20%) (67%) (30%) 






hare 

[20] 

rabbit 

[19] 

goat 

[16] 

chamois 

[6] 

rock 

goat 

[8] 

hedgehog 

[12] 

lizard 

[3] 

marmot 

[5] 

pig 

[5] 

sheep 

[51] 

wild 

pig 

[10]

P. pleiospora to leporine (hare and rabbit) dung 

(83%). My data from Italy partly confirm 

Lundqvist's, as 62% of P. myriasporarecords 

are from equine dung (none from bovine) and 

the percentage of P. pleiospora records from 

leporine dung is lower (55% versus 83%). 

After Lundqvist (1972), P. myriaspora has frequently 

(56%) been recorded from bovine and 

equine dung (Lundqvist 1973, Jahn 1993, 

Wang 1994, 2000, Leenurm 1998), and P. 

pleiospora from leporine (56%) (Dennis 1981, 

Lundqvist 1973, 1981, Muroi & Udagawa 

1984, Barrasa 1985, Valldosera 1991, Jahn 

1993, Lorenzo & Havrylenko 2001, Beyer 

2004, Hu et al. 2006, Moyne & Petit 2006, 

Richardson, 2008b). 

Schizothecium Corda (Table 50) 

In Italy Schizothecium is one of the commonest 

pyrenomycetous genera on dung, both 

in the field and in damp chambers (9% of records), 

and it shows a wide substrate tolerance 

with a slightly higer frequency of occurrence 

on cattle dung. S. conicum is very frequent in 

Italy (4% of all ascomycete records from dung) 

and the third commonest pyrenomycete s.l. 

(9% of pyrenomycete and 55% of Schizothecium 

records), following Sordaria fimicola 

and Podospora decipiens. It grows on a variety 

of dungs, but it has a marked preference for 

bovine (44% of frequency in damp chambers). 

Its relationship with caprine dung is quite 

strange, as it grows on it with a medium 

frequency (26%), but with the highest on rock 

goat dung (50%). 

I have found about 250 records worldwide 

of this cosmopolitan species, 50% from 

equine (Fuckel 1869, Griffiths 1901, Piontelli 

et al. 1981, Lorenzo & Havrylenko 2001) and 

bovine dung (Cooke 1876, Marchal 1883, Goi 

dànich 1932, Meyer 1941, Munk 1948, 1957, 

Breton 1965, García-Zorrón 1975, Barrasa 

1985, Barrasa & Soláns 1989), 18% from ovine 

dung (Schmidt 1912, Guarro Artigas 1983, Loh 

meyer 1995, Wang 2000, Richardson 2004b, 

2005, Welt & Heine 2006a). S. vesticola is 

another common coprophilous species in Italy, 

like elsewhere. In my country it has the highest 

frequency on caprine (26% of occurrence in 

damp chambers), cervine (22%) and ovine 

(20%) dung. More than 200 records worldwide 

partly disagree with mine, as 29% of them are 


from ovine dung (Barrasa 1985, Richardson 

2004b, 2005, Welt & Heine 2006a), 25% from 

leporine (Marchal 1884a, Tóth 1967 Caillet & 

Moyne 1984, Bell & Mahoney 1995, Lorenzo 

& Havrylenko 2001), only 9% from caprine 

(De Meulder 2007c, Richardson 2008a), and 

8% from cervine dung (Tóth 1965, Moravec 

1968, Kruys & Ericson 2008). 

Strattonia Cif. (Table 51) 

Lundqvist (1972) emended the genus 

Strattonia and recognised three terricolous and 

nine fimicolous species, S. insignis included. 

He revised about twenty collections of S. insignis 

from Europe and North and South America, 

and regarded it as restricted to old horse dung, 

exceptionally to roe deer. Most latter records 

(Barrasa 1985, Valldosera & Guarro 1990a, 

Valldosera 1991, Lorenzo 1992, Moyne & Petit 

2006) are from horse dung, unlike mine and 

Leenurm (1998) from elk dung. 

Tripterosporella Subram. & Lodha (Table 52) 

The genus Tripterosporella encompasses 

four taxa: T. coprophila Subram. & Lodha, T. 

pakistani (Mirza) Malloch & Cain, T. heterospora 

(Mukerji, R.N. Kumar & N. Singh) 

Doveri var. heterospora, and T. heterospora 

var. octaspora Doveri, but the first two are 

possibly synonyms (von Arx 1973a, Doveri 

2010a). All are very rare, and T. heterospora is 

only known from the type locality, the var. 

heterospora from cervine (sambur) dung 

(Mukerji et al. 1995), the var. octaspora from 

horse dung (Doveri 2010a). T. pakistani has 

been recorded from horse (Mirza 1968) and 

rabbit (Mukerji et al. 1995) dung, T. coprophila 

from horse (Abdullah & Rattan 1978), sheep, 

and elephant (Subramanian & Lodha 1968, 

Khan & Krug 1994) dung. 

Zopfiella G. Winter (Table 53) 

In Italy almost all collections of Zopfiella 

have been isolated from horse dung, on which 

the genus as a whole has a high frequency of 

occurrence. 

Lundqvist (1969b) monographed Tripterospora 

Cain (= Zopfiella) and verified several 

records worldwide of Z. erostrata and some of 

the rarer Z. longicaudata. I have found a good 

many world records of Z. erostrata proving, 

together with Lundqvist's records, this species 

371

Table 50 Records from Italy of coprophilous Schizothecium on different dung types (n° of dung samples in damp chamber cultures, in square 

brackets)*°. 

372 

cattle 

[55] 

S. aloides 4+3 

(5%) 

S. conicum 7+24 

(44%) 

donkey 

[6] 


horse 

[63] 

deer 

[24] 

fallow 

deer 

[10] 

roe 

deer 

[37] 

fox 

[7] 

hare 

[20] 

1+1 2+18 1+4 

5 

1 1 1+4 

4 1 4+9 1 1 

(17%) (29) (17%) 

(13%) 

(5%) (5%) (25%) 

(50%) (20%) (18%) 

S. dakotense 2 (4%) 1 (4%) 1 3 1 

(5%) (16%) (6%) 

S. glutinans 1 

(4%) 

S. inaequale 1 

(2%) 

S. miniglutinans 3 

3 3 

1 

2+2 

(5%) 

(5%) (12%) 

(5%) 

(4%) 

S. pilosum 1 

(2%) 

S. simile 1 

S. squamulosum 1 

(3%) 

S. tetrasporum 1+2 1 

1 1 1+7 

2 

1 

(3%) (4%) 

(14%) (5%) (37%) 

(4%) 

S. vesticola 3 

2+9 3+8 1 8 1 1 1 4 2 2 1 2+10 

1 1 

(5%) 

(14%) (33%) 

(22%) (14%) (5%) 

(25%) (33%) (25%) (20%) (20%) 

S. vratislaviense 1 

(2%) 

Total Schizothecium 


11 1 5 4 1 1 1 8 2 1 

Total schizothecium 36 1 34 18 

14 2 5 11 9 2 6 2 1 23 1 

in dampchambers** (51%) (17%) (43%) (62%) 

(32%) (14%) (25%) (42%) (44%) (33%) (62%) (20%) 

(37%) 

(40%) (38%) (33%) (47%) (38%) 

rabbit 

[19] 

goat 

[16] 

chamois 

[6] 

rock 

goat 

[8] 

marmot 

[5] 

sheep 

[51] 

mouflon 

[1] 

wild 

pig 

[10] 

wolf 

[1] 

unidentified 

animal 

[2] 





Table 51 Records from Italy of coprophilous Stattonia on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

cattle [55] pig [5] 

S. insignis 1 1 (20%) 


and frequency (%), in bold type.


Table 52 Records from Italy of coprophilous Tripterosporella on different dung types (n° of dung 


horse [63] 

T. heterospora var. Octaspora 1 (2%) 

T. pakistani 1+1 (2%) 

Total Tripterosporella in natural state 1 

Total Tripterosporella in damp chambers** 2 (3%) 

* The frequency (%) is not reported when samples of a dung type are less than five. ° Records from dung in the natural 

state, in normal type. Records from dung in damp chambers and frequency (%), in bold type. ** The frequency (%) of a 

genus as whole, on each dung type in damp chamber, results from the ratio between the total number of occurrences of 

that genus and the number of dung samples. 

Table 53 Records from Italy of coprophilous Zopfiella on different dung types (n° of dung samples 


horse 

sheep 

[63] 

[51] 

Z. erostrata 5 (8%) 

Z. longicaudata 7+10 

1 

(16%) 

(2%) 

Total Zopfiella in natural state 7 

Total Zopfiella in damp chambers 15 (22%) 1 (2%) 


bold type. 

to have a wide substrate tolerance and a slight 

preference (19% of all records) for equine dung 

(Wilson 1947, Ahmed et al. 1971, Guarro 

Artigas 1983, Richardson 2004a). Z. longicaudata, 

on the contrary, is fastidious in its 

substrate choice, as 87% of all records are from 

horse dung (Lorenzo 1990, Valldosera 1991, 

Valldosera & Guarro 1992, Jahn 1993, Caretta 

et al. 1994, Holm & Ryman 2000, Heine & 

Welt 2008). 

Zygopleurage Boedijn (Table 54). 


Zygopleurage on different dung types (n° of 



cattle horse [ goat 

[55] 63] [16] 

Z. zygospora 2+2 (4%) 1 (2%) 1 (6%) 




The genus Zygopleurage was monographed 

by Lundqvist (1969a), who verified 18 

collections of Z. zygospora out of 71 recorded 

worldwide, and noticed its tolerance for a 

variety of dungs but also a marked preference 

for bovine dung. This choice is also confirmed 

by my records from Italy (4 out of 6 from 

bovine dung) and about 50% of world records 

found by me (Spegazzini 1878, Griffiths 1901, 

Kohlman-Adamska 1965, Cailleux 1971b, 

Furuya & Udagawa 1973, García-Zorrón 1973, 

Singh & Mukerji 1979, Lundqvist 1981, Khan 

& Krug 1989, Valldsosera 1991, Soláns 1994, 

Delgado et al. 2000, Moyne & Petit 2006, Welt 

& Heine 2007, Heine & Welt 2008, Richardson 

2008b,c). 


Sordariales – Sordariaceae 

Copromyces N. Lundq. (Table 55) 

Copromyces is a very rare and strictly fimicolous 

genus, encompassing only one species 

so far. Besides mine, I know few collections 

worldwide: two from Sweden (Lundqvist 

1967) and one from Great Britain (Richardson 

1998) on rabbit dung, and two from Venezuela 

(Delgado Avila et al. 2002) on rabbit and fox 

dung. Some other collections from rabbit dung 

in U.S.A. were recorded by D.P. Mahoney (in 

Lundqvist 1989), and other two from Mallorca 

(Spain) and California (USA) were recorded by 

Lundqvist (in Richardson 1998), but the dung 

source is unknown. 

373


Copromyces on different dung types (n° of 



Cattle 

[55] 

C. bisporus 1 (2%) 




Sordaria Ces. & De Not. (Table 56) 

In Italy Sordaria is one of the commonest 

pyrenomycetous genera on dung (13% of my 

records), with a wide substrate tolerance, but a 

preference for leporine droppings in damp 

chamber cultures (51% frequency of occurrence). 

S. fimicola is the commonest Sordaria 

sp. (72% of records; 6% frequency of occurrence 

on all samples in damp chambers), with 

the highest frequency of occurrence on equine 

and caprine dung. It is widespread worldwide 

and recorded from a variety of substrates, particularly 

from dung. I have found hundreds of 

records worldwide, so I limit myself to mention 

that Lundqvist (1972) recorded more than 600 

world collections, particularly from equine 

(35%) and leporine (31%) dung. 

Unlike S. fimicola, S. macrospora is a 

fastidious species in its substrate choice, as I 

have constantly found it on leporine dung and 

88% of records worldwide (those found by me 

added to findings verified by Lundqvist 1972) 

are from this type of dung (Furuya & Udagawa 

1973, Barrasa 1985, Hilber & Hilber 1987, 

Valldosera & Guarro 1990a, Valldosera 1991, 

Jahn 1993, Lundqvist 1997, Leenurm 1998, 

Coste & Rey 2000, Moyne & Petit 2006, Favre 

2008, Richardson 2008b). 


Xylariales – Xylariaceae 

Hypocopra (Fr.) J. Kicks f. (Table 57) 

Hypocopra is infrequent in Italy, but possibly 

underestimated as its perithecia develop 

late in damp chambers and grow almost fully 

immersed in dung. It has been found on a variety 

of herbivore dungs, but it prefers leporine 

droppings (5 records out of 11). H. equorum 

has been found once on horse dung, H. aff. 

festucacea only on cervine, and H. brefeldii, H. 

leporina and H. lojkaeana only on leporine 

374 

dung. H. equorum is absolutely linked to 

equine dung also elsewhere (Phillips & 

Plowright 1876, Griffiths 1901, Lundqvist 

1960, in litt., Moreau 1953), and collections 

from other types of dung possibly refer to H. 

leporina (= H. equorum f. leporina Niessl ex 

Rehm). 

Three collections of the rare H. festucacea 

has been isolated from cervine dung (Krug 

& Cain 1974), like mine, three from sheep 

(Valldosera & Guarro 1987, Moyne & Petit 

2006, Richardson 2008b), and one from goat 

(Valldosera 1991). 

I know some records from Europe 

(Lundqvist 1981, Moyne & Petit 2006, 

Richardson 2008b) of the less infrequent H. 

brefeldii, all from leporine dung, and I know 

very few records of H. leporina (Rehm 1889, 

Eriksson 2009) and H. lojkaeana (Rehm 1888, 

Eriksson 2009) from the same substrate. 

Poronia Willd. (Table 58) 

The genus Poronia is rare in Italy and it 

has been always found in the natural state. It is 

less rare and widespread in other temperate 

regions, but less common than in 19 th century 

(Reid 1986, Whalley & Dickson 1986). 

P. punctata is linked, with rare exceptions 

(Mukerji et al. 1969, Cribb 1988), to 

equine (74% of records worldwide) (Linnaeus 

1755, de Jaczewski 1895, Dawson 1900, 

Traverso 1907, Miller 1942, Munk 1957, 

Dennis 1959, 1981, Tóth 1963, Pilát 1972, 

Moreno & Bar rasa, 1977, García Bona 1978, 

Caillet & Moy ne 1984, Barrasa 1985, Reid 

1986, Whalley & Dickson 1986, Eriksson 

1992, Calonge et al. 1993, Coste & Rey 1993, 

Matočec 2000, Fouchier et al. 2009, Gube 

2010) and bovine dung (19%) (Willdenow 

1787, Pérez-Silva 1970, Stiers 1974, García 

Bona 1978, Barrasa 1985, Hladki de Sanz 

1997). P. erici has been recorded in Australia 

from a variety of herbivore dungs (Lundqvist 

1989, Lohmeyer 1994), in Europe from 

leporine droppings (Lohmeyer & Benkert 

1988, Jalink 1992, Schavey 1994, Ves terholt 

2000), except for my collection from Italy, and 

in Central America (San Martín et al. 1998) 

from equine and unidentified dung. 

Taxonomy 

In this section I describe and discuss 13 

fimicolous species new to Italy, and I update


Table 56 Records from Italy of coprophilous Sordaria on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

bird 

[27] 

cattle 

[55] 

donkey 

[6] 


horse 

[63] 

deer 

[24] 

fallow deer 

[10] 

roe deer 

[37] 

กog [0] 

dormouse[4] 

ferret [1] 

fox 

[7] 

goose[1] 

S. fimicola 5 2 2 7+21 1 2 1+5 1 1 1 2 

6+5 4+6 4+6 3 1 1 

1 1 1 5+15 1 2+1 

(18%) (4%) (33%) (33%) (4%) (20%) (13%) 

(28%) (25%) (31%) (37%) (37%) (8%) (20%) (20%) (20%) (29%) (10%) 

S. humana 1 1 

1 

1 1 1 1+2 

1 

(4%) (2%) 

(2%) 

(3%) 

(5%) (10%) 

S. lappae 1+1 

1 1 1 

1 

(2%) 

(5%) 

(2%) 

S. macro 

3 5+4 

spora 

(15%) (21%) 

S. superba 5 

1+1 

1 2 

2 

3 

2+1 1 1+1 

3 1 1+1 

(18%) 

(2%) 

(10%) (5%) 

(28%) (15%) 

(12%) (8%) (20%) 

(6%) (10%) 

Total 

Sordaria in 

natural 

state 

9 1 3 6 10 4 2 1 1 5 3 

Total 

11 3 2 24 1 3 8 

1 1 4 1 12 13 6 4 2 2 1 1 1 

19 2 2 

Sordaria in 

(33%) (36%) (4%) (20%) (13%) 

(50%) (53%) (37%) (25%) 

damp 

chambers** 

(37%) (5%) (36%) (11%) (43%) 

(51%) (33%) (17%) (40%) (20%) (20%) (31%) (20%) 





Table 57 Records from Italy of coprophilous Hypocopra on different dung types (n° of dung samples in damp chamber cultures, in square brackets)*°. 

Cervine Leporine 

cattle 

horse 

deer 

roe deer 

hare 

rabbit 

marmot 

[55] 

[63] 

[24] 

[37] 

[20] 

[19] 

[5] 

H. antarctica 1 1 

H. brefeldii 2 

H. equorum 1 

H. aff. festucacea 1 1 (3%) 

H. leporina 1 

H. lojkaeana 1 

H. merdaria 1 (3%) 1 (20%) 

Total Hypocopra in natural state 1 1 1 3 2 

Total Hypocopra in damp chambers** 2 (5%) 1 (20%) 



hare 

[20] 

rabbit 

[19] 

goat [16] 

rock goat 

[8] 

hedgehog 

[12] 

marmot 

[5] 

mouse [2] 

pig 

[5] 

polecat 

[5] 

porcupine[1 

] 

sheep 

[51] 

wild pig 

[10] 

375 

unidentified 

animal [2]

Table 58 Records from Italy of coprophilous Poronia on different dung types (n° of dung samples 


cattle 

donkey 

[55] 

[6] 

P. erici 1 

P. punctata 1 


bold type. 

some taxa following the same nomenclatural 

order as in section “Results” 


Pleosporales – Sporormiaceae 

Sporormiella affinis (Sacc., E. Bommer & M. 

Rousseau) S.I. Ahmed & Cain, Canadian 

Journal Botany 50: 425, 1972. Figs 1–9 

Sporormia affinis Sacc., E. Bommer & 

M. Rousseau in Bommer & Rousseau, Mem. 

Soc. Roy. Bot. Belg. 25(1): 171, 1886. 

Preussia affinis (Sacc., E. Bommer & 

M. Rousseau) Valldos. & Guarro, Bol. Soc. 

Micol. Madrid 14: 82, 1990. 

= Sporormia octomera var. macrospora 

Feltgen, Piltz-Flora Luxenburg. 1, 2: 223, 

1901. 

Pseudothecia ostiolate, subglobose, 400– 

600 µm diam., membranous, glabrous, blackish, 

with a papillate, scarcely differentiated 

neck, up to 200 µm diam. Peridium twolayered, 

pseudoparenchymatous: 1) endostratum 

of pale, thin-walled, polygonal cells, up to 

25 µm diam.; 2) exostratum of dark brown, 

very thick-walled, polygonal cells, 5–12 × 4–9 

µm. Pseudoparaphyses plentiful, cylindricfiliform, 

exceeding the asci, 2.5–4 µm diam., 

narrowing at the septa, branched, containing 

several hyaline vacuoles, not inflated at their 

tips. Asci bitunicate, non-amyloid, 8-spored, 

250–270 × 25–33 µm, cylindric-claviform, 

rounded at the apex, short-stalked (stalk up to 

30 µm long). Ascospores bi- to triseriate above, 

uniseriate in the lower part of the ascus, surrounded 

by a broad gelatinous sheath, 70–75 × 

14–15 µm, cylindric-claviform, usually slightly 

curved, hyaline at first, dark brown at maturity, 

smooth, thick-walled, 8-celled, deeply constrictted 

at the transverse septa, with cells 

easily separating from each other, each with a 

diagonal germ slit. End cells subconical, longer 

than wide (11–12.2 × 9–10 µm), longer than 

the middle cells, the uppermost with a slightly 

376 

pointed apex, the lowermost with a blunt apex. 

Middle cells cylindric, wider than long, the 

third from the upper end the widest. 

Material examined – Italy, Reggio 

Emilia, Ramiseto, Pratizzano, 1200 m a.s.l., 

about ten immersed, gregarious specimens on 

hare dung, L. Levorato, 23.5.2008, 217.2– 

Ramiseto, CLSM 006.08. 

Notes – The family Sporormiaceae is 

characterised by ascoloculate, aperiphysate, perithecioid 

or cleistothecioid ascomata, fissitunicate 

asci with a scarcely developed apical 

apparatus, and dark, septate, exceptionally onecelled, 

thick-walled ascospores, often with 

germ slits and breaking into part-spores at 

maturity (Barr 2000). The Sporormiaceae, 

which multigene analyses have proved to form 

a monophyletic group (Kruys & Wedin 2009, 

Schoch et al. 2009, Zhang et al. 2009), encompass 

ten saprobic, mostly coprophilous genera 

(Kirk et al. 2008), two of which, Preussia 

Fuckel and Sporormiella, have so similar features 

that the question of their reciprocal independence 

is open for several years. Preussia 

was conceived (Fuckel 1867) as having “carbonaceous, 

non-ostiolate unevenly breaking and 

finally fragmenting perithecia, suffused with a 

soft flocculose, spurious hyphopodium, clavate, 

long-stalked, 8-spored asci, no paraphyses, 

elongate-cylindric sporidia, finally fragmenting 

into four ovoidal and angular, dark 

brown parts”. Preussia funiculata Fuckel, was 

recorded from decaying oak wood and described 

as the type species. 

Ellis & Everhart (1892) established Sporormiella 

and assigned to this genus “softcarnose 

perithecia, embedded in a flattish, 

semi-immersed, subcarnose stroma, asci and 

sporidia as in Sporormia” i.e. “elongated, 8spored 

asci, 4–20-celled sporidia, with soon 

separating, dark colored cells; mostly fimicolous”. 

Sporormiella nigropurpurea Ellis & 

Everh., a species with ostiolate ascomata and


Figs 1–9 – Sporormiella affinis. 1 Pseudothecium. 2 Detail of exoperidium. 3 Apices of bitunicate, 

immature asci. 4, 8 Mature ascus with ascospores. 5 Detail of endoperidium. 6, 7, 9 Free ascospores 

in different stages. Bars 1 = 100 μm, 2, 3, 5 = 20 μm, 4, 6 = 40 μm, 7, 8, 9 = 15 μm. 

4-celled ascospores, was recorded from dung 

and described as type. 

There are, at the present, two schools of 

thought, resulting from different interpretations 

of the original diagnoses. The former (Cain 

1961, Ahmed & Cain 1972, Barrasa & Checa 

1991, Doveri 2004a, Lumbsch & Huhndorf 

2007, Kirk et al. 2008) based on morphological 

morphological and ecological characteristics, 

regards the two genera as distinct and Sporormiella 

characterised by semi-immersed or 

immersed, ostiolate pseudothecia, cylindrical to 

cylindric-clavate asci, 4- to poly-celled spores, 

and a preferable growth on dung, with more 

than 75% of species developing on this 

substrate. These features are opposite to those 

observed in Preussia, which has superficial, 

globose, cleistothecioid pseudothecia, clavate 

to broadly clavate asci, 4-celled spores, prefe- 

rable growth on soil and decaying wood or 

textiles (von Arx & van der Aa 1987), occasional 

growth on dung. The latter, opposing 

view does not acknowledge the independence 

of Sporormiella and regards it as a later 

synonym of Preussia, as the characters utilised 

for distinguishing the two genera, for instance 

the presence or absence of an ostiolum, are 

considerably affectted by environmental factors, 

such as changes of culture conditions (von 

Arx 1973a, Guarro et al. 1997a, Chang & 

Wang 2009). This view has produced the 

recombination in Preussia of the majority of 

Sporormiella spp. (von Arx 1973a, Valldosera 

& Guarro 1990b, Guarro et al. 1997a,b, 

Abdullah et al. 1999, Arenal et al. 2004, 2005, 

2007, Chang & Wang 2009, Kruys & Wedin 

2009), but has not induced me to recombine in 

Preussia my newly established Sporormiella 

377

spp. (Doveri 2005, 2007, Doveri & Coué 

2008b), because, in my opinion, neither the 

latest and most extensive phylogenetic study on 

Sporormiaceae (Kruys & Wedin 2009) 

definitely resolves the question of synonymy. 

Sporormiella was monographed by 

Ahmed & Cain (1972), who described more 

than 60 coprophilous species and a few others 

from different substrates. Their work, although 

rather old, must be regarded as still topical. 

Since then several new species have been 

published, all briefly discussed by Doveri 

(2004a), who also updated their bibliography. 

New additional taxa were later described 

(Arenal et al. 2005, 2007, Bell 2005, Doveri 

2005, 2007, Welt & Heine 2006, Doveri & 

Couè 2008, Chang & Wang 2009, Asgari & 

Zare 2010), some as Preussia, so the genus 

Sporormiella encompasses, at present, more 

than 80 species. 

S. affinis belongs to the group of species 

with 8-celled spores, the third cell from the 

upper end being the broadest. It differs from 

the others with such features S. corynespora 

(Niessl) S.I. Ahmed & Cain; S. octomera 

(Auersw.) S.I. Ahmed & Cain; S. octonalis S.I. 

Ahmed & Cain; S. schadospora S.I. Ahmed & 

Cain) particularly in having larger spores. The 

very similar S. tomilinii O.V. Korol. can be 

distinguished from S. affinis by its somewhat 

shorter spores (55–70 µm) and narrower asci 

(20–23 µm diam., Korolyova 2000). 

S. affinis has always been recorded 

(Bommer & Russeau 1886, Cain 1934, Ahmed 

& Cain 1972, Angel & Wicklow 1975, Barrasa 

& Moreno 1980, Valldosera & Guarro 1990b) 

from leporid pellets. 

I provide an updated key to coprophilous 

species of Sporormiella with 8-celled ascospores 

from Italy: 

1) Spores with the fourth cell from the 

upper end broader than the others ......... 2 

1*) Spores with the third cell from the upper 

end the broadest ..................................... 3 

2) Spores 40–51.5 × 7.5–8.5 µm (40–49 × 

8–9, Ahmed & Cain 1972) ....... S. pascua 

2*) Spores 34–42 × 7 µm (32–36 × 5.5–6.5, 

Ahmed & Cain 1972) ........ S. minipascua 

3) Spores 38–43.5 × 8–9 µm (40–48 × 7–8, 

Ahmed & Cain 1972). End cells conic– 

378 

ovoid ..................................... S. octomera 

3*) Spores larger. End cells hemispheric or 

conic-ovoid ............................................ 4 

4) Spores 48.5–63 × 12.5–14.5 µm (48–58 

× 12–14, Ahmed & Cain 1972), with 

hemispheric end cells. Asci abruptly 

contracted towards the base .. S. octonalis 

4*) End cells subconical. Asci gently 

narrowing towards the base ................... 5 

5) Spores 52.5–60 × 9.5–10.5 µm (50–59 × 

10–11.5, Ahmed & Cain 1972) ............... 

......................................... S. corynespora 

5*) Spores 70–75 × 14–15 µm (65–80 × 12– 

15, Ahmed & Cain 1972) .......... S. affinis 

Eurotiomycetes – Eurotiomycetidae – 

Onygenales – Gymnoascaceae 

Onygenalean fungi – modern history and 

update. 

No species of Onygenales was described 

in “Fungi Fimicoli Italici” (Doveri 2004), but 

Gymnoascus reessii Baran., G. devroeyi (G.F. 

Orr) Arx (Gymnoascaceae) (Figs 10–12), 

Aphanoascus fulvescens (Cooke) Apinis, and 

Xanthothecium peruvianum (Cain) Arx & 

Samson (Onygenaceae Berk.) were later identified 

and discussed (Doveri 2006). Since then 

other onygenalean fungi have been isolated 

from dung, i.e. the species of Gymnoascaceae 

here described, and a new Neogymnomyces (N. 

virgineus ad interim), which I am about to 

publish after studying its morphological, physiological 

and molecular features. 

The Gymnoascaceae are characterised by 

usually brightly coloured ascomata, sometimes 

without a well differentiated peridium, or with 

a gymnothecial peridium of interwoven hyphae 

forming a complete or incomplete, appendiculate 

or non-appendiculate reticulum, inside 

which an interascal tissue is absent, and the 

pseudoprototunicate, usually irregularly disposed, 

asci release ascospores by deliquescence. 

The one-celled ascospores lack a gelatinous 

equipment and are oblate, very pale to brightly 

coloured, never pitted, smooth or with polar 

thickenings and/or equatorial ridges or furrows. 

Anamorphs are absent or simply arthroconidial 

(Currah 1985, Cannon & Kirk 2007). 

Unlike Onygenaceae, members of Gymnoascaceae 

are not keratinolytic (Ulfig et al.


Figs 10–12 – Gymnoascus devroeyi. 10 Gymnothecium. 11 Free, oblate ascospores in side (red 

arrow) and frontal (white arrow) view. 12 Free ascospores and 8-spored asci (red arrow) above 

encrusted and thick-walled peridial hyphae (white arrow). Bars 10 = 100 μm, 11 = 12 μm, 12 = 10 

μm. 

1998, Scott & Untereiner 2004), but some are 

keratinophilic, i.e. they have not specific 

proteolytic enzymes but utilise simpler substances 

of the early keratinic decomposition 

(Filipello Marchisio 2000, Blyskal 2009), some 

are mildly cellulolytic (Howard 2002), others 

possibly chitinolytic (Orr 1977b, Lumley and 

Currah 1995). This explains why species of 

Gymnoascaceae have been isolated from soil, 

decaying wood, plants, dung, and human tissues, 

whererarely they behave as pathogens (de 

Hoog et al. 2000, Iwen et al. 2000). 

Currah (1985) accepted Acitheca Currah, 

Arachniotus J. Schröt. (1893), Gymnascella 

Peck (1884), Gymnoascoideus G.F. Orr et al. 

(1977), and Gymnoascus s.str., in Gymnoascaceae, 

regarding Narasimhella Thirum. & P.N. 

Mathur. (1965), Petalosporus G.R. Ghosh et al. 

(1963), Plunkettomyces G.F. Orr (1977b), and 

Pseudoarachniotus Kuehn (1957) as synonyms 

of Gymnascella. He recognised three types of 

hyphal peridium, i.e. a “reticuloperidium” of 

branched and anastomosed, thick-walled, often 

appendiculate hyphae, differentiated from the 

vegetative hyphae, an “incompositoperidium” 

of scarce and uncomposed, thick-walled 

hyphae, and a “telaperidium” of thin-walled 

hyphae, scarcely differentiated from the vegetative 

hyphae. The peridial frame, together with 

the ascospore shape, ornamentation and 

arrangement inside the asci, and absence or 

possible presence of a simple mitosporic stage, 

were regarded as important features to differentiate 

genera. So, according to Currah 

(1985), Acitheca has a reticuloperidium with 

pointed appendages, smooth ascospores, and an 

unknown anamorph, Arachniotus an unappendiculate, 

sometimes absent telaperidium, bival- 

379

vate ascospores (with an equatorial groove 

bordered by two ridges), and no anamorph, 

Gymnascella an unappendiculate telaperidium, 

sometimes with intervals of thick-walled 

hyphae, ascospores smooth or with an equatorial 

groove or ridge and/or polar thickenings, 

and no or arthroconidial anamorphs, Gymnoascoideus, 

a reticuloperidium lacking appendages, 

smooth ascospores with a petaloid 

arrangement inside the asci, and an arthroconidial 

anamorph, Gymnoascus s.str., an appendiculate 

reticuloperidium, smooth asco-spores, 

and no anamorph. 

Von Arx (1986) stated that the presence 

or absence of a peridium and peridial appendages 

are inadequate to circumscribe genera in 

Gymnoascaceae, as some species can develop a 

normal peridium in natural conditions and 

completely lack it in subcultures. Starting from 

this assumption, he conceived Gymnoascus in a 

much broader sense than Currah (1985) and 

emended it, including Acitheca, Arachniotus, 

Gymnascella, and Gymnoascoideus in this 

genus. He also accepted Narasimhella with its 

often stipitate ascomata, unequally bivalvate, 

hyaline rather than pigmented ascospores, and 

asci not born from croziers. 

Although von Arx (1987) attempted a 

new classification of the Gymnoascaceae, 

based on ascospore morphology, Currah’s 

(1985) systematics was widely followed for 

many years and re-examined with the introduction 

of molecular techniques and phylogenetic 

studies. 

Unlike the assumption by Currah (1985, 

1994) that Gymnoascaceae are polyphyletic, 

data of Bowman et al. (1996) and Sugiyama et 

al. (1999) from small subunit rDNA genes 

sequences were consistent with a monophyletic 

descendance of this family. 

Subsequent studies (Sugiyama & 

Mikawa 2001, Sugiyama et al. 2002), based on 

large subunit ribosomal DNA sequences, confirmed 

the monophyletic nature of the Gymnoascaceae, 

but also proved that within a major 

clade, species are grouped in four subclades, 

Gymnascella is not monophyletic, and “the 

ascospore morphology and phylogenetic structure 

are incongruent”. 

These latest results have induced 

Lumbsch & Huhndorf (2007) to keep independent 

the genera accepted by Currah (1985) and 

380 

also include in Gymnoascaceae Narasimhella, 

Kraurogymnocarpa Udagawa & Uchiyama 

(1999), Mallochia Arx & Samson (1986), and 

Orromyces Sur & G.R. Ghosh (Ghosh & Sur 

1985). Kraurogymnocarpa and Mallochia have 

bivalvate, respectively aculeate-tuberculate and 

spiny ascospores, an atypical feature in Gymnoascaceae, 

the former has also a reticuloperidium 

with pointed appendages, the latter lacks 

peridial hyphae. According to Sigler et al. 

(1998) the monotypic Orromyces, with its 

pitted ascospores and spiralate peridial appendages 

is a synonym of Uncinocarpus queenslandicus 

(Apinis & R.G. Rees) Sigler in 

Onygenaceae. 

A phylogenetic analysis of the ITS region 

and 5.8S ribosomal DNA gene sequences (Solé 

et al. 2002) proved that the majority of collections 

isolated in pure culture, and classified by 

Currah (1985) and von Arx (1986) as Arachniotus, 

Gymnascella, Gymnoascoideus, Gymnoascus, 

and Narasimhella, forms a monophyletic 

clade. Molecular data, in combination 

with morphological features, supported the proposal 

to unify these taxa under the prior genus 

Gymnoascus, to which a key was also provided 

(Solé et al. 2002). 

After Solé et al. (2002), I conceive Gymnoascus 

in a broad sense, and under this name I 

classify my Italian collections of Gymnoascaceae. 

I also provide the following key to genera 

of Gymnoascaceae, and to species of Gymnoascus 

s.l. from Italy: 

1) Peridium a complete net of thick-walled 

hyphae (reticuloperidium) with pointed 

appendages ............................................ 2 

1*) Peridium of thin- or thick-walled hyphae. 

Appendages, when present, different in 

shape. Gymnoascus s.l. .......................... 3 

2) Ascospores aculeate-tuberculate, bivalvate 

......................... Kraurogymnocarpa 

2*) Ascospores smooth .................... Acitheca 

3) Peridium a complete net of thick-walled 

hyphae. Ascospores 3V.5–4 × 3.5–4 × 

2.5–3 µm ................................................ 4 

3*) Peridial hyphae thin-walled (telaperidium), 

or thick-walled and forming an 

incomplete net (incompositoperidium). 

Ascospores larger .................................. 5

4) Peridium with hooked appendages. 

Ascospores smooth ................... G. reessii 

4*) Peridium not appendiculate. Ascospores 

with an equatorial furrow lined with two 

parallel ridges ............................ G. ruber 

5) Ascospores smooth, 4.5–5 × 4.5–5 × 3– 

3.5 µm. Hyphae forming an incompositoperidium 

................................ G. devroeyi 

5*) Ascospores ornamented. Hyphae forming 

both a telaperidium and an incompositoperidium 

................................................. 6 

6) Ascospores 4.5–6 × 4.5–6 × 3.5–4.5 µm, 

smooth, with an equatorial rim ................ 

.............................................. G. littoralis 

6*) Ascospores 6–7.5 × 5.5–6.5 × 3.5–4.5 

µm, rough, with an equatorial rim and 

polar thickenings ............ G. dankaliensis 

Gymnoascus dankaliensis (Castell. ex J.F.H. 

Beyma) Arx, Persoonia 13: 177, 1986. 

Figs 13–15 

≡ Trichophyton dankaliense Castell., J. 

Trop. Med. Hyg. 40: 315, 1937 (nom. inval., 

art. 34, 36 ICBN). 

≡ Arachniotus dankaliensis Castell. ex 

J.F.H. Beyma, Antonie van Leeuwenhoek 8: 

107, 1942. 

≡ Gymnascella dankaliensis (Castell. ex 

J.F.H. Beyma) Currah, Mycotaxon 24: 77, 

1985. 

= Pseudoarachniotus roseus Kuehn, 

Mycologia 49: 695, 1957 (fide von Arx, 1971). 

= Arachniotus flavoluteus Kuehn & G.F. 

Orr, Mycologia 51: 864, 1959 (fide Currah, 

1985). 

= Waldemaria pernambucensis Bat. et 

al., Atas Inst. Micol. Univ. Recife 1: 8, 1960 

(fide von Arx, 1971). 

= Pseudoarachniotus flavus Thirum. & 

P.N. Mathur, in Mathur & Thirumalachar, 

Mycopath. Mycol. appl. 40(2): 99, 1970 (fide 

von Arx, 1971). 

= Pseudoarachniotus halophilus Panwar 

et al., in Mathur & Thirumalachar, Mycopath. 

Mycol. appl. 40(2): 100, 1970 (fide von Arx, 

1971). 

= Pseudoarachniotus terrestris Thirum. 

& P.N. Mathur, in Mathur & Thirumalachar, 


von Arx, 1971). 


= Pseudoarachniotus thirumalacharii 

P.N. Mathur, in Mathur & Thirumalachar, 


von Arx, 1971). 

= Pseudoarachniotus flavoluteus (Kuehn 

& G.F. Orr) G.F. Orr et al., Mycologia 69: 154, 

1977 (fide Currah, 1985). 

Ascomata globose or subglobose, yellow 

to bright orange, downy gymnothecia, 80–150 

µm diam., rarely covered with scarce, white, 

aerial hyphae. Peridium a wide mesh of thin, 

branched, septate hyphae, 2–7 µm diam., sometimes 

slightly swollen at the septa, finely 

encrusted with a yellowish orange crystalline 

material. Well differentiated appendage absent. 

Asci ephemeral, globose or broadly ellipsoidal, 

8-spored, thin-walled, 14–17 × 10–14 µm. Ascospores 

6–7.5 × 5.5–6.5 × 3.5–4.5 µm, oblate, 

globose in frontal view, ellipsoid-fusiform with 

a broad equatorial ridge and two polar thickenings 

in side view, thick-walled, yellowish, 

asperulate, conglobate inside the asci, lacking a 

petaloid arrangement. Anamorph not detected. 

Material examined – Italy, Livorno, 

Montioni, 0 m a.s.l., usually gregarious, 

sometimes crowded, superficial specimens, on 

sheep dung in a damp chamber culture, F. 

Doveri, 18.3.2009, 306.3–Suvereto, CLSM 

005.09; Grosseto, Suvereto, Azienda Agricola 

“La Suveraia”, 0 m a.s.l., on horse dung in a 

damp chamber culture, F. Doveri, 18.3.2009, 

306.3–Suvereto, CLSM 005.09 bis; Livorno, 

Bibbona, Fattoria di Faltona, 50 m, on sheep 

dung in a damp chamber culture, F. Doveri, 

18.3.09, 295.3–Sassa, CLSM 005.09 ter; Pisa, 

Volterra, Villamagna, podere Vallicella, 200 

m, on pig dung in a damp chamber culture, F. 

Doveri, 13.4.09, 285.3–Lajatico, CLSM 005.09 

quarter; Pisa, Volterra, Villamagna, podere 

Vallicella, 200 m, on cattle dung in a damp 

chamber culture, F. Doveri, 13.4.09, 285.3– 

Lajatico, CLSM 005.09 penta. 

Notes – G. dankaliensis is characterised 

by a telaperidium, and comparatively large and 

rough ascospores with a broad equatorial ridge 

and polar thickenings. 

G. marginisporus (Kuehn & G.F. Orr) 

Solé et al. has similar ascospores which, however, 

are smaller (2.2–3 × 4–4.4 µm, Kuehn & 

Orr 1963) and lack polar thickenings. 

G. citrinus (Massee & E.S. Salmon) Arx 

has ascospores with polar and equatorial 

381

Figs 13–15 – Gymnoascus dankaliensis. 13 Gymnothecium. 14 Free, oblate ascospores and 8spored 

asci. 15 Free ascospores mixed with end peridial hyphae. Bars 13 = 50 μm, 14, 15 = 20 μm. 

thickenings, like G. dankaliensis, but also 

lemon-yellow ascomata and smooth, slightly 

smaller ascospores with a narrower and much 

less marked equatorial rim (Currah 1985). 

G. dankaliensis has been isolated from 

several substrates, often from different kinds of 

dung (Kuehn & Orr 1959, Mirza & Nasir 1968, 

Currah 1985, Valldosera & Guarro 1988, 

Elshafie 2005). It was proved to be keratinophilic 

(Caretta et al. 1992, Cugnani 2000) and 

occasionally a weak and tardy keratinolytic 

(Ulfig et al. 1998). 

Gymnoascus littoralis (G.F. Orr) Arx, Persoonia 

13: 179, 1986. Figs. 16–18 

≡ Plunkettomyces littoralis G.F. Orr, 

Mycotaxon 6: 34, 1977. 

≡ Arachniotus littoralis (G.F. Orr) Arx, 

Persoonia 9: 397, 1977. 

≡ Gymnascella littoralis (G.F. Orr) 

Currah, Mycotaxon 24: 87, 1985. 

382 

Ascomata globose or subglobose, pale 

yellow to bright orange, downy gymnothecia, 

50–450 µm diam., rarely covered with scarce, 

white, aerial hyphae, with a darker orange, 

much less filamentous inner part. Peridium a 

wide mesh of thin- to thick-walled, smooth or 

asperulate, hyaline to pale yellowish or orange, 

branched, septate hyphae, 2–4 µm diam., with 

usually rounded ends. Racket hyphae (swollen 

at the septa) present but scarce. Well differentiated 

appendages absent. Asci ephemeral, 

globose or broadly ellipsoidal, 8-spored, thinwalled, 

9–11 × 7–10 µm. Ascospores 4.5–6 × 

4.5–6 × 3.5–4.5 µm, oblate, globose in frontal 

view, ellipsoid-fusiform with a narrow equatorial 

rim (0.3–0.5 µm, rarely up to 1 µm diam.) 

in side view, thick-walled, yellowish, smooth, 

conglobate inside the asci, lacking a petaloid 

arrangement. Anamorph not detected. 

Material examined – Italy, Reggio 

Emilia, Ramiseto, Pratizzano, 1200 m a.s.l., 

usually gregarious, often crowded, superficial


Figs 16–18 – Gymnoascus littoralis. 16 Free, oblate ascospores and asci mixed with peridial 

hyphae. 17 Asci with ascospores in different stages. 18 Gymnothecia. Bars 16, 17 = 10 μm, 18 = 

150 μm. 

specimens, on badger dung in a damp chamber 

culture, G. Robich, 24.5.2008, 217.2-Ramiseto, 

CLSM 008.08. 

Notes – Comparatively large and smooth 

ascospores with a narrow equatorial rim, and 

thin- to thick-walled peridial hypae characterise 

G. littoralis. Ascospores with a marked 

equatorial rim are also observable in G. marginisporus 

and G. punctatus (B.G. Dutta & G.R. 

Ghosh) Arx. Both have, however, rough rather 

than smooth ascospores, smaller in the former 

(see above, under G. dankaliensis), and 

provided with a broader equatorial rim in the 

latter, attaining one third of the spore width 

(Currah 1985). 

The majority of Gymnoascus species 

have been isolated from dung, litter, soil, and 

fresh water polluted with animal dejections 

(Currah 1985, Ulfig et al. 1998) but, before my 

Italian collection, G. littoralis had been isolated 

only from beached chitinous materials (Orr 

1977a) or other substrates likely containing 

chitin (Lumley & Currah 1995, Ulfig et al. 

1998), giving birth to the hypothesis of its 

chitinolytic activity (Lumley & Currah 1995). 

Techniques for isolating chitinophilic 

fungi and proving their chitinolytic ability are 

known and have been utilised for many years 

(De Boer et al. 1999, Shubakov & 

Kucheryavykh 2004, Swiontek-Brzezinska et 

al. 2007) but, as far as I know, G. littoralis has 

never been isolated with specific techniques, 

neither its capability to degrade chitin has been 

tested. 

Chitin is a natural polysaccaride, essential 

component of the exoskeleton of arthropodes, 

both crustaceans and insects, radula of 

mollusks, and beaks of cephalopods (Jeuniaux 

383

et al. 1993). Variable amounts of chitin are present 

on any kind of dung samples, both fresh 

and dried, in natural conditions or in damp 

chambers, particularly linked to the shells of 

beetles or coats of colonising mites and worms, 

as remnants of dead animals or sheddings. So 

the hypothesis of a chitinolytic activity can be 

extended to my collection of G. littoralis from 

dung as well as to other collections of Gymnoascus 

spp. from this substrate. The chitinolytic 

hypothesis is also supported by my experience 

in cultivating coprophilous fungi in 

damp chambers, according to which all collections 

of Gymnoascus spp. from dung have a 

tardy development in comparison with other 

families of Ascomycota Berk. This slowness 

can been explained, in my opinion, by lack of 

nutritional competition between Gymnoascaceae 

and other fungi of this coenosis, which 

early develop by feeding on simpler and more 

degradable substances. Materials containing 

chitin are later available, when, since about the 

second week, the life cycle of worms and insects 

makes for the way down in damp chamber, 

and Gymnoascus spp. begins to appear. 

From now on, G. littoralis can be called 

occasional or facultative coprophilous, as it 

optionally grows on dung and does not need to 

pass through the digestive tract of an host in 

order to germinate (Massee & Salmon 1901, 

Webster 1970, Wicklow 1992). 

Gymnoascus ruber Tiegh., Bull. Soc. Bot. Fr. 

24: 159, 1877. Figs. 19–20 

≡ Arachniotus ruber (Tiegh.) J. Schröt., 

384 

in Cohn, Krypt.-Fl. Schlesien 3: 210, 1893. 

≡ Pseudoarachniotus ruber (Tiegh.) G.F. 

Orr et al., Mycologia 69: 153, 1977. 

≡ Pseudoarachniotus trochleosporus 

Kuehn & G.F. Orr, in Orr & Kuehn, Mycologia 

64: 58, 1972 (fide Currah, 1985). 

Ascomata gymnothecial, globose to pulvinate, 

dull orange, sometimes with pinkish 

shades, mealy, 30–100 µm diam. Peridial 

hyphae scarce, sometimes absent, ephemeral, 

soon collapsed, smooth, hyaline, septate, 1.5–4 

µm diam. Asci ephemeral, globose or broadly 

ellipsoidal, 8-spored, thin-walled, 7.5–9 × 6.5– 

7.5 µm. Ascospores 3.5–4(–4.5) × 3.5–4 × 2.5– 

3 µm, oblate, globose in frontal view, ellipsoidal 

with a deep equatorial furrow bordered by 

two ridges (“pulley-wheel”-shaped, Currah 

1985) in side view, pale yellowish orange, 

smooth, conglobate inside the asci, lacking a 

petaloid arrangement. Anamorph: not detected. 


Bibbona, Fattoria di Faltona, 50 m a.s.l., about 

ten isolated or gregarious, superficial specimens, 

on cattle dung in a damp chamber culture, 

F. Doveri, 18.3.2009, 295.3-Sassa, CLSM 

003.09. 

Notes – Orange to red gymnothecia, and 

small and pulley-wheel-shaped ascospores are 

the main features of G. ruber, partly shared 

with G. confluens Sartory & Bainier, which has 

ascospores similar in size, but with a shallow 

depression not lined with ridges (Currah 1985). 

G. ruber has been isolated from several 

substrates, often from different kinds of dung 

Figs 19–20 – Gymnoascus ruber. 19 Gymnothecium. 20 Free, oblate ascospores and 8-spored asci. 

Bars 19 = 40 μm, 20 = 15 μm.

(van Tieghem 1877, Schroeter 1893, Massee 

1895, Massee & Salmon 1901, 1902, Apinis 

1964, Currah 1985). 


– Pezizaceae 

Iodophanus difformis (P. Karst.) Kimbr., 

Luck-Allen & Cain, Am. J. Bot. 56(10): 1198, 

1969. Figs 21–29 

≡ Ascobolus testaceus var. difformis P. 

Karst., Syn. Pez. Ascob. Fenn. 43, 1861. 

≡ Ascobolus difformis (P. Karst.) Nyl., 

Notis. Faun. Fl. Fenn. 10: 85, 1869. 

≡ Ascophanus difformis (P. Karst.) 

Boud., Ann. Sci. Natur. Bot. V, 10: 252, 1869. 

≡ Peziza difformis (P. Karst.) P. Karst., 

Bidr. Finl. Natur. Folk. 19: 61, 1871. 

Apothecia cleistohymenial, sessile, membranous-gelatinous, 

pale yellowish brown, 

globose in the early stages, often becoming 

pulvinate, 0.5–0.7 mm diam., with a hardly 

differentiated, slightly darker margin. Disc 

slightly convex, papillate owing to the protruding 

asci, the same colour as the outer surface. 

The latter smooth, with abundant basal hyphoid 

hairs. Subhymenium a textura angularis of 

polygonal cells, up to 10 µm diam. Medullary 

excipulum a textura intricata of hyaline, cylindric 

or ellipsoidal hyphae, 5–9 µm diam., 

narrowing at the septa. Ectal excipulum a 

textura globulosa-angularis of thick-walled, 

yellowish cells, up to 35 µm diam., scarcely 

elongated towards the apothecial margin. Many 

septate, thick-walled hyphoid hairs, 2–4 µm 

diam., with an enlarged base, are observable at 

the base of apothecia. Paraphyses exceeding 

the asci, 3–6 µm diam., simple or sometimes 

branched at the base, containing abundant 

yellowish pigments, septate, straight or slightly 

curved at the apex, non or slightly inflated at 

the tips. Asci 175–220 × 20–30 µm, 8-spored, 

weakly but diffusely amyloid, operculate, congophilous, 

cylindric-claviform, thick-walled, 

short-stalked, rounded at the apex. Ascospores 

uni- to irregularly biseriate, (16–) 16.5–19 (– 

19.5) × (9.5–) 10–11.5 µm, ellipsoidal (Q = 

1.54–1.88; average Q = 1.69), symmetrical, 

each with a hardly observable gelatinous 

sheath, hyaline, without oil droplets or gaseous 

bubbles, thick-walled in the early stages, roun- 


dish at the ends, smooth, with a wrinkled, 

cyanophilous episporium. 


Bibbona, Fattoria di Faltona, 50 m, about ten 

gregarious or isolated, superficial specimens on 

old cattle dung in a damp chamber culture, F. 

Doveri,18.03.2009,295.3-Sassa,CLSM 004.09. 

Notes – The genus Iodophanus Korf was 

originally placed (Kimbrough & Korf 1967, 

Kimbrough et al., 1969) in tribus Pezizae Fr. 

(Pezizaceae Dumort.) on account of its amyloid 

asci, cyanophilic episporic markings, and an 

Oedocephalum Preuss anamorph. Despite subsequent 

systematic changes, it finally was confirmed 

in Pezizaceae by molecular phylogenetic 

studies (Landvik et al 1997, 1998, Hansen et 

al. 2001, 2005), which also proved this family 

to be monophyletic (Hansen et al. 2005, 

Hansen & Pfister 2006). 

The latest worldwide monograph on Iodophanus 

dates back to more than forty years 

ago (Kimbrough et al. 1969). Nine taxa were 

described and placed in a key, which was later 

updated (Doveri 2004a) with the addition of six 

new species. Since then no new species have 

been published. 

I. difformis is characterised by yellowish 

apothecia and paraphyses with yellow contents, 

and it is unique in lacking episporic, cyanophilic 

markings, although its smooth epispore 

strongly stains all the same in cotton blue. As 

previously observed (Kimbrough et al. 1969), 

and also noticed by me in my collection, the 

ascospores wrinkle in some solvents and appear 

to be transversely furrowed. 

Most Iodophanus sp. are coprophilous, 

but I. difformis has been isolated from several 

substrates, particularly from textiles and paper 

(Karsten 1861, 1870, Kimbrough et al. 1969). I 

also have found records of this seemingly rare 

species from horse (Prokhorov 1989b), sheep, 

and donkey (Abdullah & Alutbi 1993) dung. 


– Pyronemataceae 

Orbicula parietina (Schrad.) S. Hughes, 

Mycological Papers 42: 1, 1951. Figs 30–42 

≡ Didymium parietinum Schrad., Nova 

genera plantarum: 24, 1797. 

385

Figs 21–24 – Iodophanus difformis. 21 Overall view of hymenium. 22 (Congo red) Cells of 

subymenium (white arrow) with bases of asci (red arrow) and paraphyses (black arrow). 23 

Cleistohymenial apothecium. 24 (Congo red) Hyphoid hairs (arrow) arising from the outermost 

cells of ectal excipulum. Bars 21 = 100 μm, 22, 24 = 20 μm, 23 = 500 μm. 

Figs 25–29 – Iodophanus difformis. 25 (Melzer) Apices of paraphyses (white arrow) between 8spored 

asci. 26, 29 Free ascospores above apices of paraphyses and asci. 27 Immature asci. 28 

(Congo red) Mature ascus with ascospores. Bars 25, 28, 29 = 18 μm, 26 = 12 μm, 27 = 40 μm. 

386

≡ Lycogala parietinum (Schrad.) Fr., 

Syst. Mycol. 3: 83, 1829. 

≡ Mycogala parietinum (Schrad.) Sacc., 

Syll. Fung. 3: 185, 1884. 

≡ Anixia parietina (Schrad.) Lindau, in 

Engler & Prantl, Nat. Pflanzenfam., Teil. I , 1: 

334, 1897. 

≡ Licea bicolor Pers., Syn. Meth. Fung.: 

195, 1801 (nom. nov.). 

≡ Tubulina bicolor (Pers.) Poir., in 

Lamarck, Enc. Meth. Bot. 8: 131, 1808. 

= Anixia truncigena Hoffm., Icon. 

Analyt. Fung. 3: 70, 1863. 

= Anixia spadicea Fuckel, Jb. Nassau. 

Ver. Naturk. 23–24: 91, 1870. 

= Sphaeria cyclospora Cooke, Pop. Sci. 

Rev. 10: 12, 1871. 

= Orbicula cyclospora (Cooke) Cooke, 

Handb. Brit. Fung. 2: 926, 1871. 

= Anixia cyclospora (Cooke) Sacc., Syll. 

Fung. 1: 36, 1882. 

= Chaetomium glabrum Berk. in Berk. & 

Broome, Ann. Mag. Nat. Hist. Ser. 4, 11: 349, 

1873. 

= Orbicula perichaenoides Cooke, 

Grevillea 8 (no. 45): 10, 1879. 

Ascomata cleistothecioid, 400–800 µm 

diam., 200–300 µm high, subglobose to pulvinate, 

flattened or slightly concave at their 

bases, semi-membranous, brown, violet dotted, 

glabrous in the upper portion, with hyphoid 

hairs in the lower, with yellowish contents. 

Peridium friable, two-layered: endostratum a 

textura globulosa-angularis with pale, thinwalled, 

roundish or polygonal cells, 8–20 µm 

diam.; exostratum a textura epidermoidea, angularis 

at intervals, of irregularly lobed, thickwalled, 

dark brown cells, 4–15 µm diam., interspaced 

with an amorphous, purplish pigment. 

Hyphoid hairs arising from the outermost exoperidial 

cells, brown, darker at the base, thickwalled, 

wavy, sometimes fasciculate, sparsely 

septate, more densely septate at their base, 

sometimes swollen at the septa, 4–17 µm diam. 

Subhymenium placed at the cleistothecial base, 

formed of small, hyaline, thin–walled cells. 

Paraphyses numerous, mixed with asci and 

usually exceeding them, 2–2.5 µm diam., non– 

or slightly enlarged at the apex (up to 4 µm 

diam.), sraight or somewhat curved, septate, 

often branched in the lower portion, sometimes 

anastomosed, filled with a few pigments. Asci 


numerous, unitunicate, non-amyloid, inoperculate, 

8-spored, cylindrical, 100–130 × 11–14 

µm, thin-walled, long-stalked, rounded at the 

apex, evanescent. Ascospores regularly or 

sometimes irregularly uniseriate, oblate, 8–10.5 

× 8–10.5 × 7–9 µm, globose to subglobose in 

frontal view, broadly ellipsoidal in side view, 

hyaline to pale yellowish, smooth, thickwalled, 

forming a powdery mass at maturity, 

when asci disintegrate. 

Material examined – Italy, Vicenza, 

Tonezza del Cimone, Altopiano dei Fiorentini, 

1450 m a.s.l., more than fifty scattered or crowded, 

superficial specimens on hare dung in a 

damp chamber culture, A. Bizzi, 13.7.2008, 

082.3–Arsiero, CLSM 002.10. 

Notes – The monospecific genus Orbicula 

Cooke was at first accomodated (Malloch 

& Cain 1971a, Malloch & Benny 1973, Benny 

& Kimbrough 1980), together with Cleistothelebolus 

Malloch & Cain and Lasiobolidium 

Malloch & Cain, in Eoterfeziaceae G.F. Atk., a 

family of plectomycetous fungi with an uncertain 

position in Ascomycota, characterised by 

cleistothecial ascomata, globose, thin-walled 

and evanescent asci, one-celled, oblate or ellipsoidal 

ascospores, and an unknown anamorph 

(Kirk et al. 2001, Cannon & Kirk 2007). 

Orbicula and Lasiobolidium were later 

transferred (Jeng & Krug 1976) to tribe 

Theleboleae Korf of Pyronemataceae Corda 

(Pezizales J. Schröt.), which is the equivalent 

of the current Thelebolaceae Eckblad, where 

also Coprotiella Jeng & J.C. Krug, and most 

genera previously placed in Eoterfeziaceae, 

were accomodated. 

Currently Orbicula nests, with Lasiobolidium 

and Pseudombrophila Boud., in a well 

defined clade of Pyronemataceae (Hansen & 

Pfister 2006, Perry et al. 2007), a family mostly 

including apothecioid discomycetes, but also a 

minority of cleistothecial fungi evolved from 

an apothecial ancestor (Hansen et al. 2005). As 

its epithet means, Lasiobolidium orbiculoides 

is similar to Orbicula parietina, but differs 

from the latter in having a simpler, one-layered 

peridium, widespread, sometimes coiled hairs 

rather than hyphoid hairs restricted to the 

peridial base, and slightly larger ascospores on 

average (10–14 × 9–12 µm, Malloch & Benny 

1973, versus 7–13 × 8–10 µm, Hughes 1951), 

although a collection of O. parietina associated 

387

Figs 30–34 – Orbicula parietina. 30 Cleistothecioid ascomata. 31, 32, 33 Details of exoperidium 

(32 = Melzer). 34 Detail of endoperidium (Melzer). Bars 30 = 150 μm, 31, 32 = 30 μm, 33 = 15 μm, 

34 = 20 μm. 

with oat kernels has ascospores 12–16 × 10–14 

µm (Campbell et al. 1991). 

The psychrophilic (von Arx 1981, 

Campbell et al. 1991, Richardson 2004b) O. 

parietina has often been isolated from several 

kinds of dung (Marchal 1884b, Udagawa & 

Furuya 1972, Bokhary 1985, Richardson 

2004b, Moyne & Petit 2006), but also from 

decaying plants, bark and paper, straw, 

compost and similar substrates (Marchal 

1884b, Hughes 1951, Dennis 1981). 


Microascales – Microascaceae 

Microascaceae – An update. 

The Microascaceae are a monophyletic 

family (Berbee & Taylor 1992, Hausner et al. 

1993, Spatafora & Blackwell 1994, Lee & 

Hanlin 1999), characterised by cellular, dark, 

often immersed, perithecioid or cleistothecioid 

388 

ascomata, single or catenulate, globose to clavate, 

evanescent asci, absence of an interascal 

tissue, and small, one-celled, dextrinoid, pale to 

copper coloured ascospores, with one or two or 

no germ pores. The anamorph is aleurio-, annello-, 

arthro-conidial or absent (Luttrell 1951, 

Malloch 1970, Benny & Kimbrough 1980, Arx 

et al. 1988, Barr 1990, Cannon & Kirk 2007). 

They behave as saprophytes on soil, dung, and 

rotting plants, sometimes as agents of animal 

and human infections (de Hoog et al. 2000, 

Issakainen et al. 2003, Rainer & de Hoog 

2006), particular immunocompromised patiens 

(Baddley et al. 2000, Mohammedi et al. 2004). 

Benny & Kimbrough (1980) also erected the 

family Pithoascaceae, inclusive of Pithoascus 

Arx and Faurelina Locq.-Lin., very close to 

Microascaceae but with constantly fusoid to 

navicular spores, lacking germ pores, and 

arthroconidial or no anamorphs. Nowadays the 

Pithoascaceae are regarded as a later synonym


Figs 35–42 – Orbicula parietina (35, 36, 38, 40, 41, 42 Congo red). 35 Longitudinal section of 

ascoma. 36, 38 Paraphyses above immature asci. 37 Hyphoid hairs. 39, 40, 41, 42 Asci with 

ascospores in different stages. Bars 35 = 200 μm, 36, 37, 42 = 40 μm, 38, 39, 40, 41 = 20 μm. 

of Microascaceae (Cannon & Kirk 2007, 

Lumbsch & Huhndorf 2007, Kirk et al. 2008), 

Faurelina has been transferred in Chadefaudiellaceae 

Faurel & Schotter ex Benny & 

Kimbr., and Pithoascus is considered a later 

synonym of Microascus Zukal by Lumbsch & 

Huhndorf (2007), in agreement with Abbott et 

al. (2002), but an independent genus by Kirk et 

al. (2008), the latter following Arx (1973b, 

1975b, 1978), Arx et al. (1988), Eriksson & 

Hawksworth (1998), Barr (1990). 

The family Microascaceae includes the 

following theleomorphic genera: Canariomyces 

Arx (1984), Enterocarpus Locq.-Lin. 

(1977), Kernia Nieuwl.; Lophotrichus R.K. 

Benj. (1949), Microascus Zukal (1885), 

Petriella Curzi (1930b), Pithoascus Arx 

(1973b), Pseudallescheria Negr. & I. Fisch., 

Petriellopsis Gilgado et al. 

The genus Petriellopsis was recently 

erected (2007) to accommodate Pseudallescheria 

africana (Arx & G. Franz) Gilgado et 

al., which molecular studies (Issakainen et al. 

1999, 2003, Rainer & de Hoog 2006, Gilgado 

et al. 2007) proved not to nest with the other 

Pseudallescheria spp. 

I provide the following key to genera of 

Microascaceae, which must be regarded as 

supplementary to Arx (1978) and Arx et al. 

(1988): 

1) Ascospores asymmetrical, with inconspicuous 

or no germ pores. Ascomata 

smooth or with hyphal hairs at most. .... 2 

389

1*) Ascospores symmetrical or asymmetrical, 

with 1 or 2 germ pores. Ascomata 

smooth or with well differentiated hairs 

or setae. .................................................. 3 

2) Ascomata not ostiolate. Peridium of textura 

epidermoidea. Asci clavate. Ascospores 

broadly ellipsoidal. Anamorph 

Scedosporium and Graphium. ................. 

............................................. Petriellopsis 

2*) Ascomata non-ostiolate or with an 

ostiolate papilla. Peridium of textura 

angularis or epidermoidea. Asci broadly 

clavate to globose. Ascospores elongated, 

usually more than twice as long as broad, 

variable in shape, reniform to cylindrical. 

Anamorph Geniculosporium, Scedosporium, 

Scopulariopsis or absent. ............... 

............................................... Pithoascus 

3) Peridium of textura epidermoidea. Ascospores 

with 2 germ pores, 1 at each end. 

Anamorph Graphium, Scedosporium, 

Scopulariopsis. ...................................... 4 

3*) Peridium of textura angularis. Ascospores 

with 1 or 2 germ pores. .................... 5 

4) Ascomata globose, non-ostiolate, glabrous. 

Ascospores ellipsodal to broadly 

fusiform, symmetrical. ............................. 

..................................... Pseudallescheria 

4*) Ascomata often ampulliform with an 

ostiolate beak or papilla, surrounded with 

differentiated hairs or setae. Ascospores 

asymmetrical, usually lunate. .... Petriella 

5) Ascomata with a non-ostiolate papilla, 

surrounded with a tuft of dark hairs, and 

with an inner skein of dark hyphae (capillitium). 

Ascospores ovoidal or ellipsoiddal, 

symmetrical, with 1 or 2 germ pores. 

Anamorph unknown. ......... Enterocarpus 

5*) Capillitium absent .................................. 6 

6) Ascospores with 2 germ pores, rarely 

asymmetrical. Ascomata non-ostiolate or 

with a hairy, ostiolate beak or papilla. No 

anamorph. ............................. Lophotricus 

6*) Ascospores with 1 germ pore, symmetrical 

or asymmetrical. Anamorph usually 

present. ................................................... 7 

7) Ascomata with an ostiolate, hairy, long 

beaked or sometimes with an ostiolate 

390 

papilla. Ascospores asymmetrical, variable 

in shape, triangular to reniform or 

lunate. Anamorph Scopulariopsis, Wardomyces, 

Wardomycopsis. .... Microascus 

7*) Ascomata non-ostiolate. Ascospores 

usually symmetrical. .............................. 8 

8) Ascomata globose, glabrous. Ascospores 

comparatively dark, greyish brown, 

ellipsoidal to broadly fusiform, with 

attenuated ends. Anamorph Chrysonilialike. 

................................... Canariomyces 

8*) Ascomata globose or angular, then with 

hairy angles. Ascospores ellipsoidal or 

ovoidal or rarely asymmetrical (reniform), 

straw-coloured or pale brown to 

pale reddish brown. Anamorph as above 

or Graphium, or absent. ................ Kernia 

Kernia cauquensis Calviello, Rev. Mus. Arg. 

Cie. Nat. 5(12): 240, 1979. Figs 43–48 

Ascomata cleistothecioid, globose or subglobose, 

sometimes hemispherical with a flattened 

base, black shining, smooth, membranous, 

80–140 µm diam. Peridium two-layered, pseudoparenchymatous, 

the inner layer of pale, 

polygonal, comparatively thin-walled cells, 6– 

14 × 6–11 µm, the outer layer of dark brown, 

thick-walled, polygonal cells (textura angularis), 

3–10 µm diam. Exceptionally single or 

sparse hairs can be oberved, short, brown, 

thick-walled, septate, 2 µm diam., with a bulbous 

base, 3–3.5 µm diam. Interascal tissue 

absent. Asci irregularly disposed, evanescent, 

unitunicate, inoperculate, non-amyloid, (6–8spored, 

10–15 × 9–11 µm, sessile, globose or 

broadly ellipsoidal to broadly clavate. Ascospores 

conglobate, 5–6 × 3.5–4.5 µm, broadly 

ellipsoidal or ovoidal, (Q = 1.25–1.66; 

averageQ = 1.40), equilateral, hyaline and 

dextrinoid at first, with one to several droplets, 

thick-walled, becoming olive-pale brown, with 

an apical germ pore, usually with a de Bary 

bubble in aqueous media. Anamorph not 

observed. 


Montioni, 0 m a.s.l., dozens of gregarious or 

isolated specimens, superficial, but fully covered 

with a white aerial mycelium, on sheep 


18.03.2009, 306.3-Suvereto, CLSM 006.09; 

Vercelli, Alagna, Alpe Pianalunga, 2000 m


Figs 43–48 – Kernia cauquensis. 43 Cleistothecia on dung (white arrows). 44 Cleistothecium in 

water. 45 Asci with ascospores in different stages. 46 Detail of exoperidium. 47 Free and clustered 

ascospores inside the asci. 48 Immature asci. Bars 43 = 200 μm, 44, 46 = 50 μm, 45, 47 = 15 μm, 48 

= 20 μm. 

a.s.l., on dung of unidentified animal in a damp 

chamber culture, A. Bizzi, 27.08.2008, 071.2- 

Alagna, CLSM 006.09 bis. 

Notes – The latest extensive work on 

sordariaceous fungi without ascospore ejaculation 

(von Arx et al. 1988) recognised a group 

of species including K. cauquensis, K. ovata 

(C. Booth) Malloch & Cain, K. pachypleura 

Malloch & Cain, K. retardata Udagawa & T. 

Muroi, and K. setadispersa Locq.-Lin., characterrised 

by globose or subglobose ascomata and 

distinctly pigmented, ovoidal or ellipsoidal 

ascospores. 

Since then only one new spicies, K. peruviana 

Udagawa & Furuya, has been published, 

characterised by ascospores variable in shape, 

reniform, roughly triangular, sometimes ellip- 

soidal or ovoid (Udagawa & Furuya 1988), 

similar to those of K. hyppocrepida Malloch & 

Cain, but smaller (3–4.5 × 3–3.5 µm). 

C. cauquensis is also characterised by 

usually glabrous ascomata, ascospores with a 

single, apical germ pore, and a Scopulariopsis 

Bainier anamorph. It is similar to K. pachypleura, 

which differs, however, in having ascospores 

with two germ pores, a much thicker 

(37–46 µm) peridium (Malloch & Cain 1971b), 

and sometimes 4-spored asci (Delgado Avila et 

al. 2001b). The former was isolated only once 

from avutard droppings (Calviello 1979), while 

the latter would seem commoner, as it was 

found on elephant (Malloch & Cain 1971b), 

deer, monkey, parrot, tortoise, rabbit (Delgado 

Avila et al. 2001b) goat, buffalo, and cow 

391

(Chang & Wang 2008) dung. K. ovata, isolated 

once from seeds, differs from K. cauquensis in 

lacking an anamorph and having longer asci 

and ascospores (6–9 × 4.5–5.5 vs. 5–6.8 × 4– 

5.3 µm) with a subapical germ pore (Booth 

1964, Malloch & Cain 1973). 

K. setadispersa has an unknown anamorph; 

isolated from goat dung, it differs from 

K. cauquensis in having ascomata covered with 

10–30, dark, long hairs, and slightly narrower 

(3.5–4.5 µm) ascospores (Locquin-Linard 

1980). The teleomorphs of K. retardata, 

isolated from soil (Udagawa & Muroi 1981), 

rotten wood and skunk dung (Lumley et al. 

2000), have, in my opinion, morphological features 

fully matching K. cauquensis, including 

the pale coloured ascospores, but the colonies 

of the former, unlike the latter, slowly develop 

in common media, and K. retardata has a 

slightly different Scopulariopsis anamorph, 

e.g., somewhat wider conidia. 

I have called “affinis” cauquensis my 

collection of Kernia because unfortunately I 

could not isolate it in pure culture and study its 

possible anamorph. Besides I have preferred to 

call it “affinis” cauquensis rather than “affinis” 

retardata because I have exceptionally observed 

ascomata with sparse brown hairs, and 

ascomata with such feature (“ascomata glabrous, 

sometimes with sparse, short hairs) were 

described in the protologue of K. cauquensis 

(Calviello 1979). 

A key to the recognised species of Kernia 

Nieuwl. was provided by me (Doveri 2004a). 

Lophotrichus bartlettii (Massee & E.S. 

Salmon) Malloch & Cain, Can. J. Bot. 49: 866, 

1971. Figs 49–57 

≡ Magnusia bartlettii Massee & E.S. 

Salmon, Ann. Bot. 15: 333, 1901. 

≡ Kernia bartlettii (Massee & E.S. 

Salmon) R.K. Benj., Aliso 3: 344, 1956. 

= Lophotrichus brevirostratus L.M. 

Ames, Monogr. Chaetomiaceae: 52, 1963 (fide 

Malloch & Cain, 1971). 

= Kernia bifurcotricha A.S. Saxena & 

Mukerji, Trans. Brit. Mycol. Soc. 54: 146, 

1970 (fide Arx et al., 1988). 

Ascomata perithecioid, subglobose, 250– 

280 µm diam., blackish, semi-membranous, 

with an apical papilla surrounded by a tuft of 

dark hairs, and with paler sparse hairs. 

392 

Peridium some layers of very thick-walled, 

polygonal cells (textura angularis), 5–8 × 4–6 

µm, the outermost dark brown, the innermost 

paler and with thinner walls. Apical hairs 500– 

1000 µm long, 5–6 µm diam., up to 10 µm 

diam. at their base, smooth, thick-walled (about 

2 µm), dark greyish brown, densely septate, 

wavy, sometimes circinate. Sparse hairs paler 

and thinner, occasionally bifurcate. Asci soon 

vanishing, broadly clavate to ovoidal or saccate, 

usually with a short stalk, thin-walled, 8spored, 

unitunicate, non-amyloid, 20–24 × 12– 

18 µm. Ascospores 8–8.5 × 6–6.5 µm (Q = 

1.23–1.45; average Q = 1.32), irregularly biseriate 

to conglobate, dextrinoid in the early 

stages, often with a gaseous bubble, fairly 

thick-walled, hyaline at first, becoming yellowish, 

finally olive-brown, broadly fusiform or 

ellipsoidal, with apiculate ends, each with a 

germ pore. Anamorph not observed. 

Material examined – Italy, Livorno, Bibbona, 

Fattoria di Faltona, 50 m a.s.l., dozens of 

semi-immersed specimens on sheep dung in a 


295.3-Sassa, CLSM 007.07 bis. 

Notes – The genus Lophotrichus R.K. 

Benj. includes coprophilous species both with 

hairy ostiolate and glabrous non-ostiolate ascomata, 

a peridium of textura angularis, pale to 

reddish brown, ellipsoidal to broadly fusiform, 

sometimes asymmetrical ascospores with two 

germ pores, and no anamorph. 

Von Arx et al. (1988) recognised six 

Lophotrichus spp., most (L. ampullus R.K. 

Benj., L. geniculosporus Locq.-Lin., L. martinii 

R.K. Benj., L. plumbescens Morinaga et al.) 

distinguishable from L. bartlettii by having 

ascomata with a well differentiated beak 

(Benjamin 1949, von Arx et al. 1988), L. 

geniculisporus and L. plumbescens also by 

their smaller ascospores (respectively 6–8 × 4.5 

µm, Locquin-Linard 1986, 6–7.5 × 5–6 µm, 

von Arx et al. 1988, versus 8–12 × 5–7.5 µm, 

Massee & Salmon 1901, von Arx et al. 1988). 

L. macrosporus (Faurel & Locq.-Lin.) 

Arx et al., with its globose, non-ostiolate ascomata, 

often covered with some apical hyphal 

hairs, resembles L. bartlettii when the latter 

develops non-ostiolate instead of the usual 

papillate ascomata with well differentiated, 

brown hairs around the ostiole. L. macrosporus, 

however can be always distinguished by its


Figs 49–54 – Lophotrichus bartlettii. 49 Free or clustered ascospores inside evanescent asci. 50, 52 

Details of apical hairs. 51 Free ascospores. 53 Detail of exoperidium. 54 Three free ascospores and 

asci in different stages. Bars 49, 51 = 20 μm, 50, 54 = 30 μm, 52 = 60 μm, 53 = 15 μm. 

larger ascospores (13–14 × 9 µm., Locquin- 

Linard 1977). 

After von Arx et al. (1988), no new 

species of Lophotrichus was published, but a 

recombination, Lophotrichus fimeti (Arx et al.) 

Gilgado et al., was stated. Gilgado et al. (2007) 

validly published this name, although they incorrectly 

cited the basionym (art. 33.4, 33.5 

ICBN 2006), which is Petriellidium fimeti Arx 

et al. (in von Arx 1978), not Pseudallescheria 

fimeti (von Arx et al.) McGinnis et al (1982). 

The genus Petriellidium Malloch, however, 

must be regarded (McGinnis et al. 1982) as a 

later synonym of Pseudallescheria Negr. and I. 

Fisch. Gilgado et al. (2007) transferred 

Pseudallescheria fimeti to the genus Lophotrichus 

based on morphological features and 

phylogenetic analyses, the latter proving the 

genus Pseudallescheria to be polyphyletic 

(Issakainen et al. 1999, Rainer & de Hoog 

2006, Gilgado et al. 2007) and P. fimeti not to 

cluster with other Pseudallescheria spp. but to 

nest with Lophotrichus plumbescens. The main 

morphological features of L. fimeti are typical 

of a Lophotrichus species (see my discussion 

above) and contrast with Pseudallescheria, 

which has a peridium of textura epidermoidea 

and a Scedosporium Sacc. ex Castell & Chalm 

or Graphium Corda anamorph (McGinnis et al. 

1982). Both Graphium anamorph and chlamydospores 

were observed (Arx 1978) in the 

original strain (CBS 129.78) of Petriellidium 

fimeti, but this living culture was later proved 

(McGinnis et al. 1982) to be a mixture of P. 

fimeti and Pithoascus langeronii Arx, and to 

develop peridia of a textura angularis 

393

Figs 55–60 – Lophotrichus bartlettii. 55 Squashed perithecium with apical hairs. 56, 57 Ascospores 

– Enterocarpus grenotii 58 Squashed cleistothecium with apical hairs and capillitium (arrow). 59, 

60 Ascospores. Bars 55, 58 = 125 μm, 56, 57, 59, 60 = 20 μm. 

(not epidermoidea, as stated in the protologue) 

and no anamorph (Gilgado et al. 2007). 

L. fimeti has spherical, non-ostiolate 

ascomata, covered with sparse, brown, hyphoid 

hairs and ellipsoidal ascospores, 11–13 × 8–10 

µm (von Arx 1978), so it can be distinguished 

from the non-ostiolate forms of L. bartlettii by 

its somewhat larger ascospores, but it is hardly 

distinguishable from L. macrosporus which, in 

my opinion, has only somewhat smaller asci 

(30–40 × 25 versus 40–70 × 18–25 µm). 

Enterocarpus grenotii Locq.-Lin. was 

regarded as a later synonym of L. bartlettii 

(von Arx 1981, von Arx et al. 1988, Valldosera 

1991), possibly because both have papillate 

ascomata with a tuft of hairs, a peridium of 

textura angularis, ascospores very similar in 

shape and size with two germ pores, and no 

anamorph. I think, however, this synonymy 

394 

was stated incorrectly, as the study of L. 

bartlettii type culture was only wished (von 

Arx 1981), and I personally observed (Doveri 

et al. 1997) non-ostiolate ascomata and a 

capillitium in E. grenotii (Figs 58–65), which 

are typical features of Enterocarpus Locq.- 

Linard, recognised at the present as an 

independent genus (Lumbsch & Huhndorf 

2007, Kirk et al. 2008). Besides I do not think 

that the numerous mycologists who studied L. 

bartlettii (Ames 1963, Tornabene & Davis 

1966, Asad & Ahmad 1968, Seth 1971, Ahmed 

et al. 1971, Furuya & Udagawa 1973, Pathak & 

Agrawal 1974, Valldosera & Guarro 1984a, 

Valldosera 1991, Wang 1992, Horie & Li 

1997, Delgado Avila et al. 2001b), overlooked 

and did not describe the presence of a 

capillitium. Both species are coprophilous, but 

E. grenotii is very rare and it has been isolated


Figs 61–65 – Enterocarpus grenotii. 61 Detail of exoperidium. 62 Immature ascospores above 

exoperidial cells. 63 Hyphae of capillitium (white arrow). 64 The whole capillitium (arrow) beneath 

the bases of apical hairs. Portion of exoperidium on the right. 65 Basal and medium portion of 

apical hairs, and exoperidial cells. Bars 61 = 20 µm, 62 = 25 µm, 63 = 40 µm, 64, 65 = 60 µm. 

from fox (Locquin-Linard 1977) and horse 

(Doveri et al. 1997) dung, while L. bartlettii is 

quite common and found on various kinds of 

dung (Furuya & Udagawa 1973, Horie & Li 

1997) 

Pithoascus intermedius (C.W. Emmons & 

B.O. Dodge) Arx, Proc. Konink. Ned. Akad. 

Wetensch., ser. C, Biol. Med. Sci. 76: 292, 

1973. Figs 66–71 

≡ Microascus intermedius C.W. Emmons 

& B.O. Dodge, Mycologia 23: 324, 1931. 

395

Ascomata 120–200 µm diam., often 

perithecioid, with a minute, apical papilla, or 

cleistothecioid, spherical to subglobose, black, 

opaque, subcoriaceous, seemingly smooth, but 

covered all over with sparse, very short, 

hyphoid hairs. Peridium pseudoparenchymatous, 

dark brown, layered, a textura angularis 

of thick-walled, polygonal cells, 5–11 × 3–7 

µm, the innermost cells paler and slightly 

larger. Hyphoid hairs subhyaline or brown, 

septate, 2–3 µm diam., Asci 8-spored, irregularly 

disposed, non-amyloid, inoperculate, 

ephemeral, non-fasciculate, broadly ellipsoidal 

or clavate, or doliiform, even subglobose, 11– 

15 × 7–9 µm. Interascal tissue absent. Ascospores 

strongly dextrinoid in the early stages, 

conglobate inside the ascus, 5–6 × 2–2.5 µm, 

cymbiform (navicular) in frontal view, subtriangular 

(plano-convex), sometimes subreniform 

(slightly concavo-convex) in side view, 

with slightly pointed ends, thin-walled, hyaline 

at first, pale yellow when mature, lacking oil 

drops, de Bary bubbles and germ pores. Anamorph 

not observed. 

Material examined – Italy, Pisa, Volterra, 

Villamagna, podere Vallicella, 200 m a.s.l., 

more than fifty superficial or usually immersed 

specimens on pig dung in a damp chamber 

culture, usually crowded, tending to form 

crusts, covered with a white, aerial mycelium, 

F. Doveri, 13.4.2009, 285.3-Lajatico, CLSM 

008.09. 

Notes – In my key to genera of Microascaceae, 

I have mentioned the main differentiating 

features between Microascus and 

Pithoascus, indirectly recognising their independence 

at genus rank, as originally stated by 

von Arx (1973b, 1975b, 1978) and followed by 

others (Valmaseda et al. 1987, Arx et al. 1988, 

Kirk et al. 2008). In the original conception 

Pithoascus has non-ostiolate or small papillate 

ascomata, narrower and paler spores lacking 

germ pores, no conidial state, and slower growing 

colonies. 

The independence of Pithoascus from 

Microascus was later questioned, when some 

taxa were noticed to possess ascoma and spore 

features typical of the former, a conidial state 

and growth rates of the latter (Malloch & 

Hubart 1987), or when the association between 

M. intermedius and a Scopulariopsis anamorph 

was proved (Roberts 1975, Domsch et al. 

396 

1980), and a reduced anamorphic stage was 

observed in M. intermedius (Abbott et al. 

2002). 

Lumbsch & Huhndorf (2007) regard 

Pithoascus as a later synonym of Microascus, 

although molecular studies (Issakainen et al. 

2003) have proved Pithoascus intermedius and 

P. nidicola (Massee & E.S. Salmon) Arx to 

cluster in a definite subclade inside the Microascus 

s. l. main clade, leaving the matter open 

whether to split or redefine the genus Microascus. 

If conceived in a broad sense, i.e. inclusive 

of Pithoascus, Microascus has papillate to 

long-beaked, rarely cleistothecioid ascomata 

with a pseudoparenchymatous peridium of 

angular (textura angularis) or exceptionally 

epidermoid (textura epidermoidea) cells, and 

asymmetrical, naviculate, (sub) triangular (plano-convex), 

reniform (concavo-convex), or 

heart-shaped ascospores with one or no germ 

pore. The anamorph is often Scopulariopsis 

Bainier (Zukal 1885, Curzi 1930a, 1931, 

Barron et al. 1961, Udagawa 1962, von Arx 

1975b, Abbott et al. 1998, Abbott & Sigler 

2001). Species of Microascus are usually saprobic 

on soil (Mathur & Thirumalachar 1962, 

Sage et al. 1995, Mouchacca 1999) dung(Zukal 

1885, Massee & Salmon 1901, 1902, Tóth 

1965, Dal Vesco et al. 1967, Saxena & Mukerji 

1973, Piontelli et al. 1981, Valldosera 1991), 

and rotting wood (Lumley et al. 2000), 

sometimes pathogenic for plants (Morton & 

Smith 1963), humans, and animals (Jones 

1936, Barron et al. 1961, Baddley et al. 2000), 

insects included (Skou 1973). They have a 

cellulolytic (Verona et al. 1967) and proteolytic 

(Malloch & Hubart 1987) activity, which explains 

the high frequency of occurrence on 

dung (Lumley et al. 2000). 

P. intermedius is characterised by usually 

papillate and ostiolate ascomata, a perium of 

textura angularis, and small, navicular ascospores, 

(Emmons & Dodge 1931, von Arx 

1973b, von Arx et al. 1988). It has often been 

compared (Barron et al. 1961, Udagawa 1962, 

Morton & Smith 1963, von Arx 1973b, von 

Arx et al. 1988, Abbott et al. 2002) with P. 

nidicola and P. stoveri Arx, and distinguished 

from both by its shorter (4.5–6 versus 6–8 µm) 

ascospores, usually plano-convex rather than 

concavo-convex in lateral view, from P. stoveri


Figs 66–71 – Pithoascus intermedius. 66 Semi-immersed ascomata. 67 Detail of exoperidium. 68, 

71 Free ascospores and asci in different stages. 69, 70 Strongly dextrinoid ascospores, free or 

clustered inside asci. Bars 66 = 100 μm, 67, 69, 70 = 15 μm, 68, 71 = 20 μm. 

also by usually ostiolate rather than nonostiolate 

ascomata, from P. nidicola by the 

ability to develop a reduced anamorph. 

P. intermedius has been isolated from 

rotting plants (Emmons & Dodge 1931, Barron 

et al. 1961, von Arx 1973b), soil (Emmons in 

Barron et al. 1961), rarely from dung (Barron 

et al. 1961). 

397


Sordariales – Chaetomiaceae 

Chaetomidium fimeti (Fuckel) Sacc., Syll. 

Fung. 1: 39, 1882. Figs 72–80 

≡ Chetomium fimeti Fuckel, Jahrb. Ver. 

Naturk. Herzog. Nassau 15: 64, 1860. 

≡ Thielavia fimeti (Fuckel) Malloch & 

Cain, Mycologia 65: 1064, 1973. 

≡ Chaetomidium chlorochaetum Speg., 

An. Mus. Nac. Buenos Aires, ser. III, 12: 336, 

1909 (fide Calviello, 1978). 

Cleistothecia globose, 300–350 µm 

diam., dark brown, membranous, wholly 

covered with comparatively short, greenish 

hairs, and a whorl of long, blackish, basal hairs. 

Peridium layered: 1) endostratum a textura 

angularis of hyaline, thin-walled, polygonal 

cells; 2) exostratum some layers of dark brown, 

thick-walled, polygonal cells (textura angularis), 

5–8 µm diam., disposed in a petaloid 

pattern around the basal hairs. Greenish hairs 

strongly encrusted, thin-walled, flexuous, 

densely septate, tapering upwards, 2.5–4.5 µm 

diam., up to 350 µm long. Blackish hairs 

flexuous, smooth, septate, sometimes narrowing 

at the septa, with pointed tips and 

elarged, sometimes bulbous bases, thick-walled 

(1 µm or more), 4.5–6 µm diam., up to 1500 

µm long. Paraphyses not observed. Asci 

fasciculate, clavate, 8-spored, long-stalked, 65– 

75 × 17–20 µm. Ascospores irregularly 

biseriate, thick-walled, limoniform, sometimes 

subglobose in frontal view, bilaterally flattened 

so ellipsoidal in side view, (11–) 12–14 (–16) × 

9–11 (–11.5) × 7.5–8 µm (Q = 1.09–1.55; 

average Q = 1.28), hyaline at first, pale brown 

at maturity, biapiculate, sometimes deformed, 

with one apical germ pore. Conidia lacking. 

Material examined – Italy, Pisa, Chiecinella, 

70 m a.s.l., about twenty superficial, 

gregarious specimens, late maturing (at 32–35 

days) on horse dung in a damp chamber 

culture, F. Doveri, 13.4.09, 274.3-Montopoli in 

Val d’Arno, CLSM 023.04 bis. 

Notes – The genus Thielavia Zopf, inclusive 

of Chaetomidium, was monographed by 

Malloch & Cain (1973), who stated that the 

presence of differentiated hairs in Chaetomidium 

and glabrous cleistothecia in Thielavia 

are not a difference enough to warrant their 

independence at genus rank, as species with 

intermediate forms are known. 

398 

Von Arx (1975a) did not attach the same 

importance to the presence of differentiated 

hairs and regarded Thielavia as having 

glabrous, setose or tomentose peridia with a 

textura epidermoidea, and fusiform, clavate, 

obovate or ellipsoidal ascospores, and Chaetomidium, 

the non-ostiolate counterpart of Chaetomium, 

as having hairy or setose, pseudoparenchymatous 

cleistothecia, and limoniform, 

laterally flattened ascospores. 

Von Arx et al. (1988) kept the same 

conception of Thielavia and Chaetomidium, but 

assigned both limoniform and broadly fusiform 

ascospores to the latter. Their key to Chaetomidium 

species is based on ascospore shape 

and peridial frame (textura angularis or textura 

epidermoidea). 

The main morphological features of the 

genus Chaetomidium were described by Doveri 

(2004a), who also provided a key to the species 

with a cephalothecoid peridium. 

Two new Chaetomidium spp. were later 

published, C. galaicum Stchigel & Guarro and 

C. triangulare Stchigel & Guarro, isolated 

from granite rock and soil. The former is similar 

to C. khodense Cano et al. (1993) and to C. 

megasporum Doveri et al. (1998a) on account 

of its cephalothecoid peridium and ellipsoidfusiform 

ascospores, but differs from C. khodense 

in having less hairy ascomata and longer 

ascospores (14–19 × 5–7 µm vs. 11–13 × 6.5–7 

µm, Stchigel et al. 2004), from C. megasporum 

in having smooth hairs and smaller ascospores 

(19–21.5 × 11–13 µm in C. megasporum); C. 

triangulare belongs to the group lacking a 

cephalothecoid peridium and can be easily 

distinguished from all the others by its glabrous 

ascomata and triangular ascospores. 

The latest key to Chaetomidium spp. was 

provided by Stchigel et al. (2004). Studies of 

developmental morphology (Greif & Currah 

2007) showed that Aporothielavia leptoderma 

(C. Booth) Malloch & Cain has a cephalothecoid 

peridium and dehiscence very similar to 

Chaetomidium arxii Benny (1980), and it must 

be recombined in Chaetomidium leptoderma 

(C. Booth) Greif & Currah. It differs from C. 

arxii in having sparse, shorter, and wavy peridial 

hairs, and narrower ascospores (14.5–16.5 

× 5.5–7 µm vs. 10.5–19 × 7.5–12.5 µm, 

(Greif& Currah 2007). It also differs from all 

the other Chaetomidium spp. in having knob-


Figs 72–77 – Chaetomidium fimeti. 72, 73 Cleistothecium. 74 Petaloid arrangement of exoperidial 

cells around the hair bases (white arrow). 75 Detail of exoperidium. 76 Lower portion of blackish, 

basal hairs (red arrow), and greenish hairs (black arrow). 77 Detail of exoperidium with hairs. Bars 

72 = 400 μm, 73 = 200 μm, 74 = 30 μm, 75 = 40 μm, 76 = 20 μm, 77 = 70 μm. 

like protrusions on the peridial cells. 

Twelve Chaetomidium spp. have been 

recognised so far, but phylogenetic analyses 

(Greif et al. 2009) prove that the genus is 

polyphyletic and must be restricted to C. fimeti 

and C. subfimeti, being the other species scattered 

through the Chaetomiaceae G. Winter and 

Lasiosphaeriaceae Nannf. 

C. fimeti, the type of Chaetomidium, is 

characterised by limoniform, laterally flattened 

ascospores, and a non–cephalothecoid peridium. 

Unlike others (Whiteside 1962), I have 

not been able to observe filaments of sterile 

cells in centrum of my collection. Chaetomidium 

subfimeti Seth has all morphological 

features similar to C. fimeti, but smaller ascospores 

(8–10 × 6–7 µm, Seth 1967). 

C. fimeti has been isolated from horse 

(Fuckel 1860), rabbit (Munk 1957, Seth 1968), 

cock (Munk 1957), and unspecified dung 

(Malloch & Cain 1973), and from manure 

(Winter 1887). It must be regarded as facultatively 

coprophilous, as it was also recorded 

from nests of free-living birds (Hubálek 1974), 

decaying plants and wood (Winter 1887, 

Bainier 1909, Spegazzini 1909, Calviello 

1978), seeds, and soil (Malloch & Cain 1973). 

Chaetomium murorum Corda, Icon. Fung. 1: 

24, 1837. Figs 81–91 

399

Figs 78–80 – Chaetomidium fimeti. 78, 79 Limoniform, bilaterally flattened ascospores in different 

stages. 80 Immature asci. Bars 78 = 15 μm, 79, 80 = 20 μm. 

= Chaetomium tomentosum Preuss, 

Linnaea 24: 99, 1851. 

= Bolacotricha grisea Berk. & Broome, 

Ann. Mag. nat. Hist., Ser. 2: 7, 1851. 

= Chaetomium griseum (Berk. & 

Broome) Cooke, Grevillea 1: 175, 1873. 

= Chaetomium humanum P. Karst., Not. 

Fauna Flora Fenn. 8: 193, 1882. 

= Chaetomium macrosporum Sacc. & 

Penz., Michelia 2: 591, 1882. 

= Chaetomium contortum Bainier, Bull. 

Soc. Mycol. Fr. 25: 205, 1910. 

= Chaetomium pampaninii Cif., Bull. 

Soc. Bot. Ital. 1923: 98, 1923. 

= Chaetomium elongatum Czerepan., 

Notul. syst. Sect. cryptog. Inst. bot. Acad. Sci. 

U.S.S.R. 15: 80, 1962. 

400 

Colonies on Emerson YpSs (Yeast 

protein Soluble starch Agar) attaining 70 mm 

diam. at 12 d, with a white aerial mycelium in 

vague concentric rings, without exudates. 

Reverse olivaceous. 

Perithecia 170–200 µm diam., globose or 

rarely subglobose, exceptionally ovoidal, dark 

greyish brown, membranous, wholly hairy, 

with an ostiole variable in size, sometimes 

scarcely visible. Peridium two-layered: 1) 

endostratum a textura angularis of pale brown 

polygonal cells, somewhat larger, 8–13 × 7–10 

µm, and with thinner walls than the exoperidial 

cells; 2) exostratum darker brown, with a complex 

frame: a textura epidermoidea, with hyphae 

2–4 µm diam., interspaced with a textura 

angularis (variable in percentage) of polygonal 

cells, 5–9 × 4–8 µm, sometimes with a textura


Figs 81–86 – Chaetomium murorum. 81 Petaloid arrangement of peridial hyphae around the hair 

bases (arrow). 82 Basal portion of apical hairs. 83 Peridial detail of textura angularis. 84 Peridial 

detail of textura epidermoidea. 85 Circinate apex of apical hair. 86 Detail of apical hairs. Bars 81, 

84, 85, 86 = 20 μm, 82 = 12 μm, 83 = 15 μm. 

cephalothecoidea (petaloid) around the hairs. 

Basal hairs hyphoid, 2.5–3.5 µm diam., dense, 

thin-walled, pale brown, flexuous, septate, 

smooth. Lateral hairs 100–300 × 2–4 µm, 

thick-walled, rigid, straight in the basal portion, 

flexuous above, tapering upwards, smooth to 

encrusted, pale brown, paler at their tips, septate, 

with an enlarged, often bulbous base and a 

slightly pointed apex. Apical hairs 450–1000 × 

5–6 µm, thick-walled, sometimes smooth, but 

also covered with a refractive crystallinematerial, 

poly-septate, brown, but paler and 

with a scarce number of septa in the apical 

portion, straight at the base, flexuous above 

and often open-circinate at their blunt or slight- 

ly pointed apex. Basal cell enlarged, often bulbous, 

10–15 µm long, 8–11 µm wide, often 

with germs of roots. Paler apical hairs present 

in different percentages, sometimes absent, 

thinner-walled, septate, wavy, 3–4 µm diam., 

often more encrusted than the darker hairs, but 

becoming smooth owing to the easily detachable 

refractive encrustations. Paraphyses 

ephemeral, cylindrical, somewhat narrowing at 

the septa, 3–4 µm diam. Asci unitunicate, fasciculate, 

clavate, 8-spored, 50–60 × 15–17 µm, 

fairly long-stalked. Ascospores irregularly biseriate 

or conglobate, 13–16 (–16.5) × (7–) 

7.5–9 (–10) µm, symmetrical, ellipsoidal to 

narrowly elipsoidal (Q = 1.55–2.00; average 

401

Figs 87–91 – Chaetomium murorum. 87, 88, 90 Ascospores. 89 Lateral hairs. 91 Immature asci. 

Bars 87 = 20 μm, 88, 90, 91 = 25 μm, 89 = 40 μm. 

Q = 1.73), with attenuated ends and one apical 

germ pore, sometimes with one slightly apiculate 

pole opposite to the germ pore, hyaline 

at first, becoming brown, often collapsing to 

form a longitudinal furrow. Anamorph absent. 

Material examined – Italia, Vicenza, 

Villaga-Monte Tondo, 250 m a.s.l., about ten 

gregarious, superficial specimens, on hedgehog 

dung in a damp chamber culture, A. Bizzi, 

19.8.07, 125.2–Longare, CLSM 010.08; Asti, 

Agliano Terme, 250 m a.s.l., on bat dung in a 

damp chamber culture, M. Filippa, 20.6.05, 

193.1-Canelli, CLSM 010.08 bis; Vicenza, 

Lumignano, Monte Brosimo, 100 m a.s.l., on 

dormouse dung in a damp chamber culture, A. 

Bizzi, 24.11.08, 125.2-Longare, CLSM 010.08 

ter; Livorno, Montioni, 0 m a.s.l., on rabbit 


18.3.09, 306.3-Suvereto, CLSM 010.08 quater; 

402 

Livorno, Bibbona, Fattoria di Faltona, 50 m 

a.s.l., on sheep dung in a damp chamber 

culture, F. Doveri, 18.3.09, 295.3-Sassa, 

CLSM 010.08 penta; Pisa, Orciatico, 50 m 

a.s.l., on sheep dung in a damp chamber 

culture, F. Doveri, 13.4.09, 285.3-Lajatico, 

CLSM 010.08 esa; Pisa, Volterra, Villamagna, 

Podere Vallicella, 200 m a.s.l., on cattle dung 

in a damp chamber culture, F. Doveri, 13.4.09, 

285.3-Lajatico, CLSM 010.08 epta; on sheep 

dung in a damp chamber culture, CLSM 

010.08 octa; Vicenza, Tonezza del Cimone, 

Altopiano dei Fiorentini, 1450 m a.s.l., on hare 

dung in a damp chamber culture, A. Bizzi, 

13.7.08, 082.3-Arsiero, CLSM 010.08 ena; 

Vicenza, Solagna, Campeggia, 980 m a.s.l., on 

mouflon dung in a damp chamber culture, A. 

Bizzi, 2.6.09, 083.3-Monte Grappa, CLSM 

010.08 deca; Vicenza, Longare, Lumignano,

Monte Brosimo, 350 m a.s.l., on snail dung in a 

damp chamber culture, A. Bizzi, 11.6.09, 

125.2-Longare, CLSM 010.08-XI; on 

dormouse dung in a damp chamber culture, 

22.6.09, CLSM 010.08-XII; Novara, Druogno, 

Santa Maria, 850 m a.s.l., on sheep dung in a 


116.1-Bellinzago, CLSM 010.08-XIII. 

Chaetomium ancistrocladum Udagawa & 

Cain, Can. J. Bot. 47: 1943, 1969. Figs 92–100 

Perithecia subglobose to broadly ellipsoidal, 

280–350 × 250–280 µm, dark greyish 

brown, membranous, wholly hairy, rounded or 

sometimes slightly pointed at their base, with a 

scarcely visible ostiole. Peridium two layered: 

endostratum a textura angularis of pale brown, 

thin-walled, polygonal cells, 10–20 × 8–15 µm; 

exostratum a textura cephalothecoidea of 

wavy, dark brown, thick-walled, short hyphae, 

2–4 µm diam., radially arranged around the 

hair bases (petaloid arrangement). Basal hairs 

hyphoid, 2–4 µm diam., thin-walled, pale 

brown to almost hyaline, wavy, septate, 

smooth. Lateral hairs intergrading to the basal 

ones, thick-walled, rigid, usually straight or 

sometimes slightly wavy, tapering upwards, 

smooth, brown, paler at their tips, septate, with 

an enlarged base, somewhat narrower than the 

apical hairs. Apical hairs 4–7 µm diam., up to 

600 µm long, parallel, thick-walled (up to 1 

µm), usually smooth in the upper portion, 

finely encrusted towards the base, poly-septate, 

greyish brown, paler at the apex, straight or 

slightly curved at maturity, usually open circinate 

at their blunt or slightly pointed apex, enlarged 

at the base up to 10 µm. Paraphyses 

ephemeral, thin-walled, septate, 4–8 µm diam., 

slightly enlarged at their tips. Asci unitunicate, 

fasciculate, clavate, 8-spored, 80–110 × 18–20 

µm, long-stalked. Ascospores biseriate in the 

upper portion of ascus, uniseriate in the lower, 

hyaline at first, becoming greyish brown, (15–) 

15.5–17.5 (–19) × (8.5–) 9–10.5 (–11) µm, 

ellipsoidal to narrowly ellipsoidal (Q = 1.45– 

2.05; average Q = 1.73), sometimes slightly 

inaequilateral, with attenuated, often apiculate 

to subumbonate ends, with one apical, 

sometimes prominent, large (about 1 µm diam.) 

germ pore and a hyaline small spot at the 

opposite end. Anamorph not detected. 


Material examined – Italy, Vicenza, 

Lumignano, Monte Brosimo, 100 m a.s.l., on 

dormouse dung in a damp chamber culture, A. 

Bizzi, 24.11.08, 125.2-Longare, CLSM 009.08; 

22.6.09, CLSM 009.08 bis. 

Notes – C. ancistrocladum is close to C. 

murorum and its relatives, i.e. to the group of 

species characterised by comparatively large 

ascomata (> 150 µm diam.), clavate asci, nondextrinoid, 

medium-large (11–20 µm long), 

ellipsoidal ascospores, with attenuated ends 

and one apical germ pore. I have called C. 

ancistrocladum my collection because its 

features fully match those described in the 

protologue (Udagawa & Cain 1969) except for 

the absence of a second kind of short, coiled 

hairs around the ostiole, concealed by the spore 

mass at maturity, not observed by me even in 

immature specimens, when spores are not 

formed or not extruded yet. The peridial frame 

of C. ancistrocladum was not described in the 

protologue, but von Arx et al. (1986) affirmed 

“it was described as cephalothecoid” and considered 

this feature as distinctive inside the 

murorum-group. I cannot know to whom they 

assigned this description, because I have not 

found any record of this species but that of the 

original diagnosis. 

C. piluliferum J. Daniels differs from 

other species of the group (C. murorum, C. ancistrocladum, 

C. circinatum Chivers) particularly 

in having abundant rather than scarce or 

absent botryose aleurioconidia (von Arx et al. 

1986). Besides it differs from C. ancistrocladum 

in having a variable peridial frame, ranging 

from intricata to epidermoidea, sometimes 

angularis or cephalothecoidea at intervals, 

flexuous to circinate and somewhat narrower 

apical hairs (Daniels 1961, Mazzucchetti 1965, 

Seth 1970, Arx et al. 1986, Bell 2005), shorter 

asci (up to 60 µm long), and slightly smaller 

ascospores on average (13–)16 (–17) × 8–10 

µm; 7–8 µm broad in one collection from 

rotting wood (Lumley et al. 2000). 

C. murorum resembles C. piluliferum in 

most respects, except for lacking a botryose 

anamorph (von Arx et al. 1986), but I can 

recognise additional differences deduced from 

literature, i.e. more flexuous apical hairs 

(Winter 1887, Palliser 1910, Stratton 1921, 

Greathouse & Ames 1945, Munk 1957, Seth 

403

Figs 92–95 – Chaetomium ancistrocladum. 92 Ascomata on dung (arrows). 93 Circinate tip of 

apical hair. 94 Apical hairs. 95 Bases of apical hairs. Bars 92 = 600 μm, 93 = 20 μm, 94 = 100 μm, 

95 = 40 μm. 

1968, 1984, Ahmad & Sultana 1973, 

Valldosera & Guarro 1984b, Arx et al. 1986, 

Bell 2005) and usually ascospores with three 

darker longitudinal bands (Arx et al. 1986) in 

C. murorum. These features are also useful to 

distinguish C. murorum from C. ancistrocladum. 

C. circinatum differs from C. ancistrocladum 

in having a peridium with a textura 

intricata (Millner 1975), flexuous apical hairs 

with closed circinate, 2–3 times recurved tips, 

shorter asci, and somewhat smaller spores on 

average (Skolko & Groves 1953, Ames 1963, 

404 

Seth 1970, Ahmed et al. 1971, Lorenzo 1993). 

Mine is the second record worldwide of C. 

ancistrocladum, coming from dung like the 

original record and most collections of C. 

piluliferum (Marchal 1885, Caretta et al. 1994, 

Caretta & Piontelli 1996, Bell 2005), C. circinatum 

(Ahmed et al. 1971, Millner 1975, 

Lorenzo 1993) and C. murorum (Karsten 1888, 

Massee & Salmon 1901, 1902, Bainier 1909, 

Palliser 1910, Stratton 1921, Ames 1963, Tóth 

1963, 1965, 1967, Mirza & Nasir 1968, Seth 

1968, 1970, Calviello 1971, Ahmad & 

Sultana1973, Hubálek 1974, Matsushima 1975,


Figs 96–100 – Chaetomium ancistrocladum. 96, 100 Ascospores. 97 Cephalothecoid peridium. 98 

Petaloid plates of textura cephalothecoidea. 99 Immature asci. Bars 96 = 15 μm, 97 = 200 μm, 98 = 

20 μm, 99 = 50 μm, 100 = 25 μm. 

Mill ner 1975, Carter & Khan 1982, Barrasa 

1985, Bokhary & Parvez 1986, Valldosera 

1991, Eriksson 1992, Caretta et al. 1994, 1998, 

Caretta & Piontelli 1996, Beyer 1999, Richard 

son 2004, Elshafie 2005, Piontelli et al. 2006). 

A brief update on the genus Chaetomium 

I previously provided (Doveri, 2008a) an 

update on Chaetomium Kunze, describing 

some coprophilous species new to Italy, and 

placing them in key. In the same work I 

provided an updated version of von Arx et al. 

(1986) key to Chaetomium species, in which I 

inserted all new taxa described after their 

monograph on the genus. 

Since then only one new species, C. 

heterothallicum Mahoney, has been described 

(in preparation), isolated from river otter dung 

and characterised by a Scopulariopsis-like 

anamorph, heterothallism, and absolute need of 

crustacean chitin for developing in axenic 

culture. 


Xylariales – Xylariaceae 

Hypocopra equorum (Fuckel) G. Winter in 

Rabenhorst, Rabenh. Krypt.-Fl. 1 (2): 178, 

1887. Figs 101–118 

Hypoxilon equorum Fuckel, Fungi 

Rhenani: 1058, 1864. 

405

Figs 101–104 – Hypocopra equorum. 101, 102, 103, 104 Immersed perithecia with emerging necks 

surrounded by clypeate stromata (endostromata = white arrows, ectostromata = red arrows), 

embedded within a hyphal mat (blue arrows). Bars 101, 104 = 1 mm, 102, 103 = 2 mm. 

Figs 105–108 – Hypocopra equorum. 105 Perithecial neck. 106 Detail of the hyphal mat. 107 

Upper portion of a perithecial neck. 108 Detail of stroma. Bars 105 = 70 μm, 106, 108 = 20 μm, 107 

= 15 μm. 

406


Figs 109–112 – Hypocopra equorum. 109 (Melzer) Apical portion of asci with uniseriate 

ascospores (germ slit = white arrow). Early amyloid reaction of the apical apparatus (red arrow). 

110 (Melzer) Dextrinoid change of the apical apparatus (blue arrow). 111 Paraphyses. 112 Apical 

portion of asci (methyl blue). Bars 109, 112 = 18 μm, 110, 111 = 20 μm. 

Coprolepa equorum Fuckel, Jb. Nassau 

Ver. Naturk. 23–24: 240, 1869. 

Sordaria equorum G. Winter, Abhand. 

Naturf. Gesellsch. Halle 13 (1): 13, 1873. 

Stromata embedded within a hyphal mat, 

clypeate to widely spreading on the substrate, 

thick, suberose, dark grey outside, strawcoloured 

inside, often revolute and crown- or 

collar-shaped around the ostiole, containing a 

single or usually several perithecia, made up of 

dark brown (the innermost much paler), thickwalled, 

polygonal cells, 4–7 µm diam. Hyphal 

mat whitish at first, becoming greyish brown, 

dense, loosening with age, formed of brown, 

thick-walled, densely septate, branched and 

anastomosed hyphae, 1.5–3.5 µm diam., with 

short ends blunt at their tips. Perithecia 

subglobose, 800–900 × 650–750 µm, membranous, 

smooth, pale yellow, with a papillate, 

subcylindrical, coriaceous, somewhat rough, 

blackish neck, wider than high, 100–120 × 

200–220 µm. Peridium seemingly two-layered, 

with an exostratum typically formed of pale 

pigmented, septate, elongated, thin-walled 

cells, 2–5 µm diam., arranged in compressed 

rows, and an endostratum of pale, thin-walled, 

cylindrical to polygonal cells, 7–15 µm diam. 

Exoperidial cells scarcely observable at the 

neck, as covered by a carbonaceous material, 

cylindrical towards the ostiole, 3.5–5 µm 

407

Figs 113–118 – Hypocopra equorum. 113, 114 Ascospores surrounded by a gelatinous perisporium 

(arrows). 115, 116, 117, 118 Detail of asci with uniseriate ascospores in different stages (germ slit = 

arrow). Bars 113, 115 = 20 μm, 114, 116, 117 = 30 μm, 118 = 15 μm. 

diam., septate, with an enlarged, papillate end. 

Paraphyses simplex or branched, exceeding the 

asci, cylindric, septate, 9–12 µm diam. at the 

base, tapering upwards, containing many hyaline 

vacuoles. Asci 8-spored, 200–260 × 20–25 

µm, cylindric, roundish or slightly pointed at 

the apex, short-stalked. Apical apparatus cylindrical, 

7–8 µm diam., weakly amyloid, pale 

blue staining at first, soon becoming pale brickred. 

Ascospores surrounded by a broad gelatinous 

sheath, obliquely to vertically uniseriate, 

(22–)25–28.5(–31) × 12–13 µm, narrowly 

ellipsoidal to subfusiform in frontal view, 

slightly to quite asymmetrical (flattened at one 

side) in lateral view, thick-walled, smooth, 

hyaline at first, becoming yellowish, finally 

dark brown, one-celled as lacking a basal 

apiculum. Each spore with a longitudinal, 

eccentric, narrow germ slit, 12–15 µm long. 

Material examined – Italy, Trento, Tres, 

Monte Corno, 1600 m a.s.l., more than one 

408 

hundred, gregarious, almost fully immersed 

(except for the neck, which protrudes above the 

stroma) specimens, on horse dung, A. Bizzi, 

4.10.08, 043.4-Cles, CLSM 002.09. 

Notes – H. equorum is characterised by 

dark, widespread stromata immersed in a 

hyphal mat, one-celled, asymmetrical, medium 

size ascospores, with short germ slits, and 

weakly amyloid to dextrinoid asci. It differs 

from H. leporina (Niessl ex Rehm) J.C. Krug 

& N. Lundq. ex Doveri (= H. equorum f. 

leporina Niesll ex Rehm) in having one-celled, 

less flattened, and larger ascospores [in H. 

leporina 20–24 × 8–10 µm (Lundqvist in 

Hansen & Knudsen 2000); 17–19 × 7.5–8.5 µm 

(Doveri 2004a)], asci with a wider apical 

cylinder, and growth on equine (Winter 1873, 

1887, Phillips & Plowright 1876, Griffiths 

1901, Lundqvist 1960) rather than on leporid 

dung. According to Lundqvist (pers. comm.) 

“it would be surprising if H. leporina could be

shown to grow on any other substrate than hare 

dung or possibly leporid dung, as well as H. 

equorum on anything but horse dung”. One 

collection of H. equorum from rabbit dung has 

been, however, described by Breton (1965), 

another recorded by Richardson (2008b) from 

hare. Other Hypocopra spp. are characterised, 

like H. equorum, by perithecia embedded 

within a hyphal mat, but most (H. anomala J.C. 

Krug & Cain, H. punicea J.C. Krug & Cain, H. 

rostrata Griffiths) can be easily distinguished 

from the latter by lacking a stroma and having 

a constantly amyloid apical apparatus. H. rostrata 

has also larger, equilateral ascospores 

(27–32 × 13–18 µm, Griffiths 1901), H. anomala 

and H. punicea smaller ascospores (18– 

23 × 9–10 µm, Krug & Cain 1974). 

H. chionopsis J.C. Krug & N. Lundq. 

(nom. inv.) is very close to H. equorum owing 

to the presence of a hyphal mat, a dark stroma 

containing one to several perithecia, a dextrinoid 

apical apparatus, and ascospores similar in 

size 24–29 × 10–12 µm (Lundqvist in Hansen 

& Knudsen 2000), but differs in its two-celled, 

strongly flattened ascospores, with a longer 

germ slit, growth on elk and roe deer dung, and 

“stromata often preceded by pink sporodochia”. 

Spores of my collection of H. equorum 

are slightly larger than those described by 

Fuckel (1869), quite larger than in Winter 

(1873, 1887, Krug & Cain 1974), but similar in 

size to those described by others (Griffiths 

1901, Cain 1934, Moreau 1953, Lohmeyer 

1995, Richardson & Watling 1997). 

Acknowledgements 

The author is particularly indebted to the 

colleagues of Department of Tree Science, 

Entomology and Plant Pathology “G. Scaramuzzi”, 

Section Plant Pathology, University of 

Pisa, for the preparation of some axenic cultures. 

He also thanks all friends who provided 

him with a share of the material subject of this 

study. 

References 

Aas O. 1992 – A world monograph of the 

genus Thecotheus (Ascomycetes, Pezizales). 

Thesis 4 Universitetet i Bergen- 

Botanisk Institutt. 


Aas O. 1978 – Koprofile Discomycetar (Ascomycetes: 

Discomycetes Operculati = Pezizales) 

i Noreg. Universitetet i Bergen- 

Botanisk Institutt. 

Abbott SP, Lumley TC, Sigler L. 2002 – Use 

of holomorph characters to delimit Microascus 

nidicola and M. soppii sp. nov., 

with notes on the genus Pithoascus. 

Mycologia 94, 362–369. 

Abbott SP, Sigler L. 2001 – Heterothallism in 

the Microascaceae demonstrated by three 

species in the Scopulariopsis brevicaulis 

series. Mycologia 93, 1211–1220. 

Abbott SP, Sigler L, Currah RS. 1998 – 

Microascus brevicaulis sp. nov., the teleomorph 

of Scopulariopsis brevicaulis, 

supports placement of Scopulariopsis 

with the Microascaceae. Mycologia 90, 

297–302. 

Abdullah SK, Al-Saadoon AH, Guarro J. 1999 

– New and interesting coprophilous ascomycetes 

from Iraq. Nova Hedwigia 69, 

211–216. 

Abdullah SK, Alutbi SD. 1993 – Additions to 

coprophilous fungi of Iraq III. Ascobolaceae: 

Pezizales. Basrah Journal of 

Science 11, 33–44. 

Abdullah SK, Alutbi SD. 1994 – Additions to 

coprophilous fungi of Iraq IV: the genera 

Coprotus and Lasiobolus: Pezizales. 

Abhath Al-Yarmouk 3, 55–63. 

Abdullah SK, Ismail ALS, Rattan SS. 1976 – 

New and interesting coprophilous fungi 

from Iraq. Nova Hedwigia 28, 241–251. 

Abdullah SK, Rattan SS. 1978 – Zygopleurage, 

Tripterosporella and Podospora (Sordariaceae: 

Pyrenomycetes) in Iraq. Mycotaxon 

7, 102–116. 

Aguirre-Acosta E, Ulloa M. 1982 (1983) – 

Primer registro en Mexico sobre la 

sucesión de hongos en el estiercol de 

vaca. Boletín de la Sociedad Mexicana de 

Micología 17, 76–88. 

Ahmad S, Sultana K. 1973 – Contributions to 

the fungi of West Pakistan XVI. Biologia 

19, 1–26. 

Ahmed SI, Cain RF. 1972 – Revision of the 

genera Sporormia and Sporormiella. 

Canadian Journal of Botany 50, 419–477. 

Ahmed SI, Ismail ALS, Abdullah SK. 1971 – 

Contribution to the fungi of Iraq II. 

409

Coprophilous fungi. Bulletin of the 

Biological Research Centre 5, 16–32. 

Ames LM. 1963 – A monograph of the Chaetomiaceae. 

The United States Army Research 

and Development Series 2. 

Angel K, Wicklow DT. 1975 – Relationships 

between coprophilous fungi and fecal 

substrates in a Colorado grassland. Mycologia 

67, 63–74. 

Apinis AE. 1964 – Revision of British Gymnoascaceae. 

Mycological Papers 96, 1–56. 

Arenal F, Platas G, Peláez F. 2004 – Variability 

of spore length in some species of the 

genus Preussia (Sporormiella). Mycotaxon 

89, 137–151. 

Arenal F, Platas G, Peláez F. 2005 – Preussia 

africana and Preussia isabellae, two new 

Preussia species based on morphological 

and molecular evidence. Fungal Diversity 

20, 1–15. 

Arenal F, Platas G, Peláez F. 2007 – A new 

endophytic species of Preussia (Sporormiaceae) 

inferred from morphological 

observations and molecular phylogenetic 

analysis. Fungal Diversity 25, 1–17. 

Arroyo I, Calonge FD, Siquier JL, Costantino 

C. 1990 – Contribución al conocimiento 

micológico de las Islas Baleares, II. 

Ascomycotina. Boletín de la Sociedad 

Micológica de Madrid 14, 49–60. 

Arx JA von. 1973a – Ostiolate and nonostiolate 

Pyrenomycetes. Proceedings Koninklijke 

Nederlandse Akademie van Wetenschappen 

76, 3, 289–296. 

Arx JA von. 1973b – The genera Petriellidium 

and Pithoascus (Microascaceae). Persoonia 

7, 367–375. 

Arx JA von. 1975a – On Thielavia and some 

similar genera of Ascomycetes. Studies 

in Mycology 8, 1–29. 

Arx JA von. 1975b – Revision of Microascus 

with the description of a new species. 

Persoonia 8, 191–197. 

Arx JA von. 1978 – Notes on Microascaceae 

with the description of two new species. 

Persoonia 10, 23–31. 

Arx JA von. 1981 – On Monilia sitophila and 

some families of Ascomycetes. Sydowia 

34, 13–29. 

Arx JA von. 1984 – Canariomyces notabilis, a 

peculiar ascomycete from the Canary 

Islands. Persoonia 12, 185–187. 

410 

Arx JA von. 1986 – The ascomycete genus 

Gymnoascus. Persoonia 13, 173–183. 

Arx JA von. 1987 – A re-evaluation of the 

Eurotiales. Persoonia 13, 273–300. 

Arx JA von, Aa HA van der. 1987 – Spororminula 

tenerifae gen. et sp. nov. Transactions 

of the British Mycological Society 

89, 117–120. 

Arx JA von, Figueras MJ, Guarro J. 1988 – 

Sordariaceous Ascomycetes without ascospore 

ejaculation. Nova Hedwigia 94, 

1–104. 

Arx JA von, Guarro J, Figueras MJ. 1986 – 

The Ascomycete genus Chaetomium. 

Nova Hedwigia 84, 1–162. 

Arx JA von, Samson RA. 1986 – Mallochia, a 

new genus of the Eurotiales. Persoonia 

13, 185–188. 

Asad F, Ahmad SI. 1968 – Coprophilous fungi 

of west Pakistan. Part II. – Karachi. 

Pakistan Journal of Scientific and Industrial 

Research 11, 284–287. 

Asgari B, Zare R. 2010 – Two new species of 

Preussia from Iran. Nova Hedwigia 90, 

533–548. 

Baddley JW, Moser SA, Sutton DA, Pappas 

PG. 2000 – Microascus cinereus (anamorph 

Scopulariopsis) brain abscess in a 

bone marrow transplant recipient. Journal 

of Clinical Microbiology 38, 395–397. 

Bainier G. 1909 – Monographie des 

Chaetomium et des Chaetomidium. 

Bulletin de la Société Mycologique de 

France 25, 191–237. 

Barr ME. 1990 – Prodromus to nonlichenized, 

pyrenomycetous members of Class 

Hymenoascomycetes. Mycotaxon 39, 43– 

184. 

Barr ME. 2000 – Notes on coprophilous 

bitunicate Ascomycetes. Mycotaxon 76, 

105–112. 

Barrasa JM. 1985 – Contribución al estudio 

taxonómico, ecológico y corológico de 

los Ascomycetes coprófilos en España. 

Universidad de Alcalá de Henares 

(unpublished degree thesis). 

Barrasa JM, Checa J. 1990 – Contribución al 

conocimiento del género Delitschia 

Auersw. en España. Revista Iberoamericana 

de Micología 7, 5–10. 

Barrasa JM, Checa J. 1991 – Dothideales del 

parque natural de Monfragüe (Cáceres).

I. Boletín de la Sociedad Micológica de 

Madrid 15, 91–102. 

Barrasa JM, Moreno G. 1980 – Contribución al 

estudio de los hongos que viven sobre 

materias fecales (2a aportación). Acta 

Botanica Malacitana 6, 111–148. 

Barrasa JM, Moreno G. 1984 – Hongos coprófilos 

de España. III. Anales del Jardín 

Botánico de Madrid 41, 3–17. 

Barrasa JM, Soláns MJ. 1989 – Introducción al 

estudio del género Schizothecium Corda 

emend. Lundq. en España. Revista Iberoamericana 


Barron GL, Cain RF, Gilman JC. 1961 – The 

genus Microascus. Canadian Journal of 

Botany 39, 1609–1631. 

Bednarczyk MA. 1974 – Materials of the 

knowledge of the coprophilous fungi in 

the Lublin region. Acta Mycologica 10, 

331–342. 

Bell A, Mahoney DP. 1995 – Coprophilous 

fungi in New Zealand. I. Podospora 

species with swollen agglutinated perithecial 

hairs. Mycologia 87, 375–396. 

Bell A. 2005 – An illustrated guide to the 

coprophilous Ascomycetes of Australia. 

CBS Biodiversity Series No 3, Utrecht. 

Benjamin RK. 1949 – Two species representing 

a new genus of the Chaetomiaceae. 

Mycologia 41, 346–354. 

Benny GL. 1980 – A second species of Chaetomidium 

with cephalothecoid peridium 

wall. Mycologia 72, 832–836. 

Benny GL, Kimbrough JW. 1980 – A synopsis 

of the orders and families of Plectomycetes 

with keys to genera. Mycotaxon 12, 

1–91. 

Berbee ML, Taylor JW. 1992 – Convergence 

in ascospore discharge mechanism 

among pyrenomycete fungi based on 18S 

ribosomal RNA gene sequence. Molecular 

Phylogenetics and Evolution 1, 59– 

71. 

Berkeley MJ, Broome CE. 1865 – Notices of 

British fungi. The Annals and Magazine 

of Natural History, ser. 3, 15, 444–452. 

Beyer W. 1999 – Ergänzungen zur Pilzflora 

von Bayreuth und Umgebung. Teil 2. 

Zeitschrift für Mykologie 65, 41–80. 

Beyer W. 2004 – Ergänzungen zur Pilzflora 

von Bayreuth und Umgebung. Teil 3. 

Zeitschrift für Mykologie 70, 207–226. 


Bezerra JL, Kimbrough JW. 1975 – The genus 

Lasiobolus (Pezizales, Ascomycetes). 

Canadian Journal of Botany 53, 1206– 

1229. 

Blyskal B. 2009 – Fungi utilizing keratinous 

substrates. International Biodeterioration 

& Biodegradation 63, 631–653. 

Bokhary HA. 1985 – Coprophilous fungus 

succession on camel dung. Arab Gulf 

Journal of Scientific Research 3, 277– 

284. 

Bokhary HA, Parvez S. 1986 – Coprophilous 

fungi of Saudi Arabia 3. Coprophilous 

fungi of cow and rabbit dung. Proceedings 

of Saudi Biological Society 9, 13– 

21. 

Bokhary HA, Parvez S, Naseef AS. 1989 – 

Coprophilous fungi of Saudi Arabia 4. 

Coprophilous fungi of horse dung. Transactions 

of the Mycological Society of 

Japan 30, 25–34. 

Bommer E, Russeau M. 1886 – Contributions à 

la Flore Mycologique de Belgique. 

Mémoirs de la Societé Botanique de 

Belgique 25, 171. 

Booth C. 1964 – Studies of Pyrenomycetes: 

VII. Mycological Papers 94, 1–16. 

Boudier E. 1912–1913 – Sur deux nouvelles 

espèces de discomycètes d’Angleterre. 

Transactions of the British Mycological 

Society 4, 62–63. 

Bowman BH, White TJ, Taylor JW. 1996 – 

Human pathogenetic fungi and their close 

nonpathogenic relatives. Molecular Phylogenetics 

and Evolution 6, 89–96. 

Breton A. 1965 – Champignons coprophiles 

des départments du Lot, du Puy-de-Dome 

et du Finistère. Bulletin de la Société 

Mycologique de France 81, 607–622. 

Bronckers R. 2002 – Trichobolus zukalii 

(Heimerl) Kimbrough, het Harig 

sinterklaasschijfje op reeënkeutels. AMK 

Mededelingen 1, 13–15. 

Brummelen J van. 1967 – A world-monograph 

of the Genera Ascobolus and Saccobolus 

(Ascomycetes, Pezizales). Persoonia, 

supplement Vol. I. Rijksherbarium, 

Leiden. 

Brummelen J van. 1981 – The genus Ascodesmis 

(Pezizales, Ascomycetes). Persoonia 

11, 333–358. 

411

Brummelen J van. 1995 – A world–monograph 

of the Genus Pseudombrophila (Pezizales, 

Ascomycotina). Libri Botanici 14. 

IHW-Verlag. 

Caillet M, Moyne G. 1983 – Les Pyrénomycètes 

de Franche-Comté. I. Bulletin de la 

Société d'Histoire Naturelle de Doubs 81, 

23–36. 

Caillet M, Moyne G. 1984–1985 – Les Pyrénomycètes 

de Franche-Comté. II. Bulletin 

de la Société d'Histoire Naturelle de 

Doubs 82, 45–63. 

Cailleux R. 1971a – Recherches sur la 

mycoflore coprophile centrafricaine. Les 

Genres Sordaria, Gelasinospora, Bombardia(Biologie-Morphologie-Systématique) 

Ecologie. Bulletin de la Société 

Mycologique de France 87, 461–567. 

Cailleux R. 1971b – Champignons stercoraux 

de République Centrafricaine. V-Podospora 

et Zygopleurage, espèces rencontrées, 

écologie. Cahiers Maboké 9, 11– 

26. 

Cain RF. 1934 – Studies of coprophilous 

Sphaeriales in Ontario. University of 

Toronto Studies, Mycological Series 38, 

1–126. 

Cain RF. 1957 – Studies of coprophilous Ascomycetes 

– VI. Species from the Hudson 

Bay area. Canadian Journal of Botany 35, 

255–265. 

Cain RF. 1961 – Studies of coprophilous Ascomycetes, 

VII. Preussia. Canadian Journal 

of Botany 39, 1633–1666. 

Cain RF, Weresub LK. 1957 – Studies of coprophilous 

Ascomycetes. V. Sphaeronemella 

fimicola. Canadian Journal of 

Botany 35, 119–131. 

Calonge FD, Menezes de Sequeira M. 2003 – 

Contribución al catálogo de los hongos 

de Madeira (Portugal). Boletín de la 

Sociedad Micológica de Madrid 27, 277– 

308. 

Calonge FD, Siquier JL, Constantino C. 1991 – 

Contribución al conocimiento micológico 

de las Islas Baleares, III. Bovista cunninghamii 

Kreisely Tuber uncinatum 

Chatin, dos nuevas citas para el catálogo 

español. Boletín de la Sociedad Micológica 

de Madrid 15, 111–122. 

Calonge FD, Siquier JL, Constantino C. 1993 – 

Contribución al conocimiento micológico 

412 

de las Islas Baleares, V. Bovista cunninghamii 

Kreisel y Tuber uncinatum Chatin, 

dos nuevas citas para el catálogo español. 

Boletín de la Sociedad Micológica de 

Madrid 15, 111–122. 

Calviello BO. 1971 – Estudio de las especies 

Argentinas del género Chaetomium. 

Revista del Museo Argentino de Ciencias 

Naturales Bernardino Rivadavia 3, 337– 

369. 

Calviello BO. 1976 – Las especies argentinas 

de “Melanospora” Cda. Revista del 

Museo Argentino de Ciencias Naturales 

Bernardino Rivadavia 5, 93–103. 

Calviello BO. 1978 – La familia Chaetomiaceae 

en Argentina. Revista del Museo 

Argentino de Ciencias Naturales Bernardino 

Rivadavia 5, 165–176. 

Calviello BO. 1979 – Contribución al estudio 

de Ascomycetes argentinos. II. Una 

nueva especie de Kernia (Microascales). 

Revista del Museo Argentino de Ciencias 

Naturales Bernardino Rivadavia 5, 239– 

243. 

Campbell C, Rodgers JD, Murray TD. 1991 – 

A psychrophilic Orbicula associated with 

oat kernels. Mycologist 5, 113–114. 

Cannon PF, Hawksworth DL. 1982 – A reevaluation 

of Melanospora Corda and 

similar Pyrenomycetes, with a revision of 

the British species. Botanical Journal of 

the Linnean Society 84, 115–160. 

Cannon PF, Kirk PM. 2007 – Fungal Families 

of the World. CABI, United Kingdom. 

Cano J, Guarro J. 1990 – The genus Aphanoascus. 

Mycological Research 94, 355–377. 

Cano J, Guarro J, ElShafie AE. 1993 – A new 

Chaetomidium from Oman. Mycotaxon 

49, 399–403. 

Caretta G, Mangiarotti AM, Piontelli E. 1992 – 

Keratinophilic fungi isolated from soil of 

Italian parks in the province of Pavia. 

European Journal of Epidemiology 8, 

330–339. 

Caretta G, Mangiarotti AM, Piontelli E. 1994 – 

Coprophilous fungi on horse, goat and 

sheep dung from Lombardia (Italy). 

Micologia Italiana 23, 11–20. 

Caretta G, Piontelli E. 1996 – Coprophilous 

fungi from confined deers in Pavia 

(Lombardia, Italy). Boletín Micológico 

11, 41–50.

Caretta G, Piontelli E, Savino E, Bulgheroni A. 

1998 – Some coprophilous fungi from 

Kenya. Mycopathologia 142, 125–134. 

Carter A, Khan RS. 1982 – New and interesting 

Chaetomium species from East 

Africa. Canadian Journal of Botany 60, 

1253–1262. 

Chang JH, Wang YZ. 2005 – The genus Cercophora 

(Lasiosphaeriaceae) in Taiwan. 

Fungal Science 20, 19–25. 

Chang JH, Wang YZ. 2008 – Two species of 

Kernia (Plectomycetes) in Taiwan. 

Taiwania 53, 414–416. 

Chang JH, Wang YZ. 2009 – The genera 

Sporormia and Preussia (Sporormiaceae, 

Pleosporales) in Taiwan. Nova Hedwigia 

88, 245–254. 

Cheype JL. 2008 – Contribution à la connaissance 

des champignons de la haute vallée 

de l’Arve (Haute-Savoie). 5 e note: “de 

belles bouses”. Bulletin Mycologique et 

Botanique Dauphiné–Savoie 191, 29–38. 

Cinto IE, Dokmetzian DA, Ranalli ME. 2007 – 

Iodophanus carneus and I. testaceus 

(Ascomycota–Pezizales): independent 

taxonomic identity or synonymy? A 

study of their morphology and isozymes. 

Boletín de la Sociedad Argentina de 

Botánica 42, 181–187. 

Coemans E. 1862 – Notice sur les Ascobolus 

de la flore belge. Bulletin de la Société 

Royale Botanique de Belgique 1, 76–91. 

Cooke MC. 1864 – The genus Ascobolus, with 

descriptions of the British species. 

Journal of Botany 2, 147–154. 

Cooke MC. 1876 – New British fungi. 

Grevillea 4, 109–114. 

Coste C, Rey H. 1993 – Contribution à l’étude 

des ascomycètes du Tarn (France, 81) – 

1ère note. Bulletin de la Société Castraise 

de Sciences Naturelles, 61–73. 



4 ème note. Bulletin de la Société Castraise 




7 ème note. Bulletin de la Société Castraise 


Cribb AB. 1988 – Fungi on kangaroo dung. In: 

Lake Broadwater. The natural history of 


an inland lake and its environs (ed. G 

Scott). Toowoomba 52–56. 

Crouan PL, Crouan HM. 1857 – Note sur 

quelques Ascobolus nouveaux et sur une 

espèce nouvelle de Vibrissea. Annales 

des Sciences Naturelles quatrième série, 

Botanique 7, 173–178. 

Crouan PL, Crouan HM. 1858 – Note sur neuf 

Ascobolus nouveaux. Annales des 

Sciences Naturelles quatrième série, 

Botanique 10, 193–199. 

Cugnani HC. 2000 – Nondermatophytic filamentous 

keratinophilic fungi and their 

role in human infection. Revista Iberoamericana 

de Micología 109–114. 

Currah RS. 1985 – Taxonomy of the Onygenales: 

Arthrodermaceae, Gymnoascaceae, 

Myxotrichaceae and Onygenaceae. Mycotaxon 

24, 1–216. 

Currah RS. 1986 – A new species of Ascodesmis 

from Alberta. Mycologia 78, 198– 

201. 

Currah RS. 1994 – Peridial morphology and 

evolution in the prototunicate ascomycetes. 

In: Ascomycetes systematics: Problems 

and perspectives in the nineties 

(ed. DL Hawksworth). Plenum Press, 

New York pp. 281–293. 

Curzi M. 1930a – Una nuova specie di Microascus. 

Bollettino della Regia Stazione di 

Patologia Vegetale 10, 302–310. 

Curzi M. 1930b – Petriella. Nuovo genere di 

pirenomicete. Bollettino della Regia Stazione 

di Patologia Vegetale 10, 380–422. 

Curzi M. 1931 – Rapporti fra i generi Microascus 

Zukal e Scopulariopsis Bainier. 

Bollettino della Regia Stazione di Patologia 

Vegetale 11, 55–60. 

Dal Vesco G, Peyronel B, Barge MT, Volpiano 

N. 1967 – Sulla micoflora dello sterco di 

coniglio. Allionia 13, 107–127. 

Daniels J. 1961 – Chaetomium piluliferum sp. 

nov., the perfect state of Botryotrichum 

piluliferum. Transactions of the British 

Mycological Society 44, 79–86. 

Dawson M. 1900 – On the biology of Poronia 

punctata. Annals of Botany 14, 245–262. 

De Boer W, Gerards S, Klein Gunnewiek PJA, 

Modderman R. 1999 – Response to the 

chitinolytic microbial community to 

chitin amendments of dune soils. Biology 

and Fertility of Soils 29, 170–177. 

413

De Meulder H. 2000a – Onderzoek naar het 

voorkomen van paddestoelen op mest 

van Galloway-runderen (deel 1). AMK 


De Meulder H. 2000b – Onderzoek naar het 

voorkomen van paddestoelen op mest 

van Galloway-runderen (deel 2). AMK 


De Meulder H. 2007a – Onderzoek naar het 

voorkomen van ascomyceten op mest van 

geit en schaap (deel 1). AMK Mededelingen 

1, 3–10. 

De Meulder H. 2007b – Onderzoek naar het 



2, 29–34. 

De Meulder H. 2007c – Onderzoek naar het 



3, 57–63. 

De Notaris G. 1849 – Micromycetes italici novi 

vel minus cogniti. Decas quinta. Memorie 

della Reale Accademia delle Scienze 

di Torino II, 10, 333–350. 

Delgado AE, Kimbrough JW, Hanlin RT. 2000 

– Zygopleurage zygospora, a new record 

from Venezuela. Mycotaxon 75, 257– 

263. 

Delgado Avila AE, Piñero Chávez AJ, 

Urdaneta García LM. 2001a – Estudios 

taxonómicos de hongos coprofílicos de la 

división Ascomicota (clase: pyrenomycetes) 

del Estado Zulia, Venezuela. Revista 

Científica FCV–LUZ 11, 247–255. 


Urdaneta García LM. 2001b – Hongos 

coprofílicos del Estado Zulia, Venezuela. 

Clases: Plectomycetes y Discomycetes. 

División Ascomicota. Revista Científica 

FCV–LUZ 11, 297–305. 


Urdaneta García LM. 2002 – Copromyces 

bisporus, a new record for Venezuela. 

Revista Científica FCV–LUZ 12, 12–14. 

Denison WC. 1963 – A preliminary study of 

the operculate cup-fungi of Costa Rica. 

Revista de Biología Tropical 11, 99–129. 

Dennis RWG. 1959 – Fungi Venezuelani: III. 

Kew Bulletin 14, 418–458. 

Dennis RWG. 1981 – British Ascomycetes. 

Sec. (ed. J Cramer). Vaduz. 

414 

Dennis RWG. 1986 – Fungi of the Hebrides. 

Royal Botanic Garden, Kew. 

Derbsch H, Schmitt JA. 1987 – Atlas der Pilze 

des Saarlandes, Teil 2, Nachweise, 

Ökologie, Vorkommen und Beischreibungen. 

Sarrebruck, Eigenverlag der 

Delattinia. 

Dissing H. 1980 – Chalazion helveticum (Pezizales), 

a new species from Graubünden, 

Switzerland. Sydowia 33, 29–32. 

Dissing H. 1992 – Notes on coprophilous Pyrenomycete 

Sporormia fimetaria. Persoonia 

14, 389–394. 

Dissing H, Raitviir A. 1973 – Discomycetes of 

middle Asia. II. Thelebolaceae, Ascobolaceae, 

Pyronemataceae and Pezizaceae 

from the Tien-Shan Mountains. Eesti 

NSV Teaduste Akadeemia Toimetised 

22, Biology 2, 124–131. 

Dissing H, Sivertsen S. 1975 – Operculate 

Discomycetes from Rana (Norway) 1. 

Chalazion sociabile gen. nov., sp. nov. 

Norwegian Journal of Botany 22, 1–4. 

Dodge BO. 1914 – Wisconsin Discomycetes. 

Transactions of the Wisconsin Academy 

of Sciences, Arts and Letters 17, II(3), 

1027–1056. 

Domsch KH, Gams W, Anderson T-H. 1980 – 

Compendium of soil fungi. Vol. 1. 

Academic Press, London. 

Donadini JC. 1979 – Le genre Peziza Linné per 

Saint–Amans (1ère Partie). Documents 

Mycologiques 9, 1–42. 

Doveri F. 2004a – Fungi Fimicoli Italici. 

A.M.B., Vicenza. 

Doveri F. 2004b – Podospora alexandri, una 

nuova specie fimicola dall’Italia. Rivista 

di Micologia 47, 211–221. 

Doveri F. 2005 – Sporormiella hololasia, a 

new hairy species from Italy. Rivista di 

Micologia 48, 31–41. 

Doveri F. 2006 – Nuove segnalazioni di 

Onygenales coprofile dall’Italia. Rivista 


Doveri F. 2007a – An updated key to 

coprophilous Pezizales and Thelebolales 

in Italy. Mycologia Montenegrina 10, 

55–82. 

Doveri F. 2007b – Sporormiella gigantea 

comb. nov., una nuova definizione di 

Sporormiella ovina s. S.I. Ahmed &

Cain, nom.amb. Rivista di Micologia 50, 

43–55. 

Doveri F. 2008a – Aggiornamento sul genere 

Chaetomium con descrizione di alcune 

specie coprofile, nuove per l’Italia. 

Pagine di Micologia 29, 1–60. 

Doveri F. 2008b – A bibliography of Podospora 

and Schizothecium, a key to the 

species, and a description of Podospora 

dasypogon newly recorded from Italy. 

Pagine di Micologia 29, 61–159. 

Doveri F. 2008c – Espèces coprophiles de 

pyrénomycètes s.l. nouvelles pour l’Italie 

après la parution des Fungi Fimicoli 

Italici. Bulletin Mycologique et Botanique 

Dauphiné-Savoie 191, 71–96. 

Doveri F. 2010a – Une nouvelle variété de 

Tripterosporella sur crottin. Une 

occasion de recombiner le pyrénomycète 

cléistothécial Cercophora heterospora 

dans le genre Tripterosporella. Bulletin 

Mycologique et Botanique Dauphiné- 

Savoie 196, 49–55. 

Doveri F. 2010b – Occurrence of coprophilous 

Agaricales in Italy, new records, and 

comparison with their European and 

extraeuropean distribution. Mycosphere 

1, 103–140. 

Doveri F, Caroti V, Cacialli G. 1997 – An 

exceptional finding from horse dung: 

Enterocarpus grenotii. Contribution to 

the study of fimicolous fungi – XXVI. 

Bollettino del Gruppo Micologico G. 

Bresadola. Nuova serie 40, 187–190. 

Doveri F, Coué B. 2008a – Sur deux taxons 

nouveaux de Thecotheus. Documents 


Doveri F, Coué B. 2008b – Sporormiella 

minutisperma, une nouvelle espèce 

coprophile, récoltée en France. Bulletin 

Mycologique et Botanique Dauphiné– 

Savoie 191, 39–44. 

Doveri F, Granito VM, Lunghini D. 2005 – 

Nuovi ritrovamenti di Coprinus s.l. 

fimicoli in Italia. New findings of 

fimicolous Coprinus s.l. in Italy. Rivista 


Doveri F, Guarro J, Cacialli G, Caroti V. 1998a 

– Contribution to the study of fimicolous 

fungi. XXVII. A new Chaetomidium 

from Italy with a cephalothecoid peridium. 

Mycotaxon 67, 427–432. 


Doveri F, Pecchia S, Sarrocco S, Minnocci A, 

Vannacci G. 2010a – Rodentomyces, a 

new hypocrealean genus from Italy. 

Fungal Diversity 42, 57–69. 

Doveri F, Sarrocco S, Pecchia S, Forti M, 

Vannacci G. 2010b – Coprinellus mitrinodulisporus, 

a new species from chamois 

dung. Mycotaxon 114, 351–360. 

Doveri F, Wang YZ, Cacialli G, Caroti V. 

1998b – Una nuova specie di Chalazion 

dalla Toscana, Italia. Contributo allo 

studio dei funghi fimicoli XXVIII. Rivista 


Eckblad FE. 1968 – The Genera of the operculate 

Discomycetes. Nytt Magasin for 

Botanikk 15, 1– 192. 

Ellis JB, Everhart BM. 1892 – The North 

American Pyrenomycetes. Newfield. 

Elshafie AE. 2005 – Coprophilous mycobiota 

of Oman. Mycotaxon 93, 355–357. 

Emmons CW, Dodge BO. 1931 – The ascocarpic 

stage of species of Scopulariopsis. 

Mycologia 23, 313–331. 

Engel H, Hanff B. 1984 – Hinweise zu einigen 

weiteren auf den Pilzfarbtafeln abgebildeten 

Arten. Pilzflora Nordwestoberfrankens 

8, 34–37. 

Engel H, Hanff B. 1985 – Hinweise zu einigen 

weiteren auf den Pilzfarbtafeln abgebildeten 

Arten. Pilzfl. Nordwestoberfrankens 

9, 54. 

Eriksson OE. 1992 – The non-lichenized pyrenomycetes 

of Sweden. SBT, Lund. 

Eriksson OE. 2009 – The non-lichenized ascomycetes 

of Sweden. KBC-tryckeriet, 

Umeå University, Sweden. 

Eriksson OE, Hawksworth DL. 1998 – Outline 

of the Ascomycetes – 1998. Systema 

Ascomycetum 16, 83–296. 

Faurel L, Schotter G. 1965 – Notes Mycologiques. 

IV. Champignons coprophiles du 

Sahara central et notamment de la 

Tefedest. Revue de Mycologie, Paris 30, 

141–164. 

Favre A. 2008 – Champignons coprophiles de 

la réserve naturelle du delta de la Dranse. 

Bulletin Mycologique et Botanique 

Dauphiné-Savoie 191, 23–28. 

Filipello Marchisio V. 2000 – Keratinophilic 

fungi: their role in nature and degradation 

of keratinic substrates. Revista Iberoamericana 

de Micología, 86–92. 

415

Fouchier F, Maurice JP, Roux P. 2009 – 

Poronia punctata (L. : Fr.) Rabenh.: 

récoltes portugaise et française. Bulletin 

semestriel de la Fédération des Associations 

Mycologiques Méditerranénnes, 

N.S. 35, 31–38. 

Fuckel KWGL. 1860. Enumeratio Fungorum 

Nassoviae. Ser. I. Jahrbücher Vereins 

Naturkunde Herzogthum Nassau 15, 1– 

123. 

Fuckel KWGL. 1866 – Über rheinische 

Ascobolus-arten. Hedwigia 5, 1–5. 

Fuckel KWGL. 1867 – Fungi Rhenani, cent. 

18. Hedwigia 6, 174–175. 

Fuckel KWGL. 1869 – Symbolae 

Mycologicae. Beiträge zur Kenntniss der 

Rheinischen Pilze. Jahrbüchern des 

Nassauischen Vereins für Naturkunde 

23–24. Wiesbaden. 

Furuya K, Udagawa S. 1973 – Coprophilous 

Pyrenomycetes from Japan III. Transactions 

of the Mycological Society of Japan 

14, 7–30. 

García Alonso F. 2007 – Ascomycetes de la 

zona de Penyagolosa (Castelló) II. 

Butlletí de la Sociedad Micológica 

Valenciana 12, 65–82. 

García Bona IM. 1978 – Macromicetos nuevos 

o interesantes de Navarra. II. Anales del 

Instituto Botánico A.J. Cavanilles 35, 

49–78. 

García-Zorrón N. 1973 – Sphaeriales coprófilos 

del Uruguay. Revista Biológica del 

Uruguay 1, 79– 96. 

García-Zorrón N. 1975 – Sphaeriales coprófilos 

del Uruguay II. Revista Biológica del 

Uruguay 3, 153–160. 

Ghosh GR, Orr GF, Kuehn HH. 1963 – A new 

genus of the Gymnoascaceae with a 

rudimentary peridium. Mycopathologia 

et Mycologia Applicata 21, 36–44. 

Ghosh GR, Sur B. 1985 – Keratinophilic fungi 

from Orissa, India, III–Two new records. 

Kavaka 12, 63–69. 

Gilgado F, Gené J, Cano J, Guarro J. 2007 – 

Reclassification of Graphium tectonae as 

Parascedosporium tectonae gen. nov., 

comb. nov., Pseudallescheria africana as 

Petriellopsis africana gen. nov., comb. 

nov. and Pseudallescheria fimeti as 

Lophotrichus fimeti comb. nov. Interna- 

416 

tional Journal of Systematic and Evolutionary 

Microbiology 57, 2171–2178. 

Goidànich G. 1932 – Enumerazione dei funghi 

della provincia di Bologna. VI centuria. 

Malpighia 32, 339–362. 

Greathouse GA, Ames LM. 1945 – Fabric 

deterioration by thirteen described and 

three new species of Chaetomium. Mycologia 

37, 138–155. 

Green E. 1931 – Observations on certain Ascobolaceae. 

Transactions of the British 

Mycological Society 15, 321–332. 

Greif MD, Currah RS. 2007 – Development 

and dehiscence of the cephalothecoid 

peridium in Aporothielavia leptoderma 

shows it belongs to Chaetomidium. 


Greif MD, Stchigel AM, Miller AN, Huhndorf 

SM. 2009 – A re-evaluation of genus 

Chaetomidium based on molecular and 

morphological characters. Mycologia 

101, 554–564. 

Griffiths D. 1901 – The North American 

Sordariaceae. Memoirs of the Torrey 

Botanical Club 11, 1–134. 

Griffiths D, Seaver FJ. 1910 – Fimetariaceae. 

In North American Flora 3(1), The New 

York Botanical Garden. 

Guarro Artigas J. 1983 – Hongos coprófilos 

aislados en Cataluña. Ascomycetes. 

Anales del Jardín Botánico de Madrid 39, 

229–245. 

Guarro J, Abdullah SK, Gené J, Al-Saadoon 

AH. 1997a – A new species of Preussia 

from submerged plant debris. Mycological 

Research 101, 305–308. 

Guarro J, Al-Saadoon AH, Abdullah SK. 

1997b – Two new coprophilous species 

of Preussia (Ascomycota) from Iraq. 


Gube M. 2010 – Poronia punctata (L. : Fr.) Fr. 

in Thüringen. Zeitschrift für Mykologie 

76, 59–66. 

Häffner J. 1986 – Rezente Ascomycetenfunde 

III – Dungbewohner, Gärfutter- und 

Nadelstreubesiedler. Einführung in die 

Gattung Saccobolus. Mitteilungsblatt der 

Arbeitsgemeinschaft Pilzkunde 

Niederrhein 4, 106–129. 

Häffner J, Christan J. 1991 – Rezente Ascomycetenfunde 

X. Trichophaea velenovskyi 

(Vacek in Svrček) comb. nov. - neu für

Deutschland! Zeitschrift für Mykologie 

57, 161–165. 

Hansen K, Laessøe T, Pfister DH. 2000 – 

Phylogenetic diversity in the core group 

of Peziza inferred from ITS sequences 

and morphology. Mycological Research 

106, 879–902. 

Hansen K, Laessøe T, Pfister DH. 2001 – 

Phylogenetics of the Pezizaceae, with an 

emphasis on Peziza. Mycologia 93, 958– 

990. 

Hansen K, LoBuglio KF, Pfister DH. 2005 – 

Evolutionary relationships of the cupfungus 

genus Peziza and Pezizaceae 

inferred from multiple nuclear genes: 

PRB2, β-tubulin, and LSU rDNA. 

Molecular Phylogenetics and Evolution 

36, 1–23. 

Hansen K, Perry BA, Pfister DH. 2005 – 

Phylogenetic origins of two cleistothecial 

fungi, Orbicula parietina and Lasiobolidium 

orbiculoides, within the operculate 

discomycetes. Mycologia 97, 1023–1033. 

Hansen K, Pfister DH. 2006 – Systematics of 

the Pezizomycetes: the operculate discomycetes. 

Mycologia 98, 1029–1040. 

Hansen L, Knudsen H. 2000 – Nordic Macromycetes 

1. Ascomycetes. Nordsvamp, 

Copenhagen. 

Hansen PB, Lange C, Petersen JH, Vesterholt 

J. 1998 – Nøgler til coprofile svampe 

(unpublished, 73 pp). 

Hausner G, Reid J, Klassen GR. 1993 – On the 

phylogeny of Ophiostoma, Ceratocystis 

s.s., and Microascus, and relationships 

within Ophiostoma based on partial 

ribosomal DNA sequences. Canadian 


Heimerl A. 1889 – Die niederösterreichischen 

Ascoboleen. Fünfzehnter Jahresbericht 

der k. k. Ober-Realschule im Bezirke 

Sechshaus bei Wien 15, 1–32. 

Heine N, Welt P. 2008 – Beiträge zur Kenntnis 

coprophiler Pilze (4). Coprophile Pilze 

im Hartschimmelgebiet bei Andechs und 

weitere bayerische Dungpilzfunde. 

Mycologia Bavarica 10, 63–83. 

Hilber O, Hilber R. 1987 – Pyrenomyceten der 

Dünen rund um Darmstadt. Beiträge zur 

Kenntnis der Pilze Mitteleuropas 3, 485– 

494. 


Hladki de Sanz A. 1997 – Contribución al 

estudio de las Xylariaceae (Xylariales– 

Ascomycotina) de la República 

Argentina. I. El género Poronia. Lilloa 

39, 35–41. 

Holm L, Ryman S. 2000 – Fungi exsiccati 

Suecici, praesertim Upsalienses. Thunbergia 

30, 1–21. 

Hoog GS de, Göttlich E, Platas G, Genilloud 

O, Leotta G, Brummelen J van. 2005 – 

Evolution, taxonomy and ecology of the 

genus Thelebolus in Antarctica. Studies 


Hoog GS de, Guarro J, Gené J, Figueras MJ. 

2000 – Atlas of Clinical Fungi, 2nd edn. 

Centraalbureau voor Schimmelcultures, 

Utrecht. 

Horie Y, Li D. 1997 – Five interesting Ascomycetes 

from herbal drugs. Mycoscience 

38, 287–295. 

Howard DH. 2002 – Pathogenic fungi in 

humans and animals, 2nd edn. Marcel 

Dekker, New York. 

Hu DM, Cai L, Hyde KD, Zhang KQ. 2006 – 

The genera Podospora and Schizothecium 

from mainland China. Cryptogamie, 

Mycologie 27, 89–109. 

Hubálek Z. 1974 – Dispersal of fungi of the 

family Chaetomiaceae by free-living 

birds. I. A survey of records. Česká 

Mykologie 28, 65–79. 

Hughes SJ. 1951 – Studies on micro-fungi. 

VIII. Orbicula and Lilliputia. Mycological 

Papers 42, 1–27. 

Issakainen J, Jalava J, Hyvönen J, Sahlberg N, 

Pirnes T, Campbell CK. 2003 – Relationships 

of Scopulariopsis based on LSU 

rDNA sequences. Medical Mycology 41, 

31–42. 

Issakainen J, Jalava J, Saari J, Campbell CK. 

1999 – Relationship of Scedosporium 

prolificans with Petriella confirmed by 

partial LSU rDNA sequences. Mycological 

Research 103, 1179–1184. 

Iwen PC, Sigler L, Tarantolo S, Sutton DA, 

Rinaldi MG, Lackner RP, McCarthy DI, 

Hinrichs SH. 2000 – Pulmonary infection 

caused by Gymnascella hyalinispora in a 

patient with acute myelogenous leukemia. 

Journal of Clinical Microbiology 38, 

375–381. 

417

Jaczewski A de. 1895 – Les Xylariées de la 

Suisse. Bulletin de la Société Mycologique 

de France 11, 108–137. 

Jahn E. 1993 – Zur Häufigkeit von Arten der 

Sordariaceae s.l. in Norddeutschland. 

Kieler Notizen 22, 32–51. 

Jahn E. 1997 – Koprophile Pezizales von 

Dungkulturen aus dem norddeutschen 

Raum und aus verschiedenen Teilen der 

Erde. Zeitschrift für Mykologie 63, 133– 

148. 

Jalink L. 1992 – Poronia erici, een kleine soort 

speldeprik–zwam in Nederland. Coolia 

35, 95–98. 

Jamoni PG. 1998 – Un nuovo discomicete 

coprofilo in Valsesia: Trichophaea subalpina 

spec. nov. Funghi e Ambiente 77, 

13–17. 

Jeamjitt O, Manoch L, Visarathanonth N, 

Chamswarng C, Watling R, Sharples G, 

Kijjoa A. 2007 – Coprophilous ascomycetes 

in Thailand. Mycotaxon 100, 115– 

136. 

Jeng RS, Krug JC. 1976 – Coprotiella, a new 

cleistocarpous genus of the Pyronemataceae 

with ascospores possessing de Bary 

bubbles. Mycotaxon 4, 545–550. 

Jeuniaux J, Voss–Foucart MF, Bussers JC. 

1993 – La production de chitine par les 

crustacés dans les écosystèmes marins. 

Aquatic Living Resources 6, 331–341. 

Jones PM. 1936 – A New species of Microascus 

with a Scopulariopsis stage. Mycologia 

28, 503–509. 

Jülich W. 1968 – Über coprophile Pilze Mitteuropas 

I. Verhandlungen des Botanischen 

Vereins für die Provinz Brandenburg und 

die Angrenzenden Lander 105, 34–43. 

Karsten PA. 1861 – Synopsis Pezizarum et 

Ascobolorum Fenniae. Helsingfors. 

Karsten PA. 1870 – Monographia Ascobolorum 

Fenniae. Notiser ur Sällskapets 

Fauna et Flora Fennica Förhandlingar 11, 

197–210. 

Karsten PA. 1888 – Symbolae ad Mycologiam 

Fennicam. XXVII. Meddelanden af 

Societas pro Fauna et Flora Fennica 16, 

33–36. 

Khan RS, Cain RF. 1979 – The genera Sporormiella 

and Sporormia in East Africa. 


1186. 

418 

Khan RS, Krug JC. 1989 – New records of the 

Sordariaceae from East Africa. Mycologia 

81, 862–869. 

Khan RS, Krug JC. 1994 – A synopsis of the 

coprophilous Ascomycetes of East 

Africa. In: Proceedings of the XIII Plenary 

Meeting AETFAT (eds JH Seyani, AC 

Chikuni). Malawi 755–772. 

Kimbrough JW. 1966 – Structure and development 

of Trichobolus zukalii. Mycologia 

58, 289–306. 

Kimbrough JW, Korf RP. 1967 – A synopsis of 

the genera and species of the tribe 

Theleboleae (Pseudoascoboleae). American 


Kimbrough JW, Luck-Allen ER, Cain RF. 

1969 – Iodophanus, the pezizae segregate 

of Ascophanus (Pezizales). American 


Kimbrough JW, Luck-Allen ER, Cain RF. 

1972 – North American species of 

Coprotus (Thelebolaceae: Pezizales). 

Canadian Journal of Botany 50, 957–971. 

Kirk PM, Cannon PF, David JC, Stalpers JA. 

2001 – Dictionary of the Fungi, 9 th edn. 

CAB International, Wallingfor. 

Kirk PM, Cannon PF, Minter DW, Stalpers JA. 

2008 – Dictionary of the Fungi. 10 th edn. 

CABI Europe, United Kingdom. 

Kohlman-Adamska A. 1965 – Niektóre grzyby 

koprofilne z okolic Warszawy. Acta 

Micologica 1, 77–103. 

Korf RP, Dirig R. 2009 – Discomycetes 

exsiccati. Fascicles 5 and 6. Mycotaxon 

107, 25–34. 

Korf RP, Zhuang WY. 1991a – A preliminary 

discomycete flora of Macaronesia: part 

12, Pyronematineae, and Pezizineae, 

Ascobolaceae. Mycotaxon 40, 307–318. 

Korf RP, Zhuang WY. 1991b – A preliminary 

discomycete flora of Macaronesia: part 

16, Otideaceae, Scutellinioideae. Mycotaxon 

40, 79–106. 

Korolyova OV. 2000 – New Ascomycete 

species Sporormiella tomilinii 

Korolyova. Mikologiya i Fitopatologiya 

34, 11–13. 

Kristiansen R. 1990 – New records of species 

of the genus Chalazion (Pezizales) in 

Norway with emphasis on their ecology. 

Agarica 10–11, 83–97.

Kristiansen R. 1994 – Ascodesmis (Pezizales) i 

Norge, en sjelden koprofil slekt eller bare 

oversett ? Agarica 13, 87–100. 

Krug JC. 1995 – The genus Fimetariella. Canadian 


Krug JC, Cain RF. 1974 – New species of 

Hypocopra (Xylariaceae). Canadian 


Krug JC, Khan RS. 1989 – New records and 

new species of Podospora from East 

Africa. Canadian Journal of Botany 67, 

1174–1182. 

Krug JC, Scott JA. 1994 – The genus Bombardioidea. 

Canadian Journal of Botany 72, 

1302–1310. 

Kruys Å, Ericson L. – 2008 Species richness of 

coprophilous ascomycetes in relation to 

variable food intake by herbivores. 

Fungal Diversity 30, 73–81. 

Kruys Å, Wedin M. 2009 – Phylogenetic 

relationships and an assessment of traditionally 

used taxonomic characters in the 

Sporormiaceae (Pleosporales, Dothideomycetes, 

Ascomycota), utilising multigene 

phylogenies. Systematics and Biodiversity 

7, 465–478. 

Kuehn HH. 1957 – Observations on Gymnoascaceae. 

V. Developmental morphology of 

two species representing a new genus of 

the Gymnoascaceae. Mycologia 49, 694– 

706. 

Kuehn HH, Orr GF. 1959 – Observations on 

Gymnoascaceae. VI. A new species of 

Arachniotus. Mycologia 51, 864–870. 

Kuehn HH, Orr GF. 1963 – Observations on 

Gymnoascaceae. IX. A new species of 

Pseudoarachniotus from soil-burial beds. 

Mycopathologia et Mycologia Applicata 

29, 257–264. 

Landvik S, Egger KN, Schumacher T. 1997 – 

Towards a subordinal classification of the 

Pezizales (Ascomycota): phylogenetic 

analyses of SSU rDNA sequences. 

Nordic Journal of Botany 17, 403–418. 

Landvik S, Kristiansen R, Schumacher T. 1998 

– Phylogenetic and structural studies in 

the Thelebolaceae (Ascomycota). Mycoscience 

39, 49–56. 

Larsen K. 1970 – The genus Saccobolus in 

Denmark. Botanisk Tidsskrift 65, 371– 

389. 


Larsen K. 1971 – Danish endocoprophilous 

fungi, and their sequence of occurrence. 

Botanisk Tidsskrift 66, 1–32. 

Lee S, Hanlin RT. 1999 – Phylogenetic relationships 

of Chaetomium and similar genera 

based on ribosomal DNA sequences. 

Mycologia 91, 434–442. 

Leenurm K. 1998 – New records of coprophilous 

Ascomycetes in Estonia. Agarica 

24/25, 155–167. 

Linnaeus C von. 1755 – Flora Suecica exhibens 

Plantas per Regnum Sueciae Crescentes. 

Editio secunda. Stockholmiae. 

Locquin-Linard M. 1977 – A propos des genres 

non ostiolés placés dans la famille des 

Microascaceae (Ascomycètes). Création 

d’un nouveau genre: Enterocarpus. 

Revue de Mycologie 41, 509–523. 

Locquin-Linard M. 1980 – Kernia setadispersa, 

nouvelle espèce de la famille des 

Microascaceae (Ascomycetes). Cryptogamie, 

Mycologie 1, 29–32. 

Locquin-Linard M. 1986 – Lophotrichus geniculosporus 

Locquin-Linard, nouvelle 

espèce de Microascales (Ascomycète 

Ascohyménien) coprophile du N.-E. Africain. 

Cryptogamie, Mycologie 7, 31–36. 

Lohmeyer TR. 1994 – New European and 

Australian records of Poronia erici 

Lohmeyer & Benkert, and a fairy tale 

concerning their possible relationship. 

The Mycologist 8, 16–20. 

Lohmeyer TR. 1995 – Pilze auf Helgoland. Zur 

Mykologie einer Ferieninsel in der Nordsee. 

Teil 1: Ascomyceten. Zeitschrift für 

Mykologie 61, 79–121. 

Lohmeyer TR, Benkert D. 1988 – Poronia 

erici - eine neue Art der Xylariales 

(Ascomycetes). Zeitschrift für Mykologie 

54, 93–102. 

Lorenzo LE. 1990 – Contribución al estudio de 

Pyrenomycetes “sensu lato” (Ascomycotina) 

coprófilos del Parque Nacional 

Nahuel Huapi (Argentina) II. Boletín de 

la Sociedad Argentina de Botánica 26, 

187–195. 

Lorenzo LE. 1992 – Contribución al estudio de 

Pyrenomycetes “sensu lato” (Ascomycotina) 

coprófilos del Parque Nacional 

Nahuel Huapi (Argentina) III. Boletín de 

la Sociedad Argentina de Botánica 28, 

183–193. 

419

Lorenzo LE. 1993 – Especies coprófilas de la 

familia Chaetomiaceae en la Patagonia 

(Argentina). Boletín de la Sociedad 

Argentina de Botánica 29, 159–172. 

Lorenzo LE, Havrylenko M. 2001 – The 

genera Arnium and Podospora from 

Argentina. Mycologia 93, 1221–1230. 

Lumbsch HT, Huhndorf SM. 2007 – Outline of 

Ascomycota – 2007. Myconet 13, 1–58. 

Lumley TC, Abbott SP, Currah RS. 2000 – 

Microscopic Ascomycetes isolated from 

rotting wood in the boreal forest. 

Mycotaxon 74, 395–414. 

Lumley TC, Currah RS. 1995 – A preliminary 

account of some unusual onygenalean 

fungi from decayed wood. Reports of 

Tottori Mycological Institute 33, 7–13. 

Lundqvist N. 1960 – Coprophilous Ascomycetes 

from Northern Spain. Svensk 

Botanik Tidskrift 54, 523–529. 

Lundqvist N. 1967 – On spore ornamentation 

in the Sordariaceae, exemplified by the 

new cleistocarpous genus Copromyces. 

Arkiv für Botanik 6, 327–337. 

Lundqvist N. 1969a – Zygopleurage and Zygospermella 

(Sordariaceae s. lat., Pyrenomycetes). 

Botaniska Notiser 122, 353– 

374. 

Lundqvist N. 1969b – Tripterospora (Sordariaceae 

s. lat., Pyrenomycetes). Botaniska 

Notiser 122, 589–603. 

Lundqvist N. 1972 – Nordic Sordariaceae s. lat. 

Symbolae Botanicae Upsalienses 20, 1– 

374. 

Lundqvist N. 1973 – Studia fungorum fimi. I. 

New records of Podosporae, and a new 

species, P. papilionacea. Svensk Botanik 

Tidskrift 67, 34–52. 

Lundqvist N. 1974 – Studia fungorum fimi. II. 

New records of Arnium, and a new species, 

A. bellum. Svensk Botanik Tidskrift 

68, 289–303. 

Lundqvist N. 1981 – Fungi fimicoli exsiccati. 

Publication from the Herbarium University 

of Uppsala, Sweden 8(I–II), 1–20. 

Lundqvist N. 1989 – Fungi fimicolae exsiccati. 

Thunbergia 9, 51–75. 

Lundqvist N. 1997 – Fungi fimicolae exsiccati. 

Thunbergia 25, 76–125. 

Luttrell ES. 1951 – Taxonomy of the Pyrenomycetes. 

University of Missouri Studies 

24, 1–120. 

420 

Malaval JC. 2004 – Espèces peu citéès ou non 

répertoriées dans la région de Montpellier 

et de Bédarieux (Hérault-France). Documents 


Malloch D. 1970 – New concepts in the Microascaceae 

illustrated by two new species. 

Mycologia 62, 727–740. 

Malloch D, Benny GL. 1973 – California Ascomycetes: 

four new species and a new 

record. Mycologia 65, 648–660. 

Malloch D, Cain RF. 1971a – Four new genera 

of cleistothecial Ascomycetes with hyaline 

ascospores. Canadian Journal of 

Botany 48, 1815–1825. 

Malloch D, Cain RF. 1971b – The genus Kernia. 


867. 

Malloch D, Cain RF. 1973 – The genus Thielavia. 

Mycologia 65, 1055–1077. 

Malloch D, Hubart JM. 1987 – An undescribed 

species of Microascus from the Cave of 

Ramioul. Canadian Journal of Botany 65, 

2384–2388. 

Marasas WFO, Westhuizen GCA van der, 

Warmelo KT van, Papendorf MC 1966 – 

New and interesting records of South 

African fungi, Part V. Bothalia 9, 229– 

243. 

Marchal E. 1883 – Pyrénomycètes coprophiles 

nouveaux pour la flore Belge. Bulletin 

des Séances de la Société Belge de 

Microscopie 10, 40–43. 

Marchal E. 1884a – Champignons coprophiles 

de la Belgique I. Bulletin de la Société 


Marchal E. 1884b – Champignons coprophiles. 

III. Bulletin de la Société Royale Botanique 

de Belgique 23, 88–94. 

Marchal E. 1885 – Champignons coprophiles 

de Belgique. Bulletin de la Société 


Marchal E. 1891 – Champignons coprophiles 

de Belgique. VI. Mucorinées et 

Sphaeropsidées nouvelles. Bulletin de la 

Société Royale Botanique de Belgique 

30, 134–146. 

Massee G. 1895 – British Fungus Flora, Vol. 

IV. G. Bell & Sons, London and New 

York. 

Massee G, Salmon ES. 1901 – Researches on 

coprophilous fungi. Annals of Botany 15, 

313–357.

Massee G, Salmon ES. 1902 – Researches on 

coprophilous fungi II. Annals of Botany 

16, 57–93. 

Mathur PN, Thirumalachar MJ. 1962 – Studies 

on some Indian soil fungi II. Sydowia 16, 

46–57. 

Matočec N. 2000 – The endangered European 

species Poronia punctata (Xylariales, 

Ascomycotina), still alive and well in 

Croatia. Natura Croatica 9, 35–40. 

Matsushima T. 1975 – Icones microfungorum a 

Matsushima lectorum. Published by the 

Author, Kobe, Japan. 

Mazzucchetti G. 1965 – Microfunghi della 

cellulosa e della carta. Attività e inquadramento 

sistematico. Il genere Chaetomium. 

Ente Nazionale per la Cellulosa e 

la Carta, Roma. 

McGinnis MR, Padyhe AA, Ajello L. 1982 – 

Pseudallescheria Negroni et Fischer, 

1943 and its later synonym Petriellidium 

Malloch, 1970. Mycotaxon 14, 94–102. 

Meyer SL. 1941 – Coprophilous Ascomycetes 

of Tennessee. Journal of the Tennessee 

Academy of Science 16, 401–405. 

Miller JH. 1942 – South African Xylariaceae. 

Bothalia 4, 251–273. 

Millner PD. 1975 – Ascomycetes of Pakistan: 

Chaetomium. Biologia 21, 39–73. 

Mirza JH. 1968 – Cleistobombardia, a new 

cleistothecial genus in the family Sordariaceae. 

Mycologia 60, 704–708. 

Mirza JH, Cain RF. 1969 – Revision of the 

genus Podospora. Canadian Journal of 

Botany 47, 1999–2048. 

Mirza JH, Nasir MA. 1968 – Additions to the 

coprophilous fungi of West Pakistan. II. 


Mohammedi I, Piens MA, Audigier–Valette C, 

Gantier JC, Argaud L, Martin O, Robert 

D. 2004 – Fatal Microascus trigonosporus 

(anamorph Scopulariopsis) pneumonia 

in a bone marrow transplant recipient. 

European Journal of Clinical Microbiology 

& Infectious Diseases 23, 215–217. 

Moravec J. 2005 – A world monograph of the 

genus Cheilymenia. Libri Botanici 21, 1– 

256. 

Moravec Z. 1968 – Remarks on some coprophilous 

fungi in Norway. Česká Mykologie 

22, 301–309. 


Moreau C. 1953 – Les Genres Sordaria et 

Pleurage. P. Lechevalier, Paris. 

Moreno G, Barrasa JM. 1977 – Contribución al 

estudio de los hongos que viven sobre 

materias fecales (1a aportación). Acta 

Botanica Malacitana 3, 5–33. 

Morton FJ, Smith G. 1963. – The genera Scopulariopsis 

Bainier, Microascus Zukal, 

and Doratomyces Corda. Mycological 

Papers 8, 1–96. 

Mouchacca J. 1999 – A list of novel fungi 

described from the Middle East, mostly 

from non-soil substrata. Nova Hedwigia 

68, 149–174. 

Mouso de Cachi N, Ranalli ME. 1989 – Systematic 

and biological study of Ascobolaceae 

of Argentina XVII. Development 

and cytology of Iodophanus carneus 

(Pezizales-Ascobolaceae). Nova Hedwigia 

49, 49–59. 

Mouton V. 1886 – Ascomycètes observés aux 

environs de Liège. Bulletin de la Société 

Royale Botanique de Belgique 25, 137– 

162. 

Moyne G. 2006 – Pezizomycetes (Discomycètes 

operculés) recoltes au cours de nos 

prospections dans le departement du 

Doubs (France). Bulletin de la Société 

d'Histoire Naturelle de Doubs 91, 115– 

132. 

Moyne G, Petit J. 2006–2007 – Quelques ascomycetes 

coprophiles recoltes en France et 

en particulier dans le Department du 

Doubs. Bulletin de la Société d'Histoire 

Naturelle du Doubs 91, 49–114. 

Mukerji KG. 1970 – Fungi of Delhi: III. Some 

interesting Ascomycetes. Mycopathologia 

et Mycologia Applicata 42, 277–280. 

Mukerji KG, Bedi K, Tewari JP, Tewari I. 

1969 – Morphology of Indian species of 

Xylaria and Poronia. Phytomorphology 

19, 219–224. 

Mukerji KG, Kumar N, Singh N. 1995 – 

Studies on Indian coprophilous fungi: IV. 

Species of genera Apiosordaria and Cercophora. 

Phytomorphology 45, 87–105. 

Munk A. 1948 – Pyrenomycetes collected in 

the peninsula Mols, Jutland. Dansk Botanisk 

Arkiv 12, 1–19. 

Munk A. 1957 – Danish Pyrenomycetes. Dansk 

Botanisk Arkiv 17, 1–491. 

421

Muroi T, Udagawa S. 1984 – Some coprophilous 

Ascomycetes from Chile. In: Studies 

of Cryptogams in Southern Chile (ed H. 

Inoue) 161–167. 

Narendra DV. 1973 – Studies into coprophilous 

fungi of Maharashtra (India) III. 

Sporormia and Sporormiella: two rare 

Ascomycetes from sheep dung. Nova 

Hedwigia 24, 481–486. 

Orr GF. 1977a – A new species of Gymnoascus. 

Mycotaxon 5, 470–474. 

Orr GF. 1977b – New Gymnoascaceae. Mycotaxon 

6, 33–42. 

Orr GF, Roy K, Ghosh GR. 1977 – Gymnoascoideus, 

a new genus of the Gymnoascaceae. 

Mycotaxon 5, 459–469. 

Otani Y, Kanzawa S. 1970 – Notes on coprophilous 

Discomycetes in Japan II. Transactions 

of the Mycological Society of 

Japan 11, 43–48. 

Palliser HL. 1910 – Chaetomiaceae in: North 

American Flora 3 (1). The New York 

Botanical Garden 59–64. 

Parker AD. 1979 – Associations between 

coprophilous Ascomycetes and fecal substrates 

in Illinois. Mycologia 71, 1206– 

1214. 

Pathak RK, Agrawal SC. 1974 – Two more 

additions to Indian ascomycetes. Current 

Science 43, 440. 

Patil MS, Ghadge DN. 1987 – Studies on discomycetes-III: 

genus Ascodesmis. Indian 

Phytopathology 40, 30–35. 

Pease D. 1948 – Sphaeronemella fimicola 

(March.) emend.: some characteristics in 

culture. Mycologia 40, 114–124. 

Peck CH. 1884 – Report of the State Botanist. 

Annual Report by the Regents of the 

University of the State of New York 35, 

131–145. 

Pérez–Silva E. 1970 – Datos sobre el genero 

Poronia (Pyrenomycetes) en Mexico. 

Boletín de la Sociedad Botánica de 

Mexico 31, 139–146. 

Perry BA, Hansen K, Pfister DH. 2007 – A 

phylogenetic overview of the family Pyronemataceae 

(Ascomycota, Pezizales). 


Persoon CH. 1796 – Observationes Mycologicae, 

Pars prima. Lipsiae. 

Phillips W. 1893 – A manual of the British 

Discomycetes with description of all 

422 

species of fungi. Kegan Paul, Trench, 

Trübner & Co., London. 

Phillips W, Plowright CB. 1874 – New and 

rare British fungi. Grevillea 2, 186–189. 

Phillips W, Plowright CB. 1876 – New and 

rare British fungi. Grevillea 4, 118–124. 

Piasai O, Manoch L. 2009 – Coprophilous 

Ascomycetes from Phu Luang Wildlife 

Sanctuary and Khao Yai National Park in 

Thailand. Kasetsart Journal Natural 

Science 43, 34–40. 

Pilát A. 1972 – Contribution à l’étude des 

Basidiomycètes de la Mongolie. Bulletin 

de la Société Mycologique de France 88, 

333–356. 

Piontelli E, Alicia Toro Santa-Maria M, Caretta 

G. 1981 – Coprophilous fungi of the 

horse. Mycopathologia 74, 89–105. 

Piontelli E, Rodrigo Cruz L, Alicia Toro Santa 

Maria M. 2006 – Coprophilous fungal 

community of wild rabbit in a park of a 

hospital (Chile): a taxonomic approach. 

Boletín Micológico 21, 1–17. 

Pirotta R. 1878 – Saggio d’una monografia del 

genere Sporormia. Nuovo Giornale Botanico 

Italiano 10, 127–163. 

Prokhorov VP. 1989a – The occurrence of Thelebolaceae 

in Estonia. Proceedings of the 

Estonian Academy of Sciences, Biology 

1989, 38, 195–199. 

Prokhorov VP. 1989b – A contribution to the 

coprophilous Discomycetes in Estonia. 

Proceedings of the Estonian Academy of 

Sciences, Biology 1989, 38, 285–288. 

Rainer J, Hoog GS de. 2006 – Molecular 

taxonomy and ecology of Pseudallescheria, 

Petriella and Scedosporium prolificans 

(Microascaceae) containing opportunistic 

agents on humans. Mycological 

Research 110, 151–160. 

Raitviir A, Prokhorov V. 1988 – Discomycetes 

of Middle Asia. IV. New data on Ascobolaceae 

and Thelebolaceae. Proceedings 

of the Estonian Academy of Sciences, 

Biology 37, 215–222. 

Ramos AM, Dokmetzian DA, Ranalli ME, 

Ferreyra LI. 2008 – RAPD patterns in 

three Argentine Coprotus species: a test 

case. Mycotaxon 103, 9–19. 

Ranalli ME, Mercuri OA. 1995a – Taxonomy 

of some Argentine species of Saccobolus. 

Mycotaxon 54, 315–327.

Ranalli ME, Mercuri OA. 1995b – The conidial 

state of Selinia pulchra (Ascomycotina, 

Hypocreales). Mycotaxon 53, 109–113. 

Rattan SS, El-Buni AM. 1980 – Some new records 

of coprophilous fungi from Libya. 

II. Sydowia 33, 265–273. 

Rehm H. 1888 – Ascomyceten fasc. XIX. Hedwigia 

27, 163–175. 

Rehm H. 1889 – Ascomyceten fasc. XX. Hedwigia 

28, 347–358. 

Reid DA. 1986 – A collection of Poronia 

punctata from Surrey. Bulletin of the 

British Mycological Society 20, 58–59. 

Richardson MJ. 1972 – Coprophilous Ascomycetes 

on different dung types. Transactions 

of the British Mycological Society 

58, 37–48. 

Richardson MJ. 1998 – New and interesting 

records of coprophilous fungi. Botanical 

Journal of Scotland 50, 161–175. 

Richardson MJ. 2001a – Diversity and occurrence 

of coprophilous fungi. Mycological 

Research 105, 387–402. 

Richardson MJ. 2001b – Coprophilous fungi 

from Brazil. Brazilian Archives of Biology 

and Technology 44, 283–289. 

Richardson MJ. 2004a – Coprophilous fungi 

from Morocco. Botanical Journal of 

Scotland 56, 147–162. 

Richardson MJ. 2004b – Coprophilous fungi 

from Iceland. Acta Botanica Islandica 14, 

77–102. 

Richardson MJ. 2005 – Coprophilous fungi 

from the Faroe Islands. Fródskaparrit 53, 

67–81. 

Richardson MJ. 2007 – The distribution and 

occurrence of coprophilous Ascobolaceae. 

Mycologia Montenegrina 10, 211– 

227. 

Richardson MJ. 2008a – Coprophilous fungi 

from the Greek Aegean islands. Mycologia 

Balcanica 5, 23–32. 

Richardson MJ. 2008b – Records of French 

coprophilous fungi. Cryptogamie, Mycologie 

29, 157–177. 

Richardson MJ. 2008c – Records of coprophilous 

fungi from the Lesser Antilles 

and Puerto Rico. Caribbean Journal of 

Science 44, 206–214. 

Richardson MJ, Watling R. 1997 – Keys to 

fungi on dung. British Mycological Society, 

Stourbridge. 


Roberts RG. 1975 – The anamorph of Microascus 

intermedius Emmons & Dodge. Mycological 

Society of America Newsletter 

36, 37. 

Rossman AY, Samuels GJ, Rogerson CT, 

Lowen R. 1999 – Genera of Bionectriaceae, 

Hypocreaceae and Nectriaceae 

(Hypocreales, Ascomycetes). Studies in 

Mycology 42, 1–248. 

Roy I. 1971 – Coprophilic Ascomycota from 

Scotland I. Nova Hedwigia 19, 883–886. 

Saccardo PA. 1878 – Enumeratio Pyrenomycetum 

Hypocreaceorum hucusque cognitorum. 

Michelia 1, 277–325. 

Sage L, Steiman R, Seigle-Murandi F, Guiraud 

P. 1995 – Description and physiological 

properties of two new varieties and one 

species of Microascus from Israel. Mycotaxon 

55, 189–201. 

San Martín F, Lavin P, Perez Silva E. 1998 – 

Xylariaceae fimicolas: Xylaria equina sp. 

nov. y nuevos registros mexicanos de 

Xylaria pileiformis y Poronia erici. Acta 

Botanica Mexicana 42, 15–23. 

Saxena AS, Mukerji KG. 1973 – Fungi of 

Delhi. XVII. Three unrecorded coprophilous 

Ascomycetes. Česká Mykologie 27, 

165–168. 

Schavey J. 1994 – Poronia erici Lohmeyer & 

Benkert. Sterbeeckia 16, 30–34. 

Schavey J. 1999 – Microfungi op konijnenkeutels. 

AMK Mededelingen 1, 15–21. 

Schmidt A. 1912 – Die Verbreitung der coprophilen 

Pilze Schlesien. Breslau. 

Schoch CL, Crous PW, Groenewald JZ, 

Boehm EW, Burgess TI, de Gruyter J, de 

Hoog GS, Dixon LJ, Grube M, Gueidan 

C, Harada Y, Hatakeyama S, Hirayama 

K, Hosoya T, Huhndorf SM, Hyde KD, 

Jones EB, Kohlmeyer J, Kruys A, Li 

YM, Lücking R, Lumbsch HT, 

Marvanová L, Mbatchou JS, McVay AH, 

Miller AN, Mugambi GK, Muggia L, 

Nelsen MP, Nelson P, Owensby CA, 

Phillips AJ, Phongpaichit S, Pointing SB, 

Pujade-Renaud V, Raja HA, Plata ER, 

Robbertse B, Ruibal C, Sakayaroj J, Sano 

T, Selbmann L, Shearer CA, Shirouzu T, 

Slippers B, Suetrong S, Tanaka K, 

Volkmann-Kohlmeyer B, Wingfield MJ, 

Wood AR, Woudenberg JH, Yonezawa 

H, Zhang Y, Spatafora JW. 2009 – A 

423

class-wide phylogenetic assessment of 

Dothideomycetes. Studies in Mycology 

64, 1–15. 

Schroeter J. 1893 – Die Pilze Schlesiens. In: 

Kryptogamen-Flora von Schlesien 2 (ed. 

F Cohn). JU Kern's Verlag, Breslau 1– 

597. 

Schumacher T. 1990 – The genus Scutellinia 

(Pyronemataceae). Opera Botanica 101, 

1–107. 

Scott JA, Untereiner WA. 2004 – Determination 

of keratin degradation by fungi using 

keratin azure. Medical Mycology 42, 

239–246. 

Seaver FJ. 1928 – The North-american cupfungi 

operculates. (ed. FJ Seaver) New 

York. 

Seth HK. 1967 – Chaetomidium subfimeti sp. 

nov. from Wales. Transactions of the 

British Mycological Society 50, 45–47. 

Seth HK. 1968 – Coprophilic Ascomycota 

from Germany. Nova Hedwigia 16, 495– 

499. 

Seth HK. 1970 – A monograph of the genus 

Chaetomium. Nova Hedwigia, suppl. 37, 

1–133. 

Seth HK. 1971 – The genus Lophotrichus 

Benjamin. Nova Hedwigia 19, 591–599. 

Seth HK. 1984 – Revision of herbarium 

specimens of genus Chaetomium Kunze 

(Ascomycetes) at the Herbarium Hamburgense 

(HBG). Mitteilungen aus dem 

Institut für Allgemeine Botanik Hamburg 

19, 165–174. 

Shubakov AA, Kucheryavykh PS. 2004 – 

Chitinolytic activity of filamentous fungi. 

Applied Biochemistry and Microbiology 

40, 517–519. 

Sigler L, Flis AL, Carmichael JW. 1998 – The 

genus Uncinocarpus (Onygenaceae) and 

its synonym Brunneospora: new concepts, 

combinations and connections to 

anamorphs in Chrysosporium, and further 

evidence of relationship with Coccidioides 

immitis. Canadian Journal of Botany 

76, 1624–1636. 

Singh N, Mukerji KG. 1979 – Studies on 

Indian coprophilous fungi. I. Some rare 

records. Journal of the Indian Botanical 

Society 58, 163–167. 

Siquier JL, Lillo F. 1994 – Contribución al 

conocimiento micológico del Parque 

424 

Nacional del Archipiélago de Cabrera 

(Islas Baleares, España). Boletín de la 

Sociedad Micológica de Madrid 19, 193– 

205. 

Skolko AJ, Groves JW. 1953 – Notes on seedborne 

Fungi. VII. Chaetomium. Canadian 


Skou JP. 1973 – Microascus exsertus sp. nov. 

associated with a leaf-cutting bee, with 

considerations on relationships of species 

in the genus Microascus Zukal. Antonie 

van Leeuwenhoek 39, 529–538. 

Slupinski L. 1991 – Quelques Ascomycetes 

stercoricoles. Société d'Histoire Naturelle 

du Pays de Montbéliard 1991, 73–96. 

Soláns MJ. 1994 – Los géneros Orbicula 

Cooke, Roumegueriella Speg. y 

Zygopleurage Boedijn, novedades para el 

catálogo micológico español. Studia 

Botanica 13, 245–248. 

Solé M, Cano J, Pitarch LB, Stchigel AM, 

Guarro J. 2002 – Molecular phylogeny of 

Gymnoascus and related genera. Studies 


Spatafora JW, Blackwell M. 1994 – The polyphyletic 

origins of ophiostomatoid fungi. 


Spegazzini C. 1878 – Fungi coprophili veneti. 

Pugillus I. Michelia 1, 222–238. 

Spegazzini C. 1909 – Mycetes Argentinenses 

(ser. IV). Anales del Museo Nacional de 

Buenos Aires ser. 3, 12, 257–458. 

Spooner BM, Butterfill GB. 1999 – Coprophilous 

discomycetes from the Azores. Kew 

Bulletin 54, 541–560. 

Stchigel AM, Guarro J, Jato V, Aira MJ. 2004 

– Two new species of Chaetomidium 

(Sordariales). Studies in Mycology 50, 

215–220. 

Stiers DL. 1974 – Fine structure of ascospore 

formation in Poronia punctata. Canadian 


Stolk AC. 1955 – The genera Anixiopsis Hansen 

and Pseudeurotium van Beyma. 

Antonie van Leeuwenhoek 21, 65–79. 

Stratton R. 1921 – The Fimetariales of Ohio. 

Ohio Biological Survey 3(2), 12, 75–144. 

Subramanian CV, Lodha BC. 1968 – Two interesting 

coprophilous fungi from India. 

Current Science 37, 245–248. 

Sugiyama M, Mikawa T. 2001 – Phylogenetic 

analysis of the non-pathogenic genus

Spiromastix (Onygenaceae) and related 

onygenalean taxa based on large subunit 

ribosomal DNA sequences. Mycoscience 

42, 413–421. 

Sugiyama M, Ohara A, Mikawa T. 1999 – Molecular 

phylogeny of onygenalean fungi 

based on small subunit ribosomal DNA 

(SSU rDNA) sequences. Mycoscience 

40, 251–258. 

Sugiyama M, Summerbell RC, Mikawa T. 

2002 – Molecular phylogeny of onygenalean 

fungi based on small subunit (SSU) 

and large subunity (LSU) ribosomal 

DNA sequences. Studies in Mycology 

47, 5–23. 

Swiontek-Brzezinska M, Lalke-Porczyk E, 

Donderski W. 2007 – Chitinolytic activity 

of bacteria and fungi isolated from 

shrimp exoskeletons. Oceanological and 

Hydrobiological Studies 36, 101–111. 

Thind KS, Kaushal SC. 1978 – Three species 

of Iodophanus from western Himalaya. 

Indian Phytopathology 31, 343–347. 

Thirumalachar MJ, Mathur PN. 1965 – 

Narasimhella, a new genus of the 

Gymnoascaceae. Sydowia 19, 184–186. 

Tieghem P van. 1877 – Sur le developement de 

quelques Ascomycetes. Bulletin de la Société 

Botanique de France 24, 159–161. 

Tornabene FA, Davis JS. 1966 – Lophotrichus 

brevirostratus, a fungal species new to 

Illinois. Transactions of Illinois Academy 

of Sciences 59, 396. 

Tóth S. 1963 – Data to the knowledge on the 

coprophilous microscopic fungi in 

Hungary I. Annales Historico-Naturales 

Musei Nationalis Hungarici Pars Botanica 

55, 181–185. 

Tóth S. 1965 – Data to the knowledge on the 


Hungary II. Annales Historico-Naturales 


57, 149–157. 

Tóth S. 1967 – Data to the knowledge of the 


Hungary III. Annales Historico-Naturales 


59, 117–123. 

Traverso JB. 1907 – Flora Italica Cryptogama. 

Pars I: Fungi. Pyrenomycetae. Rocca S. 

Casciano 2, 1–701. 


Udagawa S. 1960 – A taxonomic study on the 

Japanese species of Chaetomium. Journal 

of General and Applied Microbiology 6, 

223–251. 

Udagawa S. 1962 – Microascus species new to 

the mycoflora of Japan. The Journal of 

General and Applied Microbiology 8, 

39–51. 

Udagawa S. 1980 – Some new or noteworthy 

coprophilous Pyrenomycetes from South 

America. Transactions of the Mycological 

Society of Japan 21, 283–298. 

Udagawa S, Cain RF. 1969 – Some new or 

noteworthy species of the genus Chaetomium. 

Canadian Journal of Botany 47, 

1939–1951. 

Udagawa S, Furuya K. 1972 – Notes on some 

Japanese Ascomycetes X. Transactions 

of the Mycological Society of Japan 13, 

49–56. 

Udagawa S, Furuya K. 1988 – Emericellopsis 

sphaerospora and Kernia peruviana, two 

new soil-borne cleistothecial Ascomycetes. 

Mycotaxon 33, 291–301. 

Udagawa S, Muroi T. 1979 – Coprophilous 

Pyrenomycetes from Japan V. Transactions 

of the Mycological Society of Japan 

20, 453–468. 

Udagawa S, Muroi T. 1981 – Notes on some 

Japanese ascomycetes XVI. Transactions 

of the Mycological Society of Japan 22, 

11–26. 

Udagawa S, Sugiyama Y. 1982 – New records 

and new species of ascomycetous microfungi 

from Nepal, a preliminary report on 

the expedition of 1980. Reports on the 

Cryptogamic Study in Nepal, March 

1982 (Miscellaneous Publications of the 

National Science Museum, Tokyo), 11– 

46. 

Udagawa S, Uchiyama S. 1999 – Taxonomic 

studies on new or critical fungi of nonpathogenic 

Onygenales 1. Mycoscience 

40, 277–290. 

Ulfig K, Guarro J, Cano J, Gené J, Vidal P, 

Figueras MJ, Lukasik W. 1998 – A 

preliminary study of the occurrence of 

actidione-resistant fungi in sediments of 

Catalonian river mouths (Spain). I. Keratinolytic 

fungi and related Onygenales. 

Mycopathologia 141, 143–151. 

425

Valldosera M. 1991 – Contribución al estudio 

de los hongos coprófilos: Ascomycotina, 

en España. Universitat Autonoma de 

Barcelona (unpublished degree thesis). 

Valldosera M, Guarro J. 1984a – Estudios 

sobre hongos coprófilos aislados en 

España, II. Ascomycetes. Revista Iberoamericana 


Valldosera M, Guarro J. 1984b – Estudios 

sobre hongos coprófilos aislados en 

España, IV. Ascomycetes. Revista Iberoamericana 


Valldosera M, Guarro J. 1985 – Estudios sobre 

hongos coprófilos aislados en España. III. 

Discomycetes. Boletín de la Sociedad 

Micológica Castellana 9, 37–44. 


hongos coprófilos aislados en España 

VIII. Hypocopra festucacea Krug & 

Cain. Revista Iberoamericana de Micología 

4, 85–86. 


hongos coprófilos aislados en España. 

VI. Ascomycetes. Boletín de la Sociedad 


Valldosera M, Guarro J. 1990a – Estudios sobre 

hongos coprófilos aislados en España, 

XI. Ascomycetes. Boletín de la Sociedad 


Valldosera M, Guarro J. 1990b. Estudios sobre 


XV. El Género Preussia (Sporormiella). 

Boletín de la Sociedad Micológica de 

Madrid 14, 81–94. 



XVII. Ascomycotina. Boletin de la Sociedad 


Valmaseda M, Martínez AT, Barrasa JM. 1987 

– Annellidic conidiogenesis in Pithoascus 

schumacheri and redefinition of 

Pithoascus and related fungi. Canadian 


Verona O, Picci G, Mazzucchetti G. 1967 – On 

the cellulophilic behaviour of some species 

of Microasacus. Mycopathologia 32, 

302–303. 

Vesterholt J. 2000 – Usaedvanlige danske 

svampefund. Svampe 42, 8–10. 

Vooren N van, Gaignon M. 2002 – Compterendu 

de sortie mycologique 20 avril 

2002. “A la recherche des Ascomycètes 

426 

de printemps”. Bulletin de la Société 

Linnéenne de Lyon 71, 330–334. 

Wang YZ. 1992 – New records of coprophilous 

Pyrenomycetes from Taiwan (I). Transactions 

of the Mycological Society 

ROC. 7, 18–22. 

Wang YZ. 1993 – Notes on coprophilous discomycetes 

from Taiwan. I. Bulletin of the 

National Museum of Natural Science 4, 

113–123. 

Wang YZ. 1994 – New records of coprophilous 

Pyrenomycetes from Taiwan (II). Transactions 

of the Mycological Society 

R.O.C. 9, 185–194. 

Wang YZ. 1995 – Notes on coprophilous 

discomycetes from Taiwan. II. Bulletin 

of the National Museum of Natural 

Science 5, 147–152. 

Wang YZ. 1999 – The coprophilous discomycetes 

of Taiwan. Bulletin of the National 

Museum of Natural Science 12, 49–74. 

Wang YZ. 2000 – The genus Podospora in 

Taiwan. Mycotaxon 76, 373–391. 

Wang Z, Wang YZ. 2000 – Notes on coprophilous 

discomycetes from southwestern 

China. Fungal Science 15, 125–134. 

Weber RWS, Webster J. 1997 – The coprophilous 

fungus Sphaeronemella fimicola, a 

facultative mycoparasite. Mycologist 11, 

50–51. 

Weber RWS, Webster J. 1998 – Stimulation of 

growth and reproduction of Sphaeronemella 

fimicola by other coprophilous 

fungi. Mycological Research 102, 1055– 

1061. 

Webster J. 1970 – Coprophilous fungi. Transactions 

of the British Mycological 

Society 54, 161–180. 

Welt P, Heine N. 2006a – Beiträge zur 

Kenntnis coprophiler Pilze (2): 

Dungbewohnende Pilze Thüringens: Teil 

1. Hoher Artenreichtum coprophiler Pilze 

in einem Schutzgebiet: Indicator für eine 

intakte Natur? Boletus 29, 81–92. 

Welt P, Heine N. 2006b – Beiträge zur 

Kenntnis coprophiler Pilze 3. Coprophile 

Pilzfunde im Montafon (Vorarlberg, 

Österreich). Österreichische Zeitschrift 

für Pilzkunde 15, 213–224. 

Welt P, Heine N. 2007 – Beiträge zur Kenntnis 

coprophiler Pilze (1). Teil 2: Coprophile 

Pilzfunde im Chemnitzer NSG “Um den

Eibsee” auf verchiedenen Substraten 

sowie Ergänzungen zu den Pilzfunden 

auf Angusrind-Dung. Zeitschrift für 

Mykologie 73, 213–244. 

Whalley AJS, Dickson GC. 1986 – Poronia 

punctata, a declining species? Bulletin of 

the British Mycological Society 20, 54– 

57. 

Whiteside WC. 1962 – Morphological studies 

in the Chaetomiaceae. II. Mycologia 54, 

152–159. 

Wicklow DT. 1992 – The coprophilous fungal 

community: an experimental system. In: 

The Fungal Community. Its Organization 

and Role in the Ecosystem. 2nd edn. (eds 

GC Carrol, DT Wicklow). Marcel 

Dekker Press, New York 715–728. 

Wicklow DT, Angel K, Lussenhop J. 1980 – 

Fungal community expression in lagomorph 

versus ruminant feces. Mycologia 

72, 1015–1021. 

Willdenow CL. 1787 – Florae berolinensis Prodromus. 

Berolini. 


Wilson CM. 1947 – Coprophilous Ascomycetes 

of Virginia. Mycologia 39, 374– 

377. 

Winter G. 1873 – Die deutschen Sordarien. 

Abhandlungen der Naturforschenden 

Gesellschaft zu Halle 13, 1–43. 

Winter G. 1874 – Mykologische notizen. Hedwigia 

13, 50–57. 

Winter G. 1887 – Pilze. In Dr. L. Rabenhorst’s 

Kriptogamen-Flora von Deutschland, 

Oesterreich und der Schweiz. Leipzig. 

Zhang Y, Schoch CL, Fournier J, Crous PW, 

de Gruyter J, Woudenberg JH, Hirayama 

K, Tanaka K, Pointing SB, Spatafora JW, 

Hyde KD. 2009 – Multi-locus phylogeny 

of Pleosporales: a taxonomic, ecological 

and evolutionary re-evaluation. Studies in 

Mycology 64, 85–102. 

Zukal H. 1885. Ueber einige neue Pilze, Myxomyceten 

und Bakterien. Verhandlungen 

der Zoologisch-Botanischen Gesellschaft 

in Wein 35, 333–342. 

427

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