THESIS 4

UNIVERSITETET I BERGEN - BOTANISK INSTITUTT

OLAV AAS

AWORLD - MONOGRAPH
OFTHE
GENUS THECOTHEUS (ASCOMYCETES, PEZIZALES)

BERGE:\ 1992
 1

A WORLD - 1I0HOGRAPH OF DE GEliUS 'l'HECOTHEUS (ASCOKYCBTES,

PEZIZALES)

by

OLAV AAS

A thesis submitted for the degree of

doctor scientiarum, to the Botanical
Institute, University of Bergen,
1992.
 2

ABSTRACT

The genus Thecotheus (Ascobolaceae, Pezizales) is monographed

on a world basis. Seventeen species are recognized. Three new
species of Thecotheus are described: T. inaeguilateralis Aas,
T. lundqvistii Aas, and T. uncinatus Aas. In addition, one
species of Iodophanus is described as new: I. magniverrucosus
Aas. Four new combinations are made: T. biocellatus (Petrak)
Aas, T. crustaceus (Starb.) Aas & Lundq., T. keithii (Phill.)
Aas and T. strangulatus (Vel.) Aas & Lundq. The two species T.
africanus and T. perplexans are regarded as vulnerable. T.
biocellatus and T. pallens are apparently rare species. A key
to the species is provided. Morphological and anatomical
characters are given for all species, and their ecology and
distribution are discussed.
 3

CONTENTS

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . 5
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . • . • . . . . . . . . • . • • • . . . . 6
A. Historical survey ..............••.........•..•...• 6
B. Aims of the study ................•..•••.••.••••.•. 12
Material and methods ........•...•....•••...•..••.•••••••• 13
A. Methods ...................•.....•...••.•..••.....• 13
B. Material .....................•.....••.•.•••.••••.. 16
Morphology and anatomy .............•..••...•.•••...•••.•• 17
A. Ascoci'lrp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . • . . • • . . . . 17
B. Excipulur;; ....................••••...•...••••••.••• 18
c. Asci .....•.•....................••..•••••.•.•••••. 20
D. Ascospores .................••.......•....••.••..•• 21
E. Paraphyses •...............•......•..•..••••....... 22
F. Anamorphs •..•.....•.............•...•.•..••.....•. 23
Cultures • . • . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • . • . . • . . . . 23
Ecology and distribution . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . 25
A. Substrate preference ..........•....••...•••....••• 25
1. Du:-,:; species ~ . . . . . . . . . . . •. . . ••. . . . •. •. • 25
2. Debris species ............••.••...•....•••••.• 29
B. Distribution ....••.........•......•.••.••••..••••. 29
Taxonomy • . . . . . . . . . . . . . • . . . . . . . . . . . . • . . . . . . • . • . . . . . . . . . . • • 32
Key to the accepted species .........•....•....•..•..•••.. 39
1. Thecotheus africanus Khan & Krug ...•.•..•........• 42
2. T. bi::,c:cll3tus (Petrak) Aas n. comb . . . • . . . . . . . . . . • 46
3. T. Clnereus (Cr. & Cr.) Chenant . . . . . . . . . . . . . . . . . • . 54
4. T. crustaceus (Starb.) Aas & Lundq. n. comb . . • . . . . 70
5. T. ~i~~layensis Kaushal . . . . . . . . . . . . . . . . • . . . . . . . . . • 88
6. T. holmskjoldii (E. C. Hansen) Chenant . . • . . • ~.~ ... 93
7. T . .inaequ.ilateralis· Aas sp. novo ....••..•.••..••.. 108
8. T. keithii (Phill.) Aas n. comb . . . . . . • . . . . . . . . . . . . 110
9. T. l1.JJ"lc'1vistii Aas sp. novo .........•..•....••.... 127
 4

10. T. pallens (Boud.) Kimbr 131

11. T. pelletieri (Cr. & Cr.) Boud ~ .. 133
12. T. perplexans (Faurel & Schotter) Krug & Khan •.... 149
13. T. phycophilus Pfister .................•.....••••. 152
14. T~ rivicola (Vacek) Kimbr. & Pfister ...•.••.•.••.. 160
15. T. strangulatus (Vel.) Aas & Lundq. n. comb . . . . • . . 170
16. T. uncinatus Aas sp. novo ..•...•.......•...•...... 175
17. T. viridescens E. Ludwig •.........•••...........•. 181
Appendices ......••••.•••....•..................•......•.. 183
A. Uncertain names . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . 183
Ascophanus isabellinus Clements ...•......••..... 183
Thecotheus kimbroughii Donadini •.•••..•••••••••. 186
B. Excluded taxa .................•............•••.... 188
Literature .....•............•........•...•........••.•.•• 193.
Index •.•........•..........................•...•..••..••. 209
 5

ACKNOWLEDGEMENTS

I wish particularly to express my gratitude to Professor Nils

Lundqvist, stockholm, who proposed the genus Thecotheus as the
sUbject for my thes~s. We have had several valuable
discussions in stockholm, and Lundqvist also prepared lists of
literature-records for most of the relevant taxa. In addition
he placed much material at my disposal. Likewise, I am much
indebted to Professor P. M. J0rgensen, Bergen, for
encouragement, advice and comments on the paper. Without their
participation it would not have been possible to finish this
work.

I have received valuable help and information from many

persons and institutions. I am particularly grateful to Or J.
M. Barrasa, Madrid; Or J. van Brummelen, Leiden; Or F.-E.
Eckblad, Oslo; Or S. Goebel, Lincoln, Nebraska; Or J. W.
Kimbrough, Gainsville, Florida; Or J. C. Krug, Toronto; Or Y.
Le Gal, Concarneauj Or P. Neville, Marseille; Or V. P.
Prokhorov, Moscow and Dr A. Y. Rossman, Beltsville, Maryland.

Moreover, I am indebted to the directors, curators and staff

members at the herbaria or private persons listed under
"Material", for making specimens available for loan. Special
thanks also to Mrs. M. K. Stavdal, Mr. J. Berge, Mrs. S.
Herland and Miss o. BIUcher, all Bergen, for technical
assistance with the photographs, and to the librarians at the
University Library, Bergen. I also wish to thank Or. Hilary H.
Birks, Bergen, for revision of the English text.

The work was carried out while I was working at the Botanical
Institute, or at the University Library of Bergen, or at Sogn
and Fjordane College, Sogndal. Financial support was received
from the University of Bergen, Olaf Grolle Olsens fund, Sogn
and Fjordane College, and the Nansen fund.
 6

INTRODUCTION

A. HISTORICAL SURVEY

The genus Thecotheus was established by Boudier in 1869 for

Thecotheus ~elletieri, based on Ascobolus pelletieri of the
Crouan brothers (Crouan & Crouan 1857). This operculate,
coprophilotls discomycete with 32-spored asci, was the only
species Boudier (1869) included in the new genus. The genus
was characterized chiefly by numerous long and thin
paraphyses, its abundant gelatinous matrix and ..• "surtout par
son disque erumpent, caractere qui se voit mieux dans les
jeunes individus". In this work, "Memo ire sur les Ascoboles",
Boudier included genera with large, operculate, protruding
asci in the "Ascobolei", sectio "Pezizearum". He then further
divided the "Ascobolei" into two groups, namely "Ascobolei
genuini" with coloured ascospores, and "Ascobolei spurii"
characterized by hyaline ascospores.

Together with the genera Ascophanus Boud. and Ryparobius

Boud., Boudier (op. cit.) placed the newly erected genus
Thecotheus in the group "Ascobolei spurii". Boudier (1~69:
235) mentioned that Thecotheus, owing to the large protruding
asci, was close to the genus Ascobolus (treated in "Ascobolei
genuini"), but differed from that genus in lacking epispore
pigmentati.c::. C:1 ~.he ascospores.

Karsten (1867, 1871 a, 1871 b, 1871 c) and Fuckel (1870) were

un~ware of the work of Boudier. Fuckel (op. cit.) placed
Ascobolus pel~2tieri Crouan in the Bulgariacei. Although
Karsten (1885) later included the generic names of Boudier, he
did not mentioned the genus Thecotheus. Neither Spegazzini
(1878) nor Gillet (1879) used the generic name Thecotheus.
 7

Spegazzini (op. cit.) placed the type species in the genus

Pezizula P. Karst. emend. Speg., whereas Gillet (op. cit.)
placed the type species in the genus Mollisia in the family
pezizees.

Bommer & Rousseau (1884) placed Thecotheus, then including

only the original species, as a sUbgenus of Ascobolus. Boudier
(1885) raised the Ascobolei to the rank of a family, and
divided the family Ascoboles into two groups (subfamilies),
namely Ascoboles with pigmented spores, and Pseudoascoboles
with colorless ascospores. The latter included the genus
Thecotheus.

Quelet (1886) placed Thecotheus as one of the synonyms of the

sUbgenus Ascophanus Quelet in the genus Ascobolus. Heimerl
(1889) treated Thecotheus as a section of the genus
Ascophanus, and only the type species was included in the
section. Saccardo (1889), on the other hand, treated
Thecotheus as a (?)subgenus of Ryparobius. In addition to the
type species, Ryparobius winteri March. was also placed in
this (?)subgenus.

Schroeter (1893) treated Thecotheus (type species only) as a

genus [Gattung] of the family Ascobolacei. Massee (1895)
treated Thecotheus (type species), together with Ascozonus
Renny, as a synonym of Ryparobius in the family Ascoboleae.
Rehm (1896) also considered Thecotheus (type species) as a
synonym of Rhyparobius in the Pseudoascobolae of the
Ascoboleae. Lindau (1897) placed Thecotheus in a section of
Rhyparobius, family Ascobolaceae, including only the type
species. Lindau (1912) later omitted Thecotheus and named the
type species Rhyparobius pelletieri in the genus Rhyparobius
of the. family Ascobolaceae. Nor did Oudemaris (1900) adopt the
genus Thecotheus; he named the type species as Ryparobius
pelletierii (sic!) in the Ascobola6ees. Se aver (1904) plac~d

the type species of Thecotheus, together with other

mUltispored species, within the genus Ryparobius in the family
 8

Ascobolace~.

Overton (1906) treated Thecotheus in the family Ascobolaceae.

Later, Boudie~ (1907) included another mUltispored member in
the genus, with the combination of a species originally descr-
ibed as Ryparobius winteri March. (Marchal 1885). This
species, which contains approximately 256 spores in the asci,
was later transferred to the genus Coprotus by Kimbrough
(1967). As before, Boudier (op. cit.) placed the two species
in the tribus pseudo-Ascobolees of the family Ascobolacees.
Three other species, later included in Thecotheus, were at
that ti~e by Boudier (1907) put in the genus AscophaQus of the
same tribus.

Migula (1913) followed Saccardo (1889) in considering

Thecotheus (type species) as a synonym of Rhyparobius in the
Gattung [genus] Rhyparobius, Subgenus Thecotheus, of the
family Ascobolaceae. Ramsbottom (1914) on the contrary
followed Boudier (1907).

Chenantais (1918r broadened the concept of Thecotheus with the

inclusion of the coprophilous species Ascobolus cinereus Cr.
&. Cr. and Ascophanus holmskjoldii Hans. with these transfers,
Chenantais included the first 8-spored members of a typical,
at that time, multispored group. Le Gal (1960, 1963) accepted
this transfer, but most of the later authors continued to
treat both these species in the genus Ascophanus for several
decades.

Seaver (1927, 1928) treated Thecotheus in the tribe Humarieae

of the family Pezizaceae. He accepted only the type species in
the genus (Seaver 1928). The two additional species mentioned
above which were included in the genus Thecotheus ten years
earlier by Chenantais (1918), were placed in the genus
Ascophanus of the tribe Humarieae by Seaver (1928).

Velenovsky (1934) omitted the generic name Thecotheus. He

 9

placed the type species in the genus Rhyparobius, while T.

holmskjoldii was placed in the genus Ascophanus. According to
him, both genera belonged to the family Ascoboleae. Rostrup
(1935) also' omitted the generic name Thecotheus, and placed
the only species of Thecotheus he included, Ascophanus
holmskioldii, 'in the family Pezizaceae. Svr~ek & Kubi~ka
(1961) treated Thecotheus (type species) in the subfamily
Humarioideae of the family Humariaceae.

In his studies of selected Pseudoascoboleae (= Theleboleae),

Kimbrough (1966) agreed with Chenantais (191S) and Le Gal
(1960, 1963) and included both's-spored and multispored
species in the genus Thecotheus. Kimbrough found that the
genus was distinct from the other Pseudoascoboleae, and later,
therefore, Kimbrough & Korf (1967) transferred the genus to
the tribe Pezizeae of the family Pezizaceae. This
was mainly based upon the blueing of asci in iodine, the
presence of callose-pectate markings and apiculi on the
ascospores, and the lack of spore pigmentation. Two species
were considered by the authors, namely I. pelletieri (Cr. &
Cr.) Boud. and I. cinereus (Cr. & Cr.) Chenant. Mainly on
account of the diffuse blueing of asci in iodine, Rifai (196S)
agreed with Kimbrough & Korf (1967).

Brumrnelen (1967) treated Thecotheus together with several

other genera in his new subfamily Theleboloideae Brumrn. of the
family Ascobolaceae. However, the author pointed out that the
subfamily badly needed a thorough revision. Eckblad (196S)
disagreed ~ith Kimbrough & Korf (1967) on several points,
namely that the very simple excipulum structure of Thecotheus
was not comparable to the complex type of the Peziza-group,
and that Thecotheus possessed mostly mUltispored and
protruding asci, characters quite different from any genera in
the family Pezizaceae. Eckblad (op. cit.) therefore elevated
the subfamily Theleboloideae Brurnrn. to family rank, and
replaced the genus Thecotheus in the new family Thelebolaceae
(Brumrn.) Eckbl. Three species were mentioned by Eckblad. He
 10

was later followed by Moravec (1971), Merkus (1976), Waraitch

(1977), and Prokhorov & Taslakhchyan (1987).

An important regional treatise of the genus was made by

Kimbrough (1969), dealing with morphology and taxonomy of the
North American species. Two new species, T· agranulosus and T.
apiculatus were described by him. Kimbrough also made a new
combination in the genus, based upon Ascophanus pallens, a
species of Boudier (1888). He made the transfer of this non-
coprophilous species on the basis of Boudier's illustrations
and descriptions. Five taxa were recognized by Kimbrough, who
followed Kimbrough & Korf (1967) in placing the genus in the
tribe Pezizeae of the family Pezizaceae. Similarly, Kimbrough
(1970 a, 1970 b, 1972, 1974) maintained the position of the
genus Thecotheus in the family Pezizaceae.

Korf (1972, 1973) included Thecotheus together with i. a.

Iodophanus in the tribe Iodophaneae Korf of the family
Ascobolaceae, all genera with permanently hyaline ascospores
and asci diffusely blue in iodine. His system was essentially
followed in Hirsch (1984).

with the transfer of Psilopezia rivicola, a species found on

water-soaked twigs (Vac~k 1949), pfister (1972) included
another non-coprophilous species in the genus Thecotheus.
still another new species, T. phycophilus was described from
dead herbaceous plants by Pfister (1981). He followed Korf
(1972) in placing the genus in the tribe Iodophaneae of the
Ascobolaceae. Conway (1975) also agreed with this placement of
Thecotheus. He concluded that diffusely amyloid asci, thick-
walled ascospores, bulbous excipular cells, a perisporic
sheath that encrusts the mature ascospores and several other
charact~rs, are features in common with the Ascobolaceae.
However, he indicates a relationship with the pyronemataceae
on account of the sympodial conidia.

Dennis (1978, 1968) still included Thecotheus in the tribe

 11

Pseudoascoboleae of the family Ascobolaceae, but he called

attention to the probable heterogeneity of the tribe.

Kaushal (1980) described a new coprophilous species of

Thecotheus from India, T. himalayensis. He followed Korf
(1972) in including Thecotheus in the tribe Iodophaneae of the
family Ascobolaceae. The main reasons for this were the
presence of amyloid asci, the simple excipular structure and
small apothecia.

On the basis of septal ultra structures in reproductive and

somatic tissues in the apothecia of species of Ascobolus,
Saccobolus, Thecotheus and Iodophanus, Kimbrough & Curry
(1985) decided that Thecotheus should be retained in the
family Ascobolaceae in accordance with Boudier's (1869)
original concept. One species of Thecotheus was examined,
namely T. pelletieri. The authors considered Iodophanus as a
member of the Pezizaceae. with reference to Dissing (in
litt.), Eriksson & Hawksworth (1987) considered that
Iodophanus is near the genus Thecotheus, and both should
probably be referred to the Ascobolaceae. Eriksson &
Hawksworth (1990, 1991) also retained the genus in the
Ascobolaceae.

Krug & Khan (1987) described Thecotheus africanus as a new

coprophilous species from Africa. In.addition, they made the
combination T. perplexans, based on Ascophanopsis perplexans
(Faurel & Schotter 1965 b), a dung-inhabitating species also
from Africa. Krug & Khan (op. cit.) presented a comprehensive
key to all ten known species of Thecotheus, and recognized the
genus as a member of the family Ascobolaceae.

Hohmeyer et al. (1989) made a key to 11 species of Thecotheus.

One of the species, the coprophilous T. viridescens was
described as new.

Korf & Zhuang (1991), in their preliminary discomycete flora

 12

of Macaronesia, placed Thecotheus in the family Ascobolaceae,

together with the genera Ascobolus and Saccobolus. Only two
species of Thecotheus were mentioned, namely T. holmskjoldii
and T. pelletieri.

B. AIMS OF THE STUDY

Although species of the genus have been treated by recent

authors, a critical study of the literature and herbaria
revealed a number of misinterpretations. It also became
evident that several more or less forgotten taxa in other
genera needed to be transferred to Thecotheus.

Accordingly it proved necessary to:

I. Get a better taxonomic understanding of the genus, and its

morphological characters.

Since no one has attempted a survey of this genus, and several

misunderstandings based on interpretations of insufficient
descriptions occur, it was necessary to examine the characters
carefully and discuss them.

11. Examine, as far as possible, the type specimens.

Many authors have neglected to study the original material and

have based their views on literature only. Thus many
misinterpretations have been created and have persisted until
the present. I have therefore particularly searched for and
made careful studies of type material, and have discussed them
in detail.
 13

I l l . Attach importance to ecological aspects.

Data on substrate is important for an understanding of both

the taxa and their evolution, and is ~hus of practical
taxonomic value 'at the spec~es level. Unfortunately, this
subject has been poorly treated in many pUblications.

IV. Indicate areas for further research.

I have attempted a first general classification. of the genus

and its species by classical methods. Numerical and
statistical methods in some of the critical or variable
species would certainly give additional information on
variation particularly within the species.

Experimental work, including studies of genetics, cytology and

chromosomes would further improve our knowledge. Also cultural
work, including studies of the ontogeny is needed. Since I
mainly worked with herbarium specimens, this was not possible
in this study.

IlATBRIAL AIfD IIBTlIODS

A. IIBTBODS

The majority of the examined material consists of dried

herbarium specimens. In a few cases, fruit-bodies were
preserved in various liquids, or as microscope slides. Only a
 14

small part of the investigation was carried out on living

material, including cultures.

Dried material was moistened before examination. Generally

apothecia were rehydrated in water. Occasionally the dried
herbarium species were first moistened with 96 percent
alcohol. In a few cases, parts of apothecia were soaked in 5
percent aqueous KOH, or in 4 percent aqueous NH 4 0H, and then
transferred to water.

In addition to water, microscopic examination was made in

different staining or mounting agents: Melzer's reagent, 2
percent Congo red, Sudan IV and 1 percent Cotton blue in
lacto-phenol. occasionally other dyes or media were used: acid
fuchsin, Lugols solution, Shears mounting medium, 60 percent
lactic acid, Indian ink/water and saffranin followed by fast
green. Material for permanent slides was mounted in polyvinyl-
lactophenol and the cover glass was sealed with nail polish.

Macroscopic observations of apothecia were made with a Leitz

stereomicroscope. Microscopic examinations were from squash
mounts or sections, including both free hand sections and
frozen sections. The latter sections were made with a Leitz
1310 K Freezing Microtome connected to a Kryomat 1703. As a
routine, examination of excipular tissue was prepared by
frozen sections mounted in Cotton blue in lactophenol and then
gently heated. Measurements of the different elements were,
whenever possible, obtained from material in water. Size of
ascospores, excluding ornamentations and appendages, was based
on a minimum of 10 samples from each collection examined. As
regards size of asci and paraphyses, about five different
measurements were taken from each collection. Diverging
measurements are given in parentheses.

Drawings were made by means of a Zeiss drawing tu pe mounted on

a Zeiss standard 19 ·microscope. Photomicrographs were taken
either with a Nikon camera connected to the microscope with a
 15

phototube, or with the aid of a Zeiss Axiomat microscope with

built-in came~a. For scanning electron microscope
photomicrogr?phs, a SEM T-200 miniscan microscope was used.

Examination of spores by SEM was usually done on dried

specimens rehyorated in water. As the spores contain only a
small amount of water, the final preparations were made by
air-drying the suspended material.

In one case preparation of spores for SEM examination was made

by critical point drying. The procedure mainly followed that
outlined by Cole & Samson (1979). The material was first fixed
in 6 percent glutaraldehyde in Cacodylat buffer and 1 percent
Os04 in Caco~ylat buffer (proportion 1:1) and then dehydrated
in ethanol. Thereafter the material was placed in liquid CO 2
in a Polaron's critical Point Drying pressure chamber.

After preparation, the material was transferred by adhesive

tape on to stubs and coated with a thin layer of gold-
palladium alloy in a Sputter Coater before beeing viewed in
the SEM.

Colour descriptions refer to Kornerup & Wanscher (1978) and

are all based on stereomicroscope observations. Excipular
terminology follows Starback (1895), revised by Korf (1951,
1958).

Statistic3] c3:c~lations concerning measurements of length and

width of th~ ascospores, include mean values (X) and standard
deviations (Q).
 16

B. MATERIAL

I have tried to trace all material mentioned in the

literature, ana I have also made general requests for
material. About '450 specimens (including duplic.ates) have been
studied. The examined material is deposited in the following
herbaria (abbreviations according to Holmgren et al. 1990).
Institutes visited personally are marked with an asterisk (*):

ATC~, B, BG*, BHU, BP, BPI, BR, C*, CAS, CO*, CP, CUP, DAOM,
OR, E*, FH, FLAS, G, GZU, H, IMI*, K*, L, LPS, M, MICH, MW*,
NEB, NY, 0*, PAD, PC, PREM, PRM, RO*, S*, TAA, TELA, TENN,
TRTC, UC, UPS*, US, W, WELTU, WRSL.·

Specimens were also received from private herbaria of:

J. M. Barrasa, Universidad de Alcala de Henares, Madrid,

Espana, (Barrasa); A. Bell, Victoria University of Wellington,
Wellington, New Zealand, (a slide donated to BG); J. van
Brummelen, Rijksherbarium, Leiden, Netherlands (4 slides); I.
L. Egeland, Oslo, Norway, (slide, drawings and manuscript
donated to BG); J. Haffner, Mittelhof, Germany (descriptions
and photos of T. africanus); E. Jahn, Bad Schwartau, Germany,
(also including several collections and photos donated to
herb. BG)j R. Kristiansen, Torp, Norway, (material donated to
BG)j S. Olsen, L0dingen, Norway, (material donated to BG)j V.
P. Prokhorov, Moscow State University, Moscow, Russia.

Unfortunately, several herbaria from Europe, Asia and Africa

did not reply to inquiries about loan of herbarium material.
 17

MORPHOLOGY AND ANATOMY

A. ASCOCARP

Most species of Thecotheus have mUltiascal, sessile to

subsessile apothecia on a tapering, more or less broad base.
The narrowing base is often ± sUbimmersed. Slightly
sUbstipitate apothecia are observed only in T. africanus, T.
crustaceus and T. pelletieri. However, at maturity the stipe
is often hidden by the expanded apothecium. Young apothecia
are either globose to subglobose or more or less cylindrical.
Later the shape is variable, usually dolabriform, turbinate,
or doliiform. At maturity the apothecia expand and become
discoid to shallowly cupulate, occasionally with reflexed
margin, rarely pulvinate to lenticular, 0.3 - 7 mm diameter
when fresh, up to 4 mm diameter when dry, scattered to
slightly crowded, more rarely gregarious. The lignicolous
species T. rivicola possesses the largest apothecia. A
subiculum-like base is observed in T. inaeguilateralis and T.
lundgvistii. A corresponding net-like growth of mycelium under
the fruit-bodies is easily seen in T. phycophilus. A
characteristic crust-like form of mature apothecia is found in
T. crustaceus.

Young living fruit-bodies are usually white to light grey,

occasionally with a yellow tinge. In general they become
darker during maturity and upon drying. This is the main
reason for the various colours observed within one and the
same species. In addition the pigment development is light
dependent. However, the change of colour is rather consistent
for many of them. According to species, the colour of fresh,
mature apothecia may be yellowish, yellow-brown to grey-orange
to brown-orange or grey-brown. In senescent or dry apothecia
the colour may be darker, sometimes nearly black. Rarely the
colour in fresh apothecia may be greenish grey or greyish
yellow. This colour, often recorded as "yellowish green" is
typical of T. strangulatus and T. uncinatus.
 18

In general, the external surface of young living apothecia is

smooth, usually becoming furfuraceous with age and upon
drying, in most of the species somewhat mealy due to the
finally slightly protruding ectal cells. The anchoring hyphae
are coenocytic, septate, usually branched, subhyaline to light
yellow brown, 1.5 - 3.2 ~m wide.

At first the disc is flat to slightly convex, becoming concave

at maturity and upon drying, at that time usually paler than
other parts of the apothecium, and slightly rough due to the
protruding ascus tips.

Most of the s~ecies have a varying broad, distinct margin

which usually becomes ± incurved with age and upon drying.
This is reminiscent of several species of Fimaria Vel. In old,
expanded fruit-bodies the margin may be excluded and not
visible.

B. EXCIPULUJi

All species of Thecotheus have distinct areas of

differentiated ectal and ental excipula. The medullary
excipulum is predominately composed of a textura intricata and
a patchy textura globulosa. The former consists of a ± compact
tissue of densely septate hyphae, 2 - 3 ~m wide, which run in
different dir~ctions, mostly with distinct interhyphal spaces.
The latte~ ~~= hj~phae with hyaline, globose to sUbglobose
cells, 2 - 30 ~m diameter. In several species the textura
intricata also constitutes a relatively well-defined
hypothecium layer. Towards the ectal excipulum there is a zone
of textura porrecta, with long-celled, septate, parallel-lying
hyphae. The cells start from the central growing point of the
apothecia and partly extend along the lateral walls towards
the marginal layer; partly they extend outwards into the ectal
excipulum and terminate on the excipular surface. Isolated,
 19

mature fruit-bodies with a pUlvinate shape usually have a

reduced medullary excipulum.

The ectal excipulum is several cell layers thick,

predominately of a textura angularis composed of angular to
sUbglobose cells. The outermost cells are mainly polyhedral
and elongated, subhyaline to yellow-brown, 5 - 40 x 4 - 25 ~m,

their longitudinal axes lying perpendicular to the outer

surface. Towards the margin the cells are more isodiametric.
This kind of cells is lacking in the margin. The inner part of
the ectal excipulum is composed of scattered ± subglobose
cells 5 - 35 ~m diameter, interspersed with rooting hyphae,
1.5 - 4 ~m wide, originating from the boundary between the
medullary and ectal excipula. The septate, rooting hyphae
terminate on the excipular surface, their terminal cells
usually encrusted, sUbhyaline to brownish, with uninucleate,
bulbous to more or less inflated and elongated compartments, 4
- 15 (-20) ~m wide and 5 - 25 ~m in length. Especially at the
apothecial base and along the lateral sides, the terminal
cells become darker and apparently more thick-walled when old
and dry. Towards the margin, the cells gradually transform to
paraphysis-like cells. Both the terminal bulbous cells and the
ectal excipulum cells are cyanophilic, i.e. they stain
strongly in Cotton blue. The ectal excipulum is thickest at
the central growing point, gradually becoming narrower towards
the margin.

The excipular cell size, wall thickness and wall pigmentation

varies from species to species. The pigmentation is, however,
much influenced by light intensity.
 20

C. ASCI

Most species of Thecotheus have eight-spored asci. Regular 32-

spored asci are only found in T. pelletieri. Very rarely this
species may have asci with 16 or 64 ascospores. T. phycophilus
is characterized by a spore number lower than eight. Here,
most asci have four fully developed ascospores and
accompanying small abortive ones up to a total of eight in
each ascus. Probably the same is true for T. viridescens,
which is described with four or eight ascospores in each ascus
(Hohmeyer et al. 1989). occasionally, abortive
ascospores are observed in T. crustaceus and T. lundqvistii
too.

The asci are cylindric to subcylindric or clavate to broadly

clavate, operculate, (110-) 115 - 460 (-475) x 10 - 70 (-90)
~m, gradually tapering downwards into a stern-like base, this
part varying from long and slender to short and stout. The
lower part may be rather fragile, as seen e. g. in T.
crustaceus. A characteristic bilobed base is typical of T.
strangulatus. The apices are obtuse to ± truncate. A slightly
acute ascus apex is found in T. himalayensis. strangulate asci
are a distinguishing feature for T. strangulatus and T.
uncinatus. In the latter species the strangulation is easily
seen in living asci as well as in mature, empty asci. It is
not always visible in dry material, although a slight
thickening is often observed in dry asci.

The two-layered ascus wall stains blue to diffuse blue in

Melzer's reagent. occasionally a diffuse amyloid reaction is
observed in old, dry material. The outer wall layer also
stains uniformly in Congo Red in ammonia. When mounted in the
latter reagent, a ± distinct apical ring-shaped zone is seen
especially in T. pelletieri and T. phycophilus. At maturity,
the asci markedly protrude above the hymenium in most of the
species. In the non-coprophilous T. QPycophilus, and T.
 21

rivicola, mature asci only reach or extend slightly above the

paraphyses. Thi~ probably reflects an ecological adaptation,
just as observed by Kimbrough et al~ (1969) in Iodophanus.

D. ASCOSPORES

The ascospores are non-septate and range from narrowly to

broadly ellipsoid, occasionally ellipsoid-fusiform to
subfusiform, 12 - 41 (-46) x (5.5-) 6 - 22 (-24) ~m,
symmetrical or slightly unequal-sided. In T. biocellatus,
spores with both equilateral and inequilateral shapes are
found. Rarely, spores of two sizes in separate asci were
observed in the polysporic T. pelletieri. This kind of
heterospory is briefly mentioned by Kanouse (1950). Most of
the spores are hyaline to subhyaline, sometimes appearing to
be light yellowish or greyish yellow at maturity. This may
account for Overton's (1906) observation of slightly coloured
ascospores in T. pelletieri, and Donadini's (1985) record of a
yellowish or orange-colored spore print in T. kimbroughii.

The ascospores are smooth or verruculose to verrucose.

Sculptured spores are initially smooth, and their
ornamentation largely depends on the degree of maturation.
Condensation of secondary wall material constitutes a
permanent ornamentation on the epispore. The ascospores are
bipolarly apiculate or non-apiculate. Both the apiculi and the
ornamentation stain in Cotton blue. In T. africanus and T.
perplexans the apiculi are furnished with a distinct,
projecting collarette at their base. Immature ascospores are
thick-walled, but the wall becomes thinner by the time of
spore liberation. The inner wall layer, initially up to about
4 Mm, does not stain in Cotton blue. In mature ascospores, the
thin outer wall layer stains in Cotton blue. In most species
 22

this layer becomes loose and distorted in lactic acid and when
heated in 2% KOH.

Each ascospore is surrounded by a varying thick, cyanophilous,

gelatinous perispore, which often greatly expands after spore
liberation, when mounted in water and other mounting media.
Then, generally the gelatinous envelope soon dissolves. This
is the reason why the gelatinous layer is not always observed
on free-lying old ascospores.

No oil drops are found in the sporoplasm. In most species the

ascospore cytoplasm is granUlar, which is best seen in young
condition. The spores are probably uninucleate in all species.

E. PARAPHYSES

Thecotheus species have paraphyses of either one type or of

two different types. The first one is abundant, slender,
septate, simple or branched, with the upper part generally
straight, rarely uncinate, 1.2 - 2.5 (-3) ~m wide below,
apically mostly 2 - 4 ~m wide, occasionally distinctly
enlarged up to 8 ~m in diameter.

The second type of paraphyses is scattered, unbranched,

slightly thick-walled. Below they are irregularly septate, 2.5
- 4 ~m wide, their tips aseptate, inflated, and (3-) 4 - 8 ~m

in diameter. This type, often recorded as paraphyses-like

interascal el~ments, .is found in T. africanus, T. cinereus, T.
himalayensis, T. inaeguilater~lis, T. kei~hii, T. phycophilus
and T.rivicola.

The paraphyses are hyaline throughout, or they are yellowish

to brownish pigmented. The colour, most abundant in the tips,
is located in the walls, or as pigrnented granules within.
Their upper parts are often embedded in.a gelatinous,
 23

amorphous, pigmented matrix. As indicated by Eckblad (1968),

it is often impossible to decide whether the colouring matter
is restricted to the cell wall or located inside the cells of
the paraphyses.

Shape and colour of the paraphyses are relatively stable

specific characters. However, their pigmentation is greatly
influenced by light intensity during the development of the
apothecia.

F. ANAMORPHS

Anamorphs are very rare in the Ascobolaceae. I have not

observed any during my studies, but Conway (1975) found some
in culture of T. pelletieri and T. holmskjoldii. The
uninucleate, obovate, hyaline conidia ~f T. pelletieri, 3 - 4
x 6.5 - 7 ~m, were borne on short side branches. Up to ten
conidia were produced on each conidiophore. The conidiogenous
cells are of sympodial type, an~ morphologically the anamorph
of T. pelletieri is similar to Rhinocladiella Nannf. (Conway
1975) .

CULTURES

Ascospore germination for many coprophilous species usually

occurs within the animal gut. The spores therefore require a
 24

passage through an animal or a similar artificial treatment

(i. a. Harper & Webster 1964). As far as known, the examined
species of Thecotheus are difficult to grow in culture.
Overton (1906) did not succeed in germinating the ascospores
of T. pelletieri. Dodge (1912) reported abundant germination
of spores of Thecotheus by heating them to 50 0 - 70 0 C for a
short time. Conway (1975) was not able to induce apothecial
formation from a single-spore culture of T. pelletieri. Only
very sparse fruit-body development in culture was observed in
T. holmskioldii in the investigation" by Conway (op. cit.), but
not in single-spore culture. He suggests that both species are
heterothallic.

Germination of about 10% of the ascospores in culture was

found in T. phycophilus (Pfister 1981). The germination was
bipolar. A terminal germinal pore at each end of the
ascospores was indicated by Overton (1906) in T. pelletieri.
Dodge (1912) reported germ-tubes usually at both ends of the
spore at points only slightly to one side of the ends in T.
pelletieri, but also germination from only one germ pore.

My own examination revealed bipolar germ pores in T.

crustaceus, and also slightly sub-bipolar germ-tubes in
process of formation in T. cinereus [Krieger's Fungi saxonici
No. 335 as Ascobolus (Ascophanus) incanus J. Generally,
occurrence of germ pores, in addition to localization and
number are of great systematic value.

The ascocarps develop from several ascogonia in T. pelletieri

(Overton 1906).
 25

ECOLOGY AND DISTRIBUTION

All known species of Thecotheus are saprophytic. The majority

of the species are predominantly coprophilous, and they
inhabit a great number of different mammalian dung types. Some
of the species are obviously substrate specific, preferring
one kind or a few kinds of dung (table 1).

However, dung of the different groups of animals shows very

different features. This reflects visibly a heterogenous
fungus flora on thi~ kind of substrate.· Concerning the most
common grazing animals, dung of cattle may be up to 2000 ml in
volume, i. e. up to 240 g dry weight, while pellets of sheep
range from 0.1 to 6 g dry weight (Lumaret & Kirk 1987).
Generally, cattle dung retains most of the moisture during the
summer, whereas sheep pellets may lose up to 50% of their
water content within 3 hours in summer months (Lumaret & Kirk
1987). Besides physical and chemical changes in dung, the
fungus flora is also influenced bye. g. temperature and
rainfall. Age and seasonal changes of dung involve qualities
which favour different groups of dung fungi.

More rarely, some species also inhabit different kinds of

soil, dead grasses, herbaceous plants, or twigs and wood
submerged in water.

A. SUBSTRATE PREFERENCE

1. Dung species
A total of 18 different types of dung are known as substrates
for the species of Thecotheus. Most collections are associated
with domestic animals, as compared with wild animals:
 26

Number ot tin~s on ~ung of ~omestic animals: Total 237 (325);

cow 123 (172), horse 82 (120), sheep 19 (29), goat (including
wild goat) 10 (19), donkey 2 (1), dog (3), mule 1, and camel
(1) (Kimbrough 1969).

Number of fin~s on ~ung of wil~ animals: Total 38 (48);

deer (including i. a. fallow deer, red deer, and roe deer) 23
(27), elephant 6 (18), rabbit 1 (2), moose 2, musk ox 2, ass
2, hare 1, buffalo 1, european bison (I), and rhinoceros (1)
(Merkus 1976)_ Unfortunately, all material of this last
collection from Indonesia is used up (van Brummelen in litt.
24 _VI _ 1985).

No species has as yet been found on bird droppings.

The following species are exclusively restricted to dung:

T. africanus, T. biocellatus, T. crustaceus, T. himalayensis,

T- holmskioldii, T- inaeguilateralis, T. keithii, T-
lundgvistii, T. perplexans, T. strangulatus, T. uncinatus, and
T- viridescens.

Obviously some of the dung species are rather substrate

specific, preferring one or a few particular kind of dung
(table 1). Species found less than four times are not included
below.

Species pr'eferring ~ung of elephant:

T. africanus, and T. perplexans.

Of a total of 5 (18) number of finds, all but one are

associated with elephant dung. Apparently, both species are
vulnerable as the elephant is a threatened animal.
 27

Table 1. The euh~trate preference of the included species of

Thecotheus. The numbers without parentheses mean all of my own
examined collections, whereas the numbers within parentheses
in addition includes literature records of material not seen
by me.

Species Substrate

T. africanus 5 (6) elephant 4 (5), cow 1

T. biocellatus 1 horse
T. cinereus 6 cow 4, dung mixed with sand/soil in
greenhouse 2, soil in greenhouse *
T. crustaceus 33 (48) horse 27 (36), cow 2 (8), buffalo 1,
donkey 1, elephant 1, moose 1
T. himalayensis 3 (6) goat 2 (4), deer 1, dung (1)
T. holmskjoldii 109 (145) cow 58 (78), deer 20 (23),
sheep 18 (23), goat 8 (13),
musk 2, horse 1 (2), hare 1,
rabbit 1, dog (1), donkey (1)
T. inaequileteralis 2 moose
T. keithii 14 (19) horse 14 (16), cow (1), goat (1),
dung (1)
T. lundqvistii 4 cow 3, sheep 1
T. pallens (2) soil
T. pelleti~ri 96 (173) cow 54 (85), horse 36 (66),
sheep (6), deer 1 (4), ass 2,
dog (2), rabbit (2), donkey 1,
elephant 1, mule 1, european bison
(1), goat (1), soil (1) **
T •. perplexans (12) elepJ:1ant
T. phycoph i l.!h~ 2 dead herbaceous plants
T. rivicolq 9 (11) water-soaked wood 6 (1), soil 2,
veget. debr. 1, unknown substr. (1)
T. strangulatnc; cow
T. uncinatJ.!.$. 4 horse 3, deer 1
T. virides:::en::. ]. dung

* I strongly S~5pect, as pointed out by Kimbrough (196~) that

this was mi\r .... l~ ,}cl soil ~ ** Not seen (Bucknall 1880).
 28

species preferring ~ung of horae:

T. crustaceus, T. keithii, and T. uncinatus.

T. crustaceus is known from 6 different substrates, and 82%

(75%) of the finds are restricted to horse dung.
Correspondingly, T. keithii is known from 4 different
substrates, where horse dung constitutes 100% (84%).

Species preferring dung of cow:

T. holmskjoldii, and T. pelletieri.

T. holmskjoldii is known from 10 different dung substrates

(those of different deer species are regarded as one
substrate), and out of a total of 109 (145), those collections
on cow dung constitute 53% (54%). This species also shows a
relatively high preference for dung of deer and sheep. T.
pelletieri is found on 13 different substrates, all but one
being dung. Its preference for cow dung constitutes 56% (49%)
of records, but also horse dung is a rather common substrate
for this species. Probably, T. cinereus and T. lundgvistii
also belong to this group.

Species preferring ~ung of cervi~s:

T. himalayensis, and T. inaeguilateralis.

Although scanty data exist so far, two species must be

included here. Tentatively, T. inaeguilateralis is placed in
this group, as yet only found on dung of moose. Very probably,
all Indian collections of T. himalayensis have been made on a
cervid species.
 29

2. Debris species

T. pallens, T. phycophilus, and T. rivicola~

The species in this group prefer moist places, water-soaked

wood and different kinds of rotting plants, not infrequently
associated with algal films.
T. rivicola show a preference for submerged wood and twig~.

This species exhibits a slightly variable.habitat. The water-

soaked wood includes i. a. twigs of Prunus spinosa, Alnus sp.
and an undetermined stem (monocot. species). Two collections
from England are from vegetable debris on damp soil on the
ground, and on soil on a damp, clayey path in woodland.

The habitat of T. pallens and T. phycophilus comprises soil

and vegetable debris. Both species are associated with algae
and periodically inundated areas. T. pallens is restricted to
soil which is periodically inundated. Grelet (1944) found the
fruit-bodies on moist soil among green algae in a shaded
meadow, whereas Boudier (1888) detected the species growing on
clayey soil in forest. Unfortunately, it has not been possible
to re-examine this material.

T. phycophilus is found on wet debris and dead grasses which

are overgrown by algae, i. a. oscillatoria and coccoid green
algae. Sedges, Eguisetum, and Pilea sp. form the herbaceous
ground cover in the type locality (Pfister 1981).

B. DISTRIBUTION

Although the species of Thecotheus are known from Europe,

 30

Asia, Australia, Africa, North America and South America, too

many parts ot the world are still poorly investigated and this
makes it difficult to draw reliable, final phytogeographic
conclusion~.

The areas where Tbecotheus is best known are North America,

Central- and North Europe. In particular, the most
comprehensive inventory is from Sweden. In the present case,
an extensivE use of the moist chamber technique has
undoubtedly contributed to this outcome. As emphasized by
Lundqvist (l97Z), the majority of the coprophilous species
will remain undiscovered without this indispensable method.

T. pelletieri is one of the most frequently collected species

of the genus, known from all the continents. It is well known
to most discomycetologists, and is probably the most
conspicuous of all the Thecotheus-species. Although so far
there are rather few records from the southern hemisphere, the
species covers a large and continuous area, and it may be
characterized as "cosmopolitan".

with one exception, T. holmskjoldii has its distribution in

the northern hemisphere, to jUdge from the material examined.
It is found as far north as 72°50'N. Togther with T.
pelletieri this is the most common of all the species of
Thecotheu~, u~~ t~ere are numerous literature records since it
was descrite~ 1" !r76. However, the species may have a wider
distribution pattern in the south-temperate zone. T. keithii
is also restrict~d to the north-temperate zone (Europe -Asia -
North Americ3) .

T. crustaceus shows a bipolar distribution (Europe-Asia-

America). This species is known from most of the Nordic
countries (Table 2). Tentatively, the following species so far
indicate disju~ct distribution patterns: T. cinereus (Europe-
Asia-North America), T. himalayensis and T. uncinatus (Europe-
Asian), and T. rivicola (Europe-America).
 31

T. africanus and T. perplexans exhibit a tropical equatorial

distribution pattern within the genus, limited between 6°N and
4°S. T. africanus is known from Ivory Coast, Kenya, Tanzania,
and Sri Lanka. T. perplexans is known only from Congo in
Africa. Both species may be endemic to the Palaeotropics, and
apparently they may be regarded as vulnerable. Both species
prefer dung of elephant, and this explains their distribution
pattern.

In the Nordic countries, six species of Thecotheus are known

(table 2): T. crustaceus, T. holmskjoldii, T. keithii, T.
lundqvistii, T. pelletieri, and T. uncinatus.

Table 2. Occurrence of the species of Thecotheus in the Nordic

countries.

species Norway Sweden Finland Denmark

T. crustaceus X X X X
T. holmskjoldii X X X
T. keithii X
T. lundqvistii X
T. pelletieri X X X
T. uncilJ~.:t~!E X

The following spec~es are known in one or a few collections:

T. biocellatus (Czechoslovakia), T. inaeguilateralis (Ukraine,

Russia), T. lundqvistii (Spain, Sweden), T. strangulatus
(Czechoslovakia), and T. viridescens (Germany). T.
phycophilus, only known from the type locality in U.S.A may
have been overlooked due to its special substrate.
 32

TAXONOMY

Thecotheus Boud.

Annls. sci. Nat. 5 ser. Bot. 10: 235. 1869. -- Ascobolus

subgen. Thecotheus (Boud.) Bomm. & ROuss., Bull. Soc. Roy.
Bot. Belg. 23 (1): 144. 1884. -- Ryparobius [? subgen.] 11.
Thecotheus (Boud.) Sacc., Syll. Fung. 8: 542. 1889. --
Ascophanus sect. Q Thecotheus (Boud.) Heimerl, 15 Jahresber.
k. k. Ober-Realsch. Bez. Sechshaus, Wien: 20. 1889. --
Rhyparobius sect. 11. Thecotheus (Boud.) Lindau, in Engl. &
Prantl: Die nat. Pflanzenfam. 1 (1): 190. 1896. -- Rhyparobius
subgen. Thecotheus (Boud.) Migula, in Thome's FI. Deutsch.,
0sterr., Schweiz 10 (2): 1040. 1913. -- Type species (only
original): Ascobolus pelletieri Cr. & Cr. E Thecotheus
pelletieri (Cr. & Cr.) Boud., lectotype.
Ascophanus sect. B Holmskioldia Heimerl, 15 Jahresber. k. k.
Ober-Realsch. Bez. Sechshaus, Wien: 20. 1889. -- Type species:
Ascophanus holmskioldii E. C. Hansen 5 Thecotheus holmsk;oldii
(E. C. Hansen) Chenant., lectotype.
Ascophanella Faurel & Schotter, Cah. La Maboke 3 (2): 130.
1965 b. -- Type species: Ascophanella cinerea [sic!] (Cr.)
Faurel & Schotter E Thecotheus cinereus (Cr. & Cr.) Chenant.,
holotype. [-- non Ascophanella Korf, Rapp. Comm. VIII. Congr.
Int. Bot. I 1954. Sect. 18-20: 80. 1954. Nomen nudum].
Ascophanopsis Faurel & Schotter, Cah. La Maboke 3 (2): 130.
1965 b. -- Type species: Ascophanopsis perplexans Faurel &
Schotter 5 Thecotheus perplexans (Faurel & Schotter) Krug &
Khan, lectotype selected here.

? Zukalina O. Kuntze, Rev. Gen. Plant. 2: 875. 1891. (Fide

Korf 1972, 1973); nom. novo for Gymnodiscus Zuk., Verh. Zool.
Bot. Ges. Wien 37: 44. 1887; nom. illegit.; non Gymnodiscus
Less. 1831 (Asteraceae).
 33

ETYMOLOGY: Greek e~K~, a box, a case, wO£\V, to push;

referring to the projecting asci.

Apothecia sessile, 0.3 - 7 mm diameter when fresh, up to 4 mm

diameter when dry, at" first globose to sUbglobose or
cylindrical, expanding and shallowly cupulate to discoid at
maturity, rarely pulvinate to lenticular, at first white to
light grey, becoming yellowish to brownish at maturity or when
dry. Disc flat to a little convex, turning concave when dry,
slightly roughened by the protruding ascus tips at maturity.
Margin distinct. External surface of apothecia at first
smooth, usually becoming white~furfuraceous when dry.

Medullary excipulum predominately of textura intricata and a

patchy textura globulosa, interspersed with scattered ±
isodiametric cells or more or less elongated, sub-polyhedral
cells. Towards the ectal excipulum a zone of textura porrecta.
Ectal excipulum predominately of textura angularis, the
outermost cells elongated, with their longitudinal axes
perpendicular to the outer surface. Inner part occasionally
with scattered sUbglobose cells, in between with septate,
rooting hyphae which terminate on the excipular surface, their
terminal cells with bulbous and elongated parts.

Asci normally 8- or 32- spored, rarely irregularly 4-spored,

exceptionally 16- or 64- spored, operculate, cylindric to
sUbcylindric or ± clavate, upper part rarely strangulate,
(110-) 115 - 460 (-475) x 10 - 70 (-90) ~m, walls blueing in
their entire length in Melzer's reagent. Ascospores ±
ellipsoid, occasionally sUbfusiform, equilateral or
inequilateral, predominantly hyaline to subhyaline, uniseriate
or biseriate or 4 (-5) seriate or irregularly disposed, 12 -
41 (-46) x (5.5-) 6 - 22 (- 24) ~m, bipolarly apiculate or
non-apiculate, very rarely with a collarette at each polar
apiculus base. Ascospores smooth or ornamented with punctate-
like granules or warts which stain in Cotton blue. Each
ascospore is surrounded by a gelatinous perispore. Paraphyses
 34

hyaline or pigmented and encrusted, usually slightly exceeding

the asci, of one or two types. Cl). Abundant, filiform, thin-
walled, septate, simple or branched, upper part straight,
rarely uncinate, their apices at most slightly inflated, only
rarely with distinctly enlarged tips. (2). Scattered, thick-
walled, dispersedly septate below, their tips aseptate,
simple, apically ± inflated.

Most species grow on dung of different mammals, more rarely on

soil of different kinds, and debris such as: dead grasses and
herbaceous plants, rotting twigs and wood in water, algal
coverings.

In my opinion the genus belongs to the Ascobolaceae. The main

reasons for this are:
- the amyloid asci. Although occasionally with stronger
colouring in the apex, the asci stain diffusely blue in
Melzer's reagent.
- the asci protrude beyond the hymenium at maturity.
- the ascospores are thick-walled when young.

Moreove~, the bulbous excipular cells and their predominantly

coprophilous habitat is also a general, characteristic fea~ure

in this. family. Kimbrough & Curry (l985) proposed the term

"ascoboloid septal type" for the ascal septa in all taxa of
the Ascobolaceae, where they also retained Thecotheus. These
authors regarded this septal type in reproductive tissue as a
good family indicator, equivalent to the "pezizoid septal
type" in the family Pezizaceae. Van Brummelen (l981)
recognized the "Ascobolus type" of asci for the genera
Ascobolus, Saccobolus, Thecotheus, Iodophan~s and Boudier~ .. r
agree with him in this evaluat~on.
 35

Thecotheus differs from Iodophanus i. a. by the absence of

carotenoid pigments in the paraphyses, by the thicker
gelatinous perispore, and by the absence of hyaline, flexuous
hairs on the ectal excipulum. The pigmented mature ascospores
in Ascobolus and Saccobolus are quite different from those of
Thecotheus. The genus should not be confused with coprophilous
species of Peziza. The ascospores of this genus are thin-
walled when young, and although of rather variable amyloidity,
the intense staining at the ascus tip easily distinguishes
this genus from Thecotheus. The psilopezioid genus Pachyella
with diffusely amyloid asci differs from Thecotheus in having
hyphoid hairs on the ectal excipulum and the presence of gel
tissue inside the excipulum. Usually, dry apothecia become a
thin film.

In the arrangement of infrageneric taxa, I have tentatively

recognized groups among the present material. However, too
few cultural and developmental studies have been done to
decide a final, infrageneric taxonomy of specified rank.

I. ~. &fricanus group

T. africanus, T. perplexans.

Both species are unique within the genus in possessing a

projecting collarette at the apiculus bases. T. africanus also
has an excipulum structure slightly different from that of the
other members of Thecotheus. No material has been available of
T. perplexans, and it would be of great importance to re-
examine several morphological characters of this taxon. Except
for the cOllarette, the ascospores (size, ornamentation and
form) are similar those ofT. biocellatus,
 36

T. himalayensis and T~. rivicola. The ascospores in T.

himalayehsis differ in being non-apiculate. Of these, all
except T. biocellatus have paraphyses of two types. They are
hyaline in T. africanus, pigmented in the two other species.

'The tropical distri~ution of T. africanus and T.' perplexans

may indicate that they represent an relatively old group.
Combined with their hyaline paraphyses, which is regarded as a
primitive type (cfr. Eckblad 1968), the possession of the
collarette probably represents a primitive morphological
character.

11. ~. stranqulatus group

1. strangulatus, T. uncinatus.

These species constitute a distinctive group within the genus.

They are characterized by strongly curved paraphyses,
strangulate asci, and a yellow-green colour of the fruit-
bodies, characters not found elsewhere amongst the other
species in the genus. However, their diffusely amyloid and
protruding asci, their thick-walled ascospores and bulbous
excipular cells are characters typical of the genus.

Ill. ~. rivicola group

T· phycophilus, T. rivicola.
 37

Besides their non-coprophilous habitat, this group differs

from the other species in Thecotheus in having non-protruding
asci. Their ascus-form is ± cylindrical. Characteristically
the excipulum is "multicellular", as distinct from the few
cell layers in the excipulum of the remaining species.

Non-protruding asci probably reflects an ecological adaptation

(see p. 21). However, this character, combined with possession
of cylindrical asci may also indicate a derived group (cfr.
Eckblad 1968).

In T. rivicola the paraphyses are embedded in a gelatinous

matrix. This may indicate an ecological adaptation where the
spores are discharged after the hymenium dries up.

IV. Eu-Thecotheu8 group

T. biocellatus, T. cinereus, T. crustaceus, T. himalayensis,

T· holmskjoldii, T. inaeguilateralis, T. keithii, T.
lundgvistii, T. pelletieri, T. viridesc~ns.

All members of this group are coprophilous, they have small

apothecia, smooth or ornamented ascospores which are bipolar
apiculate or non-apiculate, and they have straight paraphyses.

Paraphyses of two types are observed in T. cinereus, T.

himalayensis, T. inaeguilateralis, and T. keithii. This type
is also found in T. africanus (group I), T. phycophilus and T.
rivicola (group Ill). Thind & Kaushal (1978) found this type
[as additional interascal elements] in Iodophanus kimbroughii,
and Dumont (1971) reported the type in two species of the
inoperculate Lambertella. Within Thecotheus I have not been
 38

able to correlate this type of paraphyses with other

characters, except that they are not observed in species with
ornamented, apiculate ascospores. Nor are they observed in
group 11. I regard this kind of paraphyses as a primitive type
which has been reduced. It has apparently evolved in various
groups, and illustrates a convergent evolution.

T. pelletieri is the only known polysporic species of

Thecotheus, and is probably an advanced species. Only one type
of paraphyses is observed. Several coprophilous genera of
different groups have evolved polyspory, and the convergent
evolution towards mUltispored asci may illustrate a major
feature in the adaptation of coprophilous fungi (Kimbrough
1972) •

Conway (1975) found anamorphs in T. pelletieri and T.

holmskjoldii. The anamorphs are morphologically and
ontogenetically similar and thus indicate a close relationship
between the two species. Both species are related to T.
cinereus and T. keithii, they all have ellipsoid ascospores
and pigmented paraphyses. T. lundgvistii has ornamented and
apiculate ascospores of same type as T. holmskjoldii, and I
regard them as closely related.

The r~maining. species, 1. biocellatus, T. crustaceus , T.

himalayensis, and T. inaeguilateralis have all ± unequal-sided
spores. All but the latter have pigmented paraphyses. T.
biocellatus and T. himalayensis are related owing their
similar spore-ornamentation, although the latter is non-
apiculate. The two other species have smooth ascospores.

v. ~. pallens group

T. pallens.
 39

This terrestrial species probably represents an isolated group

within the genus. It is characterized by the large, oblong-
fusiform asccspores and by the broad, hyaline paraphyses. The
asci are clavate and protruding. No material exists of this
species, and its taxonomic position is accordingly uncertain.

I n '1'0 '.l'lIE ACCEPTED SPECIES OF 'l'JIECOTBEUS

1 Asci 8-spored .•..•......................••.•..•..•........ 5

* Asci with different spore number ..•....•....••.•.......... 2
2 Asci 4 - S-spored .........................•..•.••.••...... 3
* Asci 16 - 64-spored ..•...•.........•.......••.••••.•...... 4
3 Spores srroQth, ellipsoid, 30 - 37 x 15 - 16.5 (-18) ~m;

paraphyses of two types; on dead herbaceous plants ••.........

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • .. 13. r. phycophilus
* Spores ornamented with small extended warts, to very
fine reticulations, narrowly ellipsoid to subfusiform,
slightly inequilateral; 16 - 20 x 6 - 8 ~m;

paraphyses of one type; coprophilous .•.... 17. r. viri~escens

4 Asci 32-~pGred; spores ellipsoid, (30-) 32 - 40 (-42.5) x

(13-) 15 ." 2~ (-24) ~m; .........•.......... 11. r. pelletieri
* Asci either 16-spored; spores ellipsoid,
32 ~ 40 x 16 ~m or asci 64-spored; spores subglobose,
11 - 24.5 x 10 - 16 ~m; : :11. T. pel1etieri
5 Spores witn a projecting collarette at apiculi
base; on elephant dung ; ......•....."•..••... ~ • . . • . . • • • • . . . . 6
* Spores lacking collarettes, with or without
bipolar api~~lus; coprophilous or non-coprophilous 7
6 Spores 12 - ~6 x 7 - 9.5 ~m 1. T. africanus
 40

* Spores 20 - 22 x ~o - 12 ~m 12. ~. perplexans

7 Spores smooth .•......••.................................... 8
* Spores ornamented 13
8 Spores api.culate ....•...........•.....'.......•. '.' '. . . . 9
* Spores non-apic~late •..•.................................. 10
9 Spores inequilateral, 13 - 16 x 7 - 8.5 ~mi ••••••••••••••••••
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7. ~. inaequilateralis
* Spores symmetrical, (16-) 17 - 21 (-22) x
(7-) 8 - 10 (-11) ~m; .....................•.... 8. ~. keithii
10 Spores less than 25 ~m in length ....•....... 4. ~. crustaceus
* Spores longer than 30 ~m ....•...••...........•...••..•.... 11

11 Spores oblong fusiform; paraphyses of one type;

on clayey soil •.•.•.•...••••••.•... ~ .•••.•.••. 10. ~. pallens
* Spores ellipsoid; paraphyses of two types;
not on clayey soil •...........•..•..•......•.••..•••..•... 12
12 Asci always 8-spored, spores ellipsoid to cylindrical-
ellipsoid, (29-) 32 - 41 (-46) X (13-) 14 - 18 (-21.5) J.Lm;
mainly coprophilous ......•.............••.•••. 3. ~. cinereus
* Asci mostly 4-spored, if 8-spored then always with
some smaller abortive ascospores, spores ellipsoid
to broadly ellipsoid, 30 - 37 x 15 - 16.5 (-18) J.Lm; on
dead herbaceous plants (see 3) •........... 13. ~. phycophilus
13 Spores apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . 14
* Spores non-apiculate 17
14 On water-soaked twigs and wood •...•.......... 14. ~. rivicola
* On dung •......•.•............•..•...............•....•... 15
15 Spores (14-) 16 - 19 x 7 - 9 (-10) ~m, broad
inequilateral ............•......••.••...••• 2. ~. biocellatus
* Spores longer, symmetrical .•.••...•........•.•...•....•.. 16
16 Spores 25 - 30 x 12.5 - 14.5 ~m, ornamentations
of minute granules ....................•.... 9. ~. lundavistii
* Spores (25-) 29 - 38 (-42) x (12-) 14 - 18 (-20) ~m,
ornamentations of larger isolated warts ... 6. ~. holmskjoldii
17 Paraphyses strongly uncinate; spores regular
ellipsoid; asci strangulate 18
* Paraphyses straight; spores slightly inequilateral,
 41

sub-ellipsoid to subfusiform; asci not strangulate ....•.. 19

18 Spores (13-) 14 -" 17 x (6-) 7 - 9 (-9.5) pm • 16. ~. uncinatus
* Spores (20-) 22 - 26 (-28.5) x 11.5 - 16 pm .•••.•••••••.••...
• • • • • . • • . • . . • • • . • . . . . • . • • • • • . . • • • • • • • . . .• 15. ~. 8trangulatus
19 Spores (12-) 14 - 16 x (5;5-) 6 - 7 pm, orname~ted
with very small warts; paraphyses of two types;
asci always 8-spored ....................•. 5. ~. himalayensis
* Spores 16 - 20 x 6 - 8 pm, ornamented with small
extended warts to very fine reticulations;
paraphyses of one type; asci 4 - 8-spored (see 3*)" •.•..•.....
• • • • . . . . . . . ... . .". • . . . . . . • • • . . . . • • . • . . • • • . •. 1 i. ~. viridescens

In this survey I have paid particular attention to detailed

information of each collection. This includes i. a. ecology,
original identifications and comments, literature references and
complete data of herbarium. As far as possible, literature
information is compared with annotations of each herbarium
collection. Generally, the lists of the examined material also
record earlier determinations.

The specimen lists are arranged geographically according to an

alphabetical sequence of the countries in each continent in the
following order: Europe (including several republics of formerly
U.S.S.R.), Asia (including Russia), Australia, Africa and North and
South America. The states of U. S. A. are arranged in a similar
way. Arrangement of the provinces in Norway, Sweden and Finland
follow the traditional south to north sequence (Lid 1985).
Generally, further division (e. g. in rural districts) is also
arranged alphabetically.

In all lists recording substrate preferences, the numbers without

parentheses refer to personally examined collections. The numbers
within parentheses record all known specimens (including literature
records) .
 42

Abbreviation.!:

Ascob. = Asco~olus

Ascoph. = Ascuphanus
F. Eur. Exs. = Fungi Europaei Exsiccati
F. Fenn. Exs. = Fungi Fenniae Exsiccati
F. Fim. Exelcpl. Exs. = Fungorum Fimicolorum Exempl. Exsiccati
F. Fim. Exs. = Fungi Fimicolae Exsiccati
F. sax. Exs. = Fungi saxonici Exsiccati
Flora Boh. Mor. Exs. = Flora Bohemiae et Moraviae Exsiccata
1+ blue (of asci) = blueing the entire length in Melzer's reagent
(me) = developed in moist chamber
E. Pezizu
T. Thecotheus

1. THBCOTHBUS AFRlCABUS Khan' Kruq (Figs. 1 - 6)

Mycologia 79 (2): 200 (1987). -- Type: Tanzania, Mt. Kilimanjaro,

Loitokitok, elev. 7500 ft., 2 0 58'S, 37°28'E, on elephant dung,
16.VIII.1966 R. F. Cain, H. D. Griffin & J. C. Krug 66.1738 h
(TRTC, holotype).

ETYMOLOGV: LRtinized from the name Africa, referring to the

continent of the type locality.

Apothecia scat~ared or occasionally slightly crowded, 0.4 - 3 mm in·

diameter when fresh,.up to 1.65 mm diameter· when dry~ At first
sUbcylindric to subfusiform, then dolabriform to turbinate,
expanding at maturity, occasionally slightly substipitate, then
with more or less discoid shape.
 43

At first white, then light grey-yellow, becoming light yellow to

light orange when dry, lower part occasionally tinged with brown.
Upper part, including the hymenium, slightly white-furfuraceous
when dry. Lower part with anchoring hyphae. Disc flat to slightly
convex, turning concave in dry condition, yellow to weakly yellow-
brown, somewhat rough due to the protruding ascus tips at maturity.
Margin distinct, roundish.

Medullary excipulum up to 280 ~m thick at the central base, partly

of textura globulosa with small, hyaline, globose to sUbglobose
cells 3 - 7 ~m diameter, interspersed with some greater, more or
less polyhedral cells up to 9 x 13 ~m diameter, partly of
filamentous hyphae mostly 2 - 3 ~m diameter. The latter hyphae run
in different directions, mostly with distinct interhyphal spaces,
and compose a kind of textura intricata. Ectal excipulum up to 90
~m at the central base, of textura angularis to textura globulosa.
Inner part predominantly with sUbglobose cells 8 - 25
(-31) ~m diameter; the polyhedral cells, up to 25 x 12 ~m, are
orientated with their longitudinal axes somewhat perpendicular to
the outer surface. Towards the exterior, especially along the sides
and towards the margin layer with long-celled rooting hyphae, 3 - 5
~m wide, originating several cell-layers deep, terminating in
slightly thick-walled, light yellow-brown bulbous parts up to 11 ~m

wide and up to 25 ~m in length; the cells stain strongly in Cotton

blue. Anchoring hyphae 2 ~m wide, ~lightly thick-walled, septate,.
sUbhyaline to light yellow brown.

Asci 8-spored, cylindrical, gradually narrowing into a stem-like

base, with truncate to slightly rounded apex with a well marked
operculum, (110-) 140 - 196 x (10-) 11 - 15 jlm, p. sp. 100·- 125
jlm, staining uniformly in Congo Red in ammonia. Ascospores
ellipsoid to subfusiform, slightly inequilateral, obliquely
uniseriate, hyaline, .thick-walled, granular 'within, 12 - '16 x
7 - 9.5 jlrn (apiculi not included), x = 14.89 x 7.93 jlm (n = 35),
Q = 0.74 (length), 0.63 (width), apparently smooth at low
magnification, however, ornamented with a rather even layer of very
fine granules (oil immersion at x 1000). This is especially
 44

conspicuous in Congo Red in ammonia. Both poles with hyaline, more

or less spherical or hemispherical to slightly elongated apiculi up
to 3.5 ~m diameter, staining strongly in Cotton blue, each apiculus
with a distinct projecting collarette at the "base, 1 - 2 ~m high
and up to 3.8 ~m·broad. Each ascospore surrounded with a thick
gelatinous iayer. Both apiculi and gel"atinous layer blackening and
greatly swelling when mounted in aqueous Indian ink. paraphyses
hyaline, of two types. The one type filiform, septate, branched or
unbranched, 1.2 - 2 ~m wide, not or only slightly enlarged at their
apices; the other type septate below, upper part from 50 - 85 ~m in
length, unseptate, inflated at apex, 3 - 6 ~m wide.

Specimens examined:

SRI LANKA. Southern Province: Hambantota Distr.: Yala National

Park, southernmost part near the coast, elephant dung in dry, open
jungle, (mc), 1l.X1.1974 N. Lundqvist 8965-c (UPS). New to Asia.
IVORY COAST: elephant dung, 2.1.1988 M. Lenz (description and
photos received, in litt. 7.X1.1989, from private herbarium of J.
Haffner, Mittelhof, Germany). New to the Ivory Coast.
~ENYA. Central Prov.: Thika Distr.: c. 5 km W.S.W. of Thika: B.
Harris Farm. Alt. c. 1.500m. 1° 4' S. 37° 3' E., cow dung, (mc) ,
24.1.1970 N. Lundqvist 6650-h (UPS - slide only).
TANZANIA. ~ilimanjaro Distr.: Mt. Kilimanjaro, W. slope: E. of
Lemosho Glades, montane forest, Alt. c. 2400 m., 3° 1'- 2' S., 37°
9' E., elephant dung, (mc) , 12.1.1970 N. Lundqvist 6381-b (UPS). --
Mt. Kilimanjaro: Loitokitok. Holotype.

UNVERIFIED RECORD: ~ENYA: Krug & Khan (1987: 200) on elephant

dung.
 45

HABITAT ~D SUBSTRATE PREFERENCE: I. africanus is a fimicolous

species, known from dung of elephant 4 (5) and cow 1, in vegetation
dominated by tropical rain forests and savannas. The African
elephant (Loxod6nta africana) is today a vulnerable species,
whereas the Indian elephant (Elephas maximus) i~ an endangered
species (Semb-Johansson 1986). Accordingly, T. africanus must be
regarded as vulnerable.
DISTRIBUTION: The species is known from tropical Africa and is
also reported for the first time from Asia (from Sri Lanka). Its
tropical equatorial distribution is limited between approximately
6°N and 4°5. The species is .regarded to be end~mic to the
Palaeotropics.
PHOTOS: Krug & Khan 1987, figs 1-7.

In their original pUblication, Krug & Khan (1987) record the

ascospores as smooth. However, both in 5EM and at high
magnification under the light microscope the ascospores are clearly
ornamented with a rather even layer of very fine granules (figs.
3 - 5). This is especially easy to observe when the spores are
mounted in Congo Red in ammonia. Moreover, the paraphyses are also
of two types contrary the original description.

Together with Thecotheus perplexans, I. africanus is characterized

by its distinct projecting collarettes at the base of each
apiculus. These two species are easily separated by the size of the
ascospores. The term collarette was introduced by Faurel and
Schotter (1965 b), when they erected the new genus Ascophanopsis,
with h. perplexans as the type.

Korf (1973) considered the genus Ascophanopsis to be a synonym of

Thecotheus. He was later followed by Krug & Khan (1987), who
regarded the presence of collarettes as insufficient for separation
at generic level. Tentatively I follow these authors. It would be
of interest to examine living material, particularly regarding the
 46

number of ascospore nuclei.

Figs. 1 - 6. Thecotheus africanus. 1 - 2: Ascospores as seen under

the light microscope, showing characteristic inequilateral form.
3 - 5: SEM of ascospores showing apiculi and ornamentations
consisting of small granules. 6: Ascospores in optical section,
showing apiculi and collarettes. 1 - 6 a: from Tanzania. Lundqvist
6381-b (UPS). 6 b: from Sri Lanka. Lundqvist 8965-c (UPS). Scales:
1: 25 ~m. 3 - 4: 5 ~m. 5: 0.5 ~m. 6: 10 ~m. Photos: 1 - 2: O. Aas.
3 - 5: J. Berge.

2. THECOTHEUS BIOCELLATUS (Petrak) Aas n. comb. (Figs. 7 - 9)

Basionym: Ascophanus biocellatus Petrak, Annales Mycologici 23

(1/2): 115 (1925). -- Type: Czechoslovakia, Moravia, near Mahrish-
Weisskirchen, at the shore of lake Betschwa by Chorin, on horse
dung, IX.1924 F. Petrak 478 (W 12224 eX.herb. F. Petrak, holotype).

ETYMOLOGY: Latin bi, two; ocellatus, which has eyes; referring to

the apiculus at each pole of the spores.

Apothecia solitary, 'rarely'a few in groups, superficial, sessile on

a broad base, up to 2 mm diameter when dry, 1.5 - 4 mm diameter
when fresh, discoid, after drying with red-brown colour (8D7-8E7).
Disc flat, then slightly convex, at first milk white or white
 49

6
1

greyish, finally when dry with dark grey or grey-brown colour.

Marqin broad, slightly incurved, with more or less irregular
surface, orange to brown-orange (6B7-6C7). Hymenium up to 220 ~m

thick. Hypothecium 20 - 40 ~m thick, consisting of small, hyaline,

subglobose cells.

Medullary excipulum of varying thickness, up to 75 (-100) ~m in the

central part, consisting of hyaline, thin-walled, subglobose to
slightly polyhedral cells 3 - 14 ~m diameter. The cells are
frequently conspicuously oblong, with their long axes in different
directions. In between with filamentous hyphae, 1.8 - 2.5 ~m wide.
Ectal excipulum 35 - 65 (-70) ~m thick at the margin, somewhat
 50

thicker at the central growing part, consisting of two different

kind of cells. The innermost cells globose, yellowish to yellow-
brown, slightly thick-walled, up to 10 (-12) ~m diameter. The cells
are partly intermingled with more oblong, polyhedral cells (textura
globulosa to textura angularis), partly with hyaline, thin-walled
rooting hyphae 2 - 3 ~m diameter, which begin several cell layers
deep and terminate on the surface with subglobose to oblong,
globose bulbs. The bulbs are slightly thick-walled, light yellow to
yellow-brown, 6 - 15 ~m diameter and 12 - 20 ~m in length. This
part of excipulum cells with intercellular, amorphous, brownish
pigment. Towards margin more or less clavate, paraphyses-like
bulbous, terminating cells up to 5 ~m in diameter. Base of the
apothecia anchored to the substrate with sparingly septate,
sUbhyaline to very light yellow-brown hyphae 2 - 3 ~m diameter.
Characteristically 2-4 hyphae are often connected in parallel
groups.

Asci eight-spored, cylindrical, somewhat truncate above, gradually

tapering below, 170 - 210 x 11 - 15 ~m, p. sp. 95 - 120 ~m.

Ascospores obliquely uniseriate, broadly ellipsoid, flattened along

one side, slightly truncate towards the apices, occasionally
equilateral, ellipsoid, (14-) 16 - 19 x 7 - 9 (-10) ~m, x = 17.12 x
8. 12 ~m (n = 37), Q = 0.78 (length), 0.74 (width); each ascospore
surrounded with a thick, gelatinous envelope wh~Ch soon disappears
after spore liberation. The spore ornamentation consists of small
punctate-like warts 0.2 - 0.9 (-1.2) ~m wide. Each·ascospore with
prominent bipolar apiculus of varying shape, hemispherical, flat or
short-pointed conical, occasionally collar-like in form, 1.5 - 4 ~m

wide, 2 - 4 ~m high; outer wall layer 0.2 - 0.3 ~m staining in

Cotton blue. Paraphyses filiform, septate, branched or not, below 2
- 2.5 ~m diameter, apices irregular spherical to clavate, inflated
to (3-) 4 - 6 (-7) ~m diameter, extending 10 - 15 ~m beyond the
asci.. This upper part of the paraphyses, especially towards the
margin, is encrusted with amorphous brownish pigments.
 51

Specimens examined:

CZECHOSLOVAKIA. Moravia: Type collection (see above).

Thecotheus biocellatus is known only from the type collection and

is probably a very rare species. Unfortunately the examined
gathering consists of only half an apothecium, therefore no
material could be used for SEM of the ascospores. However, the
species is well characterized by its broad, inequilateral,
ellipsoid and apiculate ascospores, which are furnished with fine
punctate ornamentation. Ascospores with ± same form and
ornamentation are found in T. rivicola, however, the latter has i.
a. quite different paraphyses and slightly larger spores. Also T.
lundqvistii possess ascospores which are ornamented with minute
granules, but the spores are regular ellipsoid and larger than
those of T. biocellatus. Data concerning colour and size of living
apothecia as given above, are based upon the original description
of Petrak (1925).

Typification

The original herbarium sheet of this collection was named

Ascophanus "ocellatus" Petr. n. sp., but later Petrak (1925)
published it under the name ~. biocellatus. This was the only
specimen referred to in the original pUblication, and the examined
collection must therefore be the holotype.

Figs. 7 - 9. Thecotheus .biocellatus. 7: A half of an apothecium of

discoid form with broad, slightly incurved margin. 8: Apiculate,
ornamented ascospores. 9 a: Ascospores in optical section. 9 b:
Ascus with spores. Scales: 7: 1 mm. 8: 10 ~m. 9~ 24 ~m. Photos: 7:
O. Aas. 8: J. Berge.
 53

a b

9
 54

3. THECOTHEUS CINEREUS (Cr. , Cr.) Chenant. (Figs. 10 - 24)

Bull. Soc. Mycol. Fr. 34: 39 (1918). -- Basionym: Ascobo1us

cinereus Cr. & Cr., Ann. Sci. Nat. Bot. IV. 10: 194 (1858).
Ascopbanus cinereus (Cr. & Cr.) Boud., Annls. Sci. Nat. Bot. 5 ser.
10: 249 (1869).; nom. non fungus. -- Mo11isia cinerea [sic!) (Cr. &
Cr.) Gill., Champignons de France. Les Discomycetes: 134 (1879);
comb. illeg., non Mo11isia cinerea (Batsch: Fr.) P. Karst.
Ascopbane11a cinerea [sic!] (Cr. & Cr.) Faurel & Schotter, Cah. La
Maboke 3 (2): 130 (1965 b). -- Type: France, Finistere, near Brest,
on old cow dung in a marsh, autumn, s. dato, Crouan brothers (CO
A2395, holotype; PC A2336 herb. Montagne, isotype).
Tbecotbeus setisperma Le Gal, Bull. Soc. Mycol. Fr. 78: 411
(1963); nom. novo ill. (superfluous name change). -- Type: as
above.

Misapplied names: Ascobo1usjAscophanus cinereus auct. = ~.

crustaceus. -- Humarina pa11ens (Boud.) Seaver sensu Seaver, The

North American Cup-fungi (Operculates): 124 (1928); non Ascophanus
pa11ens Boud. =Thecotheus pallens (Boud.) Kimbr.

ETYMOLOGY: Latin cinereus, ash grey; referring to the colour of

apothecia as described in the original description of the Crouan
brothers.

Apotbecia scattered, 'rarely a few crowded, superficial, sessile,

0.5 - 2 (- 4 as fresh) mm diamet~r, up to 0.7"mm high; somewhat
doliiform on a broad base to more or'less discoid when dry and
immature, finally expanding and discoid, often with an irregular
external form. Upper part concolourous with the disc, lower part
light to darker grey with a brownish tint (grey-brown - 803). Disc
flat to ± convex, after drying with a depressed,. concave surface,
 55

somewhat rough and furfuraceous owing to the protruding tips of

asci. At first pale grey, when dry with brown-grey (6C2-7C2) to
orange grey (6B2) colour. Margin distinct, rough, slightly
incurved, irregular and less distinguishable on expanded mature
apothecia. Hymenium up to 340 ~m thick.

Medullary excipulum composed of a textura globulosa with hyaline,

thin-walled, globose to sUbglobose cells, 4 - 6 - 15 (- 19) ~m in
diameter, intermingled with filamentous hyphae, 2 ~m diameter,
occasionally with some ± isodiametric polyhedral cells. Towards
excipulum a zone of parallel, long-celled hyphae, comprising a
textura porrecta. Ectal excipulum up to 150 ~m thick at the central
base, about 20 - 30 ~m thick near the margin layer. The innermost
few cell-layers made up of globose to subglobose cells, 5 - 11 ~m

diameter. Outer, several cell-layer thick part, composed of a

textura angularis, with elongated polyhedral cells 8 - 22 x 6 -12
~m, their longitudinal axes lying somewhat perpendicular to the
outer surface. The outermost cells with slightly yellow-brown
pigmented walls. The exterior part consists of thick-walled, grey-
orange to brown-orange encrusted, sUbglobose to clavate terminal
cells 5 - 9 ~m wide and 6 - 8 ~m long. The rooting hyphae, which
originate several cell layers deep, consist in their lower part of
septate, long-celled hyphae, 2 ~m diameter. Towards margin a
gradually change-over to para~hyses-like cells.

Asci 8-spored, broadly clavate, dome-shaped above, occasionally

slightly truncate at apex when young, gradually tapering downwards
into a long, slender base, (220-) 250 - 320 (- 345) x 35 - 45 ~m,

p. sp. up to 195 ~m. Ascospores oblong ellipsoid or somewhat

cylindrical-ellipsoid, oc~asionally sub-ellipsoid, biseriate,
smooth, non-apiculate, (29-) 32 - 41 (-46) x (13-) 14 - 18 (-21.5)
~m, x = 35.24 x 16.46 ~m (n = 181), Q = 2.90 (length), 1.74
(width); outer 0.5 - o. 8 ~m wall layer staining in Cotton blue,
easily loosening when heated or revived in KOH. Immature ascospores
thick-walled and hyaline, becoming thin-walled at maturity. Mature
ascospores with amber yellow (4B6) to grey-yellow (4B4) colour,
inner contents slightly granular. Each spore surrounded by a thick
 56

gelatinous envelope. Paraphyses consisting of two types, the one

filiform, septate, unbranched or branched, hyaline below, this part
1.5 - 2 ~m wide, with non-inflated, hyaline apices 2 - 3.2 ~m, the
other with inflated, nearly clavate apices up to 4 - 6 (- 8) ~m

rliameter. The 'latter ones slightly thick-walled, with apices

containing brown-orange (5C4) to grey-orange (5B4) content.

Specimens examined:

FRANCE. Finistere: Holotype and isotype (see above); in Crouan &

Crouan (1867: 56) as Ascob. cinereus.
GERMANY. Y: Erzgebirge mts., [loca'lity 'unreadable], dung
[probably of cow], VIII.1912 Th. Kupka as Ascob. (incanus)
holmskjoldii Hansen (M). -- Bayern: Ober-Franken: Burgkunstadt,
dung mixed with sand and soil in hotbed in a greenhouse, 25.VI.1909
H. Ade (S - ex herb. H. Rehm as Ascoph. holmskjoldii Hans.). --
Sachsen: Dresden: Nossen, cow dung in a greenhouse, 4.VIII.1887 W.
Krieger, Krieger: E. sax. Exs. No. 335, determ. by H. Rehm as
Ascob. (Ascoph.) incanus Phill. (BPI, CUP, H, M, NY, S, W 18064 and
W 1148, WRSL); in Krieger (1888), Kimbrough (1969: 110) as T.
cinereus (Cr. & Cr.) Chenant.
CHINA. Huang - shan: Anhewi, dung [probably of cow], s. dato S.
Q. Deng 567, determ. by S. H. Ou as Ascoph. isabellinus Clements
(BPI). New to Asia.
U.S.A. New York: The New York Botanical Garden, [on manured soil]
in greenhouse, V. (s. annus) F. J. Seaver (NY); in Seaver (1928:
112) as Ascoph. isabellinus Clem., Kimbrough (1969: 110) as
holotype of Ascoph. isabellinus Clem. [sic!]. -- soil in
conservatory, (s. annus) B. o. Dodge & F. J. Seaver (NY - two
slides only); in Seaver (1942: 310) as Ascoph. isabellinus Clem.
-- Botanical Garden, sandy soil sparingly overrun with algae,
9.VII.1913 F. J. Seaver as (in herb.) Ascoph. pallens Boud.
(NY - no apothecia found, only two slides); in Seaver (1928: 124)
as Humarina pallens (Boud.) Seaver, Pfister (1982: 8 & 22) as T.
pallens (Boud.) Kimbr.
 57

UNVERIFIED RECORDS: BELGIUM: Coemans (1862: 88) as Ascob.

cinereus Crouan, cow dung. The same collection is also referred to
by Coemans (1863: 138), Kickx {1867: 477) and Lambotte (1880: 508).
However, it is difficult to decide if this collection belongs to T.
cinereus or not. The material could not be found in the herbarium
BR. -- Mouton (1886: 141) as Ascoph. cinereus Cr., 'donkey dung.
SWEDEN: Starback (1898: 216) as Ascoph.' cinereus (Cr.) Boud. from
Ostergotland. See further comments under T. crustaceus. -- MOROCCO:
Malen90n & Bertault (1967: 241) as Ascoph. cinereus, cow dung. No
description accompanies this and several other species listed by
the authors. -- ECUADOR: Patouillard & Lagerheim (1893: 146) as
Ascoph. cinereus (Cr.) Boud., horse ,dung. NO'description
accompanies,this species. The authors make reference to Boudier's
(1869)' figure of Ascoph. cinereus, which belongs to t.
holmskjoldii.

INCORRECT RECORDS: ARMENIA: Prokhorov & Taslakhcyan (1987: 147)

as T. cinereus (Cr. & Cr.) Chenant. is T. holmskjoldii. -- AUSTRIA:
Schweiger (1985: 64) as T. cinereus (Cr. & Cr.) Chenant. is T.
holmskjoldii. -- CZECHOSLOVAKIA: Rehm (1878, 1881, 1896: 1092) as
Ascob. cinereus Cr. is T. holmskjoldii. This collection also
referred to by Massee (1895: 175, see however comments under T.
crustaceus; Kimbrough (1969: 110) as T. cinereus (Cr. & Cr.)
Chenant. -- DENMARK: Hansen (1867: 272) as £. cinerea (Cr.) Karst.
is T. crustaceus. -- Lind (1913: 103) as Ascoph. cinereus (Cr.)
Boud. is T. crustaceus. -- ENGLAND: Berkeley & Broome (1865: 449)
as Ascob. cinereus Cr. is T. crustaceus.-- Rabenhorst (1865: 252)
as Ascob. cinereus Cr. is T. crustaceus. -- Cooke (1871: 731) as
Ascob. cinereus Cr. is T. crustaceus. -- Phillips (1887: 308) as
Ascoph. cinereus (Cr.) is T. crustaceus. -- Massee (1895: 175) as
Ascoph. cinereus is T. crustaceus. -- Ramsbottom (1932: 314) as
Ascob. cinereus Cr. is T. crustaceus. -- Ramsbottom & Balfour-Brown
(1951: 59) as Ascoph. cinereus (Cr.) Boud. is T. crustaceus.
Ellis & Ellis (1988: 115) as T. cinereus is T. holmskjoldii.
ESTONIA: Prokhorov (1989 b: 287) as T. cinereus (Cr. & Cr.)
Chenant. is T. holmskjoldii. FINLAND: Karsten (1867) as Ascob.
 58

cinereus Cr. is T. crustaceus. Karsten (1871 a: 205 & 1871 b:

231) as Ascob. cinereus Cr. is T. crustaceus. -- Karsten (1871 c:
59) as £. cinerea (Crouan) Karst. is T. crustaceus. -- FRANCE:
Boudier (1869: 249) as Ascoph. cinereus (Crouan) Boud. is T.
holmskjoldii.-- GERMANY: Fuckel (1870: 288) as Ascob. cinereus Cr.
is ? T. crustaceus. See comments under the latter species.
Winter (1873: 146) as Ascoph. cinereus Boud. is T. holmskjoldii.
Beyer et al. (1986: 62) and Hohmeyer et al (1989: 30) as T.
cinereus is T. holmskioldii. -- HUNGARY: Banhegyi et al. (1985:
654) as T. cinereus (Cr.) Chenant. is T. holmskjoldii. -- ITALY:
Spegazzini (1878: 236) and Vido (1879: 559) as Ascoph. cinereus
(Cr.) Boud. on dung of cow is ? T. crustaceus. -- Bizzozero (1885:
341) as Ascoph. cinereus (Cr.) Boud. is T. crustaceus. -- Saccardo
(1889: 531) as Ascoph. cinereus (Cr.) Boud. is T. crustaceus. --
NETHERLANDS: Oudemans (1900: 210) as Ascoph. cinereus (Cr. & Cr.)
Boud. is ? T. crustaceus. See comments under the latter species. --
SPAIN: Guarro (1983: 236) as T. cinereus (Cr. & Cr.) Chenant. is T.
holmskjoldii. -- Barrasa (1985) as T. cinereus (Cr. & Cr.) Chenant.
is T. holmskjoldii. -- Calonge et al. (1986: 33) as T. cinereus
(Cr. & Cr.) Chenant. is T. holmskjoldii. -- SWEDEN: Starback (1898:
216) from Uppland as Ascoph. cinereus (Cr.) Boud. is T. crustaceus.
INDIA: Kaushal (1980: 133) as I. cinereus (Cr. & Cr.) Chen. is
T. holmskjoldii. ISRAEL: Binyamini (1973: 154) as T. cinereus
CCr. & Cr.) Chenant. is T. holmskjoldii.. -- NEW ZEALAND: Bell
(1983: 33) as T. cinereus (Cr. & Cr.) Chenant. is T.
holmskioldii. -- RUSSIA: Ta~zhik: Raitviir & P~okhorov (1988: 218)
as T. cinereus (Cr. & Cr.) Chenant. is T. holmskjoldii. -- ALGERIA:
Faurel & Schotter (1965 a: 271) as Ascoph. cinereus (Crn.) Le Gal
is T. holmskjoldii. -- CANADA: Conway (1975) as T. cinereus is T.
holmskjoldii. -~ U.S.A.: Seaver 1904: 276; 1905 a: 113; 1909: 109;
1911: 60 & 1928: 118) all as Ascoph. cinereus (Cr.) Boud. is T.
crustaceus. Brenckle (1917: 277) as Ascoph. cinereus (Cr.) Boud.
is probably T. crustaceus. -~ Seaver (1942: 310) as Ascoph.
cinereus is T. keithii. Kimbrough (1969: 110) as T. cinereus
(Cr. & Cr.) Chenant. is T. holmskjoldii. -- Pfister (1982: 8) as T.
cinereus (Cr. & Cr.) Chenant. is T. crustaceus. -- BERMUDA:
Kimbrough (1969: 110) as T. cinereus (Cr. & Cr.) Chenant. is T.
 59

holmskjoldii. -- ARGENTINA: Spegazzini (1899: 308) as Ascoph.

cinereus (Cr.) Boud. is T. crustaceus.

DUBIOUS RECORDS: GERMANY: Britzelmayr & Rehm (1877: 62) as Ascob.

cinereus Cr., cow dung. I have examined this collection of
Britzelmayr from 1876 (herbarium S), however, no species of
Thecotheus were found, only some unripe fruit-bodies of Ascobolus.
-- SPAIN: AIDo & Mora (1870) and Unamuno (1941: 310) as Ascoph.
cinereus Boud., no locality mentioned, on wood and rotten twigs.
Unfortunately, this material seems to be lost, according to J. M.
Barrasa, University of Alcala de Henares, Madrid, (in litt.
27.1.1986). It does not belong here, but may be one of the
lignicolous species of Thecotheus. A reference to Unamuno's (loc.
cit.) list of species is found in Lundqvist (1960).

HABITAT AND SUBSTRATE PREFERENCE (only examined material

included): on dung of cow, or soil most probably mixed with manure.
DISTRIBUTION: France, Germany, China, U.S.A.
ILLUSTRATIONS: Crouan & Crouan 1858, pI. 13 D figs. 17-20.
Seaver 1928, pI. 11, fig. 13 as Ascoph. isabellinus Clem. -- Le Gal
1960, fig. 1 c-d.

Chenantais (1918), Le Gal (1960) and many later authors have

regarded Ascophanus holmskjoldii as conspecific with this species.
I do not agree with this irtterpretation. T. holmskioldii has i. a.
verrucose, sUbfusiform spores with rather persistent plasma
papillae, whereas T.cinereus has smooth, narrowly ellipsoid and
non-apiculate ascospores, except in a few spores in which polar
plasma accumulations are easily disolved. Obviously the two are
closely related, but distinct species.
 60

The material of Ascobolus cinereus Cr. novo spec. in the herbarium

Crouan, Concarneau, is rather poor. It consists of one envelope
contai~ing dung fragments and only two an a half fruitbodies. The
smallest one, apparently not fully ripe, has a slightly doliiform
shape with a distinctly deveioped margin. The other entire·
apothecium, 1 mm in diameter when dry, has an irregular discoid
form. According to Crouan & Crouan (1858) the apothecia measure up
to 4 mm diameter. The text on the envelope says: "Croft sur les
bouses de vache anciennes. en Automne. Environs de Brest. tres
rare." Text on the envelope of the isotype collection in herbarium
Montagne, Paris say: "croit sur les bouses, de vache, anciennes.
Dans les marais. Environs de Brest. print. aut. rare. Ex. herb.
Crouan." This latter collection is rather poor. It is greatly
fragmented and consists of two small dung fragments, each with one
more or less compressed fruitbody. Although samples of both
holotype and isotype of Ascobolus cinereus are very small, it is
nevertheless possible to understand the status of the species.

Ascobolus cinereus has probably been one of the most misunderstood

species of Thecotheus. Undoubtedly, only very few literature
records of h. cinereus have to do with the true Ascobolus cinereus.
The misinterpretations have lead in two different directions. The
first is based on Rabenhorst (1865) who distributed an English
specimen collected by C. E. Broome 1864 as "Ascobolus cinereus
Crouan" in his Fungi Europaei Exsiccati No. 783, and this lead many
authors to consider the taxon as small-spored. So did i. a. Karsten
(1867, 1871 a, 1871 b, 1871 c and 1885), Phillips (1887), Saccardo
(1899), Massee (1895), and Seaver (1905 a., 1909, 1911, 1928 and
1942). This can be explained by the fact that Crouan & Crouan
(1858) did not make any note of the size of ascospores and asci. It
would have been possible by interpolation to see that the
ascospores must have been 36 - 42 ~m in length, which is in
agreement with my own observations on the material. I have examined
the greater part of No. 783, and all belongs to Thecotheus
crustaceus.

The other line of misinterpretation started with Boudier (1869),

 61

who regarded Ascobolus cinereus to be a species with warted spores.

Boudier was later followed by the French mycologists Chenantais
(1918) and Le Gal (1960, 1963). Boudier's (1869) description and
drawings of h. cinereus belongs to T. holmskjoldii, an~ many later
authors for that reason have interpreted T. holmskjoldii as a
synonym of T. cinereus.

Also Rehm to some extent confused the interpretation of Ascobolus

cinereus. In his Ascomyceten No. 470, he distributed Ascobolus
cinereus Crouan based on a gathering of von Niessl from 1877. This
material belongs to T. holmskjoldii, and it is notable that Rehm
(1896: 1093) doubted the deter~ination of'von Niessl: "Ich vermag
den Pilz aber weder mit Crouan (Ann. sc. ,nat. IV. pag. 194, T. X,
pI. 13, V, fig. 17 - 20) noch mit Karsten (Myc. fenn. I. pag. 59)
[= T. crustaceus] noch vollstandig mit Phillips (Man. brit. Discom.
pag. 308) [= T. crustaceus], Fuckel (Symb. myc. pag. 288) [= T.
holmskjoldii] und Michelia I. pag. 236 [= T. holmskjoldii] zu
vereinigen; auch ist Rabh. Fungi europ. 783 auf Pferdekoth in
England ganz verschieden". The latter collection belongs to T.
crustaceus.

Chenantais (1918) interpreted Ascobolus cinereus to be so closely

related to Ascophanus holmskjoldii, that he regarded them as
varieties of the same species. Mainly on account of their spores he
transferred both species to the genus Thecotheus. still his species
concept which is difficult to understand, lead to further
misunderstandings.

Le Gal (1960) re-studied the type collection of Ascobolus cinereus

and argued that it should be kept in the genus Thecotheus. However,
she found some episporial remains enclosing the smooth ascospores.
Moreover, the perispore layer formed polar apiculi on each
ascospore. Also in 1963 she mentions epispore granulations. In Le
Gal's opinion such episporial remains resembled those of Ascophanus
holmskjoldii, and she therefore regarded this latter species as a
synonym of T. cinereus. I have seen the holotype and Crouan1s
original drawings which were made from fresh material, and the
 62

ascospores are completely smooth. I did not observe granulated

ascospores. Observations made by van Brummelen (1967) and N.
Lundqvist (pers. comm.) on this material of Crouan's confirm my
findings. It is therefore clear that both Chenantais and Le Gal
were mistaken.

A particularly rich and widespread collection of T. cinereus,

Krieger's Fungi saxonici No. 335 as Ascobolus (Ascophanus) incanus,
has greatly contributed to a better understanding of the species.
During SEM studies this collection, of ascospores lying on the
surface of the hymenium, I also observed some characteristic
protuberances on the spores. At regular intervals there are also
globulose, spinose bodies lying on the hyphoid threads, both
structures seemingly as outgrowths from the ascospores(figs. 14 -
17). The hyphoid threads are about 0.5 ~m in width, whereas the
diameter of the globulose bodies varies between 3 and 4 ~m, with
spines up to 0.35 ~m high. If the structures belong to the
ascospores, they must be germ tubes in process of formation. within
the pyrenomycetes, in several genera e. g. Cercophora and
Lasiosphaera, the spores can germinate directly with phialides
"(Lundqvist 1972). It is, however, hard to believe that germ tubes
can form chlamydospores. To my knowledge this has not been observed
before.

Another more likely possibility is that the structures may belong

to a parasite. Lundqvist (in litt.) has informed me that he has
found a Podospora with spores apparently attacked by some kind of
parasite, .but looking quite different to what is observed here.
Lohm~n (1942~ described a chytridiaceous parasite on different
fungal structures of several dung-inhabiting ascomycetes, among
others on ascospores of Ascophanu5 holmskjoldii. The zoosporangia
were typically sessile, occasionally short-stalked, smooth,
hyaline, ± globose, and from 7 to 36 ~m in diameter. However, the
observed structures on Krieger's collection clearly differ from
 63

those found by Lohman (1942). I have not been able to reach a final
conclusion as to their nature.

The NY collection of Seaver as Humarina pallens (Boud.) Seaver was

examined by Pfister (1981, 1982), who found it to be a distinct
species of Thecotheus. I have seen the NY material, but no
apothecia were found, only two well preserved slides could be
studied, and I did not find any ascospore which was approximately
oblong-fusiform in shape, as described and figured by Boudier
(1888). An unpublished herbarium sketch by Seaver (fig. 24) of this
particular collection shows asci with biseriate, ellipsoid
ascospores, quite different from Boudier's original drawings of
Ascophanus pallens.

Further studies, preferably of fresh material of T. cinereus are

necessary.
 64

Figs. 10 - 24. Thecotheus cinereus. 1~: Krieger 335 as. Ascobolus

incanus (S). Mature apothecium. 11: Ditto (BPI). 12 - 13: Isotypus
of Ascobolus cinereus (PC - A2336, herb~ Montagne). Ascospores.
14 - 22: Krieger 335 as Ascobolus incanus (S). 14: SEM- survey of
~scospores with hyphoid protuberances.bearing globulose bodies at
regular intervals. 15: SEM- close up of a globose structure showing
a characteristic spiny pattern. 16: SEM- close up of a young
globose structure with a spiny pattern. Note the smooth spore
surface. 17: SEM- photo of two globulose, ornamented bodies on
hyphoid threads which originate from the ascospore-surface (bottom
right). 18: Part of medullary excipulQm with globose-subglobose
cells. In between with some ± isodiametric polyhedral" cells and
intermingled filamen~ous hyphae: 19: Part of ectal excipulum. To
the left some globose terminal cells of rooting hyphae. Then a zone
of cells composing a textura angularis, their longitudinal axes
lying somewhat perpendicular to the outer surface. 20: Survey of
excipulum. Ectal excipulum layer to the left. 21: Paraphyses of
filiform type. 22: Paraphyses of thick-walled, encrusted and
inflated type. 23: Seaver V. (s. annus) as Ascophanus isabellinus
(NY). Apothecia. 24: Seaver's herbarium sketch of Humarina
(Ascophanus) pallens (NY). Scales: 10 - 11: 1 mm. 12 - 13: 20 ~m.

14: 10 ~m. 15 - 17: 1 ~m. 18: 22 ~m. 19: 16 ~m. 20 - 21: 32 ~m. 23:
0.6 mm. All photos: o. Aas.
65
67
 69

';.I~ •.

. t--r '\.'
. \' \

\ ""J' \
\ ~
~
\
\
j
\

\
\\
\ "/-~ \
.\ ./
,.' t,.

...
I

.r
\

.......... '

,..... -:.~J>". . ., ,

'~.. . ''')' .
.,~, ")

24
 70

4. THECOTHEUS CRUSTACEUS (Starb.) Aas , Lundq. n. comb.

(Figs. 25 - 38)

Basionym: Ascophanus crustaceus Starb., Bot. Not.: 216 (1898). --

Iodophanus crustaceus (starb.) Kimbr., AIDer. J. Bot. 56 (10): 1200
(1969); nom. sed non fungus. -- Type: Sweden, Uppland, Knivsta
(=Knifsta), Ledinge, on horse dung, VIII.1895 K. Starback 19 (S,
holotype).
Thecotheus agranulosus Kimbr., Mycologia 61 (1): 112 (1969). --
Type: U. S. A., Tennessee, Knoxville, on horse dung, 26.VI.1929 L.
R. Hesler 2005 (MICH, holotype).

Misapplied name: Ascobolus/Ascophanus cinereus auct.; non Cr. & Cr.

ETYMOLOGY: Latin crusta, crust; referring to the usually crust-

like apothecial form. Latin a, without, granulose, granules;
referring to the spores without granular markings.

Apothecia sl-ightly immersed, scattered or very frequently

gregarious to crowded, occasionally ± crust-like in shape, 0.2 - 2
mm diameter, up to 0.5 mm high. At first globose, soon becoming
sUbcylindrical with a flat top, then turbinate to sub-dolabriform
with a central part slightly immersed into the substrate, finally
expanding and doliiform to thick discoid, occasionally pUlvinate,
apparently sessile, however, with a varyiably thick base more or
less adpressed to the dung, with the point of attachment slightly
immersed. -The adpressed part becomes dispersedly crust-like in old,
overripe apothecia.

Apo~hecia at first dirty white to light grey, with a yellow tinge,

later grey or grey-brown (6D3) to pale brown, occasionally grey-
orange (5B5), after drying dark grey or black, then usually
 71

brown-grey (7C2) or dark brown, infrequent with a reddish to

purplish grey tint. Disc flat or slightly convex, at first
concolorous with the receptacle, then paler and apparently nearly
whitish and rough owing to the protruding tips of the mature asci.
Dry apothecia often with conspicuous concave disc. Margin broad,
slightly rough, incurved especially in dry apothecia. Bymenium up
to 240 ~m thick. Bypothecium composed of small-celled textura
globulosa, with cells 2 - 4 ~m diameter.

Medullary excipulum of varying thickness, up to 130 ~m at the

central growing point, composed of a textura globulosa with
hyaline, thin-walled globose to sUbglobose cells, 2 - 12 (-15) ~m

diameter, intermingled with filamentous hyphae 2 - 3 ~m diameter.

Towards excipulum a zone of long-celled tissue with sUb-hyaline or
slightly light brownish, thin-walled parallel hyphae, 1.5 - 2 ~m

diameter, comprising a textura porrecta, up to 30 ~m thick. The

hyphae of this layer extend into the excipulum, where they
terminate in inflated terminal cells of different shape and
pigmentation. Ectal excipulum up to 80 ~m thick at the central
growing point, up to 40 ~m thick near margin. Towards the exterior
part with long-celled, sUbcylindrical rooting hyphae 2 - 3 ~m

diameter, originating several cell layers deep. The cells ends in

thick-walled, brown encrusted, sUbglobular to slightly elongated
bulbous terminal cells 6 - 15 (-20) ~m wide ani up to 15 - 20 ~m in
length at the central base, with intercellular amorphous brown
pigments. Towards margin with siightly smaller and more thin-
walled, occasionally less pigmented terminal bulbous cells 5 - 10
~m diameter, which gradually transform to paraphyses-like terminal
cells. The inner part of excipulum composed of a textura angularis,
of polyhedral, thin-walled, hyaline cells which with age become
light brown pigmented. Especially towards the central base and
al·ong the sides of apothecia the cells are elongated, lying with
their longitUdinal axes somewhat perpendicular to the outer
surface, the cells up to 12 - 20 x (6-) 10 - 15 ~m. Towards the
margin more or less isodiametric cells. This outer layer of textura
angularis does not extend into the margin.
 72

Asci cylindrical to cylindrical clavate, with a somewhat ,narrowing

asymmetrical, and slightly truncate apex, gradually tapering
downwards, stalk fragile, 120 - 225 (-260) x 14.5 - 20 ~m, p. sp.
85 - 125 (-160) ~m, 1+ blue, however, occasionally faintly blue in
some old' examined ma~erial, each ascus wi~h eight ascospores.
Occasionally with 5 to 6 small, abortive spores in some asci.
Ascospores inequilateral, ellipsoid, occasionally slightly narrowed
at the poles, biseriate, or irregularly arranged or obliquely
uniseriate, (16-) 19 - 23 (-25) x (8-) 8.5 - 10.5 (-11) ~m, x
20.43 x 9.60 ~m (n = 408), Q = 1.23 (length), 0.70 (width),
hyaline, smooth (light microscope!). The spores are initially
thick-walled, but the wall becomes thinner by the time of spore
liberation. Mature spores with a thin (0.2 - 0.8 ~m) outer layer
that stains in Cotton blue, and an about 2 ~m thick inner layer
which remains hyaline in Cotton blue. Innermost part of mature
spores slightly granulate, occasionally with light yellow (4B4-4B5)
colour. Each ascospore surrounded by a thick gelatinous perispore
layer. Paraphyses filiform, septate, branched or unbranched,
occasionally with anastomoses, below (1.2-) 1.5 - 2.2 (-2.5) ~m

diameter, hyaline, their apices inflated, occasionally slightly

curved, (2-) 2.5 - 4.5 (-6.5) ~m diameter, with yellowish to grey-
yellow, slightly granulate content and with light yellow-brown
encrustations. They are embedded in amorphous, gelatinous brownish
pigment. The paraphyses extend 10 - 15 ~m beyond the asci.

Specimens examined:

AZERBAIJAN: in reserves Kyzyl-Agorch, buffalo dung, 27.1.1986 Y.

U. Maleeva (MW); in Prokhorov (1989 a: 97) as T. agranulosus.
DENMARK. Sjelland: The Zoological garden at Copenhagen, old horse
dung, 111.1874 E. Chr. Hansen (C - also a slide No. 198 in F. Fim.
Exempl. EXs.); in Hansen (1876: 272) as £. cinerea (Cr.) Karst.,
 73

Lind (1913: 103) as Ascop~. cinereus (Cr.) Boud. New to Denmark.

ENGLAND. Avon [somerset]: near Bath, horse dung, X.1864 C. E.
Broome in Rabenhors~: F~ Eur. Exs. No. 783 as Ascob. cinereus Cr.
(C, FH, K, M, RO ~ three collections, respectively ex herb. De
Notaris, N. A. Pedicino.and V. Cesati 'as Ascoph. cinereus (Cr.)
Boud., S - two collections, e~ herb. H. Rehmand P. Sydow, W 2295,
WRSL); in Rabenhorst (1865: 252), and Rabenhorst in Hedwigia 4
(11): 156 (1865). -- Batheaston, cow dung, 1864 C. E. Broome as
Ascob. cinereus Cr. (K - ex herb. Berkeleyanum); in (?) Berkeley &
Broome (1865: 449), Cooke (1871: 731), Ramsbottom (1932: 314),
Ramsbottom & Balfour-Brown (1951: 59) as Ascoph. cinereus (Cr.)
Boud. -- Bathford, dung [probably of horse], 22.X.1g64 s. legator,
as Ascob. cinereus Cr. -- SUffolk: southwold, horse dung,
18.VII.1980 ·M. B. Ellis & J. P. Ellis as T. apiculatus (IMI
253134a). -- surrey: Weybridge, dung [probably of horse], X.1857
s. legator, (K - ex herb. F. Currey, rec. XII.1881,; in Phillips
(1887: 308) as Ascoph. cinereus (Cr.), Massee (1895: 175). New to
England.
FINLAND. Tavastia australis: TammeIa: MustiaIa, horse dung, IX.
s. annus, P. A. Karsten in F. Fenn. Exs. No. 658 as Ascob. cinereus
Cr. (H, K, UPS); in Karsten (1867). horse dung (?), X.1866 P. A.
Karsten as Ascob. cinereus Cr. (H - ex herb. P. A. Karsten No.
2842). -- horse dung, 20.IX.1868 P. A. Karsten as £. cinerea
[sic!](cr.) Karst. (UPS); in (?) Karsten (1871c: 59). -- horse
dung, 20.IX.1868 P. A. Karsten as Ascob. cinereus Cr. (H - ex herb.
P. A. Karsten No. 2843); in (?) Karsten (1871a: 205, 1871b: 231). -
- cow dung, 1.X.1868 P. A. Karsten as Ascob. cinereus Cr. (S).
horse dung, 2.X.1868 P. A. Karsten as Ascob. cinereus Cr. (H - ex
herb. P. A. Karsten No. 2844). -- horse dung, 5.X.1868 P. A.
Karsten as Ascob. cinereus Cr. (H - ex herb. P. A. Karsten No.
2845, (UPS); in (?) Karsten (1871a: 205). -- horse dung,
20.VIII.1869 P. A. Karsten as Ascob. cinereus Cr. (H - ex herb. P.
A. Karsten No. 2846); in Karsten (1871b: 231). New to Finland.
FRANCE. Bouches - du Rhone: 4 km NNW of Saintes-Maries-de-la-Mer,
the highroad, horse dung in open landscape with pastures and
marshes, (me), 5.V.1975 N. Lundqvist 9650-a (UPS). -- Var: fIe de
Port-Cros, donkey dung, comm. J.- C. Donadini 5.111.1982 No. 2.82,
 74

isolated by J. van Brummelen 30.V1.1982 (L 6574); in Brummelen

(1984: 150) as T. agranulosus Kimbr.
NORWAY. 0stfold: Borge: near Sellebak, elephant dung from a
circus, 9.VI.1981 R. Kristiansen (BG). New to Norway.
SWEDEN. Gotland: Frojel: the horse corral at MallgArds kallmyr
(1.5 km ESE of Puser), horse dung in coniferous forest, (me),
1.VII.1988 N. Lundqvist 17225-e (S). -- Dalsland: Dals-Ed: just W
of Nybole, horse dung in pasture, 7.VIII.1961 N. Lundqvist 3134-a
(UPS). -- Vastergotland: Medelplana: RAback, old horse dung in
deciduous forest, (me), 12.VI.1960 N. Lundqvist 2430-c (UPS). --
Uppland: Knivsta (=Knifsta). Holotype. -- horse dung, VII.1895 K.
Starback 14 (S - ex herb. H. Rehm). -- Uppsala, fresh horse dung,
(me), 25.IX.1974 E.-K. Porten (UPS, S). -- Uppsala: Norra Norby,
fresh horse dung, (me), 8.11.1975 E. Gunnerbeck 2686-e (UPS).
CANADA. ontario: Nipissing Distr.: Lake Timagami: Bear Island,
horse dung, 3.VIII.1937 R. F. Cain 11571, as Ascoph. cinereus (Cr.)
Boud. (BPI, DAOM 63950 - paratype of T. apiculatus Kimbr., TRTC
11571, UC - ex herb. TRTC, W 20983 - ex herb. Petrak 41216, W 29670
- ex herb. F. Petrak 9378); in Kimbrough (1969: 111) concerning
DAOM 63950. -- horse dung, V1I.1937 R. F. Cain as Ascoph. cinereus
(Cr.) Boud. (M - ex herb. F. Petrak). -- Yukon Territory: 2 mi. NE
of Mayo, moose dung in wooded area, 1.VIII.1949 Caddy, Ballard,
Gillette and Mitchell, developed on dung in late winter 1950, det.
B. B. Kanouse as Ascoph. cinereus (Cr.) Boud. (MICH - paratype of
T. agranulosus Kimbr.).
U. S. A. Michigan: Ann Arbor, horse dung, 25.V.1925 G. B. Cummins
10 as Ascoph. cinereus (Cr.) Boud. (NY - ex herb. G. B. Cummins,
paratype of T. apiculatus Kimbr.). -- New York: New York City:
Bronx, horse dung, X1.1914 F. J. Seaver as Ascoph. cinereus (Cr.)
Boud. (NY - slide only); in Seaver (1928: 118), Pfister (1982: 8) as
T. cinereus (Cr. & Cr.) Chenant. -- Tennessee: Knoxville, horse
dung, 26.V1.1929 L. R. Hesler 2005 as Ascoph. cine~eus (Cr.) Boud.
(MIC~ -"holotype of T. agranulosus Kimbr."); in L~rsen & Denison
(1978: 72).
ARGENTINA. Buenos Aires: La Plata, horse dung, 18.IV.1888 and
22.111.1894 C. Spegazzini (LPS 43587 and 43586); both in spegazzini
(1899: 303) as Ascoph. cinereus (Cr.) Boud. New to South America.
 7S

UNVERIFIED RECORDS: GERMANY: Fuckel (1870: 288) as Ascob.

cinereus Cr. on horse dung. The author's description indicates that
the collection very probably belongs to T. crustaceus. -- ITALY:
Bizzozero (1885: 341) as Ascoph. cinereus (Cr.) Boud. on cow dung.
Bizzozero's description leave no doubt that this collection belongs
to T. crustaceus. Not found in the herbarium at Padova (in litt.
15.XII.1984 from the herb. PAD). -- Saccardo & Sydow (1902) as
Ascoph. crustaceus Starb. Probably no material was examined by the
authors. -- NETHERLANDS: Oudemans (1900: 210) as Ascoph. cinereus
(Cr. & Cr.) Boud. on cow dung. Although no accompanying
description, the included list of literature references all
indicates a small spored species. -- SWEDEN: Starback (1898: 216)
as Ascoph. cinereus (Cr.) Boud. from osterqotland: Simonstorp:
Mela, on horse dung. Very probably T. crustaceus, like another
collection examined by mee. -- Halland: Lundqvist 2421 on cow dung;
Uppland: Lundqvist 3304, 3803, 4102 and 4279, all on cow dung;
10223 on horse dung; Inqermanland: Lundqvist 2648 on cow dung; 3379
on horse dung. -- JAPAN: Otani (1973: 33) as T. agranulosus, two
collections on horse dung. This specimen could not be located in
the herbarium of TNS where Y. otani's herbarium is now housed (Dr.
Y. otani in litt. 20.111.1985 and Dr. H. Inoue, TNS, in litt.
2.IV.1985). -- NEW ZEALAND: Bell (1983: 33) as T. agranulosus, no
substrate mentioned, but Ann Bell (in litt. 25.111.1986) has kindly
informed me that she collected the fungus on dung from 'an unknown
animal from Wellington Zoo 16.VII.1971. Part of the collection is
deposited in the Plant Diseases Herbarium (D.S.I.R.), Private Bag,
Auckland with No. 29580. The remaining part of the collection is
deposited as No. 89 in her private herbarium, which now is part of
the official WELTU herbarium. I have not been able to study the
material, but J. Kimbrough who saw a sample, agreed that it was T.
agranulosus. -- U.S.A.: Seaver { 1904: 276, 1905 a: 113, 1909: 109,
1911: 60 & 1928: 118) all as Ascoph. cinereus (Cr.) Boud., on horse
dung. Probably all these-small-spored gatherings belongs to T.
crustaceus. Seaver's collection from North Dakota (1909) is-also
mentioned in Brenckle (1917: 277). -- Brenckle (1917: 277) as
Ascoph. crustaceus Starb. on horse manure from North Dakota.
Larsen & Denison (1978: 72). -- ARGENTINA: Spegazzini (1899: 308)
 76

as Ascoph. cinereus (Cr.) Boud., two collections on horse dung.

Undoubtedly, all belong, to T. crustaceus.

INCORRECT RECORD: SOUTHERN AFRICA: Le Gal 1963 as Ascophanus

crustaceus Starb. is Iodophanus magniverrucosus Aas sp. novo
(S~e appendix B).
HABITAT AND SUBSTRATE PREFERENCE: T. crustaceus is exclusively a
fimicolous species: 33 (48) finds, from dung of horse 27 (36), cow
2 (8), buffalo 1, donkey 1, elephant 1, moose 1.
DISTRIBUTION: Azerbaijan, Denmark, England, Finland, France, (?)
Germany (Ftickel 1870: 288), (?) Italy (Bizzozero 1885: 341), (?)
Netherlands (Oudemans 1900: 210), Norway, Sweden, Japan (Otani
1973: 33)', New Zealand (Bell 1983: 33), Canada, U~ S. A. and
Argentina.
ILLUSTRATIONS: Berkeley and Broome 1865, pI. 17, fig. 30 a-c
(as Ascob. cinereus Cr.) -- Seaver 1904 a, pI. 28 (as Ascoph.
cinereus (Cr.) Boud.). -- Se aver 1904 b, pI. 14, fig 1 a-d (as
Ascoph. cinereus (Cr.) Boud.). Se aver 1905, pI. 28, fig.1 (as
Ascoph. cinereus (Cr.) Boud.). -- Se aver 1928, pI. 11, fig. 11 (as
Ascoph. cinereus (Cr.) Boud.). -- Otani 1973, fig. 1 e-h ( as T.
agranulosus). -- Bell 1983, fig. 34 (as T. agranulosus).
Brurnrnelen 1984, figs. 3-4 (as T. agranulosus).
PHOTOS: Kimbrough 1969, fig. 20 (as T. apiculatus Kimbr.), figs.
24-27 as (T. agranulosus). -- Otani 1973, pI. 1 a (as T.
agranulosus).

Notes on type collection of Ascophanus crustaceus

The holotype of Ascophanus crustaceus was collected in early August

in 1895. On a handwritten accompanying sheet of paper however,
 77

Ascophanus cinereus is said to be a synonym of b. crustaceus. This

has probably been added later, owing to reference to both page
number, year and volume of the pUblication. Another collection by
Starback, also found on horse dung from Ledinge, was collected in
July 1895. This. was originally determined by Starback as Ascophanus
cinereus (Cr.) Karsten, and lat~r called "forma obscura" by Rehm.
Very probably this is the same as Starback (1898: 216) published as
Ascophanus cinereus (Cr.) Boud. I have not found any large-spored
"cinereus" in this material; it is completely homogenous, and all
apothecia belong to T. crustaceus. It is difficult to understand
what differences Starback could see between these two collections.
The bulk of both Starback's mentioned collections will be
distributed in .Fungi Fimicolae Exsiccati by Nils Lundqvist (pers.
cornrn.).

Ascophanus crustaceus was treated as a new species by the Swedish

botanist Karl Starback in 1898. In the Latin diagnosis Starback
described the apothecia as " ... sparsa vel plerumque gregaria,
interdum crustam vagam efformantia". This crust-like pattern of
growth easily characterizes the apothecial form of this species.
Another typical, macroscopic, distinguishing mark, noted also by
Kimbrough (1969), is the "Fimaria - like" incurved apothecial
margins, especially in dried apothecia. In this condition the
fruit-bodies are to a certain extent variable in shape when still
attached to the substrate. Thus, young apothecia are subcylindrical
to dolabriform with a flat top, whereas mature apothecia are more
or less discoid and apparently seated on a broad base. Starback
(op. cit.) further described the apothecia as " ..• primum pallide
fuscidula, mox nigricantia, atra". In the original description, the
asci are said to be short-stipitate, 95 - 130 x 15 - 17 ~m, with
biseriate or obliquely uniseriate ascospores, 17 - 25 x 8 - 10 ~m;

only rarely are the ascospores uniseriate. Thus, biseriate

ascospores is another character of this species. Starback also
described the ascospores with " ... membrana crassa". The spores are
 78

initially thick-walled, but the wall becomes thinner by the time of

spore liberation. However, Starback's (1898) statement about such
thick membranes has been misinterpreted by later authors, i. a.
Chenantais (1918), Le Gal (1963) and Kimbrough et al. (1969).

The ascospores, as seen under the light microscope, appear

completely smooth. Sometimes immature spores still in asci are
observed with a very small granulation which stains in Cotton blue.
This granulation is located at one or both poles, but it is not
observed on mature spores after spore liberation. Probably this is
the remainder of the plasma from the germ pores since it stains in
Cotton blue. In SEM the ascospores appear to have a slightly uneven
surface. At the poles some very small, less than 0.5 ~m high,
granulae are occasionally seen.

Chenantais (1918) treated Ascophanus crustaceus Starback as a

synonym of Ascophanus cinereus (Cr. & Cr,) Boud. and Ascophanus
holmskjoldii Hansen. In his discussion he says that "La presence ou
l'absence de granulations est en rapport avec l'age de la spore; ce
n1est qu1un caractere d'appoint. Mieux au courant de la biologie,
l'auteur aura it conclu a l'identite de son crustaceus avec cinereus
ou Holmskjoldii". with this inclusion of Starback's species,
Chenantais (1918) brought about misconceptions about this species,
as already pointed out by Kimbrough (1969). Chenantai's opinion was
also followed in U.S.A. by Seaver (1928), who included the small-
spored Ascophanus crustaceus in synonymy with h. cinereus (Crouan)
Boud.

Le Gal (1963) called attention to the erroneous interpretation in

the paper of Chenantais (1918), and for that reason she made a
description including a thorough discussion about the differences
. .
between Ascophanus crustaceus and h. cinereus~ However, her
descr~ption of b. crustaceus was completely based upon examination
of material from South Africa and not upon Starback's material,
which was not available. The ascospores were described with large
markings up to 3 ~m high on the lateral walls, and with 4.5 ~m high
 79

apiculi at the poles. These characters, among several others,

disagree with Starback's (1898) description of h. crustaceus.

On the envelope of this sample, M. Le Gal has written that the

species is undoubtedly Ascophanus crustaceus Starback. However, she
also further remarked that the material probably belonged to
Ascophanus cinereus "de Boudier". Later, Le Gal (1963) treated
these as two different species, although she indicated that the
figure of Boudier's h. cinereus (Boudier 1869) has a shape very
similar to that of the South African specimens. Boudier's h.
cinereus belongs to T. holmskjoldii (see p. 102 ).

I have examined the collection from South Africa, and it is a

species quite different from Ascophanus crustaceus. It is here
described as a new species of Iodophanus (see appendix B).

Notes on Karsten's material of Ascobolus cinereus

T. crustaceus was collected several times by the Finnish mycologist

P. A. Karsten, from Mustiala, where he was lecturer in botany at
Mustiala agricultural college. At first Karsten (1867, 1871a,
1871b) called the species Ascobolus cinereus Cr., but placed it
later (1871c) in Peziza. It should be not~d however, that the
collections in herbarium S, H 2843 and H 2844, were first labelled.
by him as Ascobolus "fuligineus", and then correcte"d to cinereus.
Karsten and probably also Starback were undoubtedly familiar with
"Ascobolus cinereus" Crouan in Rabenhorst's Fungi Europaei
Exsiccati No. 783 (1865), which belongs to Thecotheus crustaceus.
 80

Notes on the types of ~. agranulosus and ~. apiculatus

The holotype of Thecotheus agranulosusKimbroug~was sa~d to be

deposited in herbarium TENN as No. 2005 by Kimbrough (1969). This
specimen of Hesler from 1929, could however not be found there. Dr.
Ronald H. Petersen, at the University of Tennessee, has kindly
informed me (in litt. 30.X.1984) that since the Tennessee herbarium
totally burned in 1934, no pre-1934 collections exist in this
herbarium, and could therefore not have been available to
Kimbrough. However, the collection in question was found in
h~rbarium MICH. The material had been determined by Hesler as
Ascophanus.cinereus (Crouan) Boud., restudied by Kimbrough in May
1966, and by him determined as Thecotheus sp. There is no doubt
that this specimen was available to Kimbrough, and later (Kimbrough
1969) segregated by him as holotype of T. agranulosus. Kimbrough
(in. litt. 18.II.1987) has confirmed that the correct herbarium
citation should have been MICH. It clearly belongs in T.
crustaceus.

One of the paratypes of T. apiculatus Kimbr. (Kimbrough 1969), DAOM

63950, does not belong there. This collection belongs to T.
crustaceus. Dr. J. C. Krug, University of Toronto, has kindly
informed me (in litt. January 9th., 1985) that the material housed
under TRTC 11571 " ... is part of the collection that we sent some
years ago to DAOM. This then is part of the collection that is
cited by Kimbrough as DAOM 63950." The collection TRTC 11571,
originally determined by R. F. Cain as Ascophanus cinereus (Crouan)
Boud. (see under specimens examined for further information), were
annotated by Kimbrough in 1970 as Thecotheus cinereus. Distributed
parts of this collection are also deposited in the herbaria BPI
(two collections), UC and W (two collections). Each of the capsules
are imprinted "University of Toronto, Cryptogamic herbarium
No.11571."
 81

Another paratype of T. apiculatus was found in herbarium NY, namely

G. B. Cummins No. 10 from Michigan, U.S.A. under the name
Ascophanus cinereus. There is no doubt that this collection was
available to Kimbrough, although it was not annotated by him. The
specimen belongs to T. crustaceus.

In contrast to the original description of T. agranulosus

(Kimbrough 1969), the examined holotype and paratype of this were
observed with more swollen tips of paraphyses, which were covered
on the outside with amorphous, brown encrustations.
 82

Figs. 25 - 38. Thecotheus crustaceus. 25: Holotype (S). Dry, old

apothecia with incurved margin. 26: Caddy et al. (MlCH - paratype
of T. agranulosus). Young, dry, sUbcylindrical apothecia. 27:
Cummins 10 (NY - one of the paratypes of I.apiculatus). Dry,
young, subcylindrical apothecium. 28: Holotype (S). lnequilateral
ellipsoid ascospores (interference phase contrast). 29: Cain
3.VIll.1937 (BPl - one of the paratypes of T. apiculatus).
Ascospores. 30 - 31: Holotype (S). SEM photos of ascospores,
showing smooth surface except a few very small punctations. It is
not possible to see the granulations in ordinary light microscope.
32 - 34: Ellis & Ellis (lMI - 253134a). 32: Asci staining in Congo
red, showing ascospores with swollen gelatinous perispore layer.
33: Asci with operculum. 34: Ascospores mounted in Indian
ink/water. Note the swollen gelatinous perispore layer on some of
the spores. 35: Donadini (L - 6574). Part of ectal excipulum
composing a textura angularis, the cells orientated with their
longitudinal axes perpendicular' to the outer surface. The exterior
part (bottom) with globose, terminal rooting hyphae. 36 - 37:
Holotype (S). 36: Part of medullary excipulum, with globose to
sUbglobose cells, interspersed with filamentous hyphae. 37:
Excipulum from the margin, showing bulbous, slightly elongated
terminal cells. Inner part (to the right) with a zone of long-
celled textura porrecta. 38: Porten (UPS). SEM photo of tips of the
paraphyses. Scales: 25 & 27: 1 mm. 26:' 0.2. mm. 29 & 34: 25 fJm. 30-
31 & 38: 5 fJm. 32: 45 fJm. 33: 40 fJ1Tl. 35: 9 fJm.· 36 - 37: 16 fJm.
Photos: 25 - 29, 32 - 37: O. Aas. 30 - 31: J. Berge. 38: M. K.
Stavdal.
.3
Q.

25 J.W1
as
86
8"1
 88

5. THECOTHEUS HlMALAYENSIS Kausha1 (Figs. 39 - 41)

BO~. Notiser 133: 319 (1980). -- Type: Indi~, Himachal Pradesh,

Dalhousie, Panjpula, on dung of goat, 28.VIII.1974 S. C. Kaushal
2625 (PAN, holotype, not seen).

ETYMOLOGY: Latinized from the name Himalayas.

Apothecia up to 2.5 mm in diameter in fresh condition, up to 1 mm

in diameter and 0.4 mm high when dry, scattered to gregarious,
often with 2-3 fruit-bodies placed close together, sessile or with
a short, subimmersed tapering base. The apothecia are cupulate to
turbinate, occasionally doliiform, rarely obconical, becoming
discoid at maturity, regular, fleshy, external surface white,
smooth at first, later slightly white-furfuraceous, concolorous
with the disc. Disc at first white, after drying grey-orange (6B4)
to grey-red (7B3), in most cases with a slightly reddish tint,
somewhat roughened by the protruding ascal tips. Margin entire.
Hymenium up to 170 ~m thick.

Medullary excipulum about 90 ~m thick at the central growing part,

gradually reduced towards margin, composed of a more or less dense
textura intricata with up to 6 ~m broad hyphae, interspersed with
small subglobose to somewhat elongated cells. Ectal excipulum about
70 ~m thick, composed of a textura angularis, with polyhedral,
slightly elongated cells up to 24 x 17 ~m, their longitudinal axes
lying somewhat perpendicular to the outer surface; usually more or
less hyphoid cells with clavate tips at the margin and towards the
exterior part, the latter cells slightly thick-walled.
 89

Asci 8-spored, cylindrical, with apex slightly acute and base long
and narrow, (120-) 140 - 175 (-185) x 11 - 14 ~m, when fresh deep
blue in their entire length in Melzer's reagent; young asci
slightly less blue in dry material. Ascospores sUb-ellipsoid to
narrowly ellipsoid or subfusiform, slightly inequilateral,
obliquely uniseriate, sometimes biseriate with a few spores crowded
near apex, (12 -) 14 - 16 x (5.5 -) 6 - 7 ~m, x = 15.30 x 6.44 (n =
45), Q = 0.77 "(length), 0.25 (width), non-apiculate, ornamented
with very fine verrucae; each ascospore surrounded with a gelatin-
ous layer which stains in Cotton blue and in Congo red, and after
spore liberation is only occasionally observed. Paraphyses up to
1.5 ~m wide" below, very slightly enlarged above up to 2 - 2.5 ~m,
or not at all enlarged above, this part with light yellow, slightly
granulated content, thin~walled, filiform, septate, freely branched
at various levels, straight or slightly bent. Interspersed between
asci and paraphyses are additional interascal elements which are
erect, paraphyses-like, up to 7 ~m broad, slightly narrower below,
grey-yellow (4B3) to light grey-orange (5B3), simple, stout,
septate at irregular intervals, straight or slightly bent at their
apices.

Specimens examined:

POLAND. Bialystok: Bialowieza Park Narodowy, deer dung, (mc),

5.IX.1966 L. Holm (UPS). New to Europe.
INDIA. Himachal Pradesh: Dalhousie: Panchpula, on goat dung,
28.VIII.1974 S. Chander 2625 (FLAS F 50920). -- Mahasu: Narkanda:
simla, on dung of goat [or a cervid species], 18.VIII.1971 s.
Chander 2412 (FLAS F 50929); in Thind, Kaushal & Kaushal (1978:
67). Also Coprotus lacteus is found. The substrate is not cow dung
as said by the authors.
 90

UNVERIFIED RECORDS: INDIA. Himachal pradesh, Kaushal 1980: 319,

three collections, holotype and paratypes (2 goat, one unnamed
substrate) .

HABITAT AND SUBSTRATE PREFERENCE: on dung of goat 2 (4), deer 1,

and unspecified dung (1). Very probably all collections from India
are from dung of goat or a cervid species.
DISTRIBUTION: Only known from Poland and India. Tentatively, the
species has an European-Asian disjunctive distribution pattern.
ILLUSTRATIONS: Kaushal 1980, fig. 1 B-D.

Thecotheus himalayensis is well characterized by its small,

verruculose ascospores not furnished with apiculi. They are
slightly smaller than those of T. viridescens. However, the latter
species has a different spore ornamentation and usually only 4
spores in the asci. Another distinctive feature of T. himalayensis
is the additional interascal elements interspersed between the asci
and the paraphyses (Kaushal 1980). Such interascal elements are
also found in other species of Thecotheus, otherwise they are
probably not known in the Pezizales except in Iodophanus
kimbroughii (Thind & Kaushal 1978).

It has not been possible to examine material of the holotype and

the paratypes of T. himalayensis ( Kaushal 1980) from the herbarium
at PAN. However, two collections of Thecotheus sp. in the herbarium
at FLAS belong to T. himalayensis. One of them, Chander 2625 (FLAS
F 50920) was collected at the same time, at the same place and on
the same kind of substrate as that of holotype of ~. himalayensis.
Moreover, it also has the same collecting number as the holotype.
The other collection, Chander 2412 (FLAS F 5092?), accords to one
of the paratypes of T. himalayensis. Both capsules are imprinted
"Botany Department Panjab University Chandigarh (India) ,,' in
addition to "U. S. D. A., P. L 480 Project", Le. the same
91
 92

~..

/.S ""'.,~
...... ~; D S q8 ,.25.
.~ ~,:'v,., ~
..••:._.4 SIII

programme mentioned in Kaushal (1980). Thus, very probably both

collections of Chander were collected simultaneously with those of
Kaushal's.

Figs. 39 - 41. Thecotheus himalayensis. 39: Chander 2625 (FLAS F

50920). SEM photo of ascospore with subfusiform shape. 40 - 41:
Chander 2412 (FLAS F 50929) 40: SEM photo of ascospore showing
ornamentation of very small ± sUbglobose warts. 41: SEM photo
showing tips of interascal elements. To the left a bunch of asci
with spores inside. Scales: 39 - 40: 5 ~m. 41: 50 ~m. All photos:
 93

6. THECOTgEUB HOLMBKJOLDII (E. C. Hansen) Cbenant. (Figs. 42 - 46)

Bull. Soc. Mycol. Fr. ·34: 39 (1918). -- Basionym: Ascophanus

bolmskjoldii E. C. Hansen, Vidensk. Meddel. dansk naturhist. Foren.
1876: 290 (1876). -- Ascobolus bolmskjoldii (E. C. Hansen) Wint.,
Hedwigia 17: 91 (1878). -- Ascopbanella bolmskjoldii (E. C. Hansen)
Faurel & Schotter, Cah. La Maboke 3 (2): 130 (1965 b). -- Type:
Denmark, near Nestved [= N~stved], on old cow dung, 20.VI.1875 E.
C. Hansen (C, lectotype).
Asoobolus incanus Phill., Grevillea 5: 117 (1877). -- Ascophanus
incanus (Phill.) Sacc., Sylloge fungorum omnium hucusque cognitorum
8: 529 (1889). -- Type: U. S. A., California, Colfax, Placer Co.,
Blue 'Canon, Sierra Nevada Mountains, on cow dung, [winter/spring]
1876 H. W. Harkness 419 (BPI, lectotype selected here; CUP 03777
and 04619; NY; isolectotypes).
Tbecotheus cinereus Cr. & Cr. var. major Chenant., Bull. Soc.
Mycol. Fr. 34: 39 (1918); not validly pUblished. Type as for
Tbecotbeus holmskjoldii (C, lectotype).
Ascophanus bolmskioldii E. C. Hansen var. leporinus Velen.,
Monographia Discomycetum Bohemiae. Pars I: 357 (1934). -- Type:
Czechoslovakia, Bohemia, Mnichovice, Set. Anna at Strancice, on
hare dung in wet bog, 21.VII.1925 J. Velenovsky (PRM 150217,
holotype).
Ascopbanus bolmskjoldii E. C. Hansen var. caprinus Velen.,
Monographia Discomycetum Bohemiae. Pars I: 357 (1934). -- Type:
Czechoslovakia, Bohemia, near Mnichovice, on dung of goat, VI.1938
J. Velenovsky (PRM 150349, neotype selected here).
[= Tbecotheus rehmii Zukal in litt. in Rehm, Rabenhorst's
Kryptogamen-Flora Deutsch. Oesterr. Schweiz. I (3): 1092 (1896);
nomen nudum].

Misapplied name: Ascophanus cinereus (Cr. & Cr.) Boud. 1869: 249;
non Ascobolus cinereus Cr. & Cr. 1858: 194 = Tbecotheus cinereus.
 94

ORTHOGRAPHIC VARIANTS: Ascophanus holmskjioldii, Mouton 1886:

141. -- Ascophanus holmskioldii, Schroeter 1893: 53, Migula in
Thome 1913: 1039. -- Ascophanus holmskjoeldii, Petrak 1923, Weese
1927.

ETYMOLOGY: Bolmskjoldii, after Theodor Holmskj0ld (1732 - 1794),

the author of "Beata ruris otia fungis danicis impensa".

Apothecia scattered or gregarious to densely gregarious to crowded

and congested together, sessile, slightly immersed, 0.3 -2 (- 2.5)
mm diameter, 0.3 - 0.8 mm high; soft, fleshy, at first cylindrical
to dolabriform, becoming turbinate, later expanding and more or
less discoid, then with much depressed and concave disc. Regular
when young and growing apart, later often irregular due to mutual
compression and uneven substrate.

Apothecia at first white to greyish, concolorous with the hymenium,

when dry retaining the same colour or greyish with a reddish tint,
i. a. brown-grey (7C2 -9E2), red-brown (8D4) or red-grey (12B2).
Old, over-ripened apothecia sometimes dark grey or black. External
surface at first smooth,. later slightly white roughened, when dry
with vertical ridges. Lower part with white anchoring hyphae. Disc
plane or convex, somewhat concave when dry, slightly roughened by
the protruding ascal tips. At first white to dull grey, then with
varying grey-orange to brown-orange colour; retaining the same
colour or slightly fading and white-furfuraceous on drying. Margin
entire to wavy, slightly bent inwards at maturity, occasionally
somewha~ overhanging in mature, dry apothecia. Bymenium up to 350
·~m thick. Bypothecium distinct, up to 38 ~m.thick, composed of a
more or. less dense textura intricata.

Medullary excipulum mostly up to 130 ~m thick, composed of a

textura globulosa, with sUbhyaline, ± globose cells 2 - 15 ~m

diameter. The innermost part, and towards the hypothecium with

interwoven filamentous hyphae 2 - 3 ~m wide. Towards excipulum,
 95

especially along the sides, a zone of filamentous hyphae 0.8 - 1.5

~m wide, composing a textura porrecta. Ectal excipu1um up to 150 (-
170) ~m thick near the central base, up to 50 ~m thick along the
sides, composed of a textura angularis, with light brown-orange
elongated and polyhedral cells, mostly 13 - 17 (- 22) x 7 - 9 (-13)
~m, their longitudinal axes somewhat perpendicular to the outer
surface. Towards the exterior part with long-celled, rooting hyphae
1.5 - 3 ~m wide, originating several cell layers deep. Terminal
cells along the sides with thick-walled, red-brown or light brown,
encrusted inflated cells, 5 - 12 ~m wide and up to 11 ~m in length,
interspersed with intercellular amorphous brownish pigments.
Towards margin more thin-walled, light yellow-brown, slender and
elongated terminal cells, 7 - 8 ~m wide and 15 - 20 ~m in length,
gradually changing to paraphyses-like cells. Anchoring hyphae thin-
walled, septate, branched, sUbhyaline, up to 3.2 ~m wide.

Asci 8-spored, clavate to cylindrical-clavate, narrowing towards

apex, subhyaline, apex dome-shaped and obtuse, attenuated into a
short, stern-like base, (180-) 225 - 350 (-460) x (25-) 30 - 46 ~m,

p. sp. 130 - 200 ~m. Young asci with brownish colour in iodine
solution. Ascospores ellipsoid, slightly narrowing towards the ends
when young, later with more or less rounded to broadly rounded
ends, biseriate to irregular disposed or uniseriate, parallel or
obliquely placed, separate to slightly overlapping, (25-) 29 - 38
(-42) x (12-) 14 x 18 (-20) ~m (apiculi not included), smooth and
granular within when young, later. covered ~ith small callose-pectic
warts. Especially near the poles with more prominent, blunt warts
up to 1.1 ~m long. At both ends with a hyaline, obtuse, hemispheric
apiculus, up to 2 - 5 ~m long and 2 -6.5 ~m broad at the base. A
bunch of 3 - 8 thin, hyaline, acute, thread-like structures
emanates from each apiculus. The threads, up to 40 ~m in l~ngth,

are observed only on mature, living a~cospores, and they disappear

completely after spore liberation. ,Immature spores subhyaline,
thick-walled, later with a up to 0.5 ~m thin outer wall which stain
in Cotton blue, and a thicker endospore wall, 2 - 2.5 ~m broad,
which does not stain in Cotton blue. Both warts and apiculi stains
in Cotton blue. Ripe spores often with light yellow-brown granular
 96

contents; each ascospore surrounded with a thick gelatinous layer

which also encloses the apiculi. After spore liberatio~ the
gelatinous envelope often expands, but this is not always observed
on long-dried ascospores. Paraphyses filitorm, septate, branch~d or
not, hyaline. and thin-walled below, 1.5. - 2.5 ~m wide; apices
inflated, 3 - 8 ~m wide, this part encrusted with yellow-brown to
brownish pigments. The paraphyses are somewhat agglutinated at
their tips, and they project up to 25 - 35 ~m beyond the hymenial
surface.

Specimens examined:

AUSTRIA. ? Nie~er osterreich: Pressbaum at Vienna, deer dung,

(me), I.1889 A. Heimerl as I. rehmii Zukal (W - slide, ex herb.
Heimerl); in Heimerl (1889: 20), Rehm (1896: 1092).-- Steiermark:
Grazer Bergland: Hochlantsch Gipfel, Nordabhang 1600 m, roe-deer
dung, 15.IX.1982 H. Schweiger (GZU); in Schweiger (1985: 64) as I.
cinereus (Cr. & Cr.) Chenant. -- weiz: Weizklamm 700 m.a.s.l., roe-
deer dung, 10.X.1981 H. Schweiger (GZU); in Schweiger (1985: 64) as
I. cinereus.
BELGIUM. Brabant: Tervueren, fallow deer dung, s. dato, E. Bommer
& M. Rousseau (BPI, BR) ; in Bommer & Rousseau (1884: 145) as
Ascob. (Ascoph.) holmskjoldii Hansen. -- Pare de Tervueren, fallow
deer dung, V.1884 E. Marchal (BR); ? in Marchal (1884: 93) as
Ascoph. holmskjoldii Hansen. -- Tervueren, fallow deer dung, VIII.
1884 E. Marchal (PAD - ex herb. P.A. Saccardo No. 1661). --
Liege: Gomze, sheep dung, VIII. [so ann.] V. Mouton (BR); in Mouton
(1886: 141) as Ascoph. holmskjioldii [sic!] Hansen.
CYPRUS. w. Region: 3 km E of Paphos, goat dung in hills with
pastures and macchia, (me), 25.IV.1985 N. Lundqvist 15386 b (S).
New to Cyprus.
CZECHOSLOVAKIA. Bohemia: Kasar, ? goat dung, 30.V.1948 V. Vacek
(PRM 203579) . Mnichovice: Mencice, cow dung, VI.1929
J. Velenovsky as Ascoph. obscurus Velen. in herb., unpubl. name
(PRM 150370). Mnichovice: Myslin, cow dung, 3.X.1924 J.
 97

Velenovsky (PRM 651897 - substrate only). Mniehoviee: Set. Anna

at StrAn~iee, "as Aseoph. leporinus Velen. in herb. (PRM 150217 -
holotype of Aseoph. holmskjoldii var. leporinus); in Svr~ek (1979:
12.3 & 1981 a: 85). -- Mnichovice, as Ascoph. caprinus Velen. in
herb. (PRM 150349 - neotype of Ascoph. holmskjoldii var. caprinus;
in Svr~ek (1~79: 122). --. Praha-DivokA: S!fka, cow du~g, (me),

5.X.1947 M.V. Svr~ek 349-47 (PRM 651896). -- Suchdol at Praha: in

valle Tiche Udol! ap. Trojanuv mlyn, cow dung in a shaded pasture,
5.VIII.1972 Z. Pouzar (PRM 803946).-- Moravia: Mahr.-Weisskirchen:
Bartelsdorf, roe-deer dung [probably of sheep], 10.V.1923 F.
Petrak: Flora Boh. Mor. Exs. No. 1555 as Ascoph. holmskjoeldii
[sic.] Hans. (BPI, BR, C, E, K, M 11607, PRM 7695, S, U~S, W
23149 - ex herb. F. Petrak 3596); in Petrak (1924). -- Mahr.-
Weisskirchen: Bartelsdorf, roe-deer dung [probably of sheep],
V.1923 F. Petrak in Weese: Eumycetes selecti Exsiccati No. 91 as
Ascoph. holmskjoeldii [sic!] Hansen (BPI, M -two collections, UPS,
W 3380 and duplicate in ex. herb. F. Petrak 34656). -- Mahr.-
Weisskirchen, roe-deer dung, 1923 F. Petrak (B - two collections ex
herb. A. LUdwig, K). -- Near Brlinn [= Brno], cow dung, spring 1877
G. Niessl in Rehm: Ascomyceten No. 470 as Ascob. cinereus Cr. (E-
ex herb. C.E. Broome, H - ex herb. P.A. Karsten 2849 as Ascoph.
cinereus, K, M, NY - ex J.B. Ellis collection, PAD ex herb. P~A.
Saccardo 1659, S, UPS, W 1713, WRSL); in Rehm (1878, 1881, 1896),
Massee (1895: 175; see however comments under T. crustaceus),
Kimbrough (1969: 110) horse dung[sic.!] as T. cinereus (Cr. & Cr.)
Chenant. -- Brlinn (= Brno), cow dung, s. dato, G. Niessl as Ascob.
cinereus Cr. (CUP D 9587 and F3049 - ex herb. C.E. Fairman, M 41420
- ex herb. G. Niessl); in Kimbrough (1969: 110). -- Jesen!k: Horn!
udole, cow dung, (me), 18.V.1963 Z. Nenbauer (PRM 756512). --
Valtice, cow dung, (mc),13.X.1971 M. Svr~ek 1044/71 (PRM 823501).
DENMARK. s. loco, substr. et dato, E.C. Hansen (C slide) .--
Bornholm: Salomans Kapel, cow dung, 30.VII.1918 o. Rostrup
(CP); in Rostrup (1935: 22). -- Sjalland: Nestved (C - lectotype of
Ascoph. holmskjoldii, also a slide No. 215 in Fungorum Fimicolorum
Exempl. Exsicc.); in winter (1878: 91). -- Saltholm, sheep dung,
23.V.1969 C. H. Ovesen (C).
ESTONIA: Valga distr.: Llillernae, cow dung, 9.IX.1986 V. P.
 98

Prokhorov (MW); in Prokhorov (1989 b: 287) as T. cinereus (Cr. &

Cr.) Chenant.
FRANCE. s. loco (Gallia), dung [= sheep), s. dato, misit J.L.E.
Boudier (S - ex herb. A.G. Bresadola). -- Alpes-de-Haute-Provence:
Basses-Alpes: SSW of Point Sublime along the Martel Path on the W
side of George du Verdon (= 12 km SW of Castellane), sheep dung,
(me), 30.IV.1979 N. Lundqvist 11948-b (UPS). -- corsica: Canton de
Bonifacio: Golfe de Santa Manza, 1 km W of Gurgazo, old cow-dung on
the seashore, (me), 15.V.1965 N. Lundqvist 4436-d (UPS). -- Foret
de Chiavari: 1.5 km NW of Coti-Chiavari, about 550 m.a.s.l., horse
dung, 14.X.1972 R. P. Korf & V. Demoulin (CUP - ex herb. R. P.
Korf: RPK 72-142).
GEORGIA: Abchazskaya ASSR: 3 km S of Lake Ritsa (= 26 km NE of
Gagra), cow dung in montane deciduous forest, (me), 30.IX.1986 N.
Lundqvist 16389-j (S). New to Georgia.
GERMANY. Baden-Wurttemberq: Rastatt, dung [probably of cow),
I.VIII.1870 J. Schroter as Ascobolus (WRSL). --Dresden: Pirna, cow
dung, 18.X.1892 G. Wagner as Ascoph. incanus Phill.(B, S - ex herb.
H. Rehm as Ascoph. cinereus). -- Frankfurt: Serlow: nature
conservation territory "OderhSnge Mallnow", sheep dung, 7.VIII.1984
D. Benkert (BHU). --- Gera: Rudolstadt: N valley of Heilsberg, deer
dung, 20.IX.1984 D. Benkert (BHU). -- Ha11e: Galzenberg, ? cow
dung, VII.1873 G. winter as Ascoph. cinereus Boud. ( BPI - 2 colI.,
substr. only, H - ex herb. P.A. Karsten 284B, K - as Ascob.
cinereus Cr., substr. only, M 41419 - substr. only, ex herb. G.
Niessl, S - ex herb. Rehm, substr. only); in Winter (1873: 146).
Hessen: Breitscheider Wald, roe-deer dung, V.1944 A. LUdwig (B).
Niedersachsen: Luchow-Dannenberg: near Gummern, ? fallow deer dung,
(me), 25.X.1988 E. Jahn (BG - photo only). -- ditto, red deer dung,
18.IX.1988 E. Jahn (BG - photo only). -- Potsdam: Rathenow, sheep
dung, 19.VIII.1904 W. Kirschstein (B).
HUNGARY. Fejer: Vertes Mts: .Sarkanylyukvolgy Valley: near
Verteskozma, cow dung, J. Banhegyi 6.I~.1939 (BP 67213); in
Banhegyi (1940: 114). ~- Pest: Bernecebarati: in the valley
Bernecevolgy, cattle [= cow) dung, 11.IX.1966 S. T6th 9001 as T.
cinereus (Cr. & Cr.) Chen. {BP J3382). --near Tatarszentgyorgy,
cattle [= cow) dung, 15.V.1969 S. T6th 7881 as T. cinereus (Cr. &
 99

Cr.) Chen. (BP J3383). - near Veresegyhaz, rabbit dung,

18.VIII.1969 S. T6th 7823/e as T. einereus (Cr. & Cr.) Chen. (BP
J4338). -- Vas: Csakvar, sheep dung, 1.VI.1966 S. T6th 5627 (BP
49.527); in T6th (1967: 118). -- "Torokret" near Btikkzsere, cattle
[= cow) dung, 3.XI.1964 S. T6th 4735 (BP 40.106); in T6th (1965:
150). -- Rabat6tfalu near Szentgotthard, cattle [= cow] dung,
9.VII.1964 S. T6th 4706 (BP 39.965); in T6th (1965: 150). --
Veszprem: on the "Pet6hegy" Mts. near Keszthely, deer dung,
7.IV.1966 S. T6th 5631 (BP 49.510); in T6th (1967: 118).
ITALY. Trentino: Val di Sole: near Male, deer dung (culture),
3.IV.1969 J. van Brumrnelen 2716 (L).
NORWAY. Akershus: Asker: Hvalstad, cow dung, 27.IV.1961 I.L.
Egeland (BG - drawing only). New to Norway.
POLAND. Silesia: Breslau (= Wroelaw): in the zoological gardens,
dung [? goat), VII.1880 J. Schroeter (WRSL); in Schroeter (1893:
53). -- Oppeln (=Opole): Brinnitz, dung [probably of cow], VII.1880
J. Schroeter (WRSL); in Sehroeter (1893: 53).
SPAIN. Islas Baleares: Mallorca, W part, 5 km NE of Puerto
Soller, goat dung in mountain slope with macehia, (me), 12.VI.1986
N. Lundqvist 15986-e (S). -- Madrid: El Pardo, sheep dung, (me),
5.VI.1984 J.M. Barrasa 3647-72 (private herbarium of Barrasa,
Madrid - slide); in Barrasa (1985) as T. einereus (Cr. & Cr.)
Chenant. -- Valencia: Alieante: Aleoy: Gorgo, goat dung,
23.VII.1989 H. G." Unger (BG - ~aterial also wit~ fr~itbodies of
Sehizotheeium vestieola (Berk. & Br.) Lundq.).
SWEDEN. Gotland: Eksta: Lilla Karlso Isl., sheep dung, (me),
16.V.1970 R. Jaeobson (UPS). -- Hall: E of Nors, eow dung in pine
forest, (me), 28.VII.1961 N. Lundqvist 3079-b (UPS). -- Hall: Nors,
sheep dung, (me), 5.VI.1959 N. Lundqvist 2089-e (UPS). -- Rone:
Uggarda, sheep dung in a pasture, (me), 24.VII.1961 N. Lundqvist
3036 (UPS). -- Visby: Skansudd, old eow dung.on the sea-shore,
(me), 30.VII.1961 N. Lundqvist 3091-a (UPS). -- Oland: Grasgard:
Eketorp, eow dung on alvar ground," (me), l7.V.1985 N. LU~dqvist

15452-e (S). -- Kalla: at Kalla Harbour, old eow dung in a meadow,

(me), 15.VII.1960 N. Lundqvist 2586-b (UPS). -- Koping: between
'Borgholm and Kopingvik, sheep dungin a meadow, (me), 14.VII.
1960 N. Lundqvist 2570-e (UPS). -- Viekleby: E of Karlevi on the
 100

Alvar, sheep dung, (me), 28.Vlll.l97l N. Lundqvist 7285-a (UPS).

Sklne: Simris: Simrislund, old cow dung on the beach, (me),
10.V.1961 N. Lundqvist 2870-b (UPS). -- Smlland: Gamleby: NW of
Segersgarde, pld cow dung in coniferous Iorest,' (me), 12.VII.l9~0

N. Lundqvist 2556-b (UPS).-- MAnsarp: MAnsarp railway station, old

cow dung in a pasture, (me), 5.VI.1960 N. Lundqvist 2372~a(UPS). -
- Narke: Kumla: Kumla, cow dung, 25.VII.l885 L. Romell 16959 as
Ascobolus (5 - ex herb. L. Romell). -- Sodermanland: Gryt: S of
Anhammar, cow dung in oakhill with hazel, (me), 14.IX.1985 N.
Lundqvist 15789-e (S). -- Uppland: Harbo: Kalvnasudden, on rocks at
Lake Tamnaren, she~p dung, (me), 4.VII.1974 N. Lundqvist 9364-a
(UPS). -- inqermanland: Ramsele: Rabbstuguforsen (= E of Nassjo),
old cow dung in pine forest, (me), 18.VI.1962 N. Lundqvist 3443-a
(UPS). -- ytterlannas: Vastertorp, old cow dung in coniferous
forest, (me), 20.VIII.1961 N. Lundqvist 3165-b (UPS). --Jamtland:
Braeke: Mordviken, eow dung, (me), 15.VIIl.1960 N. Lundqvist 2778-a
(UPS). -- Hammerdal, FyrAs, old eow dung, (me), 14.VIII.1960 N.
Lundqvist 2776-a (UPS). -- Norrbotten: Harparanda: in a meadow at
Torne River, old cow dung, (me), 4.VIII.1960 N. Lundqvist 2673-c
(UPS). New to Sweden.
SWITZERLAND. Graubunden ?: Scarl ?, deer dung, 1.IX.
1970 R.W.G. Dennis & E. MUller as T. cinereus Cr. (R - slide). New
to Switzerland.
UKRAINE: Poltava distr.: Komsomolsk-on-Dniepr, cow dung, s. date,
V. P. Prokhorov (MW).
YUGOSLAVIA. Medjugorje, sheep dung, 12.X.1989 M. A. Jahn (BG).
-- Croatia: S of Senj at Zrnovnica, cow dung in karst landscape
with shrubby pastures, (me), 4.V.1972 N. Lundqvist 7566-f (UPS). --
Velebit Mts.: Mt. Budim: E of Karlobag, cow dung in karst landscape
with grass heaths with scattered groves, (me), 5.V.1972 N.
Lundqvist 7607-b (UPS). New to Yugoslavia.
INDIA. Himachal Pradesh: Jandhri Ghat: Dalhousie, cow dung in
mixed forest, 25.VIl.1966 K.S. Waraitch 2094 (BPI, C); in Waraitch
(1977: 4) as T. holmskjoldii (Hans.) Eckbl. -- Jammu , ~ashmir:

Pahlgam: Bisaran, cow dung in open place, 31.VIlI.1967 K. s.

Waraitch 2255 (BPl); in Waraitch (1977: 4). -- Uttar Pradesh: Dhobi
Ghat: Mussoorie, cow dung, l5.VIII.l952 L.R. Batra (CUP INll - card
 101

only, no material received from Batra at Panjab Univ. Herbarium),

in Batra (1954: 46); Batra & Batra (1963: 137).
ISRAEL. Golan: Masaada Forest, goat dung, 7.X.1971 N. Binyamini
71a53 (TELA); 'in Binyamini (1973: 154) as T. cinereus (Cr. & Cr.)
Chen.
NEW ZEALAND. Canterbury (South Island): 20 km WNW of Springfield
at NE shore of Lake Lyndon, cow dung, (mc), 25.IX.1980 L. Tibell
8989-c (UPS- slide). -- at top end of Stoney River, deer dung,
(mc), 30.1.1976 J. Grossman (WELTU F/267 - slide); in Bell (1983:
33) as T. cinereus (Cr. & Cr.) Chen. -- at Turton's Stream ±
3900'top end of Catchment above Comyns Stream (S. Island), deer
dung, (mc after 1 months incubation), 1.VII.1975 J. Grossman, det.
A. Bell (Herb. No. 251) as Iodophanus durbanensis (FLAS F 52000 -
det. J. Kimbrough as T. cinereus).
CANADA. British Columbia: Salt Spring Island, deer dung, X.1969
J. W. Paden 728 & J. W. Kimbrough as T. cinereus (Cr. & Cr.) Chen.
(FLAS F 48885). Very probably this is the same material referred to
in Conway (1975), a culture of T. cinereus deposited in the
American Type Culture Collection (ATCC 26793).
GREENLAND. Ella Island in Kong Oscars Fjord, 72°50'N 25°W (= 100
km NW of Mesters Vig): near Oxe sea, old musk ox dung in a marsh,
6.VIII.1982 S. Sivertsen & H. Dissing Gr. 82.113 and Gr. 82.115
(C). -- near Lange sea, old musk dung, 30.VII.1983 H. G0tzsche & H.
Dissing Gr. 83.59 (C). New to Greenland.
U.S.A. California: Colfax: Lectotype and isolectotypes of
Ascobolus incanus Phill. (see above); in Seaver (1928: 112),
Kimbrough (1969: 110, as regards CUP 03777). -- Los Angeles Co.:
Santa Catalina Isl.: S of Little Harbour, cow dung, (mc), 5.IV.1966
R. Santesson 17297-h(UPS). -- Idaho: Tie Creek: Tetonia, cow dung,
22.VI.1969 K. H. McKnight and L. B. McKnight 10314 (BPI 763). --
Oregon: Hood River, cow dung, 30.VIII.1932 J. R. Kienholz K24 (NY)i
in Seaver (1942: 310) as Ascoph. holmskjoldii, Kimbrough (1969:
110) as T. cinereus (Cr. & Cr.) Chenant. -- Utah: 1 1/2 mi N of
Robinson Lake, cow dung, 19.VI.1964 J. W. Paden (FLAS F 47962); in
Kimbrough (1969: 110) as T. cinereus (Cr. & Cr.) Boud.
BERMUDA. Hungry Bay, goat dung, 14.1.1926 F.J. Seaver & H.H.
Whetzel (NY - Bermuda fungi No. 28, 2 colI.); in
 102

Kimbrough (1969:110) as T. cinereus (Cr. & Cr.) Chenant.

CHILE. Linares: close to Panim~vida, cow dung, (me), 14.V.1969 w.
Lazo (C). New to Chile.

UNVERIFIED RECORDS: ARMENIA: Prokhorov & Taslakhchyan (1987:

147) as T. cinereus (Cr. & Cr.) Chenant., on cow dung. -- AUSTRIA:
Heimerl (1889: 20) on cow dung. BULGARIA: Fakirova (1967: 226)
on goat dung, (1968: 142) roe deer dung, (1970: 186) on sheep dung,
(1974: 10) on dung; all as Ascoph. holmskjoldii Hans. --
CZECHOSLOVAKIA: Velenovsky (1934: 357) as Ascoph. holmskjoldii var.
caprinus, on goat dung. original colI., not known to be in
existence. -- Velenovsky (1934: 357) on cow and horse dung.
Smarda (1944: 2) on dung. Also referred in Svr~ek (1981 a: 85).
DENMARK: Hansen (1876: 290) on old cow dung. -- Lind (1913: 103) on
cow dung. -- Rostrup (1916: 19) on deer dung. -- Rostrup (1935: 22)
on cow dung. -- ENGLAND: Ellis & Ellis (1988: 115) as T. cinereus,
on dung. -- ESTONIA: Prokhorov (1989 b: 287) as T. cinereus (Cr. &
Cr.) Chenant., 2 colI. respectively on dung and on cow dung. --
FRANCE: Boudier (1869: 249) as Ascoph. cinereus (Cr.) Boud. on cow
dung. -- GERMANY: Beyer et al. (1986: 62) as T. cinereus (Crouan)
Chenant. on ? sheep dung. Also refe~red to in Hohmeyer et al.
(1989: 30). -- HUNGARY: B~nhegyi (1942 b: 269) on cow dung. -- T6th
(1967: 118) on cattle dung. The specimen is also mentioned in
B~nhegyi et al. (1985: 654) as T. cinereus (Cr.) Chenant. -- ITALY:
Bizzozero (1885: 341) on cow dung. This collection could not be
found in the herbarium at Padova, Italy (The Curator of the
herbarium (PAD), in litt. 15.XII.1984). -- MADEIRA: Korf & Zhuang
(1991: 317), two collections on goat dung. -- POLAND: Schroeter
(1893:· 53), herbivore dung. ~- Schmidt (1912: 16) on sheep and deer
dung. -- Kohlman-Adamska (1965: 92) on cow dung. -- Chmiel (1977:
88) on cow dung. -- ROMANIA: B~nhegyi (1942 a: 40) ·on cow dung. --
SPAIN: Bertault (1982: 11) on wild goat (Capra hircus) dung. --
Guarro (1983:.236) as T. cinereus (Cr. & Cr.) Chenant., on sheep
dung. -- Honrubia et al. (1983: 56) on cow dung. -- Calonge et al.
(1986:33) as T. cinereus (Cr. & Cr.) Chenant., on cow dung. --
 103

SWEDEN: Got1an~: Lundqvist 3062 and 3037. 61an~: Lundqvist 2564 and
2569. Upp1an~: Lundqvist 2860. Jamt1an~: Lundqvist 3414.
Norrbotten: Lundqvist 2676-a. All on cow dung. -- UKRAINE:
Girzitska (1929: 66) on cow dung. -- NEW ZEALAND: Bell (in litt.
19.VIII.1985) as T. cinereus, on sheep dung. -- RUSSIA: Tadzhik:
Raitviir & Prokhorov (1988: 218) as T. cinereus, on donkey dung.
ALGERIA: Faurel & Schotter (1965 a: 271) as Ascoph. cinereus (Crn.)
Le Gal, on dog dung. -- MOROCCO: Malenyon & Bertault (1967: 241) on
cow dung. -- U.S.A.: Dodge (1912: 152) on dung (1) from New York. -
- Seaver (1928: 112) on cow dung from New York, Pfister (1982: 7)
records it as T. ? cinereus (Cr. & Cr.) Chenant. -- Lohman (1942:
106) on cow dung from Indiana. -- Kimbrough 1969: 103, fig. 11
(CUP-D 958), fig. 16 (TRTC 40118), both as T. cinereus. -- BERMUDA:
Seaver (1942: 310) on rabbit dung.

DUBIOUS RECORD: LUXEMBOURG: Feltgen (1899: 30) on cow dung. Very

large ascospores: 50-60 x 20-27 ~m.

HABITAT AND SUBSTRATE PREFERENCE: T. holmskjoldii is exclusively

a fimicolous species: 109 (145) finds, from dung of cow 58 (78),
deer 20 (23) [including fallow deer 6~ roe deer 5 (6), and red deer
1], sheep 18 (23), goat 8 (13) [including wild goat (Capra hircus),
(1)], musk ox 2, horse 1 (2), hare 1, rabbit· 1, dog (1), donkey
(1) .
DISTRIBUTION: Armenia, Austria, Belgium, BUlgaria (Fakirova 1967,
1968, 1970, 1974), Cyprus, Czechoslovakia, Denmark, England (Ellis
& Ellis 1988), Estonia, France, Georgia, Germany, Hungary, Italy,
Madeira (~orf'& Zhuang 1991), Norway, Poland, Romania (Banhegyi
1942 a), spain, sweden', Switzerland, Ukraine, Yugoslavia, India,
Israel, New Zealand, Russia (Raitviir ~ Prokhorov1988), Algeria
(Faurel & Schotter 1965 a), Morocco (Malenyon & Bertault 1967),
Canada, Greenland, U.S.A., Bermuda, ~nd Chile.
ILLUSTRATIONS: Boudier 1869 (as Ascoph. cinereus (Cr. & Cr.)
Boud.), plo 11, fig 37, 1-6. -- Hansen 1876, plo 6, figs. 1 - 8. -
 104

- Phillips 1877 (as Ascob. incanus), pI. 88, fig. 10 a - e .

Rehm 1896, p. 1079, figs 1-2 (originaldrawing by Zukal). --
Chenantais 1918, pI. 3, fig. 1 a-f. -- Seaver 1928, pI. 11, fig. 8.
-- Velenovsky 1934, tab. 3, fig. 10. Drawings both of Ascoph.
holmskjoldii and A. holmskjoldii var. caprinus~ -- Lohman 1942 (as
Ascoph. holmskjoldii), fig. 15. -- Faur~l & Schotter 1965 a (as
Ascoph .. cinereus (Crn.) Le Gal), fig. 2 a-b. --Kohlman-Adamska
1965, fig. 10 b & d. -- Kimbrough 1969, figs.11, 16, (both as T.
cinereus), Binyamini 1973 (as T. cinereus), fig. 7. -- Waraitch
1977, figs. 10-11. -- Kaushal 1980 (as T. cinereus (Cr. & Cr.)
Chen.), fig. 1 a. -- Bell 1983 (as T. cinereus), fig. 34. -- Guarro
1983 (as T. cinereus (Cr. & Cr.) Chenant.), fig. 8 a-d. -- Honrubia
et al 1983, fig 3 a-d. -- Schweiger 1985 (as T. cinereus (Cr. &
Cr.) Chenant.), fig. 31. -- Beyer et al. 1986 (as T. cinereus),
fig. on page 62. -- Calonge et al. 1986 (as T. cinereus), fig. 6 a-
b. -- Ellis & Ellis 1988, pI. 36, fig. 360 (as T. cinereus (Crouan)
Chenant. -- Prokhorov 1989 b, fig. [1] (as T. cinereus). -- Korf &
Zhuang 1991, fig. on page 317.
PHOTOS: Kimbrough 1969 (as T. cinereus) figs. 10 - 19. --
Kimbrough 1972 (as T. cinereus) pI. 19, figs 44-45. -- Conway 1975,
figs. 7 - 11 '(as T. cinereus). -- Beyer et al. 1986 (as T.
cinereus) pI. 44: 163. Also referred to in Hohmeyer et al. (1989:
30) .

Thecotheus holmskjoldii is characterized by the large, apiculate

ascospores, where the epispore is covered with isolated warts of
different sizes, the largest near the poles. In living apothecia,
the apiculi bear a bundle of hyaline, mucilaginous, projecting
thread-like structures, which completely disappear in water mounts
after spore liberation. They are not preserved in dried material.

T. holmskjoldii has by many authors been treated as a synonym of T.

cinereus, certainly a result of Le Gal's concept (1960, 1963).
Previously, Chenantais (1918) had considered T. holmskioldii as a
variety major of T. cinereus. Also Kimbrough (1969) incorrectly
 105

synonyrnized T. holmskjoldii with T. cinereus.

There has been some misunderstanding concerning th~ author citation

of ~. holmskjoldii. Chenantais's (1918) combination is undoubtedly
valid and correct. Eckblad (1968), however, interpreted the
combination T. holmskjoldii (Hans.) Chenant. as not validly
pUblished, and he made the new combination T. holmskjoldii (Hans.)
Eckbl. This opinion of Eckblad has later been followed by, amongst
others, Waraitch (1977), Bertault (1982), Honrubia et al. (1983)
and Donadini (1985).

Notes on type collections

The lectotype of Ascophanus holmskjoldii consists of four apothecia

without substrate, sealed in a small glass·tube. On an attached
sheet of paper is handwritten "Ascophanus novo spec." A slide No.
215 of the same material mounted in glycerine is also preserved in
the book Fungorum Fimicolorum Exempl. Exsicc.

Kimbrough (1969) says that the holotype of A. incanus is a CUP -

collection No. D 3777. However, this rather poor collection is only
a part of the original material of A. incanus. There exsist four
more collections of Harkness' No. 419, and by far the best is the
one deposited at BPl which is annotated Ascobolus incanus Phill. n.
sp. with a short handwritten description signed by Phillips. The
capsule has printed on it "California Academy of Sciences No. 21",
as the bulk of Harkness' herbarium was originally deposited at
CAS (see Hawksworth 1974 and Stafleu & Cowan 1979). Later, all
specimens of fungi in this herbarium were transferred to BPl at
Beltsville, (Stafleu (1981), B. Bartholomew, collection manager at
CAS, in litt.). I chose the BPl material as lectotype.
 106

The other three collections of No. 419 (CUP-D 3777 ex herb. E. J.

Durand 13-183; CUP-D 4619 ex herb. E. J. Durand 13-184; NY ex herb.
J. B. Ellis) are all poor and fragmentary. They are considered to
be isolectotypes.

Svr~ek (1979) says that lectotype of Ascophanus holmskjoldii E. C.

Hansen var caprinus Vel. is a PRM material No. 150349. This is a
Velenovsky collection from June 1938, determined by him as
Ascophanus caprinus Velen., but it is obviously not original
material and can not be a lectotype as stated by Svr~ek. No other
collections has been found in PRM. Contrary to Svr~ek I found one
apothecium in this material which consists of a few broken
fragments of goat dung. The material matches the description well
and represents T. holmskjoldii. Most probably, the original
material or duplicates of the variety have been lost or destroyed,
and I select PRM 150349 as a neotype.

Figs. 42 - 46. Thecotheus holmskjoldii. 42: Greenland. Sivertsen &

Dissing 82.115 (C). Young, cylindrical apothecium. 43:
Czechoslovakia. Nenbauer (PRM 756512). Mature, discoid apothecium.
44: France. Lundqvist 4436-d (UPS). Ascospores showing
ornamentations of small warts. 45 - 4~: Lect~type of Ascoph.
holmskjoldii (C). 45: Ascus with biseriateascospores. 46: Asci
with spores and paraphyses. Note the bipolar apiculus at each
ascospore. Scales: 42: 0.6 mm. 43: 1.5 mm. 44: 30 ~m. 45: 50 ~m.

Photos: 42 - 44: O. Aas. 45 - 46: J. Berge.

 108
7. THECOTHEUS INAEQUILATERALIS Aas sp. novo (Figs. 47 - 48)

ETYMOLOGY: Latin inaequalis, uneven, unequal; lateralis, lateral,

at the side; referring to the somewhat inequilateral form of the
ascospores.

Apothecia sessilia, usque ad 0.5 mm diam.; prime globulosum vel

pyriforme, ad extremum discoideum, prime cinereo-album vel
ochraceo-cinereum, in sicco vi~idescens. Ex~ipulum medullarium
texturae globosae. Excipulum externum texturae angularis, hyphis
radicantes cellulis tUberosis, usque ad 7 ~m diam. terminantes.
Asci octospori, cylindrici, 140 - 175 x (10-) 12 - 15 ~m,

parietibus iodo caerulescentibus. Ascosporae obliquae uniseriatae,

ellipsoideae, inaequilateralis, 13 - 16 x 7 - 8.5 ~m, laeves,
apiculis polaribus. Paraphyses typorum duorum.
Fimicola.

Type: Russia, Moscow distr., Novo-Konstantinovo, on dung of moose,

21.VIII.1982, V. P. Prokhorov (MW, holotype).

Apothecia scattered to gregarious, sessile, often with a well

developed sUbiculum, up to 0.5 mm in diameter in dry condition;
soft, fleshy, at first globular to pyriform, then doliiform, later
expanding and more or less discoid. Apothecia at first pale grey
(-B1), then somewhat ochre grey, becoming olive grey (102) when
dry. Lower part with white anchoring hyphae, comprising a well
developed subiculum. Disc concave when dry, slightly roughened by
the protruding ascus tips; becoming white-furfuraceous on drying.
Marqin distinct.
 109

Medullary excipulum of textura globulosa, with 9lobose to

subglobose cells up to 12 ~m diameter, interspersed with hyphoid
cells. Ectal excipulum of textura anqularis, the cells up to 16 x 8
~m, with their longitudinal axes more or less perpendicular to the
outer surface; rooting hyphae 1.5 - 2.5 ~m wide originate several
cell layers deep, terminating in subglobose to clavat~-bulbous

terminal cells up to 7 ~m wide, becoming encrusted with greyish to

greyish brown pigments.

Asci 8-spored, cylindrical, narrowing below, apices ± obtuse to

slightly truncate, 140 - 175 x (10-) 12 - 15 ~m, p. sp. up to 105
~m. Ascospores ~llipsoid, inequilateral, obliquely uniseriate, 13 -
16 x 7 - 8.5 ~m, smooth. Each spore with hyaline, obtuse, mostly
.hemispherical bipolar apiculus, 1 - 2 x 2 - 3 ~m. The asco$pores
are surrounded with a gelatinous layer. Paraphyses of two types,
the one filiform, septate, hyaline, branched or not, 1.5 - 2 ~m

diameter below and at their apices, the other one, septate, hyaline
and 3 - 6 ~m wide.

Specimens examined:

UKRAINE. Kiev distr.: Vyshgorod, moose dung, s. date V. P.

Prokhorov (MW).
RUSSIA. Moscow distr.: Holotype (see above).

The few finds known so far do not permit any sure conclusions about
the ecology and distribution of the species.

T. inaeguilateralis is characterized by its small, smooth,

inequilateral ascospores furnished with bipolar apiculus. The same
form of spores are found in T. biocellatus, but this species has
ornamented and larger ascospores. Moreover, the apothecia of T.
 110

inaeguilateralis are easily recognized by its well developed

subiculum in addition to the colour of the fruit-bodies.

The collection from Ukraine has been published as T. agranulosus

(Prokhorov 1989 a).

Figs. 47 - 48. Thecotheus inaeguilateralis. Holotype (MW).

Ascospores. Scales: 20 ~m. Photos: J. Berge.

8. THECOTHEUS KEITHII (Phill.) Aas n. comb. (Figs. 49 - 67)

Basionym: Peziza keithii Phill. in Stevenson, Mycologia scotica:

308 (1879). -- Humaria keithii (Phill.) Sacc., Sylloge fungorum
omnium hucusque cognitorum. 8: 123 (1889). -- Ascophanus keithii
(Phill.) Boud.: Ramsb., Trans. Br. Mycol. Soc. 4 (2): 369 (1914). -
- Type: Scotland, Grampian, Moray, Forres, on horse dung, XI. 1878
J. Keith (K, neotype selected here).
? Ascophanus appendiculatus A. Schmidt, Die Verbreitung der
coprophilen Pilze Schlesiens: 30 (1912). -- Type: Poland, near
Breslau [= Wroclaw], Oswitz, on horse dung; not known to be in
existence.
Thecotheus apiculatus Kimbrough, Mycologia 61 (1): 110 (1969). --
Type: u. S. A., Oregon, Hood River, on dung of horse, 28.X.1932 J.
R. Kienholz K 64, (NY, lectotype selected here).
111
 112

Misappli~d name: Ascobolus albicans Fuckel sensu Karsten, Fungi

Fenn. exs. No. 762 (1868); non Ascobolus albicans Fuckel, Hedwigia
5 (1): 3 (1866); Fungi rhen. No. 1855 (1866).

ETYMOLOGY: Latin apiculus,"small terminal point; for its

apiculate ascospores. Latin appendiculatus, appendiculate, with
small appendages; for the protuberances of the spores. Keithii,
after the original collector, Rev. James Keith, who for many years
was minister in the parish of Forres, Scotland.

Apothecia scattered or gregarious, sessile to slightly immersed, up

to 3.1 mm diameter when re-hydrated, [cups 5 - 8 mm across
according to original description of Peziza keithii] when dry up to
1.6 mm diameter and 0.5 mm high. At first globose to subglobose,
then cylindrical to pyriform or turbinate to doliiform, later
expanding and more or less discoid, when over-ripe broad discoid,
apparently sessile on a broad base. Dry, young apothecia often
cylindrical in shape.

Apothecia white to light white grey or yellowish when fresh, later

light orange (6A4) to dull greyish orange (6B4) to brownish orange
(7C4), becoming reddish brown when dry, towards base with darker
pigmentation. Exterior slightly white-furfuraceus as dry. Disc flat
to ± convex, after drying turning concave and somewhat undulating,
roughened by the protruding ascus tips. Marqin obtuse to distinct,
broad, slightly incurved, with somewhat uneven and irregular
surface. Hymenium up to 250 ~m thick.

Medullary excipulum up to 120 ~m thick at the central base, with

small, hyaline, thin-walled, globose to subglobose cells 2.5 - 11
~m in diameter, composing a textura globulosa, interspersed with
scattered larger, more elongated cells up to 20 ~m diameter, in
addition to filamentous septate hyphae. This layer is limited
towards ectal excipulum by a zone of textura porrecta. Ectal
 113

excipu1um about 60 ~m broad at the central base, narrowing towards

the margin. Inner part predominately of a textura angularis with
cells mostly 12 - 20 (-24) x 6 -11 (-17) ~m, the cells are hyaline
to slightly light yellow, and orientated with their longitudinal
axes somewhat aslant to the surface, intermingled with scattered
subglobose cells. Along the sides with isodiametric c~lls 9 - 20 ~m
diameter. Towards margin the cells are more or less isodiametric,
occasionally epidermoid, with interspersed globose to sUbglobose
cells. The exterior part of the excipulum terminates in slightly
thick-walled, globose to elongated, light yellow-brown encrusted
cells, 4 - 12 ~m wide and up to 20 ~m in length, towards margin
more elongated to subclavate bulbs up to 7 x 17 ~m, the bulbs
originating several cell layers deep, their. rooting hyphae 1.5 -
2.5 (-3) ~m in diameter. Apothecia anchored to the substrate with
scattered septate, light yellow brown hyphae 2.5 - 3 ~m diameter.

Asci eight spored, cylindric, slightly narrowing towards apex,

rounded above, 150 - 250 x (12 -) 14 - 21 ~m, p. sp. up to 150 ~m,

Ascospores regular, ellipsoid to narrow ellipsoid, with rounded

ends, obliquely uniseriate, smooth, (16-) 17 - 21 (-22) x (7-)
7.5 - 10 (-11) ~m, occasionally light yellowish at maturity, each
spore surrounded by a thick gelatinous layer. Ascospores with
bipolar apiculus of somewhat variable form: long and narrow, or
hemispherical to pulvinate to papillate to acute, even pad-like,
not always visible on dry material or on ascospores in water.
Apiculi as well as outer thin spore wall layer stain strongly in
Cotton blue. Paraphyses hyaline below, filiform, septate, branched
or unbranched, below 1.2 - 2 ~m diameter, only slightly inflated up
to 3 ~m at their apices, this part with light yellowish granulated
content (484), ± encrusted, and embedded in a yellowish to light
yellowish brown (583) pigmented matrix; interspersed with broader
clavate paraphyses up to 5 ~m at their apices. This latter kind of
paraphyses have fewer septa, are slightly thick-walled, and their
apical parts possess yellowish brown encrustations. Paraphyses
extend 10 - 15 ~m beyond the asci.
 114

specimens examined:

ENGLAND. Surrey: near Esher: West End Common, horse dung, (me),
11.X.1981 B. Spooner as T. cf. apiculatus (K). New to England.
FINLAND. Tavastia australis: Forssa: Mustiala, horse dung, VIII.
s. date, P. A. Karsten: Fung. Fenn. Exs.; Cent. 8 No. 762 as Ascob.
albicans Fuck. (NY - ex herb. G. Massee, UPS); in Karsten 1871 a:
205; 1871 b: 231; 1871 c: 59 (as ~ albicans (Fuck.) Karst.); 1885:
121 (as Ascoph. albicans (Fuck.) Karst.) New to Finland.
SCOTLAND. Grampian: Type collection (see above); in Keith (1879:
11); nomen nudum.; Phillips & Plowright (1879: 100); Phillips
(1887: 98); Massee (1895: 420).
SWEDEN. Gotland: Hejnum: S of the church in a pasture, horse
dung, (me), 4.VI.1959 N. Lundqvist 2074-e (UPS - slide). -- Oland:
Boda: Grankullavik in shore meadow, horse dung, (me), 29.VIII.1959
N. Lundqvist 2292-d (UPS - slide only). -- Skane: Orkened:
Rumpeboda, horse dung in pasture, (me), 7.VI.1960 N. Lundqvist
2382-b (UPS). -- Dalsland: Dals-Ed par.: 1 km N of Grorud, horse
dung, (me after 19 days), 7.VIII.1961 N. Lundqvist 3135-a (UPS).
Sodermanland: Gryt: 1 km ESE of Gryt, horse dung in pasture, (me),
15.IX.1985 N. Lundqvist 15812-a (S). -- Katrineholm, horse dung of
horse in pasture, (me), 17.VII.1960 N. Lundqvist 2619-b (UPS). --
uppland:.Jumkil: W of Fagelbo, horse dung, (me), 10.V.1963 N.
Lundqvist 3820 (UPS). -- Arentuna: S of Granby, horse dung, (me),
7.IX.1977 N. Lundqvist 11046-a (UPS). -- Halsinqland: Mo: near the
church, horse dung, (me), 15.VIII.1960 N. Lundqvist 2786-d (UPS -
slide only). -- Inqermanland: ytterlannas: Bursjo chalets, horse
dung in pasture, (me), 27.VII.1959 N. Lundqvist 2234-e (UPS - slide
only). New to Sweden.
CANADA. ontario: Nipissing Distr.: Lake Timagami: Gordon Island,
horse dung, '22.VI.1933 R.·F. cain as Ryparobius sexdecimsporu5
(Cr.) Sacc. (TRTC 40020 - substrate only); in Kimbrough (1969: 111)
as holotype of T. apiculatus.
U.S.A. oregon: Type collection (see above); in Seaver (1942: 310)
as Ascoph. cinereus, Larsen & Denison (1978: 72).
 115

UNVERIFIED RECORDS: POLAND: near Breslau [= wroclaw], Schmidt

(1912: 30); type collection of Ascoph. appendiculatus (see
above). -- SCOTLAND. Moray: Forres, Boyd (1913: 72) as Humaria
keithii (Phill.) Sacc.; a literature record very probably founded
on the original collection of Keith. -- Pertshire: Balquhidder:
Kirkton Glen, Roger (1968: 472), as Ascoph. keithii, horse dung.
Orkney, Dennis & Spooner (1992: 500) as ~ apiculatus, horse
dung. -- JAPAN: Minoura & Yamada (1976: 325) on dung of cow and
goat, as T. apiculatus. Unfortunately, it has not been possible to
see this material, as no reply has been received on a loan request.
-- [formerly] u. S. S. R.: Prokhorov (1989 a) as T. apiculatus.

INCORRECT RECORDS: ENGLAND: Clark 1980 as T. apiculatus

[tentative determination] is T. rivicola. -- CANADA: Kimbrough 1969
(DAOM 63950) as T. apiculatus is T. crustaceus. -- U.S.A.:
Michigan, Kimbrough 1969 (NYBG) as T. apiculatus is T. crustaceus.

HABITAT AND SUBSTRATE PREFERENCE: Thecotheus keithii is an

exclusively fimicolous species, with 14 verified records on dung of
horse and also reported on dung of cow and goat (Minoura & Yamada
1976 as T. apiculatus), and from an unknown substrate (Prokhorov
1989 a).
DISTRIBUTION: England, Finland, Poland [as Ascophanus
appendiculatus], Scotland, Sweden, Japan (Minoura & Yamada 1976),
[formerly] U.S.S.R. (Prokhorov 1989 a), Canada and U.S.A. The
species is only known from the north-temperate zone, with records
from Europe, Asia and North America.
ILLUSTRATIONS: Minoura & Yamada 1976, fig. 1 a-d as T.
apiculatus. -- Ellis & Ellis 1988, plo 36, fig. 359 ·a.s T.
apiculatus.
PHOTOS: Kirnbrough 1969, figs. 21-23 as T. apiculatus. -- Minoura
& Yarnada, plo 1, figs. 9-12 as T. apiculatus.
 116

Thecotheus keithii is characterized by its small, symmetrical and

smooth spores which are furnish~d with bipolar apiculus. The form
of the apiculi are somewhat variable, from long and narrow to
smaller papillate. This was also observed by Minoura & Yamada
(1976) in their description of T. apiculatus.

The apiculi are not always visible in crush mounts in water, and
may be missing after strong heating in various reagents. The
nonapiculate spores of T. keithii should not be mistaken for those
of T. crustaceus, the latter differingin their inequilateral form.

Peziza keithii has been a·more or less forgotten species •. In. the
original report, Phillips (in stevenson 1879] remarks that this
species "has an outline much commoner amongst the Ascoboli than the
Peziz~, being thick and fleshy, with the hymenium only slightly
depressed, the form of a flattened sphere." The author briefly
mentions a relationship to "Ascophanum", but later it was regarded
to be a member of the genus Humaria (Saccardo 1889). with the
transfer to the genus Ascophanus (Boudier in litt. to Ramsbottom]
(Ramsbottom 1914), the species has for a long time been treated in
that genus, among others by Ramsbottom & Balfour-Browne (1951).

In recent years, Dennis (1968, 1978 and 1981) suggested that the
species seems to be only a narrow-spored variant of Iodophanus
carneus. He was later followed by Cannon at al. (1985). In
connection with the studies of types of coprophilous discomycetes,
Dr. J. van Brummelen kindly (in litt. 24.VI.1985) informed me that
Peziza keithii belongs to the genus Thecotheus, and that Thecotheu5
keithii would be an older name for Thecotheus apiculatus Kimbr.

Notes on type specimen

 117

Peziza keithii

Both Keith (1879) and Stevenson (1879) said that originally the
species was collected by Keith at Waterford in September, but
without mentioning any year. However, the label of the only packet
by Keith filed under Peziza (Humaria) keithii i~ the Kew herbarium·
(D. A. Reid in litt. 27.111.1987), is annotated November 1878, the
locality given is Forres [i. e. about 1~ km SE of Waterford]. It
originates from herb. mycol. M. C. Cooke 1885. Thus, presumably no
material in Phillips' or Keith's herbaria is in existance. The
collection examined. is in good accord with the original
description, and I have therefore designated the collection as a
neotype~

A9cophanu9 appendicu1atu9

The original collection of A. Schmidt from Oswitz, Poland, is lost

(in litt. 15.XI.1985 from WRSL). Both herbarium and types of this
German mycologist are unknown (Stafleu & Cowan 1985). Moreover, N.
Lundqvist (in litt.) has also informed me about a letter to him
from Dr. Krzysztof Rostanski 26.4.1965 (Wroclaw, BRSL): The
collections of A. Schmidt are not located in this herbarium, and it
is unknown where his herbarium could be deposited. Lundqvist (1973)
says that Schmidt's herbarium possibly perished in Berlin during
World War 11.

Alfred Schmidt's description of A. appendiculatus runs:

" Apothecien gelblichbraun, alt rotlichbraun, birnformig oder

verkehrt keglig, deutlich berandet, Scheibe flach oder gewolbt, 0,5
- 1,4 mm Om., 430 - 480 ~ hoch. Hulle aus gelblichen, rundlichen,
am oberen Rande quergezogen Zellen. Schlauche weit, bis 50 ~,

herausragend, zylindrisch oder zylindrisch-keulig, gerade oder

gebogen, oben abgerundet, mit einem Deckel aufspringend, nach unten
 118

allmahlich in einen Stiel libergehend, 210 - 250 ~ lang, 15,5 - 17,5

~ breit, sporenflihrender Teil 105 - 150 ~ lang, achtsporig, durch
Jodlosung nur undeutlich blau gefarbt. Sporen schrag einreihig
liegend, elliptisch,glatt, einzellig, farblos, 17,5 - 19,5 ~ lang,
8,5 - 9,5 ~ breit, an beiden Enden mit einem sehr kleinen, 1 - 2 ~

langen, kappenforrnigen Korper, auBerdem mit deutlicher

Gallerthtille. Paraphysen unverzweigt oder astig, farblos oder
besonders nach oben zu schwach gelblich, septiert, nach oben wenig
breiter, bisweilen an der Spitze bis 5 ~ dick. - Auf Pferdemisti
Mai. Oswitz."

Although the author did not make any illustrations, the description
of the species is sufficiently detailed to identify the species.

Thecotheus apiculatus

Kimbrough (1969) designated the collection housed in herbarium TRTC

40020 as holotype to T. apiculatus. This is a gathering on horse
dung from Canada: ontario: Nipissing Distr.: Lake Timagami: Gordon
Island, from June 22. 1933, collected by R. F. Cain. The type could
not be found, but John C. Krug at TRTC kindly traced it. The
collection TRTC 40020 had been annotated by Kimbrough in 1970 as
Coprotus duplus sp. ·nov., but in their pUblication on Coprotus by
Kimbrough, Luck-AlIen & Cain (1972), they do not cite the
collection under that name. I have seen TRTC 40020. The fungus
which grows on small fragments of horse dung, was originally
determined by R. F. Cain as Ryparobius sexdecimsporus (Cr.) Sacc.
Unfortunately the material is used up, as no specimens neither of
Thecotheus nor of Coprotus could be found, and a lectotypification
therefore is necessary.
 119

Ascobolus albicans Fuckel sensu Karsten.

Karsten's exsiccate No. 762 named Ascobolus albicans Fuckel, proved

to belong in Thecotheus. It is quite different from Fuckel's
original collection as distributed in "Fungi rhenani exsiccati a
Leopoldo Fuckel collecti" as No. 1855 (B, G, M, and S). The asci of
Fuckel's species do not stain blue in iodine, and the uniseriate,
ellipsoid ascospores, 12 - 14 x 7 - 8 ~m are ornamented with an
incomplete network. Moreover, the apothecia are hairy, and the
substrate is dominated by sand mixed with mosses, needles and other
organic remains rather than by horse dung (compare original
description by Fuckel 1866: 3). It has been regarded as a species
of Neottiella (Saccardo 1889, Boudier 1907); Lasiobolus (Saccardo
1889) and Lachnea (Rehm 1896), but van Brummelen (in litt.
24.VI.1985) regards it belonging to the genus Leucoscypha Boud.

Rehm (1896: 1068) was right when he doubted Karsten's

interpretation of Ascobolus albicans. Nevertheless, Karsten (1871
a: 206; 1871 c: 59) noted that the species resembled Ascobolus
cinereus Crouan. However, all examined collections of Karsten named
as A. cinereus belong to T. crustaceus, and the author did not make
any note of apiculate ascospores in Ascobolus albicans.

On Karsten's material in NY (ex herb. G. Massee) the name

Lasiobolus karsteni Massee is found, probably an unpublished
herbarium name. According to Pfister (1985), Karsten's exsiccate is
also distributed in the herbaria: FH, ?BPI, H and BM (list).
 120

Figs. 49 - 67. Thecotheus keithii. 49 - 56: Kienholz K 64 (NY -

lectotype, of T. apiculatus). 49 - 50: Narrow ellipsoid asco~pores,

rounded at their apices, their bipolar apiculus long and narrow.

51: Ascospores with variable form of .ap~culus, from long and narrow
to short and papillate. 52: Section of a mature apothecium showing
discoid form, and anchoring hyphae a: the lower part. 53 - 56:
sections of apothecium. 53: Margin layer with rooting hyphae ending
into slightly elongated bulbs (to the left). Then a zone of cells
composing a textura porrecta. 54: Lower part of excipulum (upside
down). To the right, cells of ectal excipulum composing a textura
angularis, the cells oriented with their longitudinal axes
perpendicular to the surface. Towards medullary excipulum (lower
left) a zone of textura porrecta. 55: Excipulum'from the central
part. At ~he bottom cells from the ectal excipulum composing a
textura angularis. Above the zone of textura porrecta is a part of
medullary excipulum with cells composing a textura globulosa. 56:
outermost, central part of ectal excipulum (below and left) with
globose bulbs of the rooting hyphae. 57 - 59: Neotype of Peziza
keithii (K). 57 - 58: Regular ellipsoid to narrow ellipsoid
ascospores. Note the long and narrow apiculus in 58. 59: Dry,
immature, sUbcylindrical apothecium. 60 - 62: Karsten: Fungi
Fenniae exsiccati No. 762 as Ascobolus albicans (UPS). 60: Two
apothecia, the lower right with doliiform to discoid form, the
other doliiform to turbinate. 61: Tips of paraphyses as seen i SEM-
photo. 62: SEM-photo of part of an ascospore with apiculus. 63 -
65: Sweden. Lundqvist 11046-a (UPS). 63: Ascus with oblique,
uniseriate ascospores. Note the variable form of spore apiculi. 64:
Section of the medullary excipulum, with globose to sUbglobose
cells composing a textura globulosa, interspersed with elongated
cells and filamentous hyphae. 65: Cells from the lower part of the
ectal excipulum, composing a textura angularis. The cells are
oriented with their longitudinal axes perpendicular to the
surface. Outermost with globose bulbs of the rooting hyphae.
66 - 67: England. Spooner (K). Ascospores with variable form of
their apiculi. Scales: 58 & 66 - 67: 18 ~m. 59: 1.3 mm. 60: 0.7 mm.
61 - 62: 5 ~m. Photos: 49 - 56 & 59 - 60 & 63 - 67: O. Aas. 57 -
58: J. Berge. 61 - 62: M. K. Stavdal.
121
124
125
 127

to THECOTHEUS LUNDQVIBTII ~as Sp. novo (Figs. 68 - 71)

~potheeia solitaria vel aggregata, sessilia, in sieeo 0.2 - 1

(- 1.5) mm diarn.; prime cylindricum, maturitate doliiforrne vel
turbinatum, denique ± discoideum, colore prima ex-albido,
deinde brunneo-aurantiaco vel pallide brunneo, plerumque
dilute lilacioo suffuso. Asci octospori, parietibus omnls iodo
caerulescentibus, 270 - 300 x 21 - 32 (-45) ~m. Aseosporae
uniseriatae, late ellipsoideae, 25 - 30 x 12.5 - 14.5 ~m,

granis minutis ornatae, apiculatae. Paraphyses simplices,

apicibus 5 (-7) ~m latis.
Fimicola.

Apothecia scattered or crowded, superficial, usually with a

subiculum-like base, 0.2 - 1 (-1.5) mm diameter in dry
condition, slightly rough and furfuraceous. At first
cylindrical, then expanding and becoming doliform to
turbinate, finally ± discoid.

Apothecia at first Whitish, when dry becoming brown-orange

(6C4) to light brown (7E5), usually with a lilaceous tinge and
a medium grey (IEI) lower part. Disc at first flat to ±
convex, becoming concave when dry at maturity, slightly rough
and white- furfuraceus owing to the protrUding ascus tips. At
first Whitish, darkening with age, when dry usually becoming
greyish orange (6B5). Margin entire.

Medullary excipulum forming a textura globulosa to teXtura

angular is, with ± subglobose to occasionally· polyhedral cells
varying from 5 - 12 ~m in diameter. Towards the hypotheciurn
usually with polyhedral cells, towards the ectal excipulum a
zone of long-eel led, septate, parallel-lying .hyphae forming a
textura porrecta. Ectal excipulum with ± subglobose to
 128

polyhedral, isodiametric cells, 10 - 18 ~m diameter, composing

a textura angularis. Towards the outer surface with slightly
elongated cells, their longitudinal axes perpendicular to the
outer surface. Rooting hyphae at the central base SUbglobose
to elongated, their bulbous, terminal cells up to 12 ~m wide
and up to 17 ~m in length, towards margin with more elongated,
subclavate bulbous cells up to 7 ~m in diameter. Apothecia
anchored to the substrate with hyaline, septate hyphae, 2 -
3.5 ~m in diameter.

Asci 8-spored, cylindrical to cylindrical-clavate, rounded

above, tapering towards the base, 270 - 300 x 21 - 32 (-45)
~m. Ascospores hyaline, broadly ellipsoid, uniseriate, smooth
and thick-walled when young, at·maturity thin-walled,
ornamented with minute granules, 25 - 30 x 12.5 - 14.5 ~m, and
with subglobose to hemispherical bipolar apiculus up to 2.5 ~m

long and up to 3 ~m broad. occasionally a few abortive

ascospores occur in some asci. Each ascospore surrounded with
a gelatinous envelope. Paraphyses hyaline, filiform, septate,
1.5 - 2.5 ~m wide below, with apices up to 5 (-7) ~m in
diameter, exceeding the asci by 15 - 30 ~m at maturity.

Specimens examined:

SPAIN. Caceres: Parque Nacional Monfrague, cow dung,

22.XI.1985 J. M. Barrasa 9366-82 (Private herbarium of
Barrasa, Madrid).
SWEDEN. Gotland: Eksta: Lilla Karlso Island, sheep dung,
(me), 30.VI.1988 N. Lundqvist 17195-b (S - Holotype). --
Oland: Vickleby: on the Great Alvar (Helianthemum heath), old
cow dung, 3.VIII.1961 N. Lundqvist 3115-a (UPS). --
Sodermanland: Aspo: Aspo Island, N part, cow dung in marshy
meadow, (me), 30.V.1959 N. Lundqvist 2023-e (UPS).
 129

The species is characterized by subiculum-like apothecia, form

and size of the ascospores, and the hyaline paraphyses.
Probably it is closely related to T. holmskjoldii, as the
apotheciaL form, but not size, is very similar to that
species. T. holmskjoldii, h0wever, does not have a subiculum-
like base. A similar subiculum is also observed in T.
inaequilateralis. T. lundqvistii differs from T. holmskjoldii
in having a different composition of medullary excipulum,
smaller asci and ascospores, a different form of the asci, and
a different ascospore arrangement. The thread-like structures
emanating from the ascospore apiculi in T. holmskjoldii are
not observed in living material of T. lundqvistii. Another
distinction is the always hya~ine paraphyses in T.
lundqvistii, as compared with the brownish encrusted
paraphyses of T. holmskjoldii.

T. lundqvistii is a fimicolous species so far known on dung of

cow and sheep. It is only known from Spain and Sweden. The
material is too limited to draw any conclusion about its
distribution pattern.

Figs. 68 - 71. Thecotheus lundqvistii. 68 - 69: Holotype (S).

68: cylindrical asci with ascospores and paraphyses. 69: Dome-
shaped tips of asci indicating the zone where the operculum
breaks off. 70 - 71: Spain. Barrasa 9366-82. 70: Ascospores
with bipolar apiculus. 71: SEM photo of a part of an ascospore
showing ornamentation of minute granules. An apiculus to the
right. Scales: 68: 50 ~m. 69 - 70: 20 ~m. 71: 5 ~m. Photos: J.
Berge.
 131

10. THECOTHEUS PALLENS (Bou~.) Kimbr.

Mycologia 61 (1): 112 (1969). -- Basionym: Ascophanus pallens

Boud., Bull. Soc. Bot. France (Session Cryptog. 1887) 34 (9): XLVII
(1888); Boud. Bull. Soc. Mycol. France 4: XLVII (1888). -- Type:
France, Paris, near Ecouen, in a shady forest on clayey soil in an
area which is periodically inundated, IX 1887 Boudier, not known to
exist; (Boud. ibid., pI. II, fig II a - h., lectotype selected
here) .

ETYMOLOGY: Latin palli~us, pale, light-coloured; referring to the

whitish to grey-white colour of the fruit-bodies.

Boudier's original description of Ascophanus pallens runs:

"Minutus, 1 - 3 millim. latus, albidus aut albido-cinereus.

Receptaculum hemisph~ricum aut lenticulare, sub-marginatum, extus
leve, hymenio pIano aut convexo, thecis prominentibus longe
papillato~ Paraphys~ hyalin~, s~pius continu~, ad apicem·
incrassat~, ad basim divis~ et septat~, apice 6 - 8 ~ crass~. Thec~

maxim~, ampl~, clavat~, octospor~

290 - 350 ~ long., 35 - 45-
crass~, operculo lato aliquoties umbonato. Spor~ maxim~, oblongo-
fusiformes, hyalin~, intus nullo modo granulos~, long. 40 - 45,
latit. 15 - 20."

UNVERIFIED RECORDS: FRANCE: Boudier (1888), type collection. Also

mentioned in Boudier (1904 - 1911) 4: No. 470; Boudier (1907: 77);
Grelet (1944: 93); Kimbrough (1969: 113); Grelet (1944: 93), a
collection from Savigne collected 4.VIII.1914 on moist soil among
green algae in a shaded meadow area which was periodically
 132

inundated; not known to exist.

INCORRECT RECORD: U.S.A.: Seaver (1928: 124) as Humarina pallens

(Boud.) Seaver is Thecotheus cinereus. This collection is also
mentioned in Pfister (1981: 1001, 1982: 8 & 22) as Thecotheus
pallens (Boud~) Kimbr.

HABITAT AND SUBSTRATE PREFERENCE: On periodically inundated soil.

DISTRIBUTION: Only known from two localities in France. I reg~rd

it as a rare species.
ILLUSTRATIONS: Boudier 1888, pI. 2, fig. 2. -- Boudier (1904 -
1911), 2: pI. 414 a-j. -- Chadefaud 1960, fig. 395 no. 12.

Thecotheus pallens is characterized by the large, oblong-fusiforme

ascospores. T. cinereus has spores of about the same size, but
cylindrical-ellipsoid.

Unfortunately, neither the type nor the collection of Grelet (1944)

are known to exist (Pfister 1981 concerning Boudier's collection,
and R. Cailleux, herbarium PC, Paris in litt. 4. XI. 1985). In
accordance with Article 7.5 in the ICBN (1988), Boudier's excellent
illustration is selected as lectotype of T. pallens. So far, I have
chosen to follow the opinion of Kimbrough (1969), and refer it to
Thecotheus, although there are still unanswered questions
concerning its position. Freshly collected material is needed.

Boudier does not say anything about the reaction of asci in iodine.
This is striking since he generally records this. Grelet (1944), on
the other hand, states the asci to be iodine-negative.

It is well known that the measurements of microscopic structures in

Boudier's later descriptions are too large (Maire 1917, 1928 and
Brummelen 1969). By means of Brummelen's (1969) adjusted measuring
 133

scale used on Boudier's (1888) drawings, the size of the ascospores

are found to be within a range of 36 - 42 x 13 - 17.5 ~m, whereas
the paraphyses are 1.5 ~m broad below and up to 9 ~m wide at their
apices.

Of all the material I have examined during this work, I only once
came across a collection with true fusiform ascospores. This is a
gathering (slide only) of B. o. J. Dodge from 1911 from Bermuda,
deposited in the herbarium NY. No substrate is indicated, and the
slide is labeled "Ascophanus crossed with a Sordaria. A hybrid n.
sp.?" Th~ ascospores measure 50 - 60 x 20 - 28 ~m. It is difficult
to decide whether this is a member of Thecotheus, and no species in
this genus is known to have such large spores.

11. THECOTHEUS PELLETIERI (cr. , cr.)Boud. (Figs. 72 - 80)

Annls. Sci. Nat. Bot. 5 ser. 10: 236 (1869). -- Basionym:

Ascobo1us pe11etieri Cr. & Cr., Ann. Sci. Nat. Bot. IV. 7: 173
(1857). -- Pezizula pe1letieri (Cr. & Cr.) Speg., Michelia 1 (2):
238 (1878). -- Ryparobius pe11etieri (Cr. & Cr.) Sacc. in Vido,
Michelia 1 (5): 605 (1879). -- Mo11isia pe11etieri (Cr. & Cr.)
Gill., Champignons de France. Les Discomycetes. 1: 135 (1879).
Ascophanus pel1etieri (Cr. & Cr.) Quelet, Enchiridion fungorum in
Europa media et praesertim in Gallia vigentium: 295 (1886). --
Type: France, Finistere, near Brest, on old dung of cow in a marsh,
20.VI.1857 Crouan (PC A2353, lectotype (part of type); not seen.

Misapplied name: Ascobo1us solms-laubachii [laubachi] Rabenhorst

1862. See appendix.
 134

ETYMOLOGY: Latinized from the name Pelletier: The Crouan

brother's (1857) dedicated this interesting and - to them - rare
species to commander Pelletier from Morlaix, a competent
illustrator of cryptogams.

Apothecia scattered to gregarious to crowded, sessile or with a

short immersed stern-like base, 0.2 - 3 mm diameter, up to 1.5 (-2)
mm high; closed till the spores are mature (angiocarpic
development). At first globose or shortly cylindrical to obconic,
becoming dolabriform to cylindrically doliiform, finally turbinate
or broadly discoid. Dry apothecia often sub-cylindrical to
cylindrical turbinate, with vertical ridges owing to the large
asci. Apothecia at first white, soon becoming dirty white to
greyish, with age tinged with brown or red, finally on drying with
darker brown or red greyish colour; at first smooth, after drying
somewhat dirty white, minutely mealy or pruinose on the surface
owing to the slightly protruding ectal cells. Dry apothecia
moistened in water mostly with pale yellow to light orange colour.
Anchoring hyphae more or less distinct, sUbhyaline, thin-walled,
septate and branched, up to 2.5 ~m wide. Disc plane or convex,
after drying occasionally slightly concave, mostly regular,
sometimes irregular due to mut~al compression, soft, fleshy. After
drying characteristically white-mealy to furfuraceous and roughened
by the protruding tips of asci. Margin narrow, entire to wavy.

Medullary excipulum subhyaline, of varying thickness, up to 280 ~m

at the central growing part, composed of a textura globulosa, with

thin-walled loosely arranged globose to subglobose cells, (10-)
15 - 30 ~m diameter. The cells are intermingled with scattered,
septate, filamentous hyphae. Ectal excipulum about 60 - 80 ~m thick
along the sides, composed of two types of cells. Innermost cells
composed of a textura angularis, with somewhat elongated,
polyhedral, slightly thick-walled, sUbhyaline cells, 8 - 17 x 14 -
25 ~m, with their longitudinal axes perpendicular to the surface.
 135

Interspersed are filamentous hyphae, 2 - 4 ~m wide, which originate

several cell-layers deep and terminate on the surface. Along the
sides on the surface, the tips of these terminal cells are thick-
walled, inflated up to 5 - 10 ~m diameter, encrusted with yellowish
to light brownish pigments.

Asci comparatively few, protruding, hyaline, clavate to broadly

clavate, 32-spored, rarely 16- or 64- spored, apex obtuse and with
a faint ring in Congo red, dome-shaped, below usually with a short
stem-like base, slightly thick-walled when immature, becoming
thinner with age, 260 - 460 (-475) x 41 - 70 (-90) ~m, with stalk
20 - 90 ~m long, p. sp. 240 -350 ~m, the I+ reaction decreasing
with age. Ascospores non-apiculate, smooth, hyaline, ellipsoid to
broadly ellipsoid, slightly narrowed at the ends, 2- to 4- (5)
seriate or irregularly disposed, mostly 2- seriate below, (30-)
32 - 40 (-42.5) x (13-) 15 - 22 (-24) ~m. In asci with 64
ascospores, the spores are sUbglobose, 17 - 24.5 x 10 - 16 ~m.

Ascospores in the 16-spored asci are 32 - 40 x 16 ~m. The spores

are granular within when young, and with several different-sized
vacuoles, initially thick-walled, becoming thin-walled by the time
of spore liberation. The wall is composed of three wall layers, a
thick inner layer (up to 4 ~m in immature spores, mostly 2 - 2.5 ~m

at maturity, not stained in Cotton blue), a thin (0.5 ~m thick)

outer layer that stains in Cotton ~lue. The thin outer wall layer
becomes loose and distorted in heated 2% KOH. Such ascospores may
appear wrinkled or slightly warty. Rarely very short polar
granulations occur that stain in Cotton blue. Each ascospore is
surrounded by a thick, up to 6 ~m broad, gelatinous perispore.
Paraphyses slender, septate, simple or branched especially in lower
parts, hyaline below, (1-) 1.5 - 2.5 (- 3) ~m wide, enlarged above
to (3.5-) 4 - 8 ~m diameter, and there slightly encrusted with
brownish or brownish-red pigments. The paraphyses normally reach
the same level as the asci, occasionally prot~uding up to 15 ~m.
 136

Specimens examined:

AUSTRIA. steiermark: the vicinity of cilli (south steiermark),

cow dung, (me), XI.1888 H. Zukal (W - s~ide, ex herb. A. Heimerl,
Wien); in Heimerl (1889: 20), Rehm (1896: 1100). -- Grazer
Bergland: Graz-Maria Trost: Gestlit bei GH. Gruber, horse dung,
X.1982 J. Haffellner (GZU); in Schweiger (1985: 66). --
Schladminger Tauern: Kleinsolktal slidl. Grobming: Kleinsolk-
Obertal: Sturnrnerkessel about 1400 m.a.s.l., horse dung,
30.VIII.1983 H. Schweiger (GZU); in Schweiger (1985: 66). --
Schladminger Tauern: Solktal slidl. Grobming: Kleinsolk-Obertal:
Sacherseealm about 1050 m.a.s.l., cow dung, 26.X.1982 H. Mayrhofer
(GZU); in Schweiger (1985: 66). -- Weiz: st. Kathrein Zeil: Weg
Eder-Eibisberg about 1180 m.a.s.l., cow dung, 10.X.1981 H.
Schweiger (GZU); inSchweiger (1985: 66).
AZORES. Terceira: Angra do Heroismo, ass dung, (me), 27.VI.1975
R. W. G. Dennis (K); in Dennis et al. (1977: 95), Korf & Zhuang
(1991: 318).
BELGIUM. F1andria [= Oost-V1aanderen]: near Gandavum [=Gent], cow
dung, s. dato E. Coemans in Rabenhorst: F. Eur. exs. No. 167 as
Ascob. pelletieri Cr. (C, CUP, G, K - ex herb. M. C. Cooke, L, RO -
ex herb. De Notaris, RO - ex herb. V. Cesati as Ryparobius
pelletieri (Cr.) Sacc., S - ex herb. P. Sydow, S - ex herb. H.
Rehm, UPS, WRSL)i in Coemans (1862: 88, on old cow dung in summer
1859-1862 at Petit-Beguinage near Gand, also referred to in Coemans
1863: 138), Kickx (1867: 477), Lambotte (1880: 507) and Patouillard
(1883: 74).
CZECHOSLOVAKIA. Bohemia: Mnichovice: Myslfn, horse dung,
30.IX.1924 J. Velenovsky (PRM 147300); in Velenovsky (1934: 363). -
- Mnichovice, old cow dung, VIII.1925 J. Velenovsky (PRM 152924);
in Velenovsky (1934: 363). -- Mnichovice, old horse dung, VIII.1926
J. Velenovsky (PRM 152925); ? in Velenovsky (1934: 363). --
Mnichovice, old cow dung, VIII.1938 J. Velenovsky (PRM 152927).
Rychory Mts.: Hornf Marsov, dung, 4.VI.1962 M. Svr~ek and J.
Kubi~ka (PRM 568504). -- Vrabsko: near timelice at piscinam
"Nerestec", horse dung, 2.IX.1963 M. Svr~ek 925/63 (PRM 816081).
 137

Moravia: Mahr.-Weisskirchen: Welka, cow dung, 21.IX.1912 F. Petrak:

Flora Boh. Mor. exs. (11:1) No. 231 as Rhyparobius pelletieri (Cr.)
Rehm (E, FH, M, PRM 742690, S - ex herb. H. Rehm and ex herb. P.
Sydow) .
DENMARK. s. loco, cow dung, 5.X.1875 E. C. Hansen (C - also a
capsule and a slide No. 228 in F. Fim •. Exempl.- Exs. -p. 10 as T.
pelletieri (Cr.) Boud.). -- s. loco, horse dung, s. date I. Egeland
402 not saved (BG - description and drawing).
ENGLAND. North Yorkshire: Scarborough, horse dung, s. dato [G.
Massee] (NY - drawing only, ex herb. G. Massee); in Phillips and
Plowright (1881: 69 as Ascob. (T.) pelletieri Cronan (sic!]),
Phillips (1887: 297 as T. pelletieri (Cr.) Boud.), Massee (1895:
180 as Ryparobius pelletieri Sacc.), Massee & Cross land (1905:
295) •
FRANCE. Finistere: near Brest," on horse dung and old cow dung in
marshes and meadows, spring, autumn" s. dato, Crouan (CO A2390c -
syntype, part of type, RO - ex herbaria of Crouan and De Notaris,
syntypes, part of type); in Crouan & Crouan (1867: 56), Brummelen
(1967: 231 as regards A2390c). -- near Brest, horse dung,
30.VI.1858 Crouan (CO A2390b). -- near Brest, horse dung, 9.X.1866
Crouan (CO A2390a). -- Maine-et-Loire: Angers, dung [probably of
horse], VI.1900, misit D. Gaillard [probably a collection belonging
to herb. Boudier] (PC). -- Vosqes: in vaivre Forest, old cow dung,
16.IX:1878 L. Quelet (UPS - ex herb. L. Quelet); in Quelet (1879:
236), Rehm (1896: 1100).
GERMANY. Gera: Greiz: Berga at Elster: near Clodra, cow dung in
pasture, 22.VIII.1974 E. Paechnatz (BHU). -- Bessen: Oestrich
(Nassau), old horse dung, autumn, s. dato, L. Fuckel: Fungi Rhenani
Exsiccati a Leopoldo Fuckel Collecti No. 2376 as Ascob. pelletieri
Cr. (FH - ex herb. L. Fuckel also ex herb. Barbey-Boissier No. 1307
as Ryparobius pelletieri (Cr.) Sacc., K - ex herb. C. E. Broome,
S - 3 colI.: ex herb. Fuckel, ex herb. Barbey-Boissier No. 1307, ex
herb. H. Rehm). -- Niedersachsen: LUchow-Dannenberg: near Gurnrnern,
red deer dung, 18.IX.1988 E. Jahn (BG - photo only, asci with both
32 and 16 ascospores). -- ditto: Pevestorf, cow dung, 15.VII.1989
E. Jahn (BG). -- Nordrhein-westtalen: MUnster, horse dung, (me),
spring 1889 o. Brefeldt 374 (S - ex herb. H. Rehm). -- Rheinland -
 138

Pfalz: Eifel: near Gerolstein: Heiligenstein, isolated from horse

dung, 5.X.1971 J. van Brummelen 2446 (L).
NORWAY. 0stfold: Hvaler: Kirk0Y: 0rdal, cow dung in pasture land,
6.XI.1983 R. Kristiansen (BG). -- ditto: KrAker0Y: Bj0rnevAgen, cow
dung in an oak-grove with scattered pines, dog roses and juniper,
16.IX.1982 R. Kristiansen (BG). -- Oslo: Bygd0Y, cow dung, (me),
16.111.1955 F.-E. Eckblad (0); in Eckblad (1956: 228), Eckblad
(1968: 25), Aas (1978: 194), Sivertsen (1982: 41). -- Buskerud:
R0yken: Slemrnestad, cow dung, 29.IX.1963 I. Egeland 422 (BG -
slide). -- ditto, 30.IX.1962 I. Egeland 294 not saved, description
and drawings in (BG). -- Hordaland: Tysnes: near Espevik, horse
dung, 30.VIII.1985 S. Olsen (BG). -- Sogn og Fiordane: Hyllestad:
Leirvik i Sogn: at the lake Handalsvatnet, horse dung in
pastureland, 10.X.1980 O. Aas 60/80 (BG). -- Nord-Tr,ndelag:
Inder0y: R0ra, old cow dung, (me - after four weeks) ,17.VI.1962 N.
Lundqvist 3433-b (UPS); in Aas (1978: 194), Sivertsen (1982: 41).
POLAND. Silesia: Breslau, herbivor dung [probably of horse), (IX
?)1886 J. Schroeter (WRSL - ex herb. J. Sehroeter)i in Sehroeter
(1893: 53). -- Breslau, elephant dung in the zoologieal gardens,
3.VII.1886 J. Sehroeter (WRSL - ex herb. J. Schroeter); in
Sehroeter (1893: 53), Rehm (1896: 1100).
SPAIN. vizcaya: Pena Leeanda, eow dung, (me), 28.XI.1980 J. M.
Barrasa 3142-72 (private herbarium of Barrasa, Madrid - slides); in
Barrasa (1985).
SWEDEN. Gotland: VAskinde: W of Sjalso near the seashore, horse
dung, (me), 2.VI.1959 N. Lundqvist 2034-k (UPS). -- Smaland:
Bjorko: N of Rodjenas, horse dung in pasture, (me), 17.VII.1960 N.
Lundqvist 2616-b (UPS). -- Morlunda: Morlunda, old eow dung in a
pasture, (me - after 1 month), 6.VI.1962 N. Lundqvist 3352-g (UPS).
-- Bohuslan: Hede: 2 km N of the N point of Lake Karnsjon, old eow
dung in a pasture, (me - after 1 month), 7.VIII.19~1 N. Lundqvist
3132-d (UPS). -- Vastergotland: Osterplana: osterplana near the
ehurch, old cow dung in ealehareous heath, (me), 13.VI.1960 N.
Lundqvist 2443-k (UPS). -- Sodermanland: Aspo: on the N shore of
Aspo Island at the br~dge, eow dung, (me), 30.V.1959 N. Lundqvist
2023-j (UPS). -- Uppland: Balinge: 1 km N of Marsta, very old cow
dung in spruce forest, IB.VIII.1963 4066-k (UPS). -- Haga:
 139

Torslunda, horse dung, (me), 26.X.1967 E. Gunnerbeck 1333-a

(UPS). -- Laby: Vadbacka, old cow dung, (me - after 20 days),
15.IV.1962 N. Lundqvist 3302-c (UPS). -- Lena: 1 km NE of
Storvreta, old cow dung in pasture in the wood, (me), 7.IX.1968 N.
Lundqvist 6209-d (UPS - slide). -- Jamtland: Morsil: SE of Jarpen,
old cow dung in coniferous forest, (me), 16.VI.1962 N. Lundqvist
3425-b (UPS). -- Norrbotten: Pajala: W of Erkheikki, old cow dung
in pine forest, (me), 4.VIII.1960 N. Lundqvist 2676-a (UPS). New to
Sweden.
CHINA. Kwangsi: Loh-hoh-Tsuen (Ling Yuin Hsien), cow dung,
14.V.1933 S. Y. Cheo (FH). New to China.
INDIA. Himachal Pradesh: Dalhousie, cow dung, X.1955 L. R. Batra
157 (CUP IN35); in Batra and Batra (1963: 145). -- Jammu and
Kashmir: Bhadarwah: Doda, dung in coniferous forest, 2.X.1966 K. S.
Waraitch 2166 (BPI, C); in Waraitch (1977: 1, 3), Kaushal (1980:
319). -- ut tar Pradesh: Mussoorie: Mossy Falls, cow dung,
3.VIII.1960 L. R. Batra 1014 (CUP IN175). -- Mussoorie: The Park,
cow dung, 1.VIII.1960 L. R. Batra 897 (CUP IN189 & CUP IN176); in
Batra and Batra (1963: 145). -- west Bengal: Darjeeling: Algara
hill: Kalimpong [Calcutta: Precidency College in herb.!), cow dung,
9.V.1967 A. K. Kar and K. P. Pal PCC 5 (CUP-IN 436 - card only,=
CUP 49519); in Kar & Pal (1968: 1091).
RUSSIA. Moscow: Moscow city.: at Moscow river, cow dung,
29.X.1981 V. P. Prokhorov (MW). -~ ditto: Moscow city: horse dung
from circu~, 20.IV.1984 V. P. Prokhorov (MW). -- ditto: New
Yerusalem distr., cow dung, 1.X.1920 [legator unreadable) (MW).
Sakhalin isl.: Starodubsk, dung [probably of horse), 28.VIII.1970
B. Kullman and A. Raitviir 62130 (TAA). New to Russia.
AUSTRALIA. New South Wales: Sydney, cow dung, 22.VI.1966 C. G.
Carre (K). New to Australia.
CANADA. ontario: Charlton, horse dung, 29.VII.1931 R. F. Cain
(CUP 47649). -- Maple, cow dung, 22.IX.1940 R. F. Cain (CUP 47648).
-- York Co., cow dung, 2.X.1936 R. F. Cain (CUP 47650). -- Quebec:.·
Montreal, lIe Jesus, Ste. Dorothee, horse dung, 26.VIII.1941 R. F.
Cain (CUP 47603, TRTC 12370); in Kimbrough (1966), Kimbrough & Korf
(1967): photo of CUP 47603, Kimbrough (1969: 109). -- Ste.
Catharine, cow dung, 25.VIII.1938 R. F . .Cain (CUP 47656 - ~lide);
 140

in Mains, Overholts & Pomrnerleau (1939: 730).

U. S. A. s. loco, dung, VII.1906 Harper (FH). -- Massachusetts:
Clayton, cow dung, 13.1X.1892 J. L. Zabriskie 433 (NY - slide, ex
herb. J. B. tllis).-- Michigan: Ann Arbor, dung, (me), 3.111.1949
A. H. Smith (M1CH). --Mc. Coy's Wood: Stockbridge, horse dung
14.X.1928 G. B. eummins 133 (NY - ex herb. G. B. Cummins). -- New
Hampshire: Hanover, dung [probably of cow], 26 and 27 VIII.1937 F.
J. Seaver (CUP 47653, NY). -- Thorn Hill: Balsam Hollow, dung
[probably of cow], IX.1901 R. Thaxter (FH - ex herb. R. Thaxter).
-- New York: Mc. Lean: Lloyd Cornell Preserve, dung [probably of
cow], 25.V.1964 A. SAnchez and R. ? Korf (CUP 47749). -- Staten
Island: Pennington, dung, 18.VI.1913 F. J. Seaver (NY - slide). --
North Carolina: Cullowhee, cow dung, VI.1887 R. Thaxter (FH - ex
herb. R. Thaxter, FH - ex herb. W. G. Farlow). -- Highlands, cow
dung, 30.VII.1931 s. collector 1316 det. F. J. Seaver as T.
pelletieri (CUP 47654 - slide, NY). -- Ohio: oberlin, on laboratory
horse dung cultures, 13.11.1895 F. D. Kelsey 95/1 as Ascob.
furfuraceus (NY - ex herb. J. B. Ellis). -- Preston G., cow dung
s. ann. & date A. P. Morgan 14 (NY - ex herb. G. Massee)i in
Morgan (1902: 182). -- oregon: Hood River, horse dung, 2.IX.1932 J.
R. Kienholz K33 and K34 (CUP 47657 - slide, NY)i in Seaver (1942:
312). -- Mt. Hood, horse dung, 2.X.1922 L. E. Wehmeyer (CUP 47652,
FH, MICH, UC - formerly Univ. of Michigan). -- Tennessee: Burbank,
cow dung, VIII.1896 R. Thaxter (FH -ex herb. R. Thaxter). -- near
Decatur (Meigs County), cow dung, 23.111.1941 L. R. Hesler (MICH -
ex herb. TENN 13381; asci with 32 and 64 ascospores, TENN 13381). -
- near Hopewell (Knox County), cow dung, 15.VI.1941 L. R. Hesler
(TENN 13696); in Meyer (1941: 403). -- Wisconsin: Algoma, cow dung,
IX.1909 B. o. Dodge 612 (S - ex herb. H. Rehm). -- Madison, old
horse dung in woods, VI1.1906 J. B. Overton, det. Rehm ~ Harperia
overtonii Rehm n. sp.; nom. inval. (FH - also inclUding copy of
letter of Rehm to R. A. Harper, dated July, 21. 1906, concerning
description of this "new" genus and species, S - ex herb. H. Rehm,
two colI.).
BERMUDA. Paget: Agricultural Experiment Station, horse dung,
29.X1.-14.X11. 1912 S. Brown, N. L. Britton and F. J. Seaver 1463-
6, 1566 and 1596 slide (NY, CUP 47655 slide); all in Seaver (1916:
 141

506; 1928: 148); Pfister (1982). -- Paget: Agricultural Experiment

station, horse dung, 13. & 25.1.1926 F. J. Seaver and H. H. Whetzel
22 & 55 (CUP 34622 & CUP 34632, NY).
DOMINICAN REPUBLIC. st. Paul: along road between Springfield
Estate and Corona Estate, dung (? horse), 23.V1.1970 R. P. Korf et
al. (CUP D0163). -- Springfield Plantation: 7 miles from Roseau:
near Bee House, on dung between woods in orchard, 27.VI.1970 R. P.
Korf et al. (CUP D0219). New to Dominican Republic.
JAMAICA. st. Andrew Parish: Vicinity of Dick's Pond: W of Hardwar
Gap near Holywell Recreation Area and Wag Water River, elev. 2,800-
3,000 feet, mule dung, 10.1.1971 R. P. Korf et al. as ~. sp. (CUP-
MJ250, CUP-MJ251 see Pfister 1974: 366). -- Traveller's rest:
Silver Hill Gap on the border of Portland and st. Andrew Parish,
cow dung, 8.1.1971 R. P. Korf et al. as~. sp. (CUP-MJ99). -- Trail
between Holywell and source of Wag Water River, cow dung, 20.1.1971
R. P. Korf et al. (CUP-MJ677); in Pfister (1974: 366). --st. Thomas
Parish: Trail between Barretts Gap and Corn Puss Gap, cow dung,
15.1.1971 R. P. Korf et al. in Korf and Gruff: Discomycetae
exsiccate (CUP MJ519). -- ditto (CUP MJ506).
MEXICO. Mexico City (Ciudad de Mexico): Mt. Popocatepete, burro
(= donkey) dung, 10.VII1.1961 R. F. Cain (CUP 47651 substrate
only). New to Mexico.
PUERTO RICO. Along Rte. 105 at km. 23.0, Maricao Forest Reserve,
horse dung, 13.V1.1970 R. P. Korf et al. (CUP PR4068).
BRAZIL. Rio Grande do Su1: Sao Leopoldo, old dung (probably
of cow), 1929 J. E. Rick (FH - 3 coll.). New to Brazil.

UNVERIFIED RECORDS: BELGIUM: Bommer & Rousseau (1884: 144), cow

dung. -- Maistriau & Ronflette (1884: 163), cow dung. -- BULGARIA:
Fakirova (1974: 10), dung. -- CZECHOSLOVAKIA: Smarda (1944: 8). --
Svr~ek & Kubi~ka (1961: 35) horse dung. -- DENMARK: Larsen (1971:

7), fallow deer dung. --ENGLAND: Bucknall (1880: 68), a collection

from Bristol, on soil. -- Phillips & Plowright (1881: 69) &
Phillips (1887: 298), a collection of C. Bucknall from Bristol, s.
 142

substr., (me). -- Massee (1895: 180), with reference to collections

on dung of horse, cow, sheep and dog in England. -- Dennis (1968:
65), on dung of cow and horse. --FRANCE: three collections
mentioned in Boudier (1869: 236). The author says that the species
is most frequent on old cow dung, infrequent on dung of horse,
sheep, and rare on dung of rabbit. -- Probably the same reference
in Gueguen (1908: 46). -- Guernisac (1879: 46), horse and cow dung.
-- Grelet (1944:80), horse and cow (3) dung. -- GERMANY: Fuckel
(1866: 2; 1870: 288), old dog dung. -- Engel & Svr~ek (1983: 59),
s. substrat. -- E. Jahn, Bad Schwartau, BDR (unpubl., in litt.
4.XI.1989) with following collections kept in his private
herbarium, dung of: horse (5), sheep (2), fallow deer (2) and
european bison (1). -- HUNGARY: Banhegyi (1940: 115), cow dung.
- Banhegyi (1942 b: 269), horse dung. This collection is also
mentioned in Banhegyi et al. (1985: 654). -- ITALY: spegazzini
(1878: 238) & vido (1879: 595 & 605) as Pezizula pelletieri (Cr.)
Speg., horse dung. -- Bizzozero (1885: 341), horse dung. --
LITHUANIA: Krejwis6wna (1936: 22) as Rhyparobius pelletieri, rabbit
dung. -- LUXEMBOURG: Feltgen (1899: 29 & 375), cow dung (3).
NETHERLANDS: Oudemans (1900: 210), as Ryparobius pelletierii [sic!]
(Cr.) Sacc., cow dung. -- POLAND: Schroeter (1893: 53) cow dung. --
Eichlera (1904: 27) as Rhyparobius pelletieri (Crouan) Rehm, cow
dung. -- Szab6 (1905: 18) sheep dung. -- Eichlera (1907: 15), horse
dung. -- Schmidt (1912: 15) as Rhyparobius pelletieri (Cr.) Sacc.,
dung of cow, goat and sheep. -- Kohlman-Adamska (1965: 92), cow and
horse dung ..-- Bednarczyk (1974: 332 & 338), cow and horse dung. --
Chmiel (1977: 88), cow dung (2). -- ROMANIA: Racovitza (1945: 18),
horse dung (2). Also mentioned in Bontea (1953: 493) as Rhyparobius
pelletieri (Cr.) Sacc. -- SWEDEN: The following collections of N.
Lundqvist ( in litt.), all on horse dung: Skane: 3371.
vasterqotland: 2426. Narke: 2361. Sodermanland: i548. J&mtland:
3392. -- UKRAINE: Girzitska (1929: 66) as Rhyparobius pelletieri,
horse dung. -- INDIA: Batra (1954), cow dung. -- JAPAN: otani
(1973: 31), cow (3) and horse (2) dung. -- CANADA: Seaver 1942:
312. -- ontario, cow dung; in Kimbrough (1969: 109). -- U. S. A.:
Overton (1906: 466) horse dung. --·conway (1975: 242) horse dung. -
- Larsen & Denison (1978: 72). -- Kimbrough & Curry (1985: 220),
 143

two collections, no substrate mentioned. -- Florida, cow dung; in

Kimbrough (1969: 109); Samuelson (1978: 1862). -- Iowa: Iowa City,
cow dung, (me),; in Seaver 1904: 277; 1905 a: 113; 1905 b). -- New
York: on dung; in Dodge (1912: 152), Kimbrough (1969: 109), cow
dung. -- North Dakota: Seaver (1909: 109; 1928: 148) on dung.
Seaver's (1909) collection also mentioned in Brenckle (1917:
292). -- Oregon: Snyder (1938: 11), horse or cow dung. --
virginia: Wilson (1947: 375) horse dung. -- ARGENTINA: Spegazzini
(1880: 169) as Ryparobius pelletieri (Cr.) Speg., cow and horse
dung. -- Spegazzini (1899: 310) as Ryparobius pelletieri (Cr.)
Sacc., cow dung. -- PUERTO RICO: Pfister (1974: 366), cow dung.

HABITAT AND SUBSTRATE PREFERENCE: T. pelletieri is, with one

exception, a fimicolous species: 96 (173) finds, from dung of cow
54 (85), horse 36 (66), sheep (6), deer 1 (4) [including red deer 1
and fallow deer (3)], ass 2, dog (2), rabbit (2), burro (= donkey)
1, elephant 1, mule 1, european bison (1), goat (1), and once on
soil (Bucknall 1880). The species prefers cow dung, but is also
frequently found on one or two additional types of dung.
DISTRIBUTION: Austria, Azores, Belgium, BUlgaria (Fakirova 1974),
czechoslovakia, Denmark, England, France, Germany, Hungary
(BAnhegyi 1940, 1942 b, and BAnhegyi et al. 1985), Italy
(Spegazzini 1878, Vido'1879, and Bizzozero 1885), Lithuania
(Krejwis6wna 1936), Luxembourg (Feltgen 1899), Netherlands
(Oudemans 1900), Norway, Poland, Romania (Racovitza 1945, and
Bontea 1953), Spain, Sweden, Ukraine (Girzitska 1929), China,
India, Japan (Otani 1973), U.S.S.R., Australia, Canada, U.S.A.,
Bermuda, Dominican Republic, Jamaica, Mexico, Puerto Rico,
Argentina (Spegazzini 1880, and 1899), Brazil.

T. pelletieri is perhaps the most frequently collected species of

the genus, known from all the continents. It may be characterized
as a "cosmopolitan" species.
 144

ILLUSTRATIONS: Crouan & Crouan 1857 and 1867, pI. 4 A, figs. 1-4.
-- Boudier 1869, pI. 9:22, figs. 1-9. -- Phillips & Plowright 1881,
plo 158. -- Patouillard 1883, fig. 172 a-d. -- Phillips 1887, pI.
9, fig. 56, a-f. -- Massee 1895, fig. 38. -- Lindau 1896, fig. 152
g-j (after Boudier). -- Rehm 1896, p. 1082, figs. 1-4 (after
Boudier).• -- Seaver 1904 b, pI. 17, figs 1 a-e. -~ Seaver 1905, plo
33, fig. 1 a-e, the same fig. as in Seaver 1904 b. -- Overton 1906,
pls. 19 - 20. (Pl. 19, fig 1 also in Greis 1943: fig. 116 G). --
Dodge 1912, plo 10, fig. 1 a-b. -- Lindau 1912, fig. 354. -- Migula
1913, pI. 163, figs. 1-4. -- Seaver 1928, plo 12, fig.7. -- Snyder
1938, pI. 1, fig 1 b. -- Le Gal 1960, fig. 2 D. -- Kohlman-Adamska
1965, fig. 11, a-b. -- Dennis 1968, plo 8 M. -- Kar & Pal 1968,
fig. 1 g, j-k. -- Otani 1973, fig. 1, a-d. -- Waraitch 1977, figs.
8-~. -- Dennis 1978 & 1981, pI. 11 M. -- Engel & Svr~ek 1983, fig.
p. 59. -- Banhegyi et al. 1985, fig. 343 C, a-c. -- Schweiger 1985,
figs. 32-33. -- Ellis & Ellis 1988, pI. 36, fig. 361.
PHOTOS: Snyder 1938, pI. 1, fig. 1 a. -- Kimbrough 1966, fig. 2,
i & j. -- Kimbrough & Korf 1967, fig. 1, j. -- Kimbrough 1969,
figs. 1-9. -- Kirnbrough 1972, plo 19, fig. 43. -- Otani 1973, plo
1, b-c. -- Conway 1975, figs. 5-6, 12-13, 14-23. -- Samuelson 1978,
figs. 29-35. -- Engel & Svr~ek 1983, pI. 14, fig. 041. This photo
also referred to in Hohmeyer et al. (1989: 29). -- Kimbrough &
Curry 1985, figs. 8, 18 and 22.

Boudier (1869) based the genus Thecotheus on this species only. It

is the only polysporous species in the genus. Although normally
with 32 spores per ascus, the spore number varies. Asci with both
16 and 64 spores have been observed, even within the same fruit-
body.

Heimerl (1889) has by most of the later authors erroneously been

cited as the combinator of this species name into the genus
Ascophanus. However, the correct reference is that of Quelet
(1886). In the same way, Rehrn (1896) has been described to as the
 145

combinator of the species into the genus Rhyparobius [sic.] by a

few authors. However, the combination was first made by Saccardo
(Vido 1879) which well antedates that of Rehm.

Notes on type material

In the herbarium at Laboratoire de Biologie Marine du College de

France, Concarneau (CO), the material of Ascobolus pelletieri
consists of one envelope containing 4 sheets and three collections.
The statement of Le Gal (1960) that the envelope contains only two
collections is therefore not correct. Information about the two
latter are: "Croit sur les crottins de cheval 30 Juin 1858" and "
Sur crotin de cheval le 9.8bre 1866." The third collection which
contains loose apothecia is without text, but the text on the
envelope matches that in the original description (Crouan & Crouan
1857): "Crolt sur les crottins du cheval et les bouses de vache
anciennes dans les marais et les prairies aux environs de Brest.
Printemps, automne. Rare." Identical, rather rich material of this
latter collection was also found in RO. The envelope contains glued
apothecia.

This third collection very probably represents the type material of

Ascobolus pelletieri Crouan. It is not possible to decide whether
the substrate is cow dung or horse dung, nor the year and season of
the collection. It is not possible to designate a holotype based
upon Le Gal's (1960) observations, as done by Kimbrough (1969).

According to van Brumrnelen (1967) a part of the type material is

also deposited in herbarium pc. This particular material has been
selected as lectotype by van Brummelen (op. cit.), accompanied by
the following information: ~Crouan, on cow dung in a marsh, near
Brest, Finistere, France, 20.VI.1857 (lectotype, PC-A2353)." I have
 146

not seen this particular collection since it could not be found at

herbarium PC (in litt. from Museum National d'Histoire Naturelle,
laboratoire de cryptogamie, Paris, dated 4. November 1985).
However, in this case I follow van Brummelen in his choice of
lectotype.

Figs. 72 - 80. Thecotheus pelletieri. 72 - 73: Norway. Olsen (BG).

72: Young, still closed, sUbglobose to doliiform living apothecia.
73: Mature, doliiform, living apothecium with strongly protruding
asci. 74 - 75: Czechoslovakia. Svr~ek & Kubi~ka (PRM 568504). 74:
Old, overripe apothecium with discoid form. 75: Dry, young, unripe,
cylindrical apothecium. 76 - 77: Norway. Aas 68/80 (BG). 76: Dome-
shaped tip of ascus with a faint ring in Congo red mounting,
indicating the zone where the operculum breaks off. 77: Ascospores
surrounded by thick,' swollen gelatinous perispore. 78: Azores.
Dennis (K). 'Broad,_ clavate ascus with 32 ascospores. 79: Germany.
Jahn (BG). Asci from one and the same apothecium. The one contains
32 spores, each spore about 25 x 15 ~m,'which is somewhat smaller
than in normally 32 spored asci. The other ascus contains ~omewhat

greater 16 ascospores, the spores up to 40 x 16 ~m. The left ascus

with a characteristic indentation at the operculum. 80: Azores.
Dennis (K). Upper part (from outside) of an apothecium showing
paraphyses. Scales: 72 - 73: 2 mm. 74: 1.3 mm. 75: 0.7 mm. 76 &
78 ~ 79: 60 ~m. 77: 35 ~m. 80: 40 ~m. Photos: 72 - 78 & 80: O. Aas.
79: E. Jahn.
147
 148

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'\l
ai:-
 .4.

12. THECOTHEUS PERPLEXANS (FaUrel , 8cbotter) ~rug , Khan

Mycologia 79 (2): 200 (1987). -- Basionyrn: Ascophanopsis perplexans

Faurel & Schotter, Cah. La Maboke 3 (2): 130 (1965 b). -- Type:
Congo, Vallee de la Loamba, on dung of elephant, 20.VII.1956 L.
Gauthier, not known to exist; (Faurel & Schotter ibid., fig 3 a -
c, lectotype selected here).

ETYMOLOGY: Latin perplexu9, perplexing.

 150

The following description of the species is translated from the

original paper of Faurel & Schotter (1965 b).

Receptacle sparse, superficial, sessile, whitish, fleshy,

transparent, glabrous, at first globulose or doliiform with a
concave discoid outline, marginate, then flattened with slightly
convex disc, without margin, slightly roughened by the protruding
ascus tips, 0.5 - 1 mm diameter. Hymenium white, about 200 ~m

thick; asci narrow, cylindrical, 160 - 200 x 10 - 12 ~m, tapering

towards the base, p. sp. about 100 ~m long. Spores oblique
uniseriate, ellipsoid, hyaline, smooth, thin-walled, ornarnented at
both poles with a subspherical to ± ovoid apical wart, at the base
surrounded with a cylindrical collarettej size of the spore body 20
- 22 x 10 - 12 ~mj size of the cylindrical collarette 4 - 5 ~m in
diameter and about 2 ~m in lengthj size of the apical warts 4 - 6
~m in diameter and 4 - 5 x 6 - 7 ~m in length.

UNVERIFIED RECORDS: CONGO: All records in Faurel & Schotter (1965

b: 130), elephant dung from Vall~e de la Loamba, 20.VII.1956 L.
Gauthier. -- Piste de Divenie, elephant dung, 4.IX.1956 L.
Gauthier. -- Region de Mouyanzi, ten collections on elephant dung,
autumn 1956 L. Gauthier.

HABITAT AND SUBSTRATE PREFERENCE: I. perplexans .is exclusively a

fimicoious species, with all records on dung of elephant. The
vegetation is dominated by tropical rain forest and savanna. I
regard the species as vulnerable.
DISTRIBUTION: All known records of the species are tram the
equatorial African republic Congo. Its distribution is
~pproximately limited between 5°N and 5°S r and constitutes a
tropical equatorial distribution pattern. I regard the species to
be endemic to the Palaeotropics.
ILLUSTRATIONS: Faurel & Schotter 1965 h, fig. 3 a-c as
Ascophanopsis perplexans.
 '"
This species has never been re-examined. It is the type of the
genus Ascophanopsis Faurel & Schotter, a genus erected on the basis
of the cylindrical collarettes at the apiculus base. Kart (1973)
considered Ascophanopsis as a synonym of Thecotheus, and he was
later followed by Krug & Khan (1987). The authors did not regard
the collarettes as a generic character. I. africanus also possesses
the same kind of collarettes.

According to Faurel & Schotter (1965 b), all collections referred

to are deposited in Faculte des Sciences d'Alger (AL). However, no
material of A. perplexans could be found in this herbarium, nor at
PC in Paris. The collections of the species therefore, appear to be
lost or mislaid. It is well known that material and collections of
L. Faurel, said to be placed in Paris (PC), are difficult to trace
(Lundqvist 1980). In accordance with Article 7.5 in the IeBN
(1988), Faurel & Schotter's original illustration is selected a
lectotype of Thecotheus perplexans.

As for T. africanus, I have chosen to regard Ascophanopsis

perplexans as a member of Thecotheus. Fresh material is needed to
critically examine its status and characters. Nothing is known
about the excipular structure, paraphyses and ascospore surface.
Both species are certainly closely related, and can be
distinguished by the size of the ascospores. However, within a
species, ascospore size is generally variable, dependent .on the
cegree of maturity. At this ·moment is impossible to estimate the
normal range of spore variation in T. perplexans.
 152

13. THECOTHEUS PHYCOPHILUS Pfister (Figs. 81 - 92)

Mycologia 73: 1001 (1981). -- Type: U.S.A., Minnesota, Lake

. Itasca, University of Minnesota Field 'Station, on dead grasses
and herbaceous plants overrun by algae, on low spot near
campus, D. H. Pfister, ~. L. 'Pfister and M. Palm 1.VIII.1980
Mn 96 (FH, holotype).

ETYMOLOGY: Greek lyAKOE, sea-weed; latin -philus, friend of;

referring to the habitat.

Apothecia solitary, rarely a few in groups, superficial,

sessile on a broad base, up to 5 mm diameter when fresh, when
dry up to 2.5 mm diameter and up to 0.65 mm high; at first
turbinate, then expanding and cupulate, broadly attached when
fUlly mature, greyish brown (9D3). Disc flat to ± convex,
undulate and somewhat rough in larger specimens. After drying
with flat to concave disc. When fresh cream to light brown
finally when dry with a slightly reddish tint (8D3 - 8E4).
Marqin broad, even to somewhat wavy when fresh, conspicuously
uneven, waved and slightly incurved after drying. Hymenium up
to 360 ~m thick.

Medullary excipulum 80 - 145 ~m thick at the central base,

consisting of different kinds of cells. Along the walls and
toward the outer boundary more or less parallel, thin-walled,
sparingly septate hyphae with diameter 2 -3 ~m composing a
textura porrecta. The hyphae terminate in ellipsoid to
subglobose terminal cells on the outside of the ectal
excipulum. Towards margin the hyphae run parallel to the
hyrnenium, terminating in narrow clavate terminal cells which
gradually change to paraphysis-like cells. The innermost part
of the medullary excipulum, especially at the central base,
 153

consists of thin-walled, septate, mostly narrow and sparingly

branched hyphae 2 - 3 ~m diameter, together with predominant,
much wider ascogenous hyphae which are intermingled with both
± subglobose and polyhedral cells varying from 3 - 15 (-20) ~m

diameter. Ectal excipulum 90 - 140 ~m thick at the 'base of

the apothecia, about 20 ~m thick at the margin. At the central
base this layer consists of globose to more or less
polyhedral, almost isodiametric cells, composing a textura
globulosa to textura angularis, with cell-diameter varying
from 9.5 - 20.5 (-25) ~m, intermingled with smaller cells. The
largest cells of the flesh, which are found at the innermost
part are ± elongated, the outermost ones mainly globose. In
the lower central base the outermost subglobosecells on the
outside give rise to thin-walled, hyaline, narrow' branched or
unbranched and sparingly septate hyphae 1.5 - 2 (-3) ~m

diameter, which bind the apothecia to the substrate. These

hyphae are easily observed macroscopically. Towards the margin
bulbous to more or less clavate, deeply rooted hyphoid cells
are predominant, with diameter of the terminal cells from 5 -
10 ~m.

Asci in most cases with 4 fully developed ascospores, although

the number of ascospores in the same fruit-body varies from
three, five, six or eight, and often with accompanying smaller
abortive ascospores. Asci broadly cylindrical, gradually
narrowing towards the base, 250 - 320 x 30 - 35 ~m; apex broad
and often conspicuously truncate with a distinct ring when
mounted in Congo Red in ammonia. Only when mature do the asci
reach, or slightly extend above the level of the paraphyses.
Ascospores irregularly obliquely uniseriate, in exceptional
cases biseriate, ellipsoid to broadly ellipsoid, smooth, 30 -
37 x 15 - 16.5 (-18) ~m, each surrounded with a gelatinous
envelope which is not visible on ascospores outside the asci.
Young ascospores with extremely thick wall. Paraphyses
filiform, septate, thin-walled, apically branched, below 1.5 -
2 ~m diameter. Apices occasionally uncinate and only slightly
inflated, 2 - 4.5 ~m diameter, this upper part partly
 1.4

encrusted on the outside with subhyaline to light brown-yellow

amorphous pigments. Between the paraphyses are interascal,
paraphyses-like elements, divided into thick-walled
compartments with brownish content, enlarged at apex from 4 to
8 ~rn, below from 2.5 - 4 ~rn diameter.

Specimens examined:

V.B.A. Minne90ta: Lake Itasca: Holotype (see above). --

Ditto, on wet debris with Pachyella babingtonii, D. H. Pfister
9.VIII.1980 MD 131 (FH - paratype); in Pfister (1981: 1003).

HABITAT AND SUBSTRATE PREFERENCE: on dead grasses overrun by

algae, on herbaceous plants and other vegetable debris. Both
collections are associated with low moist ground and the
occurrence of algae, a o. Oscillatoria sp. and coccoid green
algae. Sedges, Eguisetum sp. and pilea sp. form the herbaceous
ground cover in the type locality.
DISTRIBUTION: The species is only known from the type
locality in Minnesota, U.S.A. It may be an overlooked species
owing to its special substrate.
ILLUSTRATIONS: Pfister 1981, fig. 1.

Thecotheus phycophilus and T. pallens are the only species of

Thecotheus growing on soil and vegetable debris. Both species
are also associated with the occurrence of algae and with
periodically inundated areas. The main distinguiShing"
characters between the two species are the form, size and
number of the ascospores, and the colour of the apothecia and
paraphyses. T. pallens has oblong fusiform ascospores, whereas
those of T. phycophilus are smaller and ellipsoid to broadly
ellipsoid. The latter species is also characterized by usually
 155

4-spored asci and paraphyses of two types.

On the other hand, the form and size of ascospores of T.

phycophilus are rather similar to those of the regularly 8-
spared T. cinereus, Which also possesses paraphyses of two
kinds. However, the species are easily distinguished. Among
other things, in i. phycophilus the medullary excipulum is
dominated by narrow, 2 - 3 ~m wide hyphae, composing a textura
porrecta, whereas the corresponding layer in T. cinereus is
composed of a textura globulosa. The ± distinct, ring-formed
zone in the ascus apex when mounted in Congo Red in ammonia is
characteristic for T. phycophilus and T. pelletieri. I have
never observed this in T. cinereus.

Typically, in T. phycophilus and T. rivicola, which have the

largest apothecia in the genus, the ectal excipulum is "multi-
cellular". This in contrast to the smaller species where the
excipulum consists of fewer cell layers.

Figs. 81 - 92. Thecotheus phycophilus. 81 & 83 - 92: Holotype

(FH). 82: Paratype (FH). 81: Clusters of apothecia showing the
characteristically hyaline hyphae which bind the apothecia to
the substrate. 82: Close up of an apothecium showing the
binding hyphae. 83 - 85: Asci with different numbers of
developed ascospores. 86: Section of apothecium. 87: Section
of ectal excipulum from the lower, central part. 88 - 89:
Mature ascospores. 90: Ascus tips mounted· in Congo Red in
ammonia, each with a distinct ringformed zone. 91: .Ascospores.
The smallest, immature spores with very thick walls. 92: SEM
of a part of an ascospore showing smooth surface. Scales: 81 -
82: 1 mm. 88 - 89: 32 ~m. 90: 25 ~m. 92: 5 ~m. Photos: 81 - 82
& 86 - 89 & 91: O. Aas. 83 - 85 & 90 & 92: J. Berge.
156
157
159
 160

14. THECOTHEUS RIVICOLA (Vacek) Kimbr. , Pfister

(Figs. 93 - 105)

Bull. Torrey Bot. Club. 99 (4): 199 (1972). -- Basionyrn-:

Psilopezia rivicola Vacek, stud. Bot. Cech. 10 (4): 129
(1949). -- ~ype: Czechoslovakia, Bohemia, Vonoklasy, on
sUbmerged, decorticate twigs of Prunus spin6sa, 17.X.1948 V.
Vacek (CUP 52287, holotype).

ETYMOLOGY: Latin rFvulus, brooklet, rivulet, - cola,

dweller; referring to the water-soaked habitat.

Apothecia solitary or scattered to gregarious, superficial, at

first centrally seated, then apparently sessile and broadly
appressed to the substrate, 1 - 7 mm diameter and up to 2 mm
high when fresh, up to 4 mm diameter and 0.3 - 0.7 mm high
when dry. The apothecia are easy to remove from the substrate;
soft, fleshy, at first pulvinate to somewhat lenticular, round
in outline, Qecoming cupulate to discoid on a broad base,
concolorous with the disc. External surface, also inclUding
disc, at first smooth, becoming white furfuraceous,
occasionally dark brown roughened. Disc at first convex, in
dried condition flat to somewhat concave, especially the
central part, at first whitish to dirty white, becoming light
to medium grey or yellow-brown (5E4) to grey-brown (6E3-5E3),
when dry mostly brownish black with a red or purple tint,
rarely dark grey. Disc somewhat rough and furfuraceous,
occasionally with low bumps. Usually with greyish yellow
colour (4B5) when rehydrated in water. Harqin apparently
indistinct when young, in dried condition lobed or occasio-
nally overhanging, up to 120 ~m broad. Hypothecium composed of
a more or less dense textura intricata with scattered
interspersed cells 5-10 ~m diameter. Towards flesh with small
 161

globose cells, mostly 2-3 (-5) ~m diameter.

Medullary excipulum up to 250 ~m thick at the central base,

inner part composed of a textura globulosa, with small,
sUbhyaline, globose to sUbqlobose cells, intermixed with
filamentous, sUbhyaline, slightly thick-walled hyphae, 2 - 3
~m diameter. Towards ectal excipulum the filamentous cells"
form a textura porrecta, with septate, parallel-lying hyphae,
2 - 3 (-4) ~m wide, terminating in the hyrnenium layer with
more or less clavate, up to 5-6 ~m wide terminal cells. Ectal
excipulum up to 100 (-230) ~m thick at the central base,
consisting of many cell layers, composing a textura angularis,
with ye"llowish to yellow-brown, slightly thick-walled and
elongated cells, 15 - 30 (-40) x 7 - 12 (-25) ~m, their
longitudinal axes perpendicular to the outside of the
apothecium, interspersed with scattered, ± globose cells a -
12 (-35) ~m in diameter. Usually greatly reduced cell-size
along the lateral walls. Towards the exterior part with long-
celled rooting hyphae originating several cell-layers deep,
2 - 4 ~m wide below, their dark-walled terminal cells
inflated, 6 - 15 ~m diameter, with up to 25 ~m long terminal
part. This cell layer appears as a border composing a textura
globulosa as seen from the outside. More elongated, terminal
swellings of the rooting hyphae along the lateral walls.

Asci a-spored, cylindrical, usually slightly truncate at apex,

attenuate below, (120-) 150 - 300 x 12 - 17 (-20) ~m,
distinctly to diffusely 1+ blue. Mature asci only reach, or
slightly extend beyond the level of the paraphyses. Ascospores
thick-walled when young, ellipsoid, slightly inequilateral,
uniseriate to obliquely uniseriate, (15-) 16 - 22.5 x 7 - 9 (-
10) ~m; outer wall staining in Cotton blue, with very small,
inconspicuous irregular warts. At both spore ends a ± globular
to somewhat elongated apiculus, 2 - 5 ~m broad, 3 (-5) ~m

long, not always observed in water-mounted slides. Both warts

and apiculi stain deep blue in Cotton blue. Each ascospore
surrounded by a gelatinous sheet which does not stain in
 162

Cotton blue. Paraphyses filiform, septate, of two types; the

first one numerous, crowded, thin-walled, hyaline below, 1.2 -
2 ~m wide; apices not or only very slightly inflated, this
part occasionally slightly knotty to ramified with small
inconspicous ± yellowish granules, externally minutely
encrusted, and embedded in a gelatinous, yellowish to yellow-
brown matrix. The other type always unbranched, slightly
thick-walled, with scattered septa, upper 50 - 60 ~m aseptate,
3 - 6.5 ~m wide.

Specimens examined:

CZECHOSLOVAKIA. Bohemia: Vonoklasy. Type collection (see

above) .
ENGLAND. Hereford: Peterstow, water-soaked twigs of Alnus in
stream bed, 24.VIII.1976 W. D. Graddon 2907 (K); in Graddon
(1979: 183), Clark (1980: 52). near Worcester: Upton -
on - Severn, bare soil, 8.VIII.1982 M. C. Clark 3071 as T. cf.
apiculatus (K). -- Worcester: near Droitwich: Dunhampstead,
bare soil on a damp, clayey path in woodland, 27.VIII.1982 M.
C. Clark 3077 as T. cf. apiculatus (K). -- Norfolk: Honingham
Fen, on vegetable debris on damp soil on the ground,
6.VIII.1979 M. C. Clark (K); in Clark (1980: 52) as T.
apiculatus Kimbr.
GERMANY. Schlesvig-Holstein: Scharbeutz, MTB 1930/4, on a
water-soaked twig of ? Alnus (or? Fraxinus), also together
with Miladina lechithina, 14.VIII.1988 P. Steindl (private
herbarium of E. Jahn). New to Germany.
u. s. A. Michigan: 66 at the Missaukee-Kalkasa co. line, on
decorticated log in running water, 20.VIII.1969 N. J. smith
2279 as "Psilopezia" (FH - ex herb. Discomycetes D. H. Pfister
354 as T. rivicola. -- New York: Lloyd Cornell Preserve:
 163

McLean, on rotten wood, 11.X.1969 D. H. Pfister 324 (FH).

ECUADOR. Prove Pichincha: about 36 km from Quito: on Quito-
Tandayapa Road, elev. about 8.000 ft., on indet. monocot.
stem, 6.VIII.1975 K. P. Dumont, S. E. Carpenter, and P.
Buritica, No. 2307 Dumont-EC (NY). New to South-America.

UNVERIFIED RECORDS: ENGLAND: Hereford: Peterstow, on water-

soaked Alnus wood, W. D. Graddon 3130; in Graddon (1979: 183).
-- U.S.A.: New York: Lloyd Cornell Preserve, McLean, s. subs-
trate, 2.X.1953 R. A. Shoemaker, R. L. Shaffer & R. P. Korf
(R.P.K. 53-227 and 53-229); in Pfister (1972: 200).

HABITAT AND SUBSTRATE PREFERENCE: Most specimens inhabiting

twigs and wood in water 6 (1), more rarely on bare soil or on
vegetable debris on damp soil 3 (see however comment below).
In one collection the substrate is unknown.
DISTRIBUTION: So far the species is known from
Czechoslovakia, England, Germany, U.S.A., and Ecuador. The
distribution may indicate an European-American distribution
pattern.
ILLUSTRATIONS: Graddon 1~79, fig ... 6 a-b. -- Vacek 1949, fig.
1, 1a-1c (as Psilopezia rivicola Vacek). -- Ellis & Ellis
1985, plo 36, fig. 352.

Thecotheus rivicola is characterized by the apiculate,

o~namented ascospores, paraphyses of two types and its
occurrence on water-soaked wood. The coprophilous species T.
biocellatus has ascospores of about same size and form.
The reason why data about young specimens are rather scanty,
is because no fresh material of T. rivicola has been
 164

available to me

with some hesitation, the collection of smith from Michigan is

inciuded in T. rivicola. Its apothecia are dark grey, without
any element of brown-black or sometimes a purple tint
typically found in the other fruit-bodies. Enclosed with the
collection is an annotation by N. J. smith that the apothecia
are dull smokey grey [? = yellowish grey (3C2)] over all.
Moreover, the asci only reach up to 170 ~m in length, the
ascospores are slightly shorter than, but as broad as, mature
ascospores, ± smooth and mostly non-apiculate. It may
represent an atypical, immature specimen, as the ascospores of
T. rivicola are smooth and non-apiculate when young.

T. rivicola was at first placed in the genus Psilopezia (Vacek

1949). The author originally described it (as £. rivularis in
herb.) with non-amyloid asci and smooth ascospores. Later,
Pfister (1972) transferred the species to Thecotheus and
provided a modified description. Among other things, the
ascospores were found to be apiculate with inconspicuous,
irregular warts. Moreover, the asci were diffusely I+ for most
of their length. I found the same characters as Pfister.
Contrary to earlier descriptions, there are, however, two
kinds of paraphyses, which are easily observed in all examined
material, including the type collection.

The form of the ascospores varies greatly in the type

material, ranging between narrow ellipsoid to ± broadly
ellipsoid. Even within the same ascus the ascospore form is
obvious variable. This is the main reason why three gatherings
from England, all apparently on soil, are tentatively
classified as T. rivicola. I cannot find any convincing,
morphological character separating these specimens from the
wood-inhabiting ones. Further investigations are needed on
this sUbject.
 165

Notes on type material

No other material of T. rivicola than that in herbarium CUP

No. 52287 has been traced. It contains a sheet of paper
annotated "Specimen No. Pragae 703772", and PRM has not
replied to a request if there still might be material left of
Vacek's original collection. I therefore for the moment
believe that this is the only specimen in existance, and
regard it as the holotype.

Figs. 93 - 105. Thecotheus rivicola. 93 - 95 & 101: Holotype

(CUP). 96 - 100 & 102: England. Clark 3071 (K). 103 - 104:
England. Clark 3077 (K). 105: U.S.A. smith 2279 (FH). 93: Part
of an apothecium attached to the wood. 94: Apothecium as seen
from above. Note the concave sentral part. 95: Cross section
of an apothecium with hymenium to the left, showing
overhanging margin layer and a relatively broad central base
with binding hyphae. 96: Cross section of apothecium, showing
medullary excipulum (upper part) and ectal excipulum (lower
part). 97: Medullary excipulum, showing ± globose cells of
varying size, interspersed with filamentous hyphae. 98: Ectal
excipulum, showing polyhedral, elongated cells, their
longitudinal axes perpendicular to the outside of the
apothecium. 99: Specimen in Cotton blue in lactophenol,
showing apiculate ascospores and thick-walled paraphyses (at
arrows). 100: SEM-photo of ascospores showing bipolar apiculus
and walls with irregular warts. 101: SEM-photo showing an
ascospore and empty asci. 102 - 104: SEM-photo of ascospores
showing bipolar apiculus and ornamentations of small warts.
105: SEM-photo of ascospores. Note the very fine roughness on
the surfaces. Scales: 93: 0.5 mm. 94 - 95: 1 mm. 96: 100 ~m.
97 & 99: 50 ~m. 98: 25 ~m. 100 - 103 & 105: 5 ~m. 104: 0.5 ~m.
Photos: 93 - 99: O. Aas. 100 - 105: J. Berge.
167
168
169
 170

15. THECOTHEUS STRANGULATUB (Vel.) Aas , Lundq. n. comb.

(Figs. 106 - 108)

Baslonym: Ascophanus strangulatus Vel., Monogr. Discorn.

Bohemiae. I: 358 (1934). -- Type: Czechoslovakia, Bohemia,
Mnichovice, Bozkov, on old cow dung, 20.X.1928 J. Velenovsky
(PRM 150368, holotype).

ETYMOLOGY: Latin stranqulatus, throttled, i.e. narrowed and

then widened again; referring to the shape of apices of living
asci.
 171

Apothecia solitary, sessile or partly immersed, 0.5 - 2 mm

diameter when fresh, up to 0.9 mm diameter and 0.5 mm high
when dry; glabrous, at first subglobular, then expanding and
becoming somewhat cup-shaped and with a rather broad, central,
immersed stalk. Upper part concolorous with the disc, lower
part with brownish yellow colour (5C6). Disc concave,
slightly rough, greyish yellow (4B5-4B6). Marqin slightly
incurved, when dry yellow brown (5E7). Hymenium up to 200 ~m

thick at the central base. Hypothecium composed of hyphoid to

epidermoid cells intermingled with small globose cells up to 5
~m in diameter.

Medullary excipulum basically composed of same kind of cells

as within, but those in the flesh layer are slightly larger,
up to 8 ~m diameter together with scattered polyhedral cells,
their long axes up to 15 - 20 ~m diameter. Towards excipulum a
layer of hyaline, long-celled, parallel-lying thin-walled
cells composing a textura porrecta. This layer extends along
the sides and terminates in ± inflated, clavate cells. The
cells in this part slightly thick-walled, encrusted with
yellowish to yellowish brown pigments. Terminal cells from 4
to 9 ~m diameter. Ectal excipulum of varying thickness, up to
130 ~m thick at the central base, up to 25 ~m thick near
margin, consisting of sUbglobose cells (textura globulosa),
intermingled with slightly polyhedral, isodiametric cells
(textura angularis). Both kind of cells up to 15 ~m diameter.
Terminal cells along the sides and at the base of the
apothecia are thick-walled, light yellowish encrusted, 16 - 20
x 10 ~m diameter, their longitudinal axes perpendicular to the
surface.

Asci 8-spored, cylindrical to cylindrical-clavate, upper part

strangulate (not always seen in dry material), rounded above,
narrowing and bent towards the base, bilobed below, (80-)
120 -150 (-175) x 20 - 28 ~m, p. sp. up to 110 ~m. Ascospores
hyaline, ellipsoid, biseriate, occasionally irregularly
obliquely uniseriate, (20-) 22 - 26 (-28.5) x 11.5 - 16 ~rn,
 172

ornamented with a regular pattern of rounded warts 0.3 - 1.9

~m wide and 0.2 - 1.2 ~m high, non-apiculate. outer wall layer
(up to 0.5 ~m thick) staining in Cotton blue. Each ascospore
surrounded with a thick gelatinous envelope. Paraphyses
filiform, septate, branched or not, hyaline, 1.8 - 2 ~m

diameter below; apices only slightly dilated, 2.2 - 2~5 (-3)

~m diameter, strongly uncinate.

Specimens examined:

CZECHOSLOVAKIA. Bohemia centr.: Mnichovice: Holotype (see

above) .

HABITAT AND SUBSTRATE PREFERENCE: Only known on old cow

dung.
DISTRIBUTION: Only known from the type locality in
Czechoslovakia.
ILLUSTRATIONS: Velenovsky 1934, tab. 3 fig. 12.

As noted by Svr~ek (1979) the present type material of

Ascophanus strangulatus consists of old dung blackened on its
surface and covered in algae. I found three badly preserved
apothecia on different small dung fragments. The examined
apothecia vary from 0.5 to 0.9 mm diameter, but Velenovsky
(1934) described the apothecia (fresh ?) to be up to 2 mm in
diameter. Characteristically the colour of apothecia of this
specimen is "yellowish green". This is also true for T.
uncinatus.

Velenovsky did not say anything about ascospore ornamentation,

but contrary to Svr~ek (1979), who described the ascospores as
smooth, the examined apothecia possess ascospores ornamented
with regular rounded warts. No apiculi were observed.
 173

strangulation of the asci was observed between the second and

the third ascospore in the upper part of the asci. As also
figured by Velenovsky (op. cit.) the apex of the asci contains
a dark, sUbglobose structure.

Svr~ek (1979) says that Ascophanus strangulatus probably

belongs to the genus Thecotheus, but that the present type
material is insufficient for more exact taxonomic conclusions.

T. strangulatus is closely related to T. uncinatus. It can be

easily distinguished from the latter by the size of the
ascospores, and the epispore ornamentation. These are the only
species of Thecotheus which have strongly curved paraphyses.
owing to conformity of asci, paraphyses, ascospores and colour
of apothecia, both species would deserve establishment within
a separate section of Thecotheus.

Figs. 106 - 108. Thecotheus strangulatus. 106: Section of

apothecium. 107 - 108: Ascospores with ornaments of
conspicuous rounded warts. 107: Spores seen in interference-
phasecontrast. 108: Spores as seen in Hellfeldt phasecontrast.
Scale 107: 25 ~m. Photos: 106: O. Aas. 107 - 108: J. Berge.
17'
 175

16. THECOTHEUS UNCINATUS Aas sp. novo (Figs. 109 - 117)

ETYMOLOGY: Latin uncinatus, hooked; its paraphyses are

shaped like a hook.

Apothecia solitaria vel subdistans, sessilia, 0.2 - 1 mm

diam.; primo cylindricum, maturitate turbinatum vel
discoideum, initio album, deinde cinereo-luteum vel
luteofuscum. Asci cylindrici vel cylindrico-clavati, strangu-
lati, 115 - 160 x 10 - 15 ~m, pariete iodo caerulescenti.
Ascosporae ellipsoideae, obliquae uniseriatae vel interdum
irregulariter dispositae, (13-) 14 - 17 x (6-) 7 - 9 (-9.5)
~m, sine apiculo, minitue granulosae vel verrucatae.
Paraphyses filiformes, septatae, apicibus paulo dilatatis, 2 -
3 ~m latis.
In fimo equino crescens.
Typus: Sweden, Dalarna, Garpenberg, N of Krommetsbo, on old
horse dung in spruce forest, 31.VIII. 1973 N. Lundqvist 8671
(UPS, holotype).

Apothecia solitary or crowded, superficial but with a slightly

immersed central part, 0.2 - 1 mm diameter, up to 0.4 mm high;
glabrous to slightly rough, at first more or less cylindrical,
later expanding and ± turbinate, finally almost discoid and
depressed to the substrate. At first white, becoming greyish
yellow (4C5) when dry, lower part with a light brownish tint.
Disc at first flat to convex, then concave, when over-ripe
occasionally again with convex disc, slightly rough owing the
protruding tips of asci. Disc colour of dry, mature apothecia
occasionally turning medium grey or greyish with a reddish
brown tint. Margin entire.
 176

Medullary excipulum composed of globose to subglobose cells

(textura globulosa), the cell diameter varying from 6 to 15
~m. Towards excipulum a layer of long, parallel-lying cells, 2
- 4 ~m diameter, composing a textura porrecta. This layer of
thin-walled, hyaline.cells terminate in bulbous cells on the
margin and on the outside of the lateral walls of the
apothecia. Ectal excipulum mainly composed of polyhedral,
elongated and slightly thick-walled cells (textura angular is),
5 - 7 x 12 - 15 ~m diameter, with their longitudinal axes
perpendicular to the surface. Rooting hyphae subglobose to
elongated, slightly thick-walled, mostly 5 - 6 ~m diameter.
These bulbous cells originate several cell-layers deep.

Asci 8-spored, cylindrical to cylindrical-clavate, more or

less rounded at apex, upper part strangulated, lower part
narrowing, 115 - 160 x 10 - 15 ~m, p. sp. mostly 80 - 90 ~m.

When fresh the 1+ blue colour of the asci occasionally soon

disappears, but returns after some days. Ascospores hyaline,
ellipsoid, thick-walled, obliquely uniseriate, rarely
irregularly biseriate, (13-) 14 - 17 x (6-) 7 - 9 (-9.5) ~m, x
= 15.68 x 8.27 ~m (n = 77), Q = 1.00 (length), 0.82 (width),
non-apiculate, verruculose, slightly larger warts towards the
poles. Each ascospore totally or partly surrounded with a
thick gelatinous envelope. Paraphyses, filiform, septate,
branched or not, hyaline below, 1.2 - 2 ~m diameter, with
apices 2 - 3 ~m wide, characteristically uncinate, with light
yellowish granulated contents. The paraphyses extend up to 20
~m beyond the asci at maturity.

Specimens examined:

SWEDEN. Da1arna: Garpenberg: Type collection (see above). --

 177

Jamtland: Brunflo: Ope, horse dung in pasture, (me after 16

days), 16.VI. 1962 N. Lundqvist 3420 a (UPS). -- Frostviken:
Sjuls~sen, old horse dung in coniferous forest, (mc after 3
weeks), 17.VI.1962 N. Lundqvist 3438 b (UPS).
CHINA. Szechwan: Jiu Zhai Gou: Yuanshi Senlin, deer dung,
probably of Cervus elaphus, in mossy, mainly coniferous
forest, alto 3300 m., (me), 21.V.1989 T.-B. Engelmark (S).

HABITAT AND SUBSTRATE PREFERENCE: T. uncinatus is a

fimicolous species, preferring old horse dung. In most cases
the species is collected in coniferous forests.
DISTRIBUTION: Sweden and China. So far the species exhibits
a European-Asian disjunct distribution pattern.

This species is related to T. strangulatus, from which it can

be distinguished by the smaller ascospores, and the
verruculose spore ornamentation. In T. strangulatus the
ascospores are verrucose. Both species are characterized by
the "yellow-green", i. e. greyish yellow (4B5-4B6) colour of
dry apothecia, the strangulated upper part of the asci, and
the strongly uncinate paraphyses.

The spore size of T. uncinatus is close to T. himalayensis.

However the latter has a slightly different spore form, in
addition to paraphyses of two kinds.
 118

Figs. 109 - 117. Thecotheus uncinatus. 109: Holotype (UPS).

110 - 114: china. Engelrnark (S). 115 - 117: Sweden. Lundqvist
]438 b (UPS). 109 - 110: SEM of ascospores. 111: Asci with
ascospores. 112 - 113: Empty asci showing operculum and
strangulation. 114·- 116~ Asci, ascospores and paraphyses.
Note the strangulated asci and the uncinate paraphyses. 117:
Paraphyses. Scales: 109 - 110': 5 Jlrn. III & 116 - 117: 20 jlrn.
112 - 114: 25 pm. 115: 60 Jlrn. All photos: J. serge.
179
 180

.~---
...., " .
;!
' I
~

,.

, f
"
"
~" .~

'-,. ~ .
115
 181

17. THECOTHEUS VIRIDESCENS E. LUdwiq

Thecotheus viridescens E. Ludwig in Hohmeyer et al., Hoppea,

Denkschr. Regensb. Bot. Ges. 47: 27 (1989). -- Type: Germany,
Bayern, Nationalpark Berchtesgaden, near Schapbach, MTB
8443/2, 1000 m, on game dung, 12.VIII.1982 E. Ludwig (B,
holotype). Not seen.

The type specimen has not been available for study, but the.
original description of the species runs:

"Apothezien klein, bis maximal 1 mm breit, jung

tonnchenformig, dann kreiselformig, umgekehrt flach
kegelformig, zentral angeheftet bis sitzend; Hymenium jung
etwas vertieft, dann flach, schliesslich emporgewolbt, rauh,
anfangs hellgrau, dann graugrlinlich, sehr hell olivgrau; Rand
mit einer hervorstehenden flockigen Zahnelung; Aussenseite
glatt, gleichfarbig.
Hymenium 110-140 ~m dick; SUbhymenium bis 30 ~m dick,
bestehend aus einer regellosen textura intricata; medullares
Excipulum bis 25 ~m dick, bestehend aus einer parallel zur
Aussenseite hin orientierten textura intricata, deren Elemente
urn 2 ~m breit sind; zum Rand hin ausdlinnend; ektales Excipulum
30-60 ~m dick, aus textura angularis, deren Elemente 8-20 ~m

breit sind, gegen den Rand hin mehr und mehr in textura
porrecta libergehend. Die Zahnelung des Apothezienrandes wird
aus langgestreckt-haarartigen, dlinnwandigen, farblosen,
agglutinisierten, 3-5 ~m breiten Strukturen gebildet, die liber
den Rand hinausreichen. Asci operculat, 11-15 ~m breit,
ganzlich amyloid, pleurorhynch, vier- oder achtsporig, vor dem
Abschleudern der Sporen 10-20 ~rn liber das Hymenium
hinausragend. Paraphysen einfach, 3-4 ~rn breit, and der Spitze
leicht keulig auf 4-5 ~m erweitert. Ascosporen schmal
ellipsoid bis schmal subfusiforrn, rnanchrnal leicht asymetrisch
(16-)17-20 x (6-)6.5-7.5(-8) ~rn, ohne Oltropfen; jung relativ
 182

dickwandig, reif eher dlinnwandig; das Ornament ist sehr fein

verlangert warzig bis sehr fein reticulat an den Polen
verdichtet bis pseudoapiculat".

ILLUSTRATION: Hohmeyer et al. 1989, figs. 11-12.

Thecotheus viridescens is characterized by the form and

ornamentation of the ascospores and the colour of the
apothecia. T. himalayensis possesses approximately the same
kind of ascospore ornamentation, but otherwise has larger
ascospores and i. a. quite different paraphyses. T.
biocellatus and T. rivicola have ornamented ascospores of
about the same size as those of T. viridescens, but they are
apiculate. A varying four- to eight ascospore number is also
characteristic for T. phycophilus.
 183

APPDfD:ICES

A. UNCERTAIN NAMES

ASeOPHANUS ISABELLINUS elements

Ascophanus isabellinus Clements in Pound & elements, Bot. Surv.

Nebraska 5: 9 (1901). -- ColI. orig. on stercorate mud from
U.S.A., Nebraska, Otowanie woods; not known to exist.

ETYMOLOGY: Latin isabellinus, light leather-coloured with a

reddish tint; referring to the colour of mature apothecia. Its
origin alludes to the spanish Queen Isabella's clothes which during
a siege gradually turned a dirty colour.

Pound & Clements' original description of h. isabellinus runs:

"Ascoma humicolous, carnose, at first nearly doliiform, then convex

or explanate, naked, white, then isabel-colored, 1 - 3 mm.; asci
broad, 8-spored, stipitate, blue with iodin, 250 x 30 - 40 ~;

paraphyses of two sorts, linear, much exceeding the asci, and

clavate, only slightly longer than the asci, 3 ~ wide; spores
distichous, elliptical, hyaline, smooth, with a thick, lamellose
wall 2 - 3\ ~ wide, 27 - 30 x 12 - 13 ~.

On stercorate mUd, otowanie woods. (12109)."

 184

INCORRECT RECORDS: Seaver (1928: 112) is Thecotheus cinereus.

Seaver (1942: 310) is T. cinereus. -- Kimbrough (1969: 109), as
holotype of A. isabellinus. This collection is identical with
Seaver's (1928: 112) [= T. cinereus).

HABITAT AND SUBSTRATE PREFERENCE: On stercorate mud.

DISTRIBUTION: Only known from the type locality, at the otowanie
woods, Nebraska, V.S.A.

In spite of a rather short description, without any illustration,

there is no doubt that this taxon belongs to Thecotheus. The
species is characterized by the size and arrangement of the
ascospores, and the possession of paraphyses of two types. T.
isabellinus is probably closely related to T. cinereus and T.
phycophilus.

Pound & Clements (1901) described the mature apothecia as isabel-

coloured. The interpretation of this colour varies from yellowish,
tawny yellow, light brown, and yellowish brown to light olive
brown. However, the colour-terms used by the authors (op. cit.: 5)
conform to those listed in Saccardo's Chromotaxia (1891). This is
also the basis for my own interpretation of the colour (see
etymology, above).

From time to time A. isabellinus has been mentioned in the

literature by different authors. Saccardo & Sydow (1902: 1149) and
Farlow (1905: 275) refer only to the original paper by Pound &
Clements. In Seaver (1928: 112) an accompanying description
identical with the one by Pound & Clements (1901), including also a
figure (pI. 11, fig. 13), is found. In addition, Seaver (loc. cit.)
included New York as a locality of the species. I have examined
 185

this latter collection, from herbarium NY, and it belongs to T.

cinereus.

Later, Seaver (1942: 310) included a collection by B. o. Dodge,

also from New York~ This material unfortunately consists of two
slides only. On a handwritten sheet [? of Seaver] accompanying one
of the slides, a question mark follows "isabellinus". It is also
noted that the material is "closely related to A. incanus ". To the
other slide is added "= A. pallens Boud.?". Tentatively, this
collection is identified as T. cinereus. An unpublished collection
identified as A. isabellinus in herbarium.BPI from China also
belongs to T. cinereus.

Notes on type material

Kimbrough (1969: 109) interprets A. isabellinus as a synonym of

Thecotheus cinereus, with reference to his examination of a Seaver
collection from a greenhouse in New York Botanical Garden, May 5,
1901, referred to as holotype of Ascophanus isabellinus. The
opinion of Kimbrough (1969) was later followed by Kimbrough, Luck-
AlIen & Cain (1969) and Pfister (1982). I disagree with Kimbrough
on this point. Under no circumstances can Seaver's collection from
New York be a holotype of Clements' new species originally
collected from Nebraska. Very probably, this particular collection
of Seaver's referred to, is identical with Seaver's (1928: 112)
Ascophanus isabellinus [= T. cinereus).

Several efforts have been made to trace the type collection of A.

isabellinus. There is no indication in Pound & Clements (1901) that
any of these authors are the collector. A note from the Editorial
committee, dated March 12, 1901, in "Botanical Survey of Nebraska"
Vol. 5, page 4, says that Messrs Pound & Clements have identified
 186

the fungi, and moreover that the Committee thanks Messrs M. E.

Moore, of Springview, and J. P. Anderson, of Lamoni, Iowa, for
collections, so the material may have been in their possession.

I have also tried to trace the type collection of A. isabellinus

among the types described by F. E. Clements. In spite of extensive
search in several herbaria (B, BPI, CUP, K, MICH, NEB, NY, PAD, PC,
RM, RMS, RO, S, UCSB, and US), I have not been able to locate it.
Nor is it included in a list of " Probable types or cotypes of
fungi described by Dr. Frederic E. Clements", a list kindly
forwarded to me by Dr. A. Y. Rossman, herbarium BPI. I would
suggest further search in the following herbaria in U.S.A.: Buffalo
(BUF) , Cambridge (GH), Manhattan (KSC) and saint Paul (MIN).

THECOTHEUS KIKBROUGHII Donadini, inval.

Donadini (1985) made a brief description in French of an apiculate,

ornamented species of Thecotheus from France, i. a. characterized
by its yellowish ascospores.
Unfortunately, no measurements of ascospores, asci and paraphyses
are given in the description, which runs as follow:

"- Petite espece de-0,3 a 2 mm de diametre, de consistance ceracee,

d'abord conique, puis cylindrique, obconique, et enfin aplatie plUS
ou moins recurvee-ondulee, blanche puis ochracee grisatre; poussant
sur bouses de vache agees de plus d'un an et de moins de deux

(Alpes de Haute-Provence, 2200 m, Allos-Valcibiere, aout-septembre

 187

1983-1984) .
- Hymenium de 215 a 250 ~m.

- Asques octospores , renfles, a bleuissement parietal par l'iode.

- Paraphyses incolores, composees d'articles greles et courts en
haut, plus ou moins ampullaces en bas, uninuclees une ou deux fois
divisees.
- Spores ellipsoldales, uninucleees, contenant le plus souvent
deux petites guttules lipidiques apicales orangees (B carotene
probablement present) dans un cytoplasme finement granuleux;
apiculees: apicules de 1,5 a 2 ~m de long sur 3 a 5 ~m de large,
plus ou moins aplatis, cyanophiles; ornamentation composee
generalement de petites verrues cyanophiles; l'exospore est tres
decollee dans l'eau et souvent striee transversalement; a sec, elle
est apprimee sur la perispore, ce qui masque un peu
l'ornamentation.
- Excipulum medullaire a textura intricata avec un nombre variable
d'articles ampullaces.
- Excipulum ectal a textura globulosa surtout, de 100 a 110 ~m

d'epaisseur, dont les cellules ne depassent pas 45 x 35 ~m".

I owe Professor Pierre Neville, Marseille, Fiance (in litt.

11.VII.1990), thanks for information about the late J.-C.
Donadini's material. All his collections are deposited in the
herbarium at l'Institut de botanique de Montpellier (MPU).
Unfortunately, no reply has reached me from MPU, although the
material is most probably to be found there.

This species appears from the description not to belong to any of

the four species with apiculate, ornamented ascospores recognized
by me in Thecothus: T. biocellatus, T. holmskJoldii, T.
lundqvistii, and T. rivicola. Only T. lundqvistii has hyaline
paraphyses of ± the same type as indicated in T. kimbroughii, but
differs, apparently, in the texture of the excipulum~
 188

B. EXCLUDED TAXA

IODOPHANUS MAGNIVERRUCOSUS Aas sp. novo (Figs. 118 - 120)

ETYMOLOGY: Latin maqnus, great, large; verrucosus, warty;

for the large warts and apiculi on the ascospores.

Apothecia sessilia, solitaria vel gregaria, usque ad 1.5 mm

diam., maturitate disciformia, pallide luteofusca. Excipulum
ectalium cellulis globosis vel angulatis, marginem versus
rectangularibus vel doliiformibus. Excipulum medullarium
texturae intricatae. Asci octospori, cylindrici, 190 - 300 x
16 - 19 ~m, parietibus iodi ope caerulescentibus. Ascosporae
obliquae uniseriatae, ellipsoideae, 19 - 22 (-25) x 9.5 - 11
~m; verrucis grandibus sphaeroideis usque ad 5.5 ~m diam.
instructae, apiculo bipolari usque ad 5.5 ~m in diam. Omis
spora circumcincta tenui mucilaginoso strato. Paraphyses
filiformes, septatae, simplices vel ramosae, infra usque ad
1.5 ~m diam., apicibus dilatato usque ad 6.5 ~m diam.,
flaviguttulato.
Fimicola.
Type: Southern Africa, Transvaal, Pretoria, De Beers Rust,
Poplar grove, on cow dung collected after heavy rains,
17.11.1939 E. M. Doidge (PREM 36831, holotype).

Apothecia scattered to gregarius, sessile, 0.5 - 1. 5 mm

diameter; at first globose, then expanding and becoming
cupulate, finally discoid, concolorous with the disc. Exterior
part usually dirty white to dull yellow furfuraceous, often
 189

clothed with small globose to sUbglobose granules, dUll yellow

in dry condition. Lower part furfuraceous, concolorous with
the margin layer, often with a red tint. Disc at maturity flat
or concave, somewhat 'roughened by the protruding ascus tips;
as dry brown-grey (5D2) to light brown (5D4 - 5C4). Margin
distinct, darker than the disc, usually with greyisb brown
(6E3) colour.

Medullary excipulum of loosely interwined hyphae, 2 - 3 ~m

wide, composing a textura intricata, with lobed cells, 3 - 5

~m diameter, constricted at septa, intermingled with ±
subglobose cells up to 6 ~m diameter. Ectal excipulum
composed of atextura angularis to textura globulosa, below
with globose cells, towards margin with several cell layers of
more or less rectangular to barrel-shaped cells, 8 - 15 x 10 -
22 ~m diameter.

Asci eight-spored, 190 - 300 x 16 - 19 (- 21) ~m, cylindrical,

short-stipitate, attenuate towards the base, only slightly
protruding above the level of hyrnenium at spore liberation,
their walls strongly blue in Melzer's iOdine, especially
towards the apices. Young asci occasionally with reddish brown
colour in iodine solution. Ascospores obliquely uniseriate,
ellipsoid to slightly broadly ellipsoid, 19 - 22 (-25) x 9.5 -
11 ~m, at maturity with large, coarse warts up to 3 ~m long
and up to 5.5 ~m broad. Each ascospore with a large apiculus
up to 4 ~m long and up to 5.5 ~m broad at both poles. Both
apiculi and warts stain in heated Cotton blue. Inner spore
wall, about 1.8 ~m thick, remains unstained in Cotton blue,
whereas the c. 0.2 ~m thick outer layer stains in Cotton blue.
Each ascospore surrounded by a thin gelatinous, cyanophilous
sheath. Paraphyses filiform, septate, branched or unbranched,
sUbhyaline and up to 1.5 ~m wide below, with apices inflated
up to 6.5 ~m, containing small, slightly yellowish guttules.
 190

specimens examined:

SOUTHERN AFRICA. Transvaal: Type collection (see above).

ILLUSTRATIONS: Le Gal 1963, fig. 1 as Ascophanus crustaceus

Starb.

This is Ascophanus crustaceus Starb. sensu Le Gal, Bull. Soc.

Mycol. Fr. 78: 407 (1963). Le Gal made the description of
Ascophanus crustaceus Starb. based upon this examined
material. The authour remarked that Starback did not mentioned
any ornamentations of the ascospores. The reason, according to
Le Gal (op. cit.) was probably that Starback did not use
lactic blue. Le Gal's opinion was later followed by Kimbrough
et al. (1969). See however under Thecotheus crustaceus for
further comment.

The presence of carotenoid pigments in the paraphyses and the

callose-pectic ornamentation of the ascospores well
characterize this species as a member of Iodophanus.
Iodophanus magnigverrucosus is so far only known from the type
locality near Pretoria, Southern A!rica. The size of the
ascospores, cqmbined with the large apiculi and coarsly
verrucose markings on their walls, distinguish this species
well from other known members of the genus. Spores with large
ornaments are i. a. found in I. subgranulatus (Berk. & Curtis)
Pfister, and I. bermudensis (Seav.) Kimbr. & Korf, but the
type of ornamentation is quite different and the spores are
larger.

Figs. 118 - 120. Iodophanus magnigiverrucosus. Ascospores.

Scales: 118 - 119: 20 ~m. 120: 10 ~m. Photos: 118 - 119: J.
Berge.
 191

120
 192

Thecotheus leveillei (Crouan) Lamb., FI. mycol. BeIge, Suppl.

1.: 282 (1887).
The correct name for this should bee Ascozonus leveillei
(Crouan) Brumm. See van Brummelen (1976) for f~rther comments.

Thecotheus winteri (Marchal) Boud., Hist. Classif. Discom.

Europe: 75 (1907).

The correct name for this species is Coprotus winteri

(Marchal) Kimbr. See further comments in Kimbrough (1969) and
Kimbrough et al. (1972).

Ascozonus solms-laubachii (Rabenh.) Lundq. & Aas in ed.

Some authors have placed the more or less forgotten name

Ascobolus solms-Iaubachii [laubachi] Rabenh. in synonymy with
Thecotheus pelletieri. They include Fuckel (1866, 1870),
Spegazzini (1878), Schroeter (1893) and Feltgen (1899).
Rabenhorst distributed material of A. solms-Iaubachii in his
Fungi europaei exsiccati as No. 420 (Rabenhorst 1862). I have
examined several specimens of this number, and all contain
fruit-bodies with 64-spored asci, not 32-spored as stated in
the original diagnosis. The material is conspecific with
Ascozonus woolhopensis (Renny) Hans. In a separate paper
(Lundqvist & Aas, in prep.), the combination Ascozonus solms-
laubachii (Rabenh.) Lundq. & Aas will be made, and the name
Ascozonus woolhopensis (Renny) Hans. placed in synonym.
 193

LITERATURE

As far as possible, periodical title abbreviations follow Alkire,

L. G. jr. (ed.), 1988: Periodical title abbreviations: by
abbreviation. Vol. 1, ed. 6, 1-835. Michigan.

Aas, 0., 1978: Koprofile Discomycetar (Ascomycetes: Discomycetes

Operculati = Pezizales) i Noreg. Med s~rleg vekt pA innsamling
i Hordaland og Sogn og Fjordane. Cand. real. thesis, unpubl.,
University of Bergen, 233 pp;
Amo , Mora, M. del, 1870: Flora cryptogamica de la Peninsula
Iberica: descripci6n de las plantas acotyled6neas que crecen en
Espana y Portugal, 849 pp. Granada.
Banheqyi, J., 1940: Contributions a la connaissance des Disco-
mycetes de la Montagne vertes. - Borbasia 2: 95 - 116.
Banheqyi, J., 1942 a: Discomycetak a felszabadult Felvidekr6l. -
Bot. Koezl. 39 (1-2): 33 - 41.
Banheqyi, J., 1942 b: Discomycetak a Szekelyfoldrol. - Ibid.
39 (5): 261 - 269.
Banheqyi, J., T6th, S., Ubrizsy, G. , Voros, J., 1985: Magyar-
orszag mikroszkopikus gombainak hataroz6konyve. 2. Eumycotina
(Ascomycetes: A Discomycetest6l, Basidiomycetes, Deuteromycetes)
pp. 517 - 1151. BUdapest.
Barrasa, J. M., 1985: Estudio de los Ascomycetes copr6filos
Espana. Tesis doctoral, unpubl., Universidad de Alcala de
Henares.
Batra, L. R., 1954: Studies on the Discomycetes of Mussoorie Hills
I.- Pezizaceae. M. Sc. (Hons.) Thesis, Panjab University, India.
Batra, L. R. , Batra, S. W. T., 1963: Indian Discomycetes. - Univ.
Kans. Sci. Bull. 44: 109 - 256.
Bednarczyk, M. A., 1974: Materials of the knowledge of the copro-
philous fungi in the LUblin region. - Acta. Mycol. 10 (2): 331
- 342. (In Polish).
Bell, A., 1983: Dung fungi. An illustrated guide to coprophilous
fungi in New Zealand. Victoria Univ. Press, 88 pp.
 194

Berkeley, M. J. , Broome, C. E., 1865: XLVII.- Notices of British

Fungi. - Ann. Mag. nat. Hist. Ill. 15: 444 - 452.
Bertault, R., 1982: contribution a la flore mycologique de la
Catalogne. - Acta Bot. Barcinonensia 34: 1 - 35.
Beyer, W., Engel, H. , Hanff, B., 1986: Neue Ascomyceten-Funde 1984
(z. T. auch frliher) in Nordwestoberfranken. - Die pilzflora
Nordwestoberfrankens 9 (9) 1986: 45 - 63.
Binyamini, N., 1973: Coprophilous fungi of Israel. 11. - Israel J.
Bot. 22: 151 - 158.
Bizzozero, G., 1885: Flora Veneta crittogamica. 1. I Funghi, 572
pp. Padova.
Bommer, E. , Rousseau, M., 1884: Florule mycologique des environs
de Bruxelles. - Bull. Soc. Roy. Bot. Belg. 23 (1): 15 - 365.
Bontea, V., 1953: Ciuperci parazite ~i saprofite din Republica
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 207

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 208

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95.
 209

XHDBI

africanus 11, 16 - 17, 21 - ~2, 26 - 27, 31, 35 - 37, 39,

42 - 46, 151
agranulosus 10, 70, 72, 74 - 76, 80 - 82, 110
albicans 112, 114, 119, 120
apiculatus 10, 73 - 74, 76, 80 - 82, 110, 114 - 116, 118, 120,
162
appendiculatus 110, 115, 117
Ascobolus 6 - 7, 12, 32, 34 - 35, 42, 59, 98, 100
Ascophanella 32
Ascophanopsis 32, 45, 151
Ascophanus 6 - 8, 32, 42, 105, 116, 133
Ascozonus 7
babingtonii Pachyella 154
bermudensis Iodophanus 190
biocellatus 21, 26 - 27, 31, 35 - 38, 40, 46, 49 - 51, 109,
163, 182, 187
Boudiera 34
caprinus 97, 106
caprinus, holmskjoldii var. 93, 97, 102, 104, 106
carneus Iodophanus 116
Cercophora 62
cinerea 32, 54, 57 - 58, 72 - 73
cinerea Mollisia 54
cinereus 8 - 9, 22, 24, 27, 28, 30, 32, 37 - 38, 40, 54 - 64,
73 - 81, 93, 96 - 105, 114, 119, 132, 155, 184 - 185
Coprotus 8, 118
crustaceus 16 - 17, 20, 24, 26 - 28, 30 - 31, 37 - 38,
40, 54, 57 - 61, 70 - 82, 97, 115 - 116, 119, 190
duplus Coprotus 118
durbanensis Iodophanus 101
Fimaria 17, 77
fuligineus 79
 210

furfuraceus 140
Gymnodiscus 32
himalayensis 11, 20, 22, 26 - 28, 30, 36 - 38, 41, 88 - 92,
177, 182
holmskjoldii 8 - 9, 12, 24, 26 - 28, 30 - 32, 37 - 38, 40, 56,
57 - 63, 78 - 79, 93 - 106, 129, 187
Humaria 116
inaequilateralis 17, 22, 26 - 28, 31, 37 - 38, 40, 108 - 110,
129
incanus 24,56,62,64,93,98,101,104 - 105,185
Iodophanus 10 - 11, 21, 34, 35, 79, 190
isabellinus 56, 59, 64, 183 - 186
karsteni 119
keithii 22, 26 - 28, 30 - 31, 37 - 38, 40, 58, 110, 112 - 120
kimbroughii 11, 21, 186 - 187
kimbroughii Iodophanus 37, 90
Lachnea 119
lacteus Coprotus 89
Lambertella 37
Lasiobolus 119
Lasiosphaera 62
lechithina Miladina 162
leporinus 97
leporinus, holmskjoldii var. 93, 97
Leucoscypha 112, 119
leveill:ei 192
lundqvistii 17, 20, 26 - 28, 31, 37 - 38, 40, 51, 127 - 129,
187
magniverrucosus Iodophanus 76, 188 - 190
major, cinereus var. 93, 105
Mqllisia 7
Neottiella 119
obscura, cinereus f. 77
obscurus 96
ocellatus 51
overtonii 140
Pachyella 35
 211

pallens 10, 26, 2B - 29, 38, 40, 54, 56 - 57, 63 - 64,

131 - 133, 154, 1B5
pelletieri 6 - 7, 9, 12, 17, 20 - 21, 23 - 24, 27, 2B,
30 - 32, 37 - 39, 133 - 146, 155, 192
perplexans 11, 21, 26 - 27, 31 - 32, 35 - 36, 40, 45,
149 - 151
Peziza 9, 35, 42, 79
Pezizula 7
phycophiluS 10, 17, 20, 22, 24, 27 - 29, 31, 36 - 37, 39 -
40, 152 - 155, 182, 184
Podospora 62
Psilopezia 162, 164
rehmii 93, 96
Rhinocladiella 23
Rhyparobius 7 - 9, 32, 145
rivicola 10, 17, 21, 22, 27, 28 - 30, 36 - 37, 40, 51, 115,
155, 160 - 165, 187
rivularis 164
Ryparobius 6 - 9, 32
Saccobolus 12, 34 - 35
setisperma 54
sexdecimsporus Ryparobius 114, 118
solms-laubachii 133, 192
Sordaria 133
strangulatus 17, 20, 26 - 27, 31, 36, 41, 170 - 173, 177
sUbgranulatus Iodophanus 190
Thecotheus 6 - 12, 17 - 1B, 20, 22 - 26, 29 - 31, 32 - 34, 35,
37 - 39, 42, 45, 59 - 63, 80, 90, 116, 11B - 119,
132 - 133, 144, 151, 154, 164, 173, 184, 186 - 1B7
uncinatus 17, 20, 26 - 2B, 30 - 31, 36, 41, 172, 175 - ~78

vesticola schizothcium 99
viridescens 11, 20, 26 - 27, 31, 37, 39, 41, 90, 181 - 182
winteri 7, B, 192
woolhopensis Ascozonus 192
Zukalina 32