An illustrated guide to the

coprophilous Ascomycetes of Australia

Ann Bell

Mole End, Lower Hutt

New Zealand

Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands

Fungal Biodiversity Centre
An Institute of the Royal Netherlands Academy of Arts and Sciences

11l
 \
\ \
\

Plate J. Dade ' s " Dungscape" pa inting

v
 .... ­ ..._- .............. .. .
,­
... . . -.. -­
.-
........ -. _- ~~ . - -

M
Plate 1. A-B. Ascobolus immersus. A. Apothecium with asci. B. Ascus with spores . C. Ascobo/us quezelii. Asci with spores.
D. Podospora fimis eda perithecium. E. Podospora appendi culata perithecium. F-G . Petriella sordida. F. Perithecia with
exuding spores. G. Germinating ascospores . H-J. Sordaria macrosp ora. H. Perithecia. 1. Ascus with spores. J. Ascospores in
water. K-M. Sporormiafimetaria. K. Asci with spores. L. Ascospore bundle (phase). M. Ascospore bundle (brightfield). N.
Poronia erici stroma with perithecia. Photography by D.P. Mahoney.

VI
Plate 2. A- B. Saccobolus infeston s. A. Asci in semiperrnanent mounting. B. Asci in water. C-G . Melanospora brevirostris. C.
Perithecia exuding ascos pores. D. Per ithecia. E. Hairs channeling spores. F. Asci. G. Conidia on germinating ascospore. H-J.
Ascobo lus dadei sp.nov. H. Apothecia with emergent asci , 1. Apotheciurn and asci. J. Asc i with asco spores. K-L. Sp orormiella
herculea. K. Asci and ascospores. L. Asci and ascospores. M-N. Sporormiella plat y m era., M. Ascus with ascospores, N.
Ascospore with gerrn slits. Photography by D.P. Mahoney.

VII
Contents

Preface ix
Acknowledgements ix
Major Harry Dade (1 895-1978) 1
Summary ofDade's research on coprophilous Ascomycetes of Australia 6
Other research on Australian coprophilous fungi 6
Materials and methods 7
Collection of substrate 7
Interpretation of distribution data 8
Recording Ascomycetes 8
Measurements 9
Illustrations 9
Coprophilous Ascomycetes II
The coprophilous discomycetes ("cup fungi") 12
Cleistothecial Ascomycetes 28
Pyrenomycetes 31
Glossary of shapes and ornamentations 58
Appendix 1. Description of new species 59
Appendix 2. Fonnulae of stains used for microscopie examination 63
References 64
Illustrations 71
Index to funga1 names 171

A complete list of a1l species found together with their frequencies of occurrence and States of Australia in which
they were found will be made available on the CBS website: www.cbs.knaw.n1

Vlll
 Preface his life and weil suited his rather military appearance", He
subsequently worked for the British govemment as a field
At the l3 th Botanical Congress (Sydney 1981), 1 had the mycologist in Ghana from 1921 until 1935, at which time
pleasure of meeting ML John Walker curator offungi at the he joined the Commonwealth Mycological Institute at Kew.
Biological and Chemical Research Institute of the N.S.W. He became Assistant Director at Kew in 1940 and remained
Department of Agriculture. He informed me of the exis­ in that position until his retirement in 1962. As his friend
tence of sorne 2,000 microscope slides and assorted note­ and colleague G. Ainsworth noted, Dade was: "most
books on Australian coprophilous fungi made by the late versatile ...an expert draughtsman, competent water colour­
Major Harry Dade during his retirement years in Victoria. ist and elever cartoonist and skilful manipulator of wood
Since 1 had been involved in research into coprophilous and metal". Dade was also a keen beekeeper and an active
fungi in New Zealand, Walker thought that 1 would proba­ member of the Quekett Club, an organization devoted to aIl
bly be interested in studying the 'Dade Collection'. the intricacies of microscopy. Dade contributed over 100
ln 1982 1 spent the greater part of two months scrutiniz­ publications on a variety of subjects. His book: The anat­
ing Dade's slides and his notebooks. Walker and 1 planned omy and dissection of the Honey Bee (J962) is still consid­
to publish Dade's work, but subsequent events in the early ered to be the authorative account on the subject. Dade
to mid 1980's caused me to reluctantly put aside these retired to Australia to be near to his only son who had
plans. In 1996 1 attended the Melbourne Systematics Con­ emigrated there. During these retirement years he began his
ference heId in part to commemorate the work of Baron research into coprophilous fungi.
Ferdinand von Mueller, There 1 learned of the plan by the The first chapter of this book is devoted to those retire­
ABRS to publish the Australian Fungal Flora in sorne 64 ment years of Dade as expressed in his correspondence with
volumes. In 1998 1 was successful in obtaining a grant to Crichton and in his various mycology notebooks. 1 have
enable me to undertake research on the coprophilous also included sorne of his views and reflections on his
Ascomycetes of Australia as part of this fungal series. The earlier life aIl of which serve to iIluminate the observations
research for this book covered the years January 1998 ­ made by Ainsworth that Dade was a multifaceted and
June 2002, but since it was conducted for the most part talented person. Throughout the remainder of the book 1
alongside teaching, it amounts to sorne three years of full­ will include comments and illustrations made by Dade on
time work. This can only be considered to be a smali various species of Ascomycetes alongside my own observa­
indication of the total number of ascomycetous fungi that tions.
must abound in such a vast continent.
ln anticipation ofthis work, 1made contact with Barbara
Dade (Dade's widow), Dr John Dade (Dade's son), and ML Acknowledgements
George Crichton, a fellow mycologist who knew Dade weIl
and had corresponded with him for years. ML Crichton 1 am indebted to numerous people for their considerable
provided me with copies of aIl Dade's correspondence with help and forbearance during the preparation of this manu­
him from 1964-1975. These letters together with Dade's script. Firstly 1 wish to thank the Australian Biological
meticulous laboratory notes have enabled me to enjoy a Resources Study (ABRS), who have supported me during
comprehensive understanding of Dade's views on mycol­ this project by granting me the funds to COYer sorne of the
ogy and other events that affected his life. Reading personal equipment and consumables necessary for this work. 1 wish
letters makes one privy to all kinds of extraneous informa­ to express sincere thanks to Walter Gams, Pedro Crous,
tion that would not normally be considered as relevant to Rob Samson and aIl other staff members of the Centraalbu­
include in a publication, which is after aIl, principally a set reau voor Schimmelcultures who made the publication of
of identification keys. But since the events of our lives do this work possible and to ML Gerry Keating of Image
help shape us as scientists, 1 consider them worthy of the Services, Victoria University, Wellington, New Zealand,
teIling. 1 found it illuminating to read of his attitude to war, for preparing the scans of the original illustrations and to Dr
the human condition, environmental problems and the Michael Priest curator of fungi at the Herbarium at Orange,
political events of the time. 1 also learned of his love of for kindly arranging for the scan of Dade's "Dungscape"
small, personalized grocery shops where he could purchase (frontispiece). 1 am indebted to Barbara Dade, George
his favourite Brie, Camembert and pork pies. His sharp wit Crichton and Leigh Winsor for all their knowledge of Harry
(as evidenced by his description of the Australian meat pie) Dade, which was so freely given to me. Very special thanks
enlivened the letters *. 1 also leamed to appreciate more fully to Dr Daniel Mahoney, my husband, for his unstinting
the extraordinary good fortune, which 1 have enjoyed by not support, numerous "fungal" discussions, fine photography
having my life marred by personal involvement in two and invaluable mycological knowledge without which this
World Wars. publication would be the poorer; to Dr Nils Lunqvist, who
There are at least two brief accounts of Dade's life: his kindly sent me ail the collections of Australian coprophi­
obituary by A. Onions (Bull. Br. mycol. Soc. 13 (l), p. 24, lous ascomycetes which he possessed and for answering my
1979) and an outline of the main events of his life by G.c. constant queries as to his opinions upon various taxa. Dr
Ainsworth (Brief biographies ofBritish mycologists, 1996). Mike Richardson kindly sent me duplicates of his collected
These inform us that Dade died aged 82 in July 1978, that dung samples from Australia and generously shared his
he served in the Royal ArtilIery during the First World War, knowledge with me. Sincere thanks to Dr Tom May, curator
was wounded during the battle of the Somme (1916) and of fungi at the Herbarium, Melbourne Botanic Gardens, Pat
"achieved the title of Major, a title retained for the rest of Grey and Katy Sommerville who forwarded aIl the dried
dung collections to New Zealand. Last but not least, grate­
fui thanks to aIl those stalwart volunteers who risked the
* " ... a smear ofanonymous slime between two sheets ofcard­
board". (Letter to George Crichton 5th Oct. 1970). ribald comments oftheir friends and still collected the dung
Îx
samples for me. These include: Joan Broadberry, A.C. Disclaimers
Cochrane, Robin Corringham, Julia Davis, Cecily Falking­ While 1 fully acknowledge the help from many during the
ham, Cicely Fenton, Janet Fenton, Ron Fletcher, Alistair & production of this publication final decisions upon its
Amanda Freeman, Pat Grey, Cheryl Grgurinovic, R. Grif­ content are mine and hence any mistakes herein are exclu­
fiths, Robert Kellermeia, Greg Kirby, Sandra Lambert, sively my own.
Teresa Lebel, Jean Lightfoot, Tom May, Dave Munro,
Mark Newton, Susan Palmer, Ceri Pearce, Jennie Pearce,
Joan & Bob Rowlands, lA. Simpson, Margery Smith, Julie Ann Bell
Strudwick, Katrina Syme, Kevin Teys, Tim Wardlaw, and Mole End, 45 Gumey Road
Nigel Weston. Without their help this research would not Lower Hutt, New Zealand
have taken place. E-mail: AskUs@xtra.co.nz

x
Major Harry Arthur Dade (1895-1978) visit them regularly; l picked up an amazing assort­
ment of useful stuff. At one time l even assembled a
complete microscope from unrelated parts, afoot here,
His early life a limb there, a stage elsewhere and soon altering them
to make them fit together. At the Quekett there was
Harry Dade was born in Greenwich, London, England, another goldmine in the 'For Sale & Wanted' book, in
the only son of an only son, thus he had few relations which members wrote their wants and disposais. ln
(Barbara Dade, pers. comm.). At an early age he began those years l was building up a reserve of accessories
being interested in natural history and developed a and materials (old brass, for example) in preparation
prodigious knowledge of biology in general and a for my retirement. l also had a windfall of about [
special interest in insects and fungi. He was an accom­ 300-worth ofapparatus left to me by afriend... When l
plished artist and an excellent microscopist. ln various emigrated l left behind a truckload of stuff and now
letters to George Crichton he explains that his interest wish that l had brought it with me ... "(Letter 14th April
in biology began when he was very young: "Starting 1970).
as a boy, l snatched every opportunity to get away to Dade was eighteen when he joined the Royal Artil­
the fields, which in those days were at my doorstep, lery during the 1914-1918 war. At first he was "in the
fields now built up over a solid 30 miles farther out saddle ail day", but later involved in" pretty strenuous
from London. l made a survey ofpond life over most of trench fighting from which l still wonder how l sur­
NW Kent, and wish l still had my notes, lists and vived''. He was badly wounded in the Battle of the
maps ... " (Letter 14th April 1970). "And in my gram­ Somme. "My face was badly burned in an explosion, it
mar school l founded a natural history society and will not now tan in the sun, but only goes pink!). My
took parties out for field work ... " (Letter 23rd Decem­ eyeballs were charred and l was told that l would be
ber 1970). "know Watkins & Doncaster! as a small blind. With ail my skin offand my eyes swollen closed,
boy, trudging up a long steep staircase in the Strand, l had twice daily removals of coke with a scalpel,
just beyond Charing Cross Station! And l have a sec­ while an orderly forced my raw eyes open with his
ondhand egg-collector 's cabinet still, now used for thumbs. Could not see anything for 3 weeks, then
innumerable small tools (which l have indexed as began to see light and in time recovered completely
otherwise 1 should never find them)! This set of draw­ and had excellent sight ever since (except for old-age
ers cost me Il shillings and 6 pence! What a Heaven long sight ofcourse) ... " (Letter 4th June 1965).
that shop was for ail this kind of thing ... " (Letter 5th Dade was deeply affected by his experiences in
March 1975). "Secondhand bookshops are far from WW 1 as many participants were and spoke of these
rare in Britain or at least were not when lleft in '62. horrors many years later to his wife Barbara. At eight­
As a small boy l haunted them spending my small een he had "the awful responsibility of sending young
pocket money (a shilling a week) on prizes from the 2nd boys out to certain death, having no choice" (Barbara
3rd & 61h boxes ranged outside the shops. l had a won­ Dade, pers.comm.).
derful collection! - lncluding Quekett 's ]'1 edition After WW 1, Dade attended the Royal College of
Treatise...for 6 pence ...still have it ... " (Letter i h May Science in London and after graduating took a post as
1970). a government mycologist in Ghana in 1921. He obvi­
Dade was a good student and he was awarded many ously enjoyed his 15 years in Ghana as evidenced by
book prizes for his achievements. He was also awarded the following comments made to George Crichton:
a scholarship by the London County Council. Since his "Then in Ghana a large part of my lime was spent on
school did not teach biology as a subject at that time, trek. There were no roads and no cars for years. l was
he decided to teach himself. He was also very practical out of touch with my headquarters for weeks on end
person. Evidence for this is expressed everywhere in rarely seeing another white man: - the most wonderful
his long correspondence with his friend George Crich­ sense offreedom that is possible to imagine. l did very
ton. In Britain he amassed a great quantity of micro­ little collecting then, my interest being purely 'ap­
scopical equipment and regularly visited shops which plied'... " (Letter 4th April 1970).
stocked such items. In 1935 Dade left Ghana for the Imperial (Iater
"There are few if any sources of secondhand micro­ Commonwealth) Mycological Institute at Kew. At
scopical apparatus here (Australia). Even in London, Kew he was more or less confined to laboratory work
it is now difficult to find. Formerly this was a goldmine but he also undertook lecturing and attending his be­
of 'junk', but now most ofthe useful shops have closed loved Quekett society meetings.
down. l am told in letters that ail the secondhand stuff "For the next 20 years in England l was on the move
is being sent to the USA where it commands high ail the time in my labo And then walking a great deal in
priees! l used to know ail the micro-junk shops and the country and lecturing for 3 hours a night with
journeys to and from central London, until l was really
physically tired. After that at about 60, l was forced to
1 Watkins & Doncaster was a well-known suppl y store for micro­ slow down by lumbago or whatever it is, which immo­
scope and biological equipment in London.

1
 bilized me suddenly without warning at any distance observations of the fungi as they appeared over a
from my home. It still does and prevents me from much period of several weeks. The greater part of the dung
collecting. " (Letter 23 rct December 1970). collections were made through the "invaluable and
Early in the Second WorJd War (1939-45), Dade unflagging co-operation of George Crichton", a my­
volunteered for military service and served in the army cologist whose own interests lay in the Gasteromycetes
for: "about two years " (Barbara Dade, pers. comm.), (Orchard et al. 1996). Dade also corresponded with
after which he was recalled because his laboratory Gordon Beaton an amateur enthusiast (Orchard et al.
work was seen as more urgent. He also was in charge 1996). Dade made several references to his apprecia­
of the local Home Guard during the remaining war tion of George Crichton's diligent collecting: "This
years and became Assistant Director of Kew in 1940. jact makes me particularly grateful to you as the main
His knowledge of tropical fungi was very valuable source of my coprophil material. 1 should be stymied
during those years and he undertook research con­ without your contributions ... .. (Letter 28 1h September
cemed with deterioration of army equipment and J 965).
materials . He was also responsible for establishing the
culture collection at Kew. He described the culturing
methods, which he used at the CMI in the Herb LM.L
.•
Handbook (1960), where his clear writing and practical
attitude to life are clearJy expressed . This chapter had a
great influence upon me as a postgraduatejust embark­
ing on my own mycological career because his account
of growing fungi in culture was the clearest and most
straightforward that 1 had read . In this account he
reminds the reader that we tend to "lose our field
sense " when we enter our laboratory and become
involved with " brass and glass". He reminds us that
we know "little of micro-ecology " (still true today)
and that in nature fungi do not grow in pure culture on
"unaerated jelly, which is laced with sugars". Thus
the recipes he preferred for culturing used natural
substrates wherever possible. The secret of Dade's
writing is that he wrote to inform his audience when so
much of science seems to be written to impress. He
made laboratory work with fungi accessible and hence
strongly influenced me as a postgraduate student little
knowing that 1 would one day be writing about him.
The publication 'Class Work with Fungi ' 1966 (re­
printed 1969) which he coauthored with Jean Gunnell
after his retirement is a clearly written, beautifully
illustrated publication. Harry Dade with Duke of Edinburgh.

His retirement years "But 1 could never have done it without your help,
Dade's practicaJ skills were put to good use wh en he which has been a godsend. " (Letter 24 1h October
retired to Australia . He set aside a room for his home 1966).
laboratory, constructed shelves for books and racks for To each dung collection he assigned a number
slanting agar and bound his own books. He also under­ prefixed by the letter 'V' to distinguish this series of
took many repair tasks about the home as this excerpt fungi from his Ghana and Kew collection numbers.
from one of his letters illustrates: "Tve been veJY Although most of his dung samples were from the state
occupied with odd jobs as weil as fungi, one of them of Victoria, in fact he kept this prefix for ail his Aus­
being to French polish the sideboard top (ruined by an tralian collections made from other states also. His
accidentaI spillage of gin). The only 'French polish' 1 initial observations were collated together in six bound
could buy was a thick varnish with instructions to books (Vols I-1V), which he called his 'Rough Case
apply with a brushll! However, after diluting il with Note s ' (or sometimes his 'bench books'). In each of
meth. andfinding some suilable cotton cloth. 1 got il to these he entered the dung collection number on the far
work quite weil " (Letter 26 1h February 1966). left of the left-hand page followed by a description of
Dade studied about 1,200 collections of dung of the dung, locality and date together with the collectors
both corn mon native marsupials and introduced herbi­ name or initiais. On the right-hand page he recorded
vores. Each collection was incubated in a glass-lidded the fungi as they appeared , allotting a letter of the
container lined with moist blotting paper. He made alphabet to each new species.

2
For example: never without interest and value. In a way, my out/ook
Vl36 'Possum dung, dry- Goomalibee, G. Harris is similar to that ofthe field naturalist who watches the
26.7.63. pellets habits of birds or animais, or flowering plants. And
(a) Aspergillus candidus in 3 days on 2 or 3 especially there is the matter of range and variation. "
(b) Penicillium claviforme (Letter l3 th January 1969).
"Two or three interesting fungi have come to light
Using this notation, a slide of the Aspergillus above as weil as the equally useful increment of knowledge
would be labeled: V136a. If a slide number was fol­ and experience ofthe commoner species. Untilone has
lowed by Roman numerals (e.g. V313II), this indicated watched a common fungus going through ils paces
that a second batch of the dung collection V313 was many times one does not acquire a really sound con­
incubated on a second occasion. ception of it.i.I am sure that many descriptions are
Fungi are such numerate organisms that it is neces­ based on single observations of scanty material and
sary for mycologists to become somewhat specialized. that is why identification is sometimes difficult ... "
Dade showed a c1ear preference for Ascomycetes, ln a letter to Crichton (2nd January 1967), Dade
particularly discomycetes and he tended to ignore the made the fol1owing observations regarding ascospore
Basidiomycetes, which tumed up on his dung samples measurements: "In my view il is not only impossible to
(though there are a few slides of Coprinus amongst his get nearer than 0.5 um in practice, but il is also ridicu­
slide collection). "L've had numerous agarics mostly lous for reasons which 1 now mention. We measure in
Coprinus spp. But 1 am not interested in these order to identify. We consult published descriptions.
things ... " (Letter 1i h September 1968). These descriptions are usually based in the first in­
He sent portions of sorne of the collections to his stance on one or a few collections and they give no
long-time colleague Jean Gunnell at the Common­ idea ofrange ofvariation ...
wealth Mycological Institute. She regularly corre­ 1 hold that il is quite useless to express spore meas­
sponded with Dade and there are several references to urements (for example) in the form: 9.7-10.5 x 7.0­
her in his 'bench books' and other notes. It is through 7.6 J'm. 1 hold that il is good enough in this case to say
this collaboration with her that sorne of his fungi are instead: 'about 10 -11 x 7-8 um '. 1 search a few
assigned IMI numbers in addition to his own number­ fields, note unusually small and big spores and form
ing system. He also made sorne reference in his corre­ an idea of the rough sizes on the great majority, then
spondence to having cultured sorne of the fungi (see record the figures ofter the word 'about'. Spore di­
for example a letter dated lOth December 1964), but 1 mensions are, afler ail, only one characteristic of a
have been unable to find any further details conceming species as weil as a variable one ... The tendency to
this aspect ofhis work. split hairs should be avoided. .,
Fungi that Dade studied in sorne detai 1 were entered ..We must not in fact be in a hurry (to publish) and
into loose-Ieaf files under the names of the genera and neither you or 1 nor G.B. (Gordon Beaton) are likely to
species. Such was his meticulous nature that he then be. By piling up records of our things we are doing
made "fair copies" of the "Rough Case Notes" be­ useful work and as time goes on we shall if we live,
cause the originals usually became marred by additions have something to show for il. 1 believe that when we
and corrections. These neat versions he called his have sufficient records we shall see the light and be
"Case Books" Vol. 1 & Il However, 1 consider that it able to produce reliable data on the limits ofvariation
is the very alterations and additions to his original in some, perhaps many, species. ,,].
"Rough Case Notes ", which are especially valuable Both in his letters and his research notes there is
because in them one can trace the thought processes of ample evidence of Dade's understanding of the task of
the author. Another collection of notes he the systematist: "A pigeon hole simile is a sound one
called: "Obsn" which includes summaries of dung (referring to classification), representing the true
collections, dung preferences of certain species and situation, for the whole structure of our conception of
related ecological data. A summary of these notes taxonomy is simply afiling system! It is not a hard and
together with a list of the species he found follows this fast diagrammatic picture of a family tree type of true
chapter. relationships, but only ofwhat we guess theyare. The
Dade strongly believed in the importance ofmaking only thing about an organism that is immutable is the
many field observations of living material. He "de­ species, the name label which we attach to il is in
tested" the study of "dried mummies" in herbaria as many cases, perhaps the majority, temporary guess
attested by the fol1owing: "Though fungi have not been work. On what we regard as satisfactory new evi­
at their best, incubations have not been useless by any dence, we take the thing out of one pigeonhole and put
means. 1 always get new scraps of information about il into another. A nuisance, very oflen but the best that
quite common and well-known species or can check or we can do. Naturalists hate changes of names of
conjirm fresh ideas. 1 never feel happy about published
accounts based on the dried mummies in herbaria and
like to watch living specimens. So new incubations are 2 See section conceming Lasiobolus ciliatus for an example of the impor­
tance of careful and cumulative observations from fresh material.

3
 course, but ifwe tried to maintain names as immutable consider writing up aIl his observations to be published
we should have a greater muddle, getting worse as as a CMI Mycological Paper, although Dade had sorne
time goes on, as in an office where documents are not misgivings about this: "Ainsworth, Director of the
classified at al! .. .! am an ant-splitter though not a CM!, suggests my stuff might be published as one of
lumper..." (Letter 25th May 1965). the CM/'s Mycological Papers. 1 don 't know ifit (is) in
"1 am finding the ecology ofcoprophiles very inter­ the form usually favoured, i.e. Deadpan skeletal de­
esting and think that nobody except Bul!er has looked scriptions which are so inadequate and at the same
at the coprophiles in that light, i.e. with the eye of a time so encrusted in technicallanguage. Ali my stuffis
field naturalist as distinct from the purely taxonomie in very simple language, deliberately as free as possi­
angle ... " (Letter 24th October 1966). ble from unnecessary long words. For instance, why
Dade held in high regard those who studied the write 'furfuraceous' when you can use 'scur.fy'? or
natural world in their spare time (rather than those 'doliiform' when you can say 'barrel-shaped'?" (Let­
rd
employed specifically to do so). "Amateur work laid ter 23 November 1967).
the foundations of mycology and al! good professional He continued to record data from dung samples
mycologists at the CM1 have a great respect for ama­ sent to him by Crichton until about 1969 when he
teurs, a good tradition and they do al! that is possible wrote: "1 have begun to write up my stuff in semi-final
to assist them, understanding very weil their great form as something to work on, add to and revise. I'm a
difficulties when they have no easy access to the vast bit worried about re-drawing ail my micro-sketches for
literature and material for comparison. 1 am in exactly my blasted eyes now make work difficult... " (Letter 3rd
that state myself now, having very little previous April 1969).
knowledge of the fungi which now interest me and the In J uly 1969 he was laid low with his: "annual
CM1 cannot give me much assistance there, the discos win ter attack of so-called gastric 'flu". He also ex­
not being one of their specialties ... " (Letter 13th No­ pressed sorne suspicion as to the Librium which he had
vember 1965). been proscribed which "makes me sleep". By the end
Dade was often frustrated by being so isolated and of the year he wrote: "1 am very laboriously writing up
dependent upon others for confirmation of certain the first part of my stuff, i.e. the general ecology, tax­
identifications because he no longer had ready access onomy etc. 1t is laborious because 1 have such a mass
to the library resources which he had previously en­ of notes, which have to be boiled down, and fitted
joyed at Kew. "1 am gradually accumulating reprints together. 1 will send it to you for criticism when it is at
as part of the drive for literature... 1t is unfortunate last typed... " (Letter 8th December 1969).
that 1 never made attempt to collect books etc. years Throughout the following year, Crichton continued
ago, but 1 never dreamed that 1 would be living at any to send dung samples to Dade but during that year
distance from my old 1nstitute where they have every­ there are several references to his continued concem
thing!" (Letter 29 th April 1965). regarding being hampered by not being able to finalize
Dade was an excellent correspondent and in his retire­ sorne of his identifications (due to lack of literature
ment years exchanged letters with many well-known and/or responses to his letters from other mycologists).
mycologists including: G.C. Ainsworth, R.K. Benja­ Such problems (in addition to increasing ill-health),
min, C. Booth, J. van Brummelen, R.F. Cain, M. Le stymied the writing up of his research.
Gal and R.W.G. Dennis. He was appreciative of those "The only thing to do is to file and describe al!
who responded promptly to his letters and under­ unidentified things as: 'Taxonomie species XVII' etc.,
standably frustrated with those who did not. In his this gives them a temporary label and they can be so
relative isolation from other mycologists, letters were a described in a published list. At some future date,
valuable lifeline. He indicated on a number of occa­ somebody will name them. 1 propose to deposit al! of
sions that he fully intended to publish his records of my material in Willis 's herbarium ful!y documented. 1
Australian coprophiles', "1 have also managed to hope some day before 1 croak, to publish an annotated
bring my records up to date, both writing up fair cop­ list of the coprophiles, with notes on ecology etc .... "
ies of my 'Case Book', which 1 write in the first in­ (Letter 14th April 1970).
stance in a rough note book and rewrite after observa­ A month later he wrote: "and am rather depressed
tions cease and are amenable to consideration" (Let­ by the difficulty of getting started on writing up my
ter 13th N ovember 1965). stuff, 1 have al! the information on paper ofcourse, but
He also began the task of reorganizing the mass of can 't make up my mind about over-all treatment. And
information, which he had accumulated and produced if 1 ever do get it done, who on earth would publish
tables listing dung preferences of different fungi and it?!" (Letter s" May 1970).
other ecological data. In November 1969, Ainsworth The following year Dade was hospitalized for the
visited Dade and made the suggestion that Dade might removal of gallstones, and although he wrote of having
made a good recovery there are no further letters refer­
3 Crichton (pers. comm.) has indicated ta me that in his view Dade ring to his writing up of his research for publication.
had no serious intention ofpublishing the work, but was only He and his wife Barbara moved from Benalla to Mom­
interested in making it available ta others who might do sa.

4
ington in early ]972 where his son and his family now
resided. It is c1ear from his letters that he fully in­
tended to re-establish his laboratory in his new home.
There is a gap between ]972 and ]975 when there was
no correspondence between Dade and Crichton, be­
cause Crichton too had shifted home and also was
retiring. In addition, Dade's health had deteriorated to
the extent that further work was beyond him. The last
letters which he wrote jointly to Crichton and Beaton
in February and March ]975, dealt with the details of
making slide boxes and how to prepare good micro­
scope sIide preparations. He also mentioned the con­
tinued problem of getting certain equipment for mi­
croscopy.
On 261h July ]978 Dade died. Ali his notes, sIides
and drawings reside in perfect condition at the herba­
rium at Orange, NSW. George Crichton who lives at
Croydon near Melbourne still retains this correspon­ Harry Dade working in his home laboratory in Australia.
denee that he had with his friend. Barbara Dade died
on the 2] sr of January, 2005.

5
Summary of Dade's research on coprophilous Podospora perplexans
Ascomycetes of Australia Podospora pilosa
Podospora pleiospora
Podosporate~aspora
ln his research notes entitled 'Obsn " Dade includes a
Podospora vesticola
four-page list of sorne 900 dung collections which he
Podospora tax. sp. l
studied.The dung sarnpled include that of: possum, Podospora tax. sp. Il
rabbit, wallaby, kangaroo, wombat, cattle, horse, rat, Podospora tax. sp. III
sheep, goat, pig, dog, koala, bandicoot, emu, and deer. Podospora tax. sp. IV
Approximately 239 of these samples were of rabbit Podospora tax. sp. VI
dung representing the most frequent dung collected. Of Poronia punctata
the remainder, approxirnately 305 samples were dung Pseudeurotium ovale
of wombat, wallaby, kangaroo or possurn. Sorne 112 Pulvinula cinnabarina
collections of possum dung together with Pyronema dubium
approximately 149 collections of wallaby/kangaroo Saccharomyces (species undetermined)
Saccobolus depauperatus
dung represent the most frequently collected dung of
Saccobolus ?globuliferellus
the indigenous Australian animaIs that Dade studied. Saccobolus glaber
ln the sarne notes he gives a check list of Victorian Saccobolus minimus
fungi which includes the following ascomycetes Saccobolus versicolor
recorded on dung: Sordaria arctica
Sordaria brevicaulis
Aleuria aurantia Sordaria equina
Ascobolus amoenus Sordaria jimicola
Ascobolus~u~uraceus Sordariajimiseda
Ascobolus immersus Sordaria macrospora
Ascobolus crenulatus Sordaria spermatiophora
Ascobolus hawaiiensis Sphaeronaemella jimicola
Ascobolus (species undetermined) Sporormiella australis
Ascophanuscarneus Sporormiella intermedia
Ascophanus microsporus Sporormiella megalospora
Ascophanus argenteus Sporormiella minima
Ascophanus sexdecimsporus Sporormiella tax. sp. 1
Ascozonus woolhopensis Sporormiella tax. sp. Il
Chaetomium caprinum Sporormiella insignis
Chaetomium cochliodes Thelebolus stercoreus
Chaetomium jimicola Trichobolus zukalii
Chaetomium murorum Trichodelitschia bisporula
Chaetomium (species undetermined)
Cheilymenia coprinaria Dade also observed and narned species additiona1 to
Cheilymenia stercorea those in the above checklist and in 1982 1 studied sorne
Cheilymenia raripila ofthese slides. They include:
Cheilymenia (species undetermined)
Coniochaeta discospora
Delitschia marchalii
Coniochaeta hansenii
Gymnoascus reesii
Coniochaeta leucoplaca
Nigrosabulum globosum
Delitschia winteri
Pyxidiophora sp.
Delitschia (species undetermined)
Saccobolus kervernii
Fimetariella maxima
Lasiobolus pilosus (= ciliatus)
Leucoscypha (species undetermined) Sorne of the names in the table above have now been
Melanospora zamiae changed due to subsequent publications. Since he also
Melanospora damnosa left excellent illustrations of his observations, 1 have
Peziza anthracophila been able to match his drawings with the newer narnes.
Peziza (species undetermined) ln the chapters which follow 1 will include Dade's
Pleospora sp. observations alongside my own.
Podospora anserina
Podospora australis
Other research on Australian coprophilous fungi
Podospora aloides
Podospora curvula
Podospora decipiens Coprophilous fungi are attractive organisms and thus
Podospora jimicola they have been observed and collected by many people
Podospora leporina and it is very likely that there exists Australian
Podospora pyriformis collections of which 1 am unaware. Certainly there are

6
herbarium samples additional to Dade's data stored at upon the State in which it was collected and/or the
the Herbarium at Orange (DAR), and Cribb has distance from similar collections. For example, if it
published a number of papers on Australian dung fungi was clear that a collector had collected several samples
(see Cribb 1988, 1989, 1991, 1992, 1994, 1996a, of the same kind of animal scats from the same
1996b,1997, 1998, 1999a and 1999b). Sorne of the locality, 1 chose one or two of these in which to study
herbarium material pertaining to these publications rather than all of them. In addition four samples of
resides at the herbarium of the Plant Protection Unit, rabbit dung were obtained from Macquarie Island, a
Queensland Dept. of Primary Industries (BRlP). Other Sub-antarctic Island territory of Australia. And sent to
collections are held overseas. For example, collections me by Dr Stephen Stevenson. A total of 193 of the 300
made by Tibell and identified by Lundqvist reside in collections have thus far been studied. Upon opening
Sweden (UPS), sorne in our own herbarium, which is each of the collections, 1 compared the given
now part of the New Zealand national collection and identification of the dung sample against the
included on the national database of Landcare in illustrations in "Tracks, Scats and other Traces - A
Auckland (PDD). Many collections of fungi from field guide ta Australian Mammals" (Triggs 1996), or
Commonwealth countries were sent to the (then) "Animal Tracks" (Murie 1974). 1 also referred to:
Commonwealth Mycological Institute" and published "What Mammal is That?" (Strahan 1995), in order to
by resident mycologists (e.g. Rifai 1968). ascertain the distribution of a particular animal. Sorne
animal scats are very distinctive (e.g. scats of wombat,
brush tail possum, koala, ring tail possum, rabbit), but
Materials and methods others are not so easy to distinguish. In certain
instances, the distribution data of a particular animal in
Collection of substrate Strahan's book corroborated identification, in other
Since this research was conducted in New Zealand, instances it was extremely difficult to distinguish the
sorne considerable organization had to be undertaken scats of the larger wallabies from those of the various
in order to arrange for the collection of the dung subspecies or races of kangaroo. Thus in Table 1 the
substrate from various parts of Australia to be category: macropod includes the scats of the grey &
forwarded to me. The initial collections were made red kangaroo, wallaroo, and swamp wallaby together
through the Australian fungi-mapping scheme centred with similar sized wallabies. Of the 193 collections
at the National Herbarium, Melbourne Botanic studied 154 (81%) were of marsupial dung and the
Gardens. A brief description of the proposed project remaining 39 (19%) were scats of animais other than
was printed in the Fungimap Newsletter (no. 4, April marsupials. Of the samples thus far studied the
1997) requesting those volunteer naturalists who had distribution of dung types was as follows:
hitherto collected Basidiomycetes for earlier volumes
to switch their attention to the collection of small Table 1. Collectionsof dung types
samples of dried marsupial dung. The collectors
recorded details of the collection site (latitude & Dung type Number of
longitude), description of the local area and its collections
vegetation, date of collection and the identification of Agile wallaby/Rock wallaby 3
Bandicoot 2
the dung collected. Meanwhile in New Zealand
Bat 1
arrangements were made for the establishment of a Brumby(wild horse) 2
quarantine room in which to incubate and study the Brushtail possum 13
dung collections. Ail collections were initially Camel 1
forwarded to the Herbarium at Melbourne where Pat Cassowary 2
Grey and Katy Sommerville entered ail the data Cattle 8
pertaining to the samples on a computer programme Emu 2
after which the samples were boxed and forwarded to Fox (?) 1
New Zealand. Approximately 300 dung samples were Horse (domesticated) 2
Koala 7
collected in this manner. Upon their arrivai they were
Lizard (blue tongue) 1
placed in a refrigerator until such time as they were Macropod 89
thoroughly soaked in sterile distilled water and plated Quokka 3
in glass lidded dishes for observation. All States of Quoll (or cat) 1
Australia were sampled although of the 300 collections Rabbit 15
made the majority were from New South Wales or Rattus sp. 1
Victoria. In order to cover ail the States in the time 1 Ringtail possum 5
had available for this project, 1 selected samples based Rock ring tail possum 2
Sheepor Deer 1
Wombat 30
"Now called: CAB International, Internet: http://www.cabi­ Unknown 1
bioscience.org

7
Interpretation of distribution data pyrenomycete fungi was Podospora (209 or 22% of
collections), followed by Sporormiella (106 or Il % of
Recording the distribution of fungi is fraught with collections). Of the discomycetes the most frequently
problems, which those recording the distribution of encountered genera were: Ascobolus (98 collections),
other organisms such as mammals or many plants do Saccobolus (78 collections), Iodophanus (52
not encounter. Firstly there is the very real problem collections), and Coprotus (49 collections). However,
that mycologists do not know the full extent of an the most frequently observed species overall was
individual entity since we record reproductive Iodophanus carneus (on 47 or 5% of collections), a
structures for the most part and any individual fungus fungus which enjoys a worldwide distribution.
may produce an infinite number of reproductive Interestingly this fungus (together with Podospora
bodies. Fungal literature abounds with species whose tetraspora & Chaetomium globosum) has ascospores
distributions initially were considered to be limited, capable of withstanding desiccation for sorne 16 years
possibly because their authors felt pressed to emulate after which they will germinate to produce ascomata
the pronouncements of other biologists studying more when their substrate is rehydrated '. It would seem to
amenable organisms. There were always few be advantageous for coprophilous species to have at
systematic mycologists worldwide and currently even least one of their spore states able to survive long
those meagre numbers are dwindling. As a periods considering the somewhat itinerant nature of
consequence of this, except in certain instances', the their chosen substrate. This may in part account for the
distribution data concerning fungi more nearly follow variable germination rates of a number of species. A
the perambulations of mycologists rather than the true rich reward lies in wait for young mycologists who are
distribution of the fungi themselves. Vast tracts of the prepared to store animal scats until their retiring years!
world still remain mycologically unexplored. One of Of the recorded species 13 represent species new to
the objects of this research was in response to that very science. The descriptions of 10 of these appear in
problem, but after three years of intense study 1 have Appendix 1. One of the remaining three (Podospora
only scrutinized a mere 193 dung collections from the n.sp.), will be published later and the remaining two,
vast continent that is Australia. (Podospora petrogale & Semidelitschia nanostellata,
While monographie works are a godsend to those have already been published (Bell 1999, Bell &
of us undertaking country inventories, too many of Mahoney 2001). Of these new species, Ascobolus
them are (understandably) based upon an overreliance dadei n. sp. was the one that was found most
on dried herbarium material rather than on the study of frequently (14 collections)". It is too premature to
living collections of fungi growing on their natural surmise that these new species might be confined to
substrates. This is in part due to the fact that the dung of marsupials, considering that other
individuals constrained for time such as those engaged similarly infrequently found species such as
on Doctorate theses have largely undertaken Podospora prethopodalis (previously recorded from
monographie work. For these reasons, sorne taxa are Africa), was also recorded from a single collection of
(in my view), too narrowly defined. Interpretation of wallaby dung from the Kimberley region of Western
species is fundamental to ail statements pertaining to Australia. Thus the other 13 new species may turn up
ecological preferences or distribution. infrequently elsewhere in the world and may simply
For ail these reasons taxa, which 1 have seen on a represent species that are rare or produce their
few collections only, 1 do not consider that the data is teleomorph infrequently under laboratory conditions.
sufficient upon which to make any substantive
comment as to their distribution or substrate Recording Ascomycetes
preferences. The pyrenomycetes were most common
with 23 genera compared to the 16 genera of the One hundred and seventy-six species of Ascomycetes
discomycetes". It wou Id be tempting at this juncture to were identified during the course of this present study
surmise that under the desiccating climate of Australia, of which 13 (7.4%) represent hitherto undescribed
pyrenomycete fungi had a selective advantage were it species. Dade recorded 9 (10?) species which 1 did not
not for the fact that the most frequently recorded encounter. These included species such as Ascozonus
species was the seemingly delicate apothecia of woolhopensis, a species known to be restricted to
Iodophanus carneus! Of the 936 records of species, fruiting in the winter months (Larsen 1971, Bell 1975)
575 (61%) were of pyrenomycetes, 343 (37%) were of and five species of Cheilymenia, a genus that generally
discomycetes and 18 (± 2%) were of plectomycetes. does not develop upon laboratory-incubated dung. 1
The most frequently recorded genus of the have seen a number of species that Dade did not find,

5 Suchas thosefungi which are associated with a particular 7 Collections of marsupial dung collected in 1982and kept
host. boxed in the laboratory were rehydrated in 1998at the onset
6 These numbers include those discomycetes found by Dade ofthis study.
in addition to my own observations. 8 Dade also recorded this species several times.

8
 but there is a certain amount of serendipity in finding separate putative taxa (see for example problems met
relatively rare species (e.g. Mycoarctium, Strattonia, in genera such as Sporormiella, Delitschia and
Orbilia). Dade also spent more time carefully Sordaria). Evidence from morphology alone may be
describing individual species recording their early insufficient to fully interpret these taxa.
development through to their old age. There were
fewer monographs when he was undertaking his Illustrations
research, thus he would have spent more time puzzling
over identifications. Since a large percentage of the human brain (50% at
Semipermanent microscope sIides were made of least)!", is involved with vision and our visual system
each species each time it was recorded. Exceptions to has the capacity to process huge amounts of
this rule being when the species was exceptionally information simultaneously, it would seem that
cornmon and lor taxonomically unproblematic (e.g. information presented in a visual way might be more
Iodophanus carneus or Sordaria jimicola). Shear's easily assimilated than a written description of the
mounting medium or Cotton Blue in lactophenol was same. For reasons such as this 1 favour the use of
genera1ly used for semipermanent mounts. Recipes for copious illustrations wherever possible, including
these media can be found in Appendix 2. 1 belatedly iIIustrated keys.
discovered that the best technique to seal coverslips Details of ail species were first recorded by means
was to first ring each with c1ear fingernail base coat or of pencil drawings attached to a coyer sheet pertaining
bonder, the formulation of which is specifically to each (numbered) dung sample. Measurements of ail
designed to adhere to slightly greasy surfaces such as structures and details of ascoma colour were made at
fingernails and slide preparations. Once the base coat this stage. When a particular species had been
was dry, a final coat of c1ear nail varnish was applied observed several times these observations was pooled
around the edges. and the final illustrations made. Within a particular
Herbarium material was made whenever possible, genus, ascospores were drawn to the same scale to
collections being stored in our own herbarium which is allow for easier comparisons. Exceptions to this rule
annexed to the herbarium at Auckland (PDDt are on the few occasions when the ascospores of a
Cultures were also made of species of particular particular species were so large (relative to the other
interest. species of the genus) that the illustrations could not be
accommodated on the paper (e.g. ascospores of
Measurements Ascobolus immersus). The morphology of perithecia is
only useful for identification in a generalized way.
At least 10 ascospores of each species were taken for Two exceptions to this rule being the genera
identification purposes. Measurements are given to the Chaetomium and Cercophora where perithecial
nearest whole micron. In the majority of instances a1l morphology plays a useful role in the identification
measurements were made from water mounts although process. In ail instances 1 have endeavoured to
there is little evidence that thick-walled ascospores iIIustrate typical perithecia for each species iIIustrated.
significantly diminish in semi-permanent mounting Colour was used in those genera where colour is of
media. For sorne species described here just one particular diagnostic importance. Thus in the
sample has been found and hence the ascospore discomycetes colour is especially useful, since so
measurements recorded should be interpreted with the many diverse discomycete taxa have single-celled,
understanding that the range given is limited. hyaline ascospores. Illustrations are in black in those
Moreover many described species have not been taxa where colour is not such a significant feature for
recorded from Australia and slight variations from identification. Ali illustrations were drawn to allow for
published descriptions may reflect as yet unrecorded a 50% reduction during the final printing stage. Paper
geographical differences. For these reasons, one used for the illustrations was Saunders Waterford acid­
should interpret ascospore measurements as free hot-pressed paper 300 g/rn-, Drawing implements
approximate and one should not be surprised to find a used included Mars-Magmo disposable pens giving
sample of the same species whose ascospores may be a constant line widths of 0.25 mm, 0.35 mm and 0.5
few microns different. In most instances, species mm, together with a steel-nibbed dip pen, which gave
recognition is not merely dependent upon variations in lines of variable thickness. Watercolours were used for
ascospore size, although they do form a crucial aspect the coloured illustrations and finishing done with HB,
of identifying ascomycetes. Particular difficulties are B, 3B and 4B pencils. A smail amount of Letraset
encountered when the ascospore sizes coyer a very shading and symbols were used for scatter diagrams
broad range and there is little else upon which to and for certain ascospores (e.g. Chaetomium species).

9 This (originally private) herbarium will be finally housed 10 Information obtained from the Institute for Innovative
at Landcare in Auckland (PDD) when we cease to do re­ Blind Navigation Website http://isd.saginaw.kI2.mLus/
search. -rnobility/

9
 Coproph ilous Ascomycetes

Ascoma an apothecium. Le. a disc goblet

,
•
or pulvinate structure usually light coloured ­
(e.q. white. yellow, orange), seated on the
dung surface. Theasci are situated onthe
upper surface from whlch they may protrude at maturity

.. ~.
..,."..,
Discomycetes
(page 11)

apothecium * ~l',
l '
. -.::. "-,' -.<B.
.;"~?~
. ;-J!~
;-.-- ~""'-

Ascoma a cleistothecium: Le a closed -{..,

structure lacking any preformed opening (=ostiole i­

,A.~~·
the ascospores being dispersed by the rupture
ordlsintegrabon oftheperidium

.. ,

(e--~-'
# ...'
._ - - - -
.
f - ----}

.. o·
:7
­
~
Plectomycetes
(page2S)

•
f" ~-'-'.;- ",,--,.:, • • •
(

,
' •......... --­ ­ .

-~

.---~

cleistothecium

Ascoma a perithecium i.e. a globose or­

flask-shaped structure, usually dark atmaturity
provided wlth a pre-formed openmg (=ostiole)­
seated at theapex of a neck of variable length Pyrenomycetes
(page 31)
.:.Jj:4~~---
,_
...,..-e: -~~::=.:~~ ~ /'

... ..
~ ~"
perithecium '~.~~
Fig. 1. Picture key illustrating the morphology of ascomata. The terms discomycetes, plectomycetes and pyrenomycetes are
used here in a generalized sense and do not indicate their arrangements into classes. For further discussion see text.

10
Process white was used to break up lines or make The coprophilous discomycetes ("cup fungi")
corrections.
1 have tried to arrange the illustrations such that The discomycetes as earlier defined were a class of
species, which might cause sorne confusion, are either ascomycete fungi characterized by having cup-shaped
on the same or neighbouring pages to allow for easier sessile or stalked ascomata (apothecia). The class is no
comparisons to be made. Finally computer scans of the longer generally accepted, but is a useful term to use in
illustrations were done at 600 dpi and saved as tif files. a general descriptive sense since, regardless of
experience, aIl users can for the most part distinguish
such a shape. However, certain discomycete genera
Coprophilous Ascomycetes may cause sorne confusion (e.g. Lasiobolus,
Thelebolus, and Trichobolus) due to the fact that in
Keys their immature state the ascomata have a roof of
hyphae and thus resemble a cleistothecium.
The picture key illustrating the morphology of the Lasiobolidium is considered to be a cleistothecial
ascomata (Fig. 1) will direct the reader to the three genus closely related to Lasiobolus (see page 17 ).
major groupings used in this book, i.e.: discomycetes, Very many publications are concemed with
plectomycetes, pyrenomycetes. It must be stressed that discomycete fungi and a number of these will be
these three groupings no longer represent formai mentioned when discussing particular genera.
classes of fungi, but the terms are used in a generalized However, no discussion of these minute and beautiful
way since in an identification manual such as this, the fungi would be complete without mention of the
morphology of the ascoma is the first thing which the magnificent publication by Boudier: Icones
observer notices and thus these terms serve a very mycologicae ou iconographie des champignons de
useful practical function. Subsequent keys to the France (1904-1911). Special mention also should be
genera either take the form of a dichotomous key for made of: The Australasian Pezizales in the herbarium
the assemblage of plectomycetes, or to further synoptic of the Royal Botanic Gardens, Kew by Rifai, British
picture keys for the discomycetes and pyrenomycetes. Cup Fungi by Dennis (1978), and Fungi of
Dichotomous keys guide the reader to each species. In Switzerland: Vol J: Ascomycetes by Breitenbach &
the body of the text the genera are arranged according Krânzlin (1984).
to their families as defined by the 9th edition of the With the exception of the genus Orbilia, the
Dictionary of the Fungi (Kirk et al. 2001). Reference coprophilous discomycetes included herein are
to the artificial nature of the old class Plectomycetes operculate and belong to the Pezizales. Sorne of the
and the more recent views as to the more natural families of this order are composed of mostly
classification of this assemblage of fungi is dealt with coprophilous genera (e.g. Ascobolus and Sacco bolus in
in sorne detail where these fungi are discussed. By the Ascobolaceae). The genus Orbilia (= family
following the sequence of synoptic picture keys the Orbiliaceae) is inoperculate. Although the Orbiliaceae
reader will (hopefully) arrive at the correct genus. The were originally considered to be members of the order
dichotomous keys to the species together with the Heliotales, recent research suggests that they are not
annotated illustrations form the basis for identification. closely related to that order and form a separate order
There are no formai descriptions to species already Orbiliales, (Kirk et al. 2001). However, the genus
known to science. The illustrations contain information Orbilia as it presently stands may itself represent a
additional to that which appears in the keys since key heterogeneous assemblage of poorly related species
construction demands brevity. The formai descriptions and is in need of revision (Korf 1992).
to new species appear in Appendix 1. The discomycete genera identified during the
course of this research are differentiated in a picture
key (Fig. 2). By comparing specimens to be identified
with the boxed features as illustrated, the reader may
proceed to the pages as indicated where further keys to
species are located. Generic descriptions and keys of
the discomycetes are arranged within the alphabetical
arrangement of the families to which they are currently
considered to belong (Kirk et al. 2001). Colour
descriptions in the keys follow the notation of the
Methuen Handbook of Colour by Komerup &
Wanscher (1989), (given as K. & W. throughout
remainder ofther text).

Il
 o @

·. / ,..,..-:~ ~'

('.~
~ "\
O ll~ Ji
. ,. ~ ~: ~

'.~.
,li', ­ "

G~ ". "'

":,,"""m (p. 26)

~Iil1
Pseudom brophila (p . 25)
Thelebolus (p. 27) a
OG, ,.

. ~
>". •

Thelobous (p. 27)

~
°l
. .
l,..

~
t:.* ~f, ~·~~~
~
~
-----=.~

i
i [\
~ ii ~~
1 1
J
Pulvinula (p. 25) onV~' \\ 0 '<\ :,:

Lasiobolus (p. 19)

Saccobolus(p.16)
...
Ascozonus(p . 25)
)0 Orbilia (p. 20)

,) ~;:~

.... ' "

L ..-;" c. " .c.e., '
"""""'-. ..
1. ' -. v~
:;:.
" "z'
. 7 ;' '''J J

- ,

';
~ ~

;.::..~ '
~~ - -=-=:::

1 Peziza (p. 21)

O ." "
1
. , 1

/'~' v
- "
"

/'
"'~u' ~ s ,~:. .::~.:r 1

:·
--­Y ':--- , :>-­
\m
~ " r'q, ~.. /

~.. .... . '. '

.

O@
,
~ = t I .~ '; .' ­...
/i'­
_ . •
. -~
r :;:::.... ::-.
~ r" "\
: . : /! . '.:
~ r
.:

o
':' :
~
~
Ascobolu s tp . 13) "

'.
/ ---­
;r r..-_ ~ ,

~
,..., ~ .,

( ' \' fJ\;i/1 /l r <, -:' ' '

c\\ ('r r (1 (1 '}

~
... -: . ~. ~--~ - :;::::;

~ /.
® ,/

U"o,",", (p. " 1

Coprotus (p. 24)
Ascodesmis (p. 17)

Fig. 2. Picture key to coprophilous discornyce tes. To use this key compare salient features of the specimen with the boxed
illustrations. Upon rnaking a choice proceed ta pages indicated where the keys ta the species are Iocated.

12
Family ASCOBOLACEAE (warts) at their poles, rather than being evenly punctate
as originally described. In sorne Australian samples the
Ascobolus Pers. polar markings may fonn quite prominent warts.
ln the same letter to van Brummelen (August 25th
There are 52 known species of Ascobolus (Kirk et al. 1969), Dade described another species which he
2001). The genus is mostly comprised of coprophilous variously named Ascobolus amoenus Oudem. (syn. A.
species, many of which appear on dung within the first americanus (Cooke & Ellis) Seaver). Examples of his
two weeks of incubation. The pale yellow/luteous siides of this species include V194f, V396f, V537d,
apothecia commonly occur in large numbers and are V827b and V878a. He stated that the Australian
variable in size, in sorne species minute (ca 300-500 material was rather different conceming its ascospore
um) and in other species (e.g. Ascobolus furfuraceus) omamentation "This never, in Australia, consists of
reaching 1 cm or more diam. The walls of the fine punctate, but generally of small tubercles. These
ascospores are quite thick (circa 1um in sorne species) may be discrete, but often coalesce, though never
and hyaline when young. As they mature, a thick sufficiently to form a continuous sheet. There is a
pigmented epispore is deposited on the outsides of the tendency, often marked, for these projections to be
spores giving them their characteristic colour. The exaggerated at the poles of the spores. These features
most distinctive feature is the rose/red/vinaceus'! will be found in the micro slides which 1 send to you.
colour of the mature ascospores, which change colour The species is most commonly on the dung ofkangaroo
to umber as they age when the thick, often and wallaby, often on that of wombat and rabbit, less
omamented, epispore peels away. A pad of mucilage frequently on other dung 's. Sent to you are apothecia
sometimes surrounds (or is lateral to) the ascospores of Iwo collections preserved in 50% glycerol + 5%
but this disappears in dried specimens. The epispore formalin, which preserves their fresh appearance
may be smooth and featureless save for a few irregular better than in dried material. Slides of various
fissures (e.g. A. immersus); highly omamented due to collections are also sent. "
anastomosing parallel crevices (e.g. A. crenulatus); or There were no records in Dade's files of any reply
broken into coarse warts (e.g. A. brassicae). Because being received, but it is quite clear from his notes that
of the ageing processes of the ascospores, it is Dade considered his Australian fungus was the Ascobolus
important to secure identifications from mature and not amoenus of Oudemans (1882), and a copy of a paper by
elderly ascospores in which the epispore has started to Oudemans quoting this record resides in his notes. 1 have
disintegrate. The most complete description of species recorded this species 14 times during the course of this
of the genus is the monograph by van Brummelen current study and it does not at ail resemble Oudemans
(1967). Since his publication, a number of local Ascobolus amoenus as described by van Brummelen in
surveys appeared: Laksen (1970), Moravec (1970), his monograph where he makes the point that the type
Liou & Chen (1977), Paulsen & Dissing (1979), Bell material of Oudemans does not at aIl match Oudemans
(1983), Kaushal & Thind (1983), Dissing (1989), original description 12. 1 consider that this Australian
Wang (1 993a, 1996), Wang & van Brummelen (1997), species with its magnificent warted ascospores represents
Doveri et al. (2000b). an undescribed species. This was further confirmed by
Dade made extensive notes and drawings of the the fact that Lundqvist also sent a slide of this species to
species of Ascobolus that he encountered. He also me (Tibell 11205-b) and in his view it represented an
corresponded with van Brummelen and sent specimens undescribed species. Hence it will be described herein as
to him. In a copy of one letter he refers to his Ascobolus dadei (see Appendix 1). Other species of
observations on Ascobolus hawaiiensis mentioning Ascobolus recorded by Dade include: A. immersus, A.
sorne slight differences between the given description furfuraceus (slide V924b), A. crenulatus (slide V123a)
and the Australian collections. Dade describes the andA. albidus (slidesV210a and V341k).
ascospores of this species as: "within the range of Other records of Ascobolus from Australian
dimensions which you give for the species, but are material include: Ascobolus archeri Berk., A.
somewhat more slender and elegant than the figure in scategenus (Berk.) Brumm. (van Brummelen 1967)
your monograph". Upon checking, 1 find that Dade and A. crenulatus ( Cribb 1992).
was correct, the illustrations of A. hawaiiensis in van During this study 1 have identified the following 13
Brummelen's monograph are drawn wider than they species of Ascobolus. A scatter diagram (Fig. 23)
should be relative to their lengths, resulting in an illustrates the size ranges of the ascospores of
altogether broader looking ascospore. This observation Australian collections of those species that may cause
caused me to re-evaluate New Zealand material of A. sorne confusion due to their similarity.
hawaiiensis upon which 1 had reported in 1983. Sorne
of the ascospores of both New Zealand and Australian
material of this species may have larger punctations

II Ascospore colour ranges from 11B8 through ta 12E7 and finally 12 "Oudemans ' original description gives a deceptive impression of
6C7 as they age (K. & W.). the ascospores" (van BrummeJen 1967, p. 81).
13
 apothecial morphologv Crenulate margin

emarginate marginate sessile

apothecium apothecium apothecium

~t~
epispore
ID
ascospore

granulate punctate/warted parallel crevices fissured

Fig. 3. Glossary to terms used in the key below.

Key to the species of Ascoholus

1. Ascospores rose-coloured (12B6-8) at maturity; epispore smooth or with a

few irregular fissures 2
1. Ascospores rose-coloured (12B6-8) at maturity; epispore omamented with parallel
crevices, discrete punctations or coarse warts of darker rose 3

2. Ascospores mostly ellipsoidal, symmetrical, occasionally slightly pyriform;

epispore with a few irregular fissures; ascospores
54---67 x 30-38 um Ascobolus lmmersus Pers. : Fr. (Fig. 17)
2. Ascospores short/ellipsoidal to subglobose; epispore smooth with a few irregular
fissures; ascospores 38--44 x 32-341lm Ascobolus quezelli Faurel & Schotter (Fig. 18)

3. Ascospore omamentation consisting ofanastomosing rows ofpaler crevices, more or

less parallel or oblique to the long axis 4
3. Ascospore omamentation as even fine granulations, or coarser discrete or coalescing
punctations/warts 9

4. Many crevices in the epispore short, irregularly disposed about the axis of the ascospores 5
4. Crevices in epispore more-or-less parallel to the long axis of the ascospores 6

5. Apothecia luteous/ochreous (5B4); epispore crevices broken, mostly irregularly

disposed, short comma-shaped interspersed with isolated warts, ascospores ellipsoidal,
(19-)20-22 x (9-)10-11(-12) um Ascobolus albldus Crouan (Fig. 22)
5. Apothecial excipulum with scattered dark brown pigment; ascospores ellipsoidal,
epispore of anastomosing parallel crevices or short obI ique crevices, ascospores
32-36 x 14-15 um Ascobolus albidus "British Form" sensu van Brummelen (Fig. 22)

6. Apothecia sulphur-yellow/greenish yellow/umber, usually > 1 mm diam, often

with crenulate margin 7
6. Apothecia luteous/ochreous with scattered dark brown pigment in excipulum,
<1 mm diam, emarginate, sessile; ascospores ellipsoidal,
32-36 x 14-15 um Ascobolus albidus "British Form" sensu van Brummelen (Fig. 22)

7. Apothecia reaching 5 mm diam; ascospores ellipsoidal,

26-30 x 12-14 um Ascobolus furfuraceus Pers. : Fr. (Fig. 23)
7. Apothecia smaller, ascospores shorter 8

8. Apothecia sulphur-yellow/umber, reaching 2 mm diam, with crenulate margin, sometimes

short-stipitate; ascospores with at least 5 parallel anastomosing crevices in view; ascospores
ellipsoidal, 13-16 x 7-10 um Ascobolus erenulatus P. Karsten (Fig. 21)
8. Apothecium as above but epispore of fewer anastomosing paral1el or oblique crevices in
view; ascospores ellipsoidal, 20- 26 x 10-13 um .Ascobolus michaudii Boud. (Fig. 21)

14
9. Ascospores ellipsoidal; epispore comprising evenly disposed fine granulations;
ascospores 34- 40 x 17-19 um Ascobolus amoenus Oudem. (Fig. 17)
9. Ascospores ellipsoidal or subglobose; epispore comprising even or uneven scattered,
medium/coarse punctations (warts) which may be larger at the poles 10

10. Epispore with medium/coarse punctations sorne ofwhich coalesce, not enlarged at the poles;
fissures occasionally present II
10. Epispore with medium/coarse punctations (warts) which are frequently enlarged at the
poles on many ascospores 13

Il. Ascospores 50-62 x 28-36 um Ascobolus bistisii Gamundf & Ranalli (Fig. 18)
II. Ascospores < 50 um long 12

12. Ascospores ellipsoidal; epispore in the form of coarse punctations sorne

ofwhich coalesce occasionally also with fissures; ascospores
29-37 x 16-19 um .Ascobolus stictoideus Speg. (Fig. 19)
12. Ascospores globose, epispore coarsely punctate/warted;
ascospores ca 12 um diam Ascobolus brassicae Crouan (Fig. 20)

13. Ascospores ellipsoidal; 20-27 x 9-12 um Ascobolus hawaiiensis Brumm. (Fig. 20)
13. Ascospores ellipsoidal; (35-) 40-50(-54) x (17-) 18-22 um; epispore with coarse
punctations/warts frequently forming large blisters at the poles Ascobolus dadei sp. nov. (Fig. 19)

NOTES: Only one collection ofvery old A. furfuraceus has been seen (on cattle dung collected in Tasmania). The apothecia were in too
poor a condition to ilIustrate hence a diagram of a typical ascoma ofthis common species is ilIustrated.
Van Brummelen (1967, pp. 69, 74) states that A. quezelii Faurel & Schotter was: "..not va/id/y published: /acking indication of
type ....probab/y represents aform ofA. immersus ". But this taxon was regularly found on dung collected in Queensland in addi­
tion to A. immersus and was clearly distinguishable from it.

Saccobolus Boud. asci. Inside their thick coating they frequently are
ellipsoidal, sometimes slightly asymmetrical with blunt
There are 26 named species of Saccobolus Kirk et al. or truncated ends. Since the major key to the species is
(2001), a distinctive genus having its ascospores united based upon the ascospore arrangement as weil as the
into bundles of four or eight in each ascus. A number ascocarp colour, this aging process sometimes makes
of species are confined to dung as a substrate and their the species difficult to ascertain. For example, after
distribution is worldwide. Van Brummelen produced a sorne considerable wavering, 1 have decided that what
fine monograph to the species in 1967 and since that 1 first considered to be perhaps a sample of S. thaxteri
time a number of other authors have produced keys Brumm. or maybe S. globuliferellus Seaver, could
and country lists, e.g. Laksen 1970, Liou & Chen equally be an ageing variant of S. verrucisporus
(1977), Aas (1978), Bell 1983, Dissing 1989, Wang Brumm., since there is little in the ascospore shapes or
(1995) and Doveri et al. (2000b). sizes to separate them. Similarly, 1 could not find any
The tiny (ca 200 um) pulvinate ascomata of Sacco­ clear distinction between S. verrucisporus and S. beckii
bolus frequently appear in dense clusters. They may Heimerl (as described by van Brummelen 1967) or S.
appear sorne shade of amber to ochrous (6B8 to 6E7 reticulatus Aas (as described by Aas 1978). The 39
K. & W.) due to the pigmented paraphyses, or grey to collections of (what 1 am finally identifying as S. ver­
livid-vinaceous (6B 1 to 10E5), in which instances the rucisporus) showed every variation of ascospore or­
paraphyses are hyaline. But the most distinctive fea­ namentation from coarse but discrete warts to a more
tures are the ascospores, which appear under the dis­ or less continuous thick plating separated by fine re­
secting microscope as ellipsoid, blackish to dark vina­ ticulate fissures resembling a tortoise shell, (as iIIus­
ceous bundles (12B6), protruding above the surface of trated by Aas for S. reticulatus).
the apothecia. At higher magnification, the ascospore Dade spent sorne considerable time describing and
bundles are rose in colour (11 B6-8), although they are illustrating species of Saccobolus. He came to the
frequently described as being purp1e. The pigment conclusion that he had five species of the genus from
resides in the thick outer walls of the ascospores (the dung collected in Victoria. " One of these, S. versi­
epispore) and frequently serves to cement the 4 or 8 color occurs very commonly having been seen in hun­
ascospores together. One or two round masses of a dreds ofcollections" After studying van Brummelen's
gelatinous substance may be seen attached to the side (1967) monograph, Dade deduced that the 5 species he
of the 8 united ascospores. As the spores age, they had identified were: S. versicolor, S. glaber, S. verru­
change in colour to umber or charcoal grey (8E7 to cisporus, S. minimus and S.? globuliferellus. However,
8F2) and the thick outer walls frequently peel away. At it is also clear that he remained a little unsure of sorne
this point the ascospores may lose their previous ar­ of his identifications and on several pages of his notes
rangement and lay somewhat haphazardly within the the words appear: "Rewrite in the light of Brumm 's
15
monograph ". On one box of slides he wrote: "Check the coarser markings of the unknown species first ob­
ail and measure. S. versicolor is mostly labeled depau­ served by Cribb. There may weil be an undescribed
peratus or violaceus S. verrucisporus - variously species here, but until further evidence is forthcoming, J
labeled S. depauperatus - variously labeled . " It may am declining to designate these specimens as represent­
have been ilI health which plagued him in his later ing a new species, but simply wish to alert users of these
years that preventing him from doing any final identi­ keys to its presence. 1 have, however, illustrated a typical
fication work on this genus. 1 have seen a number of ascus of these specimens alongside the illustration of S
his slides of Saccobolus species in 1982 and made minimus (Fig. 27 0).
comments upon them to the best of my knowledge at
that time. However, now that 1 have had a lot more Table 2. Identification of Saccobolus spp from Dade's sIides
experience with Australian collections of Saccobolus, 1
think that sorne of my previous identifications might Dade's slide number Identification Present identifi­
not be the same now! Table 2 is a 1ist of the siides that (1982) cation
V102b, V127f, V688g S. verrucisporus S. verrucisporus
1 studied, my identifications in 1982 together with my V251c S. beckii S. verrucisporus
present views: (=lMII13407c),
Other records of Sacco bolus from Australia include: V493b
S minimus and S obscurus (Cribb 1994) and S verrucis­ V128a, V128d, VI28n S. versicolor S. versicolor
(=IMII04294n),
porus (Cribb 1996). Cribb (1995) also reported an inter­ V512d. V613b.
esting species of Saccobolus that he could not identify to V677a. V680e. V
species level. He found this species "in small quantifies 708c, V806b
on only one collection ofdung" (of agile wallaby). J too V602b, V799c, S. citrin us S .citrin us
have found the same species also from dung collected in VI19c(=lMl104289c), S. glaber S. glaber
VI8k 472c, 590b
Queensland. It has prominent yellow paraphyses and fits V183c. V521b S. minimus S. minimus
into van Brummelen's section Sacco bolus (p. 167, 1967). V438a. V551g, V844a S. depauperatus S. versicolor
The ascospores are of the size of S minimus but they are VI56g S. globuliferellus S. globuliferellus
quite heavily punctate, whereas van Brummelen de­
scribes them as smooth or finely punctate. However the During this current study J have identified seven
ascospores of S minimus frequently become more species of Saccobolus. Ali ascospore measurements were
roughly punctate when they age, somewhat resembling made without inclusion of the thick pigmented epispore.

0)
(~) ~
r2
~
~ n ~
or;:0 . . ~
".~- .... :.'
---::-.~~.,

Pu lvinate apothecium
-

tf):
cQJ" ~ L.::::'
N

6'. "
.i--.
Ascospore arrangement
(after van Brummelen 1967)

----.
finely punctate
•
•
coarsely punctate ------..;.~ fissured

Fig. 4. Glossary to terms used in the key below.

Key to the species of Saccobolus

1. Apothecia hemispherical, black; ascospores arranged according to pattern V,

ascospores ca Il x 5 um; paraphyses yellow Saccobolus infestans (Bat. & Pontual) Brumm. (Fig. 28)
1. Apothecia not black; ascospores arranged in another pattern 2

2. Apothecia amber to ochraceous (6B8--6E7 K. & W.); ascospores first arranged according
to pattern J, II or IV 3
2. Apothecia colourless, grey or vinaceous (8F2-l2E7); ascospores first arranged according
to pattern II 7
16
3. Mature ascospores arranged in pattern IV, with thick irregular pigment layers; individual
ascospores ca 12 x 6 um Saccobolus portoricensis Seaver (Fig. 26)
3. Ascospores arranged according to patterns 1 or III 4

4. Ascospores arranged according to pattern 1 5

4. Ascospores arranged according to pattern III, coarsely punctate, individual ascospores
12-14 x 6-9 um Saccobolus geminatus Thaxter ex Brumm. (Fig. 26)

5. Ascospores coarsely punctate, 12-18 x 7-10 um Saccobolus? minimus Velen. (Fig. 27)
5. Ascospores finely punctate 6

6. Ascospores finely punctate, 13-14 x 6-7 um Saccobolus minimus Velen. (Fig. 27)
6. Ascospores finely punctate often with fissures,
17-20 x 8-10 um Saccobolus citrinus Boud. & Torrend (Fig. 27)

7. Ascospores more or less smooth or with minute pits sometimes with fissures,
12-18 x 6-9 um Saccobolus versicolor (P. Karst.) P. Karst. (Fig. 25)
7. Ascospores at maturity coarsely punctate to adorned with interlocking plates
of a thick pigmented layer, broken by fine anastomosing fissures; ascospores
13-20 x 7-10 um Saccobolus verrucisporus Brumm. (Fig. 25)

NOTE: Van Brummelen (1967) describes S. geminatus as having colourless paraphyses. listing Thaxter's original herbarium specimen in
which the paraphyses had possibly lost their pigment in the dried state.

Thecotheus Boud.

A genus of sorne 17 species (Kirk et al. 2001). The keys to species of Thecotheus. Wang (1993b) de­
majority ofthese species are coprophilous, sorne being scribed a new species from Taiwan. Ranalli & Mercuri
cosmopolitan regarding their choices of dung types, (1998) discussed the genus as represented in Argen­
others are more substrate-specific (Aas 1992). The tina. Overton (1906) and Conway (1975) described and
genus Thecotheus is characterized by apothecia that illustrated ascocarp formation in Thecotheus. Conway
are frequently white, pale vinaceous to livid-vinaceous also described anamorphs "similar to Rhinocladiella"
(12A3-12E7). When young they are characteristically for T. pelletieri and T. cinereus. Ranalli & Mercuri
obpyriform to doliiform, becoming campanulate with also obtained a Rhinocladiella anamorph for T. pelleti­
age. They can frequently be mistaken for immature eri but not for T. cinereus.
species of Ascobolus yet to develop their characteristic
ascospore pigmentation. Kimbrough (1969), Krug &
Khan (1987), and most recently Aas (1992) provide

Key to the species of Thecotheus

1. Asci containing 8 uniseriate, ellipsoidal, hyaline, thick-walled ascospores,

19-21 x 9-10 um Thecotheus crustaceus (Starb.) Aas & N. Lundq. (Fig. 24)
1. Asci containing 32 multiseriate, ellipsoidal, hyaline, thick-walled
ascospores, 31-34 x 16-18 um Thecotheus pelletieri (Crouan & Crouan) Boud. (Fig. 24)

Family ASCODESMIDACEAE

Ascodesmis Tiegh. Lasiobolus Sace.

Ascodesmis Tiegh. comprises six species Kirk et al. This genus comprises a group of operculate coprophi­
(2001). Neither Dade nor 1 have seen this fungus from lous discomycetes whose apothecia are beset with hya­
Australian collections, but Cribb (1992, and 1998) line, aseptate, thick-walled bristles, which are acicular to
recorded Ascodesmis sphaerosporus Obrist from dung. subulate at their bases and inserted into the outer layer
The genus was monographed by Obrist (1961). More of the exipulum. The sessile apothecia may develop
recently Doveri et al. (2000b) described and iIIustrated singly or, more frequently, in groups on the dung sur­
three species from Italy, and Wang (1999) described face and their colour ranges from pale luteous to orange
two species occurring in Taiwan. (4A3--6A7). The apothecia can be quite variable in size
17
even within the same species and may be minute (ca approx. 400 x 23-26 urn, rather smaller than those given
150 urn diam) or reaching as much as 1 mm diam. for L. diversisporus (Fuckel) Sacc., but this species may
Sorne species have asci containing 8 ascospores, others exhibit variability in bristle dimensions as does L. cilia­
are polysporous. The fully mature ascospores of sorne tus.
species are distinctly pale luteous (4A3). A wrinkled These painstaking observations made by Dade from
hyaline jacket may be observed surrounding the asco­ the unhurried study of an abundance of fresh material is
spores, but this feature is not constantly visible. Accord­ but one example highlighting the importance of the
ing to Kirk et al. (2001), molecular studies indicate that study of living material rather than an overriding reli­
Lasiobolus should he placed in the family Ascodesmi­ ance upon what Dade denounced as "dried mummies" in
daceae. Previously it had been placed in the Thele­ herbaria (important though these might be). Aiso the
bolaceae (Kimbrough & Korf 1967, Eckblad 1968). ascospore arrangement within the asci is contingent
Amongst the 8-spored species there is considerable upon the state of maturity of the asci and their contained
variability as to bristle development, ranging from spores. Often apothecia are squashed for a slide prepara­
collections which sport many robust bristles to those tion when they are marginally mature (a state described
almost devoid of bristles. Since so much weight has by Dade as extraction by "Caesarian section "). This
been placed upon the features, Dade's remarks on action may affect the amount to which the asci swell in
Lasiobolus ciliatus are worth quoting here in full: water and thus also ascospore arrangements.
"The dif.ferences in overall size, breadth of asci and As a result of this variability in bristle morphology
length of setae, were first thought to indicate the exis­ and ascospore arrangement there has been considerable
tence of Iwo species, but by the time one hundred collec­ commentary as to what exactly constitutes Lasiobolus
tions had been examined the writer concluded that only ciliatus (Schmidt. : Fr.) Boud. (ascospores uniseriate),
one is involved, for by this time intermediate forms, L. cuniculi Velen. (ascospores biseriate) and L.
linking the extremes had been found The very large intermedius Bezerra & Kimb. (ascospores uni- or biseri­
apothecia do not appear in incubations and have been ate). These three species, whose ascospore measure­
seen only on dung at the time ofcollection. It is assumed ments are very similar (see Fig. 5), are reported to ex­
therefore that open field conditions are necessary for hibit differences in the morphology of their apothecial
their development. Asci in large apothecia are at first bristJes. For full discussion ofthese species and sorne of
cylindrical, 200 um or more in length and 17-22 um their synonymies the reader is directed to Rifai (1968),
broad, with short narrow bases and the ascospores are Breitenbach & Krânzlin (1984), Moravec (1971), Bez­
uniseriate. When they are maturing the asci protrude erra & Kimbrough (1975), and Yao (1996). Spooner &
partly above the upper surface of the hymenium; at this Butterfill (1999) describe and illustrate two species
stage they expand laterally to 32-35 um in breadth white obtained from dung collections made in the Azores. In
the spores migrate into the broad tip, becoming crowded the latter paper the authors indicated sorne difficulty in
and more or less regularly biseriate. Through shrinkage ascribing certain collections to either Lasiobolus inter­
of its lower part the ascus now assumes the form of a medius or L. ciliatus. They also record finding L. cu­
short broad body on a long narrow stem In dwarf apo­ niculi but state that certain of their collections may
thecia or on the shallow edges of larger apothecia the represent another (undescribed) taxon. However, a
asci may be broad from the outset" number of other authors maintain the separations be­
Dade: Rough Case Notes: Discos. 3 and booklet tween these species (see for example Prokhorov 1994,
marked: 'List' containing rough drafts of a planned pub­ Wang 1995, and 1999, Richardson & Watling 1997, and
lication. Cacialli et al. (1997).
Dade left a number of slides, excellent drawings in Other records include Rifai (1968) who lists two
addition to the written notes conceming his observations collections of L. ciliatus from Australia, both of them
on Lasiobolus. According to his notes, he recorded from cow dung, and Cribb (1992 and 1996b) who
Lasiobolus pilosus (Fr.) Boud. (syn: L ciliatus) on > 100 recorded Lasiobolus intermedius and L. microsporus.
dung samples (including dung from domesticated ani­ ln the scatter diagram (Fig. 5) boxed areas indicate the
mais as well as marsupials). He noted that the: "hairs ascospore dimensions of the species as described by
(bristles) appear to be larger on apothecia developed out Bezerra & Kimbrough. Superimposed are samples of
of doors than in incubation". Sorne of his collections measurements from six of the 30 collections 1 have
(V261a, V688c and V707a), had "minute bristles and not made from Australian material during the course of
many of them either". In 1982 1 scrutinized these slides this study. 1 am choosing to cali these collections
and sorne of the bristles were only approx. 90 um long. Lasiobolus ciliatus since this name predates both L.
Another ofhis collections (V393g) 1could not fit into any cuniculi and L. intermedius.
particular species described by Bezerra & Kimbrough
(ascospores 23-29 x 13-16 um), The bristles were

18
 -
15
 ­

~ 14
~ -

o 00
10 o
- o
00 0

9
-
a.....
L./--r----,----r--.--.--+--r---r----.---.---.----,..----.--.­
13 1 14 1 15 1 16 1 17 1 18 19 1 20 1 21 1 22 1 23 1 24 1 25 1 1

Ascospore length (IIm) ~

Fig. 5. Scatter diagram of ascospore measurements from four collections of Lasiobolus ciliatus and two of 1. diversisporus
(0-0). Boxed areas indicate ascospore dimensions ofpublished species as indicated.

As can be seen in Fig. 5, there is a "clinal contin­ can be uniseriate or multiseriate. Due to the variability
uum" (quote from Booth 1982) in ascospore dimen­ expressed, 1 have taken a whole page (Fig. 29) in
sions. The majority of the ascospores measured faU which to illustrate this species. Since my observations
between 16-22 x 10-14 um, 1 have been unable to mirror those made earlier by Dade, 1 share his opinion
distinguish these or the remaining 24 collections on to the effect that there is but one variable species in­
any other substantive features. The apothecial bristles volved here.
are variable in numbers and dimensions. At least one To date 1 have identified four species of Lasiobolus
collection seemed to be entirely devoid ofbristles until from Australian material which are keyed out below:
the apothecia were dissected and viewed under the
compound microscope. The ascospore arrangement

Key to the species of Lasiobolus

1. Ascospores < 12 um long 2
1. Ascospores > 12 um long 3

2. Ectal excipular cells vertically orientated; brist1es hyaline, smooth, ca 5 um

at the widest point; ascospores broadly ellipsoïdal, 8-9 x 5-7 um
.........................................................................Lasiobolus trichoboloides R.S. Khan & J.L.Bezerra (Fig. 31)
2. Ectal excipular ceIls horizontally orientated; most bristles punctate at their tips; ascospores
broadly ellipsoïdal to globose, 10-12 x 8-9 um, luteous (4A3) at full maturity .
..................................................................................Lasiobolus microsporus J.L.Bezerra & Kimbr. (Fig. 30)

3. Ascospores ellipsoidal, (13-)16-22 x (9-)10-14 um, luteous (4A3) when mature

.......................................................................................... Lasiobolus ciliatus ( Schmidt: Fr.) Boud. (Fig. 29)
3. Ascospores ellipsoidal, 22-25 x 12-15 um, luteous (4A3) when fully mature
............................................................................................ Lasiobolus diversisporus (Fuckel) Sacco (Fig. 30)

Family ORBILIACEAE

Orbilia Fr.

Orbilia is a genus of tiny (ca 0.2-1 mm), translucent, that substrate is also the residence of many nematodes.
inoperculate ascomycetes found on a variety of sub­ The asci and ascospores of Orbilia species are ex­
strates including dung. On dung they usually appear tremely smaU and mature extremely slowly. There are
after a very long incubation time (i.e. 2-3 months). a number of publications dealing with species of Or­
Their apothecia may be stalked or sessile and are bilia and keys including Bachman (1908), Dennis
frequently luteous to flesh/coral in colour (4A4-9A7). (1954) and Korf (1992). Spooner (1987) described the
Since their anamorphs have frequently been found to Australasian species of Orbilia listing sorne seven
be nematode-trapping fungi, it is not too surprising to species. Publications in which new species are de­
see the Orbilia teleomorph appearing on dung since scribed include Jeng & Krug (1977), Haines & Egger

19
(1982), Kohlmeyer et al. (1998), and Webster et al. of the genus which "desperately needs to be done", a
(1998). Anamorphs of Orbilia species are described in situation which remains today.
Pfister (1994) and Pfister & Liftik (1995). Korf (1992) Two species of Orbilia have been identified during
drew attention to the fact that there was no monograph the course of this present study.

Key to Orbilia

1. Apothecia sessile or on a short blunt stalk, pale luteous to saffron (4A4-4A7),

up to 1 mm diam; paraphyses stout with surface incrustations; ascospores hyaline,
acerose, 5-7 x 1.5-2 um Orbilia inflatula (P. Karsten) P. Karsten (Fig. 32)
1. Apothecia short stalked, peach to coral (7A5-7A7), up to 0.25 mm diam:
paraphyses with inflated tips, containing pigment granules and minute
incrustations; ascospores hyaline, ovoid to irregular, containing granules at
one end, 3-4 x 2-3 um Orbilia alnea Velen. (Fig. 32)

Note: Synonym: Orbi/hl alnea is a S) nonym or 0. coccinelle of Spooner ( 1987) and Breitenbach & Krânzlin (1984), see Korf (1992).

Family PEZIZACEAE

Iodophanus Korf

lodophanus Korf is a genus of sorne 14 coprophilous water and having a distinct operculum. Dade also
species with worldwide distribution having an Oedo­ commented upon the very variable col our of the apo­
cephalum anamorph (Kirk et al. 2001). The history of thecia describing sorne of them variously as "faintly
the genus together with a key to sorne ten species is to pinkish, ochraceous. pale salmon or apricot". He
be found in Kimbrough et al. (1969). Further species comrnented that the specifie epithet 'carneus' (the
were described by Kimbrough (1970), Jeng & Krug colour of meat) was quite inappropriate for this fungus.
(1977) and Thind & Kaushal (1978). lodophanus He considered that 'incarneus' (skin-coloured) would
carne us is one of the most common discomycetes be a better term. He also noted that l. carneus fre­
found on dung and 46 records of it have been made quently produces its apothecia "in masses" on the dung
during this current study. An interesting find is that as weil as on the surrounding blotting paper. Rifai
this species is able to remain viable for sorne 16 years. (1968) recorded l. carneus from soil in Adelaide.
lodophanus carneus was recorded by Dade (si ides Cribb (1988) recorded it from Queensland and he also
V183g and V526f). Originally he referred to this spe­ recorded l. granulipolaris (Cribb 1994, 1995).
cies by the earlier name: Ascophanus carneus (Pers.: Two species of lodophanus have been identified
Fr.) Boud., but changed this when the publication by during this current study. I found them to be indistin­
Kimbrough et al. (1969) became available. He made guishable on the dung, their apothecia both being flesh
the observation that the asci do not always assume a to salmon coloured (ca 6A5 K. & W.). Kimbrough et
blue colour when treated with Melzer's reagent and he al. (1969) described l. verrucosporus as having apo­
considered such tests as rather inconsequential as thecia that are "brownish when fresh, drying yellow­
compared with morphologicaI data. 1 too have found ish". In ail other respects, the Australian material
that l. carneus asci do not always stain blue in Mel­ matches the published description for this species.
zer's. However, the asci of lodophanus species are
quite distinctive being broadly saccate, swelling in

Key to the species of Iodophanus

1. Ascospores hyaline, ellipsoidal, usually evenly, finely granulate, occasionally

smooth, 20-23 x 10-13 um /odophanus carneus (Pers. : Fr.) Korf. apud Kimbr. & Korf(Fig. 33)
1. Ascospores hyaline, ellipsoidal, evenly and coarsely granulate, granulations
larger at the poles, 25-29 x 11-14 um /odophanus verrucosporus (Graff) Kimbr. (Fig. 33)

20
Peziza Fr.

Peziza Fr. is a polyphyletic genus of more than 80 found during this study were ca 12 mm diam. The
species of operculate discomycetes whose ascus key by Richardson & Watling (1997) has been used
apex stains a distinctive blue in Melzer's reagent to identify two species. A recent paper by Hansen
(Kirk et al. 2001). A few species are found fairly et al. (2001) shows that within the genus Peziza
regularly on dung. The apothecia developed upon distinct types of ascus amyloid reactions supported
incubated dung may be very variable in shape and different rDNA lineages. The two species identified
size and this factor makes determination of the here had a distinct amyloid rim around the edges of
species difficult. For example, Rifai (1968) re­ the opercula and ascus tops (Fig. 34 B & H), which
corded P. vesiculosa on various substrates includ­ in the paper by Hansen et al. indicates inclusion in
ing "manured ground" collected from Australia, the "Group IV lineage ", along with species such as
but the apothecia were up to 80 mm diam, whereas Pezizafimeti which is also coprophilous.
the examples of what 1 consider to be this species

Key to coprophilous species of Peziza

1. Apothecia stalked, buffto ochreous (6B5-6E7), apothecial surface with

coarse uneven warts; asci 8-spored; ascospores ellipsoïdal, finely and
evenly granulate 14-18 x 8-9 um Pezlza pleurota sensu Richardson & Watling (Fig. 34)
1. Apothecia stalked, buff /ochreous (6B5-6E7), apothecial surface almost
smooth; asci 8-spored; ascospores smooth, 22-24 x 11-13 um
................................................................................................................ Peziza vesiculosa Bull. : Fr. (Fig. 34)

Note: Richardson & Watling (J 997) did not give an author citationfor this species

Family PVRONEMATACEAE

Cheilymenia Boud.

Cheilymenia is characterized as usually having cies, and Doveri et al. (2000b) published descrip­
fairly conspicuous sessile apothecia that are sorne tions of the species found in Italy.
shade of red or yellow. The apothecia are beset The genus Cheilymenia most certainly is present
with stiff bristles (setae) in ail but one species (C in Australia, and Rifai (1968) recorded five species
granulata). The ascospores are hyaline and el1ip­ from Australia, collections of which reside at the
soid. ln sorne slide preparations a transparent jacket Kew herbarium (K). To date no species of Cheily­
(perispore) can be observed surrounding each asco­ menia have developed on the dried dung sent to me
spore, but generally this is not visible in many sl ide during the course of this current study. This is due
preparations and according to Yao & Spooner to the fact that this genus generally does not appear
(1996) "il may prove to have little diagnostic value on dung that is incubated in the laboratory unless
for Cheilymenia", although other authors such as the apothecia were partly formed before it was
Moravec consider the perispore as a useful adjunct col1ected. Dade observed several species of this
to the classification process. The genus has re­ genus in Australia and left copious notes and sorne
ceived much attention by taxonomists (as compared fine illustrations, which are reproduced in this
with sorne other genera of coprophilous fungi) and publication. He stated that he only found Cheily­
as a result there are a number of keys to the species menia in the field during the months of May to
and many illustrations. Boudier established the September, and he wondered how such hyaline and
genus in 1885 listing five species with Cheilymenia relatively thin-wal1ed asco spores of such fungi as
stercorea (Wigg.) Boud. designated as the type Cheilymenia, Ascophanus (= Coprotus), and
species of the genus. Since that time, several taxo­ Lasiobolus could resist desiccation during the hot­
nomie accounts have been published including Le test parts of the Australian summer. He assumed
Gal (1953), Denison (1964), Rifai (1968), Garnundi that sorne ascospores at least did manage to survive
(1972), and Dennis (1978). More recently, Moravec in the soil and dust and thus got carried up aloft on
(1984, 1988, 1989a & b, 1990a & b, 1992, 1993) the herbage when it grew the following spring. In
published extensively on the genus, Yao & Spooner the "unusually dry" autumn of 1966 he noted that
(1996) published descriptions of the British spe­ discomycete apothecia were few in number and
those that did appear were dwarfed and did not

21
 often reach the stage of sporulation. During that with ascospores in a single series. Ascospores are
autumn, he did not see any Cheilymenia species. hyaline, ellipsoidal and smooth 15-23 x 10-}4 l'm.
He records that Cheilymenia species are character­ Paraphyses are numerous and contain the orange
istic of cattle dung "appearing only uncommonly pigment which is variable in quantily. They are 2-4
on other dungs when these are protected from um broad and enlarged at their tips, which are
desiccation ". Since this genus does not grow in subglobose and usually about 7 um broad but
incubation vessels where the relative humidity is sometimes up to 12 l'm. On dung of herbivores but
high, he concluded that Cheilymenia species prefer most luxuriant on cattle dung. On other dungs the
regimes of "wet feet and relatively dry air". apothecia are formed sparsely. "
Interestingly, Cheilymenia pallida A. Bell & Dade listed sorne 14 collections of C. copri­
Dennis (1971), a species seemingly restricted to naria. In 1982 1 surveyed a number of his slides
possum dung in New Zealand (and which readily including V147a (lMI 104359a) and V239a, both of
develops under laboratory conditions), was not which match his description of C. coprinaria.
recorded by Dade even though he scrutinized many Regarding Cheilymenia stercorea (Pers.) Boud.,
samples of possum dung. 1 cannot believe that he Dade stated that it often occurred in the company
overlooked it. Hitherto 1 had considered it most of C. coprinaria, but .. was less common and seems
likely to be a fungus unwittingly imported into to be less variable judging by the fewer collections
New Zealand from Australia together with the made here, which are closely similar. It is distin­
possum. This serves, as a reminder of how little is guished by the presence of short stellate hairs in
known concerning the distribution of fungi in gen­ addition to the large setae." These observations
eral and a warning against making premature as­ match my own made in New Zealand, where this
sumptions as to their distribution patterns. species is also less corn mon than C. coprinaria, is
Dade described and illustrated five species of less variable and often occurs together with C.
Cheilymenia. As was his habit, his descriptions coprinaria. Dade's slides V362b and V404a are
were considered and thoughtfu!. The following is a good examples of the Australian material of C.
portion of his account of Cheilymenia coprinaria stercorea. He also recorded C. raripila (Phil!.)
(Cooke) Boud. " This is a very variable species in Dennis on cattle dung and his observations were as
ail ils characteristics and in their assortment. After follows: "This species is distinguished by ils very
seeing a few collections, the student may suspect small setae which may escape notice at x 30 and by
that he has several difJerent species, but after many ils larger ascospores. The apothecia are bright
collections have been examined il is found that the orange up to 1550 um diam and 850 pm in depth,
wide range of variation is continuous for each bowl-shaped in profile when weil grown. They
feature and that variation in each is independent of develop from small spherical bodies, which often
variation in the others. The apothecia are gregari­ become oblate while a depression forms on the
ous, usually crowded closely together, especially upper surface; sparse setae cover the excipulum
on cattle dung on which they form large conspicu­ near the rim. As the body expands the collarette
ous patches by both their size and (usually) bright becomes everted and is finally about 100 um wide,
orange colour. They are commonly up to 2 mm in glistening like the rest of the exciple and composed
diameter but often reach 5 mm and 9 mm has been of bladdery cells, mostly 20-45 um diam. In old
recorded. In profile they are saucer- or bowl­ age the hymenium becomes convex and the col­
shaped, attached to the substratum by a small base. larette flat. The setae range from 130-240 um in
A hyaline collarette ahout 50-60 um wide sur­ length, with simple bases, thin- walled, with few
rounds the hymenium, composed of cells of pris­ septa (0-6) and very pale, sometimes with a slight
matie texture which become swollen in age. A trace of colour. Ascospores are hyaline, ellipsoidal
fringe of setae projects from below the collarette. and smooth in a single series, (21-)23-26 x 11-17
These setae spring from the exciple and are usually l'm. Paraphyses about 3 um broad, clavate at the
stout, thick-walled and deeply coloured brown but tips, where they are 5-10 um broad, containing
may be smaller and paler; their length varies from orange pigment." Dade's slide V218b is a good
250 to nearly 1000 um and they have up to 18 example of C. raripila. Cheilymenia theleboloides
septa, rarely none. The apothecia develop from (Alb. & Schw.: Fr.) Boud. was also recorded by
small spherical bodies which become obconical Dade. He expressed sorne difficulty in assigning a
with the collarette weil formed at an early stage specifie name to these particular collections and he
and with vertical or even converging setae. As the clearly conferred with his colleague Jean Gunnell
dise expands, the collarette and setae assume a at Kew over this particular species. Although 1
horizontal posture. In age, the hymenium becomes could find no copies of any letters between them,
convex and the collarette, now with swollen ceIls, there are sorne fine drawings of Cheilymenia pul­
becomes everted. The hymenium is eaten avidly by cherrima (Crouan & Crouan.) Boud. accompanying
springtails. Asci are at first short and broad cylin­ the drawings of his C. theleboloides with the note
drical when mature and then 165-215 x 10-20 um
22
 "from Crouan 's specimens ", indicating that this smooth, and certainly Dade drew them as smooth.
was also a specifie name considered here. Certainly Dade recorded Cheilymenia granulata (Bull. : Fr.)
the description and illustrations of C. pulcherrima Moravec a few times. His slide V392a is a good
in Moravec (1992) have a striking similarity to example of this species and the ascospores on this
those of C. theleboloides in Rifai (1968) and Yao & sIide preparation do exhibit a wrinkled perispore.
Spooner (1996), although there are sorne small size Ali the species of Cheilymenia observed by him are
differences recorded for the ascospores of the se keyed below. Rifai also recorded C. coprogena
two species by these authors. Additionally, Mo­ (Berk. & Broome) Rifai from dung collected in
ravec (199ûb) describes the ascospores of C. thele­ Queensland.
boloides as having a fine ornamentation of parallel
punctation revealed when they were subjected to
SEM photography. Other authors describe them as

/
fl---'
/
\ ".
.

-,
" . . '
(
, .

apotneciurn ernarqtnate

'\
/
/
/,
J,

perispore

apolhecium marginale
bulbous bases
10brislles

rooling bases
10 bristles

Fig. 6. Glossary to terms used in the key below.

Key to the species of Cheilymenla

1. Apothecia gregarious, apricot/umber ca (SCS-SDS), emarginate, without bristles;

ascospores hyaline, ellipsoidal, 13-17 x 7.5-9 um ......Cheilymenia granulata (Bull. ; Fr.) Moravec (Fig. 37)
1. Apothecia gregarious or scattered, various shades ofyellow/orange/red, marginate, with bristles 2

2. Bristles conspicuous, robust, brown (SES), with rooting bases 3

2. Bristles inconspicuous, hyaline or pale yellow (4A4), with simple bulbous bases 4

3. Stellate hairs present on the apothecia; ascospores hyaline, ellipsoidal, 20-22 x

10-12.5/lm Cheilymenia stercorea (Wigg. : Fr.) Boud. (Fig. 36)
3. Apothecia without stellate hairs; ascospores hyaline, ellipsoidal, (15-) 16-23 x
(9-)10-12(-14) um Cheilymenia coprinaria (Cooke) Boud. (Fig. 36)

4. Ascospores hyaline, ellipsoidal, 16-22 x 1O-12/lm

Cheilymenia theleboloides (Alb. & Schw. : Fr.) Boud. (Fig. 35)
4. Ascospores hyaline, ellipsoïdal, 25-2S x 15-1S um Cheilymenla rarlplla (Phi Il.) Dennis (Fig. 35)

Note: Cheilymenia granulata was described as "gingy orange" by Dade

23
Coprotus Korf & Kimbr.

Sorne of the 19 described species of Coprotus (Kirk et Dade also recorded a few Coprotus species, which
al. 2001), are found upon dung. Many of these appear he referred to by the earlier generic name Ascophanus
after the dung has been incubating for sorne time. Boud., although he also knew that the genus was about
Characteristics of the genus include small, mostly to be changed. He made notes on three species: "Asco­
sessile, pale apothecia; white, yellow and orange being phanus 16-sporus ( C. sexdecimsporusf", which he
the predominant colours encountered. 1 do not find described as having white apothecia and ascospores
apothecial colour a particularly reliable feature with "about 12 x 7 um"; Ascophanus argenteus: "with
which to define the species. Apothecia may occur in ascospores similar to those of C. lô-sporus" and
groups or develop singly. The ascospores are hyaline, "another sp. not identified has spores }4-16 x 6-7
ellipsoid, smooth and most develop a conspicuous de um; otherwise like A. argenteus ". 1 have not been able
Bary bubble especially when mounted in a semi­ to find his slides of the latter two species, but two
permanent mounting medium such as Shear's mount­ slides (V 128q and V489b) that 1 have seen are of
ing medium. A key to the North American species was Coprotus sexdecimsporus.
developed by Kimbrough et al. (1972). Jeng & Krug Cribb (1998) recorded C. duplus, C. lacteus and C.
added to the known species (1977) as did Gené et al. niveus. [ have distinguished six species from Australia
(1993). Aas (1983) described the Norwegian species, during the course of this study. The most common
Wang (1999) produced a key and descriptions to the species encountered (from 31 dung collections) has
Taiwanese species, and Doveri et al. (2000b) described caused sorne problems regarding its identification. 1
the species found in Italy. Not all taxonomists accepted am referring to it as Coprotus cf. disculus Kimbr.,
the generic name of Coprotus (see, for example, Luck-Allen & Cain. The information presented in the
Pouzar & Svrëek 1972). Taxonomie arguments around scatter diagram (Fig. 38) illustrates the problem. The
this matter are discussed by Aas (1983). A number of ascospore dimensions straddle the given measurements
ambiguities appear in sorne of the publications; for for three species. The apothecia may be white, dilute
example the ascospore measurements of C. breviascus yellow or dilute orange (3A3-7A4). The paraphyses
in the key by Kimbrough et al. (1972) differ from are branched and frequently uncinate at their tips. For a
those given under the description of that species. Aas time 1 considered that there were two species involved
(1983) refers to C. breviascus as perhaps being syn­ here: C. disculus and C. leucopocillum, but as the
onymous with C. ochraceus. These comments together collections grew, my ability to distinguish between
with the fact that many of the species seem so closely them waned. At the extremities of the ascospore size
drawn and the apparent lack of much supporting cul­ range, the ascospores look quite different as the scatter
ture work with this genus lead me to believe that this is diagram shows, but in fact the majority of the asco­
yet another in need of revision. spores fall somewhere between.

Key to the species of Coprotus

1. Asci containing 8 ascospores 2

1. Asci containing > 8 ascospores 5

2. Ascosporesbiseriate/multiseriate. hyaline, ellipsoidal, (10-)11-15 x (6-)7-10 um;

paraphyses frequently uncinate and branched at their tips,
............................................................................. Coprotus cf. dlsculus Kimbr., Luck-Allen & Cain (Fig. 38)
2. Ascosporesuniseriate; combination ofascospore characters not as above 3

3. Paraphysesuncinate; ascospores hyaline, ellipsoidal, 7-8 x 4-5 um .

........................................................................................................C, baeosporus Jeng & lC. Krug. (Fig. 39)
3. Paraphyses more or less straight or sinuous; ascospores larger. 4

4. Ascosporeshyaline, ellipsoidal, 9-10 x 6-7 um C luteus Kimbr., Luck-Allen & Cain (Fig. 37)
4. Ascosporeshyaline, ellipsoidal, 14-16 x 8-9 um C ochraceus (Crouan & Crouan) Moravec (Fig. 39)

5. Asci containing 16 ascospores, 11-14 x 7-10 um C sexdecimsporus (Crouan & Crouan) Kimbr. (Fig. 40)
5. Asci containing > 64 ascospores, 10-11 x 6-7 um C winteri (Marchal) Kimbr. (Fig. 40)

24
Pseudombrophila Boud.

Pseudombrophila Boud. (syn: Fimaria Velen.) is a excellent illustrated account of his findings, 1 consider
genus of predominantly coprophilous discomycetes. it valuable to include here. The rabbit dung was sample
The apothecia are sessile or short-stalked, often in a V726, and there are 6 good slides of it stored (along
shade of umber, isabelline or fawn (5B6-7D6), with aIl of Dade's notes) at the herbarium (DAR) at
with/without vinaceous margins (11D5). They fruit Orange. ln 1982 1 studied one of his slides (V726 a).
infrequently and since for the most part the apothecia He identified his specimen as Pulvinula cinnabarina
range from 0.5 to more than 20.0 mm in diam, they are (Fuckel) Boud. His description is as follows: " The
unlikely to be overlooked. They are seldom recorded apothecia begun as pinking spheres surrounded by a
in the literature although a new species was described dense growth of radiating hyphae, the latter being the
by Jeng & Krug (1977), and P. virginea Svrëek & usual accompaniment of many fungi developed in
Moravec was obtained in culture by Fort & Guarro humid dishes. By the s" day of incubation the summits
(1988). Van Brummelen (1962) described and illus­ of the spheres ruptured, invagination ensued and the
trated 4 species under the earlier generic name of apothecia rapidly expanded, ending as dises or
Fimaria. Pfister (1984) produced a key to nine species plaques with lobed edges in projile like shallow dishes,
(also under the generic name Fimaria) and in a recent supported on short broad columnar bases. The invest­
monograph 19 species are recognised (van Brummelen ing hyphae disappeared from the exterior of the dies,
1995). but persisted as a grayish felt on the bases. The ex­
1 have yet to find any species ofthis genus on Aus­ cipulum was cream-coloured with pale brown fiecks.
tralian material, but Dade did and he left good slides, Hymenium fiat, becoming convex, light orange at jirst,
drawings and written descriptions of his finds. In 1982 becoming tan colour (alutaceus), jinally cinnamon
1 studied his slides (V213h, V798b, and V851c). At (cinnamomeus) as viewed in strong daylight with the
that time he called the fungus Leucoscypha sp. With­ naked eye, brighter at x30 in strong spotlight". The
out seeing fresh or dried material, it is hazardous to ascospores are uniseriate, 8 per ascus glo­
place a specifie name on Dade's specimens, but his bose/subglobose, hyaline. smooth, 11-10 x 11-12 um,
descriptions of the apothecial form and colour (repro­ collapsing on one side in lactophenol but not in water.
duced below) together with the size ranges of the While the hymenium was young, a few spores con­
ascospores (14-18 x 8-10 um) lead me to consider that tained de Bary bubbles, but these were not again seen
the species was P. theioleuca Roll. His description is and have no taxonomie signijicance. Paraphyses jila­
below and his illustrations appear in Fig. 42. mentous, 2-3 um broad, septate usually not at ail
"Young apothecia are conical, almost white except infiated at their tips, though the terminal cell or one
at the apex, which is reddish brown and from the near the apex was very slightly broader and more
outset there is here a small, clearly dejined circular deeply staining". RA.D., Feb. 1968.
depression. As growth proceeds, the upper part occu­ Dade's illustrations of Pulvinula cinnabarina are
pies most of the upper surface. The hymenium is now reproduced in Fig. 41. This (and other species of the
coloured very pale dull yellow and the edge of the genus) is to be found in Rifai (1968). Korf & Zhuang
surrounding exciple is fiecked with brown, this ap­ (1991) discuss Pulvinula and describe a new species,
pearance being produced by the expansion and break­ P. subprolata Korf & Zhuang.
ing up of the original outer cells of the exciple. In
projile the mature apothecium has the form of an urn
or egg eup, i.e. a bowl-shaped upper part on a nar­ Family THELEBOLACEAE
rower foot. In some collections it is campanulate and
the brown colour may extend ail over the surface and Ascozonus (Renny) E.C. Hansen
persist, but more usually most of the apothecium re­
mains pale and the effect on the naked eye ofthe whole Ascozonus is considered to be a member of the Thele­
body produces a pale cinnamon colour. At ail times the bolaceae. Van Brummelen (1998) described the ascus
apothecia are smooth and are extremely elegant in tip in considerable detail and Landvik et al. (1998)
their form In age the hymenium becomes convex and discuss its position as demonstrated by phylogenetic
the surface of the hymenium becomes distinctly white techniques. Ascozonus woolhopensis (Renny) E.C.
and scurfy The whole body is now much darker." Hansen is a fungus that forms its ascomata in the labo­
Discomycetes 3, Dec. 1968. ratory on dung only collected during the winter months
as recorded by several mycologists (e.g. Larsen 1971,
Bell 1975, and M. Richardson pers. comm.). 1 have not
Pulvinula Boud. seen this species on any of the Australian dung sam­
pIes incubated thus far. Dade recorded A. woolhopen­
Pulvinula is not generally considered to be a coprophi­ sis on sorne nine dung collections (on ringtail possum,
lous genus, but since Dade found a good sample of this brushtail possum, rabbit, cow, bush rat, and wallaby
genus on rabbit dung collected by Crichton and left an dung). He also noted that it was a fungus restricted to
25
fruiting during the winter months. He described the which were based upon the ascospore numbers. Wick­
apothecia as "brilliantly (almost bluish) white, with a low & Malloch (1971) started sorting out the problems
hairy fringe to the apothecia". He also recorded the of species definition in Thelebolus experienced by
ascospores as being yellowish when mature. He did Kimbrough & Korf, by culturing isolates with 8, 32,
not leave any details of the ascospore measurements of 64, 128, 256 and >500 ascospores per ascus. They
his material, but his illustrations drawn to scale c1early stated that for ail these strains the ascospore number
indicate that the species he saw was A. woolhopensis per ascus remained constant, but they did not connect
(asci containing 64 ascospores, ascospores about 12­ these findings to any type material of described spe­
13 x 3-5 1ll11). His drawings are reproduced in Fig. 41. cies. Thus from the taxonomie point of view, the prob­
lems still remain. In his presidential address to the
Mycological Society of America, Kimbrough (1981)
Mycoarctium K.P. Jain & Cain c1arified sorne of these issues and stated that Thelebo­
lus crustaceus (Fuckel) Kimb. has 64 spores per ascus
Mycoarctium was established as a new genus in the and T polysporus has 256 spores per ascus. This is at
family Thelebolaceae by Jain & Cain (1973). The odds with the account of the genus by Domsch et al.
genus is characterized by having sessile apothecia (1980) who, based on Kimbrough & Korf (1967), state
beset with thick-walled aseptate bristles, which in M that T polysporus is 128-spored. Other literature (see
ciliatum (the type species) are hooked (uncinate) at Kimbrough & Korf 1967) list additional species de­
their apices. The ascospores of M ciliatum K.P. Jain & fined by the number of asci per ascocarp in addition to
Cain are hyaline to yellowish brown (12-) 13-14(15-) the number of ascopores per ascus. As stated by
x 8-10(-11) urn and ornamented with a regular pattern Domsch et al. (1980), there still needs to be critical
of coarse, raised reticulations. The species is weil studies of these fungi to determine the stability (or
iI1ustrated in Jain & Cain's article and again in a paper otherwise) of ascus number and ascospore number
by Korf & Zhuang (1991). To date this is the only within each species, since there is little to separate
species of the genus and it has been found on dung in them on ascospore size. Regarding broader taxonomie
Colorado (USA) and Tenerife (Canary Islands). Cur­ concepts, utilizing a culture of Thelebolus stercoreus,
rently, Mycoarctium is thought to be closely related to Landvik et al. (1998) show that this species is closely
Lasiobolus (Korf & Zhuang 1991) although the asci related to Ascozonus woolhopensis and place these two
appear to be non-operculate. species in the family Thelebolaceae. Van Brummelen
In 1982 1 collected a sample of koala dung from the (1998) described ascus dehiscence in a number of
koala enclosure in Sydney. When incubated, the dung species of Thelebolus and related genera.
yielded fine specimens of a second species of My­ In his notes of 1966, Dade stated that Tode's genus
coarctium bearing globo se, reticulate ascospores 9-11 was an "ambiguous generic name - nobody can tell
um in diameter. Due to the nature of these ascospores, what fungus Tode had". Elsewhere Dade wrote: "It ls
1 am calling this species Mycoarctium sphaerosporum unfortunate that the name Thelebolus has been perpe­
n. sp. It is iIIustrated in Fig. 31 A-F, (Appendix 1). trated, for Tode 's description is quite inadequate and
unrecognisable, but has been made respectable by
Kimbrough 's emendment to fit it to the fungus ac­
Thelebolus Tode and Trichobolus (Sacc.) Kimbr. & cepted by Zukaî". Dade described two species of
Korf Thelebolus. which he recorded on dung: Thelebolus
stercoreus (Tode : Fr.) Kimbr. and Thelebolus micro­
Heinricus Julius Tode (1790) established the genus sporus (Boud.) Brumm.; the latter he referred to as
Thelebolus naming T stercoreus as the single species. Ascophanus microsporus (Berk. & Br.) E.C. Hansen,
The genus consists of number of psychrophilic species or its synonym Ascophanus subfuscus Boud. Consider­
with extremely minute ascomata (about 200 um diam), ing morphological features alone, T microsporus is
sorne species of which appear as c1eistothecial to the quite dissimilar to the other species that Kimbrough
uninitiated but are deemed to be modified apothecia (1966) placed within his emended genus Thelebolus.
(Kimbrough 1966, Kimbrough & Korf 1967). Consid­ Rather this species superficially resembles species of
erable confusion has occurred over the intervening Orbilia having clearly apothecial, translucent, apricot
years as to the circumscription of Tode's genus and to sienna ascomata, furnished with strongly capitate
still there resides sorne disagreement as to sorne of the paraphyses and numerous 8-spored asci. Corte et al.
solutions promulgated by Kimbrough & Korf (e.g. (1993) described T microsporus as psychrophilous,
Svrëek 1972). Otani & Kanzawa (1970) described and preferring a temperature of 15-20°C for growth; they
iIIustrated Thelebolus crustaceus (Fuckel) Kimbr. and listed one of Dade's collections (lMI 108525c) on
T polysporus (P.Karsten), Otani & Kanz. but did not sheep dung collected on Macquarie Island and another
mention the numbers of spores per ascus for either of (lMI 104303b) from wombat dung collected from
these species. This is unfortunate, since, as stated by Warrenbayne. 1 studied another portion of the wombat
Kimbrough & Korf, a number of species have been collection (V 150b) in 1982 together with sIides V 132c
assigned to the older synonym Ryparobius Boud. ail of and VI 32ffrom possum dung. My own two Australian
26
 collections of this fungus have been from wallaby his notes that the species (which he called T. zukalii)
dung collected in Victoria. Ali these specimens match preferred possum dung (11 collections) but that he
the description of T. microsporus. The slide V 192e had also collected it on macropod and rabbit dung
labelled "Rhyparobius polysporus" collected from (one collection of each). The latter two collections he
rabbit dung from Mount Sterling ("alpine flora") is recorded as being atypical in lacking setae. One of
what 1 have been referring to as Thelebolus stercoreus. them was subsequently cultured on sterilized possum
Kimbrough & Korf (1967) raised Saccardo's subge­ dung where it "produced setae on normal ascomata".
nus Trichobolus to full generic rank as Trichobolus Jean Gunnell, Dade's colleague at Kew, evidently
(Sacc.) Kimbr. & Korf. The genus is characterized by planned to publish details of this Australian Tricho­
its minute ascomata (ca 200-500 um diam), beset with bolus species and he wrote to her (23 rd Nov. 1966),
hyaline, sparsely septate bristles (setae). The number presumably suggesting that she should wait until the
of ascospores per ascus ranges from 8 to > 1000 be­ delayed Kimbrough & Korf publication appeared in
tween the described species (Kimbrough & Korf 1967, the American Journal of Botany in 1967. However,
Krug 1973). The measurements that Kimbrough & Gunnell apparently never pub lished her account of
Korf give for their three species of Trichobolus are as the Australian Trichobolus after the Kimbrough &
follows: Korf paper appeared.
T. zukalii - Ascospores 9.7-10.5 x 7-7.6 um New Zealand collections of Trichobolus had asco­
T. pilosus - Ascospores 10.5-11.2 x 7-7.4 um spore measurements within the range 7-12 x 7-10
T. sphaerosporus - Ascospores 9-9.6 x 8.4-9 urn um, i.e. measurements that straddled across a1I three
species of Kimbrough & Korf (Bell 1983). 1 identi­
Other differences between these three species are fied these collections as T. sphaerosporus Kimbr. and
the presence (or absence) of de Bary bubbles and the manifestly disregarded the minute size differences
number ofasci per ascocarp (1 to 3), as noted by Dade: between the three species as stated by Kimbrough &
"This (= the limitation of asci to 1-3 per ascocarp), is Korf. Ascomata of the New Zealand collections con­
unfortunate as our Australian fungus has many more tained 1-2 asci as opposed to the one ascus stated by
asci per apothecium ". Elsewhere Dade writes: Kimbrough & Korf for that species. Australian mate­
"Kimbrough separates the three spp. on very tenuous rial that 1 have examined during this study exhibited
grounds, i.e., number of asci per apothecium (1-3), ascomata containing both one and two asci. For these
really slight differences in spore dimensions and in one reasons, 1 too have doubts as to the veracity of
case the presence of de Bary bubbles, which 1 cannot Kimbrough & Korf's species distinctions of Tricho­
accept as a reliable character. (These were seen in the bolus. To date, there have been no further publica­
dry material ofT. pilosus). 1 strongly suspect that there tions on Trichobolus that might elucidate these ap­
is in fact only one species, T. zukal ii ". parent anomalies presented by the New Zealand and
The Australian material of Trichobolus certainly Australian collections, thus 1 have decided to name
bear ascomata which contain many more than 1-3 the Australian material that 1 have also collected as T.
asci, and Dade's drawings c1early show ascomata zukalii until further information should prove other­
with up to ten mature and immature asci. He stated in wise.

Key to the species of Thelebolus and Trichobolus

1. Ascomataclearly apothecial, nail-shaped, apricot/sienna (6A7-688);

many 8-sporedasci; ascospores hyaline, ellipsoidal, 7-10 x 3-5 um; paraphyses
stronglycapitate, heads bright yellow (2A8), Thelebolus microsporus (Berk. & Br.) Kimbr. (Fig. 44)
1. Ascomatanot as above, appearing as cIeistotheciawhen young;
asci multispored; paraphyses not capitate 2

2. Ascomatabeset with hyaline mostly non-septate bristles, containing 1

enormous (Le. > 200 um long), multispored ascus, or up to 12 multispored
asci; ascospores hyaline, globose, 7-12 x 7-10 um Trichobolus zukalii (Heimerl) Kimbr. (Fig. 43)
2. Ascomatawithout bristles, ascospores not as above 3

3. Ascomatacontaining 1 enormous (approx. 200 um long), ascus containing > 500

hyaline, ellipsoidal, ascospores, 5-7 x 3--4 IAm Thelebolus stercoreus (Tode. : Fr.) Kimbr. (Fig. 44)
3. Ascomatacontaining several subglobose asci, each containing > 64 ellipsoidal,
hyaline, thick-walled ascospores, each about 6 x 4 IAm
...................................................................... Thelebolus polysporus (P. Karsten) Otani & Kanzawa (Fig. 43)

Note: Trichobolus is now considered to belong to the family Ascodesmidaceae.

27
Cleistothecial Ascomycetes

Ascomycetes characterized by non-ostiolate more or The minute (ca 200 um diam) ascomata of certain
less globose ascomata used to be placed together in the species of the discomycete genus Thelebolus may be
class Plectomycetes. The artificial nature of the Plec­ considered to be cleistothecia by uninitiated observers.
tomycetes has been suspected for many years, and Chaetomium (most species of which are perithecial),
according to Malloch (in Reynolds 1981), Bessey could easily be mistaken for being cleistothecial due to
(1935) was one of the first to question its veracity in the dense vestiture of appendages that clothe the asco­
print. Later, Cain (1956) and Malloch & Cain in many mata, effectively obscuring the ostioles. Malloch (in
papers published in the early 1970's, expanded upon Reynolds 1981) drew attention to the progression of
these ideas. Malloch fully documented and consoli­ the genus Chaetomium towards the cleistothecial con­
dated his thoughts in a comprehensive paper entitled: dition. For example, the asci are scattered throughout
"Plectomycetes and their anamorphs" (in Kendrick the centrum and the ascospores are not forcibly dis­
1979) and in another entitled "The Plectomycete Cen­ charged. Sorne species of Chaetomium can also have
trum" (in ReynoldsI981). A further detailed history of cleistothecial ascomata (e.g. Chaetomium irregulare).
the Plectomycetes can be found in Benny & Contrary to the treatment of the genera Sporormiella
Kimbrough (1980). They reason that the cleistothecial and Preussia by Valldosera & Guarro (1990), von Arx
habit results from selection pressures, which favour et al. (1986) choose to continue to include the cleisto­
passive dispersal of ascospores as exemplified by thecial species of Chaetomium together with perithe­
species that inhabit soil. It is now universally recog­ cial species under the perithecial taxon Chaetomium.
nized that the Plectomycetes as originally conceptual­ Many orders and families have yet to be subjected to
ised was a highly artificial assemblage and that the sequencing techniques, thus at present there is a com­
non-ostiolate habit has evolved several times within plete mix in the literature of the traditional systematics
diverse families (Kirk et al. 2001). For sorne mycolo­ using morphological techniques (which did not always
gists the solution to this is to gradually unite various distinguish between the results of parallel evolutionary
cleistothecial taxa with their perithecial (or apothecial) forces from derived characters), alongside cladistic
counterparts. Thus Valldosera & Guarro (1990) pro­ analyses.
pose the gradual disestablishment of the genus One of the principal functions of this current publi­
Sporormiella Ellis & Everh. 1892 in favour of the cation is to make identifying Australian coprophilous
earlier cleistothecial genus Preussia Fuckel 1868. ascomycetes a relatively simple task, thereby encour­
Similarly, the perithecial genus Trichodelitschia is aging more people into the study. For this reason,
generally placed in the cleistothecial family bringing together ail cleistothecial ascomycetes under
Phaeotrichcaceae due to its great similarity with the the old artificial class Plectomycetes serves a very
genus Phaeotrichum (Cain 1956, Lundqvist 1964). practical function since ascocarp morphology is the
Berbee & Taylor (1992) propose the reinstatement of first most visible feature observed. By placing together
the old class names Plectomycetes and Pyrenomycetes the illustrations to various of the genera one can more
but the circumscription of these names would be alto­ readily appreciate the diversity of ascospores (and
gether different from the traditional notions of these hence the artificiality) of the plectomycetes rather
taxa. more than from an abstruse description of the same.
With the development of techniques employing However, in instances of monographie works, uniting
DNA analysis, attention has been given to deciphering cleistothecial species with their natural perithecial and
the natural relationships of old plectomycete genera. apothecial counterparts would seem to be a more intel­
Thus Suh & Blackwell (1999) show that the cleistothe­ lectually sound practice.
cial Pleuroascus nicholsonii is related to the inopercu­ Only ten cleistothecial fungi have been recorded
late discomycetes and that Nigrosabulum globosum is during this study. Scrutiny of dung from other than
related to the Hypocreales. Formerly these two fungi herbivores will most certainly yield further species.
were placed in the Onygenaceae and the Cephalothe­ Dade also recorded Gymnoascus reessii (siide no.
caceae, respectively. The same paper also confirms VI72n) and Nigrosabulum globosum, a fungus new to
that the (mainly) ostiolate genus Chaetomium is natu­ him (siide VI83k).
rally placed within the Sordariales.

28
Key to the c1eistothecial Ascomycetes (see aiso the discomycete genus Thelebolus)

1. Cleistothecia (gymnothecia) composed of alose network of interwoven hyaline

hyphae through which the globose asci can be readily seen 2
1. Cleistothecia with a well-defined peridium composed of textura globosa or textura angularis 3

2. Cleistothecia reaching 100 um diam, often confluent, pinkish white (9A2-3),

peridial hyphae soon disappearing; ascospores ochreous (SB6), circular and about
um diam including an outer rim in face view, elliptical and ca S um in side view,
rim extending either side Pseudoarachniotus roseus Kuehn (Fig. 4S)
2. Cleistothecia reaching 400 urn diam, apricot (6A7) in colour; peridial hyphae
persistent, robust, orange, sorne furnished with single or double blunt hooks;
ascospores hyaline, circular in face view, ca 4 um diam, elliptical in side view
ca 2-3 um Gymnoascus reessii Baranetsky (Fig. 4S)

3. Cleistothecia white; peridium of textura globosa; with scattered long hyaline,

flexuous peridial hairs; ascospores uniseriate, hyaline, more or less globose,
11-12 x 9-10 um, often with de Bary bubble Lasiobolidium orbiculoides Malloch & Benny (Fig. 46)
3. Cleistothecia black, ornamentation absent, ascospores not as above 4

4. Cleistothecia with conspicuous hairs S

4. Cleistothecia without conspicuous hairs 8

S. Cleistothecial hairs dense, entangled and even over the exposed portions of the
ascocarp, minutely spiny ca 2 um wide; ascospores olivaceous, very variable in
shape: limoniform/ovoidl broadly lunate to irregular, 14-16 x 8-10 um
.......................................................................................................... Chaetomium irregulare Sôrgel (Fig. S3)
S. Cleistothecial hairs and ascospores not as above 6

6. Cleistothecial hairs few, hyaline, smooth, tightly spiralled; ascospores rose/red

(lIAS) in mass, individual1y almost hyaline, spherical, 2-3 um diam, smooth or
irregularly & minutely granulate Pleuroascus nicholsonii Massee & Salmon (Fig. 47)
6. Cleistothecial hairs dark, sinuous not spiralled 7

7. Cleistothecia globose; robust, dark, septate hairs evenly scattered; ascospores

consisting of a dark ellipsoidal cel1 with truncate base, 9-10 x 7-8 um and a
hyaline pedicel 12 x 6 um, soon collapsing ..... Zopfiella erostrata (Griff.) S. Udagawa & K. Furuya (Fig. 49)
7. Cleistothecia ovoid, ellipsoid or irregular; robust, dark hairs emerging as two
opposing tufts; ascospores rose-red (lI AS) in mass, ellipsoidal to ovoid,
ca S x Jum Kernia nitida (Sace.) Malloch & Cain (Fig .46)

8. Asci spherical; ascospores hyaline, globose ca 3-4!lm diam, smooth, often with
minute surface debris Nigrosabulum globosum Malloch & Cain (Fig. 47)
8. Asci cylindrical, long- or short-stalked; ascospores 4-celled, cells readily separating
into I-celled units 9

9. Asci long stalked, ascospores cylindrical with tapered ends, 26-30 x 6-9 um
....................................................... ............................................Preussiafuniculata (Preuss) Fuckel (Fig. 48)
9. Asci short stalked, ascospores separated into I-celled components within ascus
ascospore portions short cylindrical 4-S x 3-4 um Preussia cylindrica Malloch & Cain (Fig. 48)

Note Cultures of Pleuroascus nicholsonii (CBS 110040) and Nigrosabulum globosum (CBS 110041) have been deposited at
the Centraalbureau voor Schimmelcultures.

29
 To use this key compare malure ascospores of the specimen with the groups iliustrated. Upon
making a choice proceed to pages indicated where the keys to the species are located.

::0 . lBfi;
'.
~.
'~.::.
',~ ,- . " "
".

t "")
,,'"
"<t''I,~· :·,":w:,r,
ùD DGD -l-1-~
00
o@o
perilhecia ,

Selinia(p. 33)
Ophiostoma (p 32)
Sphaeronemella (p 31)
GO
Phomatospora(p. 49)
Melanospora(p. 32)
perilhecium

->.
[

\j
"J
)
D
. - - perithecium Petriella (p. 31) Chaetomium(p. 35)

perilheCi,um ~'--

lie
;;
/--;:/'

AC",
~"~
I!r. "..,
,f
\"\
-, <~.;/
\
1
-­
200' 1'"tm
,.{
((
\\.."\\
Î
~ ,
\ \
\}:) Fimetariella (p. 00)
Sordaria (p. 38) \ \
'\

•
\.
\
•
li \ ·.li, \
r~.)

6 ,t i
/i7
1mm ~

" ~~~t \" ":::,

~~
peiithècium
Arnium (p. 47)
Cercophora(p. 41) :/
. ./'
Strattonia (p. 48) Bombardioidea(p. 49) Zygopleurage (p. 49)

. ~:-::~-:2J
.

~ .11
/

'-0
t :

A.t, ..
~/ .,," ------.
Coniochaeta (p. 40)

1 l'"
\t
• 1

F' j ûetitscnie(p. 52) ,

\.

...... ~./
Podospora(p 43)

') ~) ,• Trichodelitschia(p. 57)

11
, Semidelitschia(p. 57)

.- -- ,.,...
~. l',.r/
'"

"Î 5=-0
1

L;<~7';),.'
~~"
.;--"
;i~'!b-
, ~ '
-1"
•
,lA' •
i ·
.a
•

A
H
t'\!1
\
~
t , .'-= .. , CffJ
A piosordaria
(p. 43)
r~fJ

• ,i ,,'

1 .
1
i '~'"
1 !
Sporormia & Sporormiella (p. 53)
Pyxidioohora
(p. 31)
D
Hypocopra, Podosordaria & Poronia (p. 50) \ .'

Zygospermella (p. 49)

Fig. 7. Characters ofascospores and perithecia ofpyrenomycetous genera

30
Pyrenomycetes connected with that of their arthropod dispersal agents.
Dade found at least three species of Pyxidiophora and
The tenn: pyrenomycetes is employed here in a non­ he left fine illustrations, which are reproduced in Fig.
technical way to include ail Ascomycetes which have 50. In 1982 1 studied Dade's slides: V 790f, V902b,
flask-shaped ascomata opening by means of an ostiole. V944c and V430IIc (illustrated), which are ail P. arven­
For the purposes of this manual, the inclusiveness of sis (ascospores 40-50 x 3-5 um). His slide VI84f (IMI
this tenn is extremely useful since the mature ascomata 108523f) (Fig. 50 B), could not be identified to species.
as presented on the dung rarely give any elues to the The ascospores of this specimen measure 17-19 x 3-4
uninitiated observer as to their ontogeny. Thus 1 am um, with 6 um appendages at one end (Fig. 50 C). A
including both perithecial and pseudothecial species further unidentified species of Pyxidiophora iIlustrated
here. It is now universally acknowledged that sorne of by him appears in Fig. 50 D-E.
them are closely related to cleistothecial species so that
families should include both cleistothecial and perithe­
cial genera. 1 have arranged the pyrenomycete copro­ Order MICROASCALES
philes within their currently known family groupings.
Within the families 1 have arranged genera according to Family MICROASCACEAE 13
their morphological similarities, for example Sphaero­
naemella next to Ophiostoma, Cercophora next to Petriella Curzi
Podospora. The illustrations of these pyrenomycete
fungi follow the same principle. This organization has The genus Petriella Curzi was revised by Barron et al.
allowed me to place the illustrations of certain ambigu­ (1961). P. sordida (Zukal) Baum & Gilman has been
ous species, such as Podospora australis, P. gigantea found during the course of this study and isolated in
and P. petrogale, in close proximity to other genera culture by D. Mahoney. The ascospores genninated
which they resemble. This is in anticipation that further within 24 h. It freely produced both its Graphium and
research may cause these problematic species to be Scedosporium anamorphs in culture together with the
placed in other genera. Unlike discomycetes, pyreno­ more slowly developing teleomorph. The ascospores of
mycete genera often have similar ascomata but very P. sordida are light brown (5B3), fusoid/ellipsoid in one
diverse ascospores. This makes the ascospores useful view, tlattened along one side in another with rounded
characters by which to distinguish genera with relative apices, 9-12 x 5-6 urn, containing several small gut­
ease and speed. Hence a picture key to the pyrenomy­ tules at maturity (Fig. 52 A- F),
cete genera appears in Fig. 7. The reader compares the
unknown pyrenomycete with the taxa illustrated. After Sphaeronaemella P. Karst.
deciding the illustration that most closely matches the
specimen the reader should turn to the pages indicated This is a small genus of the Microascales comprising
under the drawing(s) where further keys may be located. sorne 1-5 species (Kirk et al. 2001). Sphaeronaemella
fimicola Marchal is a coprophilous species of the genus
which 1 have found regularly on dung samples from
Order PYXIDIOPHORALES New Zealand, but found once once from dung collected
in Australia. This was from koala dung collected in
1981. This sample was incubated only a few days after
Pyxidiophora Bref. & Tavel it was collected. Apart from making a slide and identify­
ing it, 1 did not make detailed drawings of the fresh
Pyxidiophora is a genus of sorne 15 species of minute material at that time. Dung collections dried and for­
pyrenomycetes. Lundqvist (1980) described and illus­ warded to me in New Zealand for this study failed to
trated a number of species. According to Kirk et al. produce this species, perhaps indicating that it does not
(2001), they are mainly coprophilous, although Black­ respond weil to long periods of desiccation. Dade re­
weil (1994) states that one can find this genus "almost corded Sphaeronaemella from Victoria noting that it
any place we look". Over the years there has been much occurs "after rain in July-August and was probably a
speculation as to where this genus fits into the taxo­ species which fruits in the cooler seasons". The illustra­
nomic hierarchy. Eriksson & Hawksworth (1993) place tions (Fig. 51 I-K) of this species are from Dade's
this genus in the non-mycelial Laboulbeniales. Black­ notebooks. He noted that the perithecia were "bright
weil (994) states that cladistic studies indicate the yellow" with long necks and the discharged ascospores
Pyxidiophora lineage resides outside the main body of are held in a "globule at the fimbriate tip of the perithe­
the pyrenomycetes and lies closer to the loculoascomy­ cial neck", In 1982 1 studied his slide (V194e =IMI
cetes and discomycetes. Pyxidiophora is quite unlike 10961 le) ofthis species.
any other perithecial fungus found on dung. The minute
(ca 80-100 J.UTl diam) translucent and transient perithe­
cia appear early during the incubation period and when
the dung is rather moist. Their Iife cycle is strongly 13 The c1eistothecial genus Kernia also belongs here.
31
 Shaeronaemella fimicola has been subsequently and including in it a number of fungi now relegated to
shown to be a species that is stimulated to grow and other genera. Cannon & Hawksworth (1982) give
fruit by the presence of other fungi which it may also historical data conceming Melanospora in addition to
parasitise (Cain & Weresub 1957, Weber & Webster providing a key to clarify the generic concepts offungi
1998). A concise account of the history of the genus can which Doguet had originally also included in Melano­
be found in Rossman et al. (1999). spora. They also provide a key, together with descrip­
tions of 12 species of Melanospora found in Britain,
and this is the most recent key available for identifica­
Order OPHIOSTOMALES tion. Using data derived from ribosomal DNA se­
quences Jones & Blackwell (1998) confirmed that
Family OPHIOSTOMATACEAE Melanospora is related to Petriella, Ceratocystis and
Microascus and all were phylogenetically close to the
Ophiostoma Syd. & P. Syd. anamorph Raffaelea, a fungus associated with ambro­
sia beetles. This suggests that several of these genera
This is a large genus of non-coprophilous ascomycetes have a propensity to utilise the metabolites of others or
found on woody substrates many species of which are have sorne level of mutualism with other organisms.
pathogenic to the plant on which they are found. A Various anamorphs have been associated with Melano­
new species of Ophiostoma was found fruiting on one spora species (see Cannon & Hawksworth 1982), but
sample of koala dung and clearly associated with the the definitive culture work on the genus has yet to be
leafy diet of that animal. D. Mahoney who subse­ done.
quently cultured it found that it produced a Sporothrix During this current study, four collections of
anamorph and he forwarded it to Wingfield and de Melanospora were identified from the dung of koala,
Beer at the University of Pretoria who have stated that rock ringtail possum, kangaroo & bettong (?). Unfor­
it is an undescribed species. It is planned to publish tunately none of them can be identified with any de­
this species at a later date. It is illustrated in Fig. 51 A­ gree of confidence using the Cannon & Hawsworth
D. key. One collection had 4-spored asci the other three
had 8-spored asci. All had very similar perithecia
Order HYPOCREALES 14 without perithecial necks. The ostiolar bristles were ca
50-200 um long. Samples of the ascospores from these
FAMILY: CERATOSTOMATACEAE collections are illustrated in Fig. 52 J, K & L. One
collection (from koala dung) was obtained in culture
Melanospora Corda by D. Mahoney. The ascospores of these collections
are ellipsoidal, symmetrical or more typically asyrn­
Melanospora is a genus of Ascomycetes characterized metrical about the long axis to varying degrees, with
by small (circa 200-300 um iam) orange to red (5A5­ two prominent polar germ pores which sometimes
5A8), translucent perithecia with or without perithecia1 protrude, giving the ascospores a limoniform shape.
necks but a1ways with a fringe of stiff hyaline brist1es The ascospores sizes were within the range 15-21 x
surrounding the ostiole. When fully mature, the brown (8-)9-12 urn (a total of 72 ascospores measured).
ascospores can be clearly seen through the perithecial Much of the emphasis in the key supplied by Cannon
wall and they are frequently discharged in a long cirrus & Hawksworth relies on the exact description of the
at maturity. The genus is mostly found on decaying shape and dimensions of the ascospores. Insofar as the
vegetation but may also be seen on incubating dung Australian collections are concerned, these shapes and
samples. There is strong evidence that species of dimensions fall between several described species
Melanospora may directly benefit from the presence of Thus these collections have ascospores rather smaller
other fungi, although the nature of the relationship is than those given for either M brevirostris (Fuckel)
unclear (see Cannon & Hawksworth 1982). Melano­ Hëhnel, M damnosa (Sace. & Berl.) G. Lindau or M
spora species will grow in pure culture but frequently zamiae Corda, although in other respects the collec­
they do so more vigorously in the company of certain tions resemble any of these three species.
other fungi (O. Mahoney pers. comm.) When growing Dade also made extensive notes and detailed draw­
on dung, Melanospora appears late and frequently as a ings of his nine collections of Melanospora. Using
cottony mycelium strung from pellet to pellet over the Mason's data (1933), he identified one of his collec­
top of other fungi; the perithecia are produced in abun­ tions as Melanospora zamiae and the other eight as M
dance on this aerial mycelium providing further evi­ damnosa. His collections in all respects match my
dence to the theory that they may at least opportunisti­ Australian collections.
cally utilize the metabolites of others. D. Mahoney (pers. comm.) has taken a particular
Doguet (1955) produced the first large and compre­ interest in the genus Melanospora for sorne time and
hensive treatise on the genus taking a very broad view has cultured a number of isolates from California and
New Zealand including the afore-mentioned Australian
14 The cleistothecial genus Nigrosabulum also belongs here. collection. He reports that there is considerable varia­
32
tion in morphology among the seven cultures that he Selinia P. Karsten
has thus far studied and none of them are easily placed
using the Cannon & Hawksworth key. As mentioned Rossman et al. (1999) place this genus in the family
by Cannon & Hawksworth themselves, there is a need Bionectriaceae within the order Hypocreales. The
to study single-ascospore cultures of Melanospora genus comprising 2 (? 4) species is characterized by
from a wide variety of substrates and geographical having perithecia embedded in a pale orange (7 A5),
areas in order to make the correct anamorph connec­ superficially developed stroma. The thick-walled,
tions and to clarify sorne of the current teleomorph hyaline, ellipsoidal ascospores each have two promi­
identification difficulties. At the current state of nent nuclei and are surrounded by a thin gelatinous
knowledge, the Australian material of Melanospora sheath. One collection of Selinia pulchra (G. Winter)
can be best assigned as Melanospora cf brevirostris Sace. has been identified during the course of this
(Fig. 52 G-I). study and Jahn (2000) identified S. africana R.S. Khan
& J.c. Krug from dung collected in Queensland. 1 also
recorded S. africana on wallaby dung in New Zealand
(Bell 1983). Cribb (1988) recorded an unknown spe­
cies of Selinia that had 32-38 um long ascpspores.
This too was probably S. africana.

Key to the species of Selinia

1. Ascospores symmetrical, ellipsoïdal, with acute ends, 32--40 x 16-20 um

.................................................................................. Selinia a/ricana R.S. Khan & J.e. Krug (not illustrated)
1. Ascopsores symmetrica1, ellipsoidal, with acute ends, 54--60 x 20-26 um
...................................................................................................... Selinia pulchra (G. Winter) Sace. (Fig. 42)

Order SORDARIALES

Family CHAETOMIACEAE

Chaetomium Kunze

Gottfried Kunze established the genus Chaetomium in Chaetomium species are found on a wide variety of
1817, Chaetomium globosum being the type of the substrates, e.g. plant remains, decaying textiles, feath­
genus. Since its establishment many other authors have ers, seeds, dried spices, and many seem to favour
contributed further species to the genus resulting in materials with high cellulose content. Many are found
sorne 300 species described by 1985 (von Arx et al. on dung although no species is exclusive to that sub­
1986). Sorne of the publications in these intervening strate. For this reason, sorne consider Chaetomium
years included substantial monographie and semi­ species not to be true coprophiles, but merely oppor­
monographie treatments including the publication of tunistic inhabiting the voided dung by chance. The
Banier (1910), Chivers (1915), Skolko & Groves fruitbody of most species of Chaetomium is a perithe­
(1953), Udagawa (1960), Ames (1963), Seth (1972), cium, although for a beginner this may not be immedi­
Cannon (1986) and von Arx, Guarro & Figueras ately obvious since the ostiole is obscured by the dense
(1986). For a full account of the history of the genus vestiture of dark terminal hairs. Sorne species have
see Chivers (1915) and for a list of the many other beaked perithecia and in these species the perithecial
authors who have variously contributed research to the nature is more convincing. A few species (e.g. Chae­
genus see von Arx et al. (1986). Typical of ail genera tomium irregulare) can be cleistothecial. The asci of
that span nearly 170 years of existence, the accompa­ Chaetomium species are evanescent and discharged
nying philosophical views changed as to what consti­ ascospores are characteristically retained in dark
tuted a species together with improvements in micros­ masses caught up in the perithecial hairs. It has been
copy. More than 300 specifie names that had accumu­ suggested that the elaborate hairs of Chaetomium spp.
lated by 1985 included many synonyms. Kirk et al. serve the function of dispersal by invertebrate animaIs
(2001), accord 81 species to the genus. The family (Cain 1956).
Chaetomiaceae is placed in the order Sordariales by Dade listed five species of Chaetomium in his notes
Lee & Hanlin (1999) based on ribosomal DNA analy­ entitled: Check List of Victoria Fungi: C. caprinum, C.
SIS. cochliodes, C. fimicola, C. murorum, and Chaetomium
sp. indet.

33
 In other notes he recorded a further unknown spe­ found these two texture types difficult to distinguish in
cies but his drawings leave no doubt that the species this genus, and so 1 have chosen to refer to them as
was C. cuniculorum. He wrote that the genus "appears textura intricatalepidermoidea in the following key
not to be a common coprophile here but when it does (Fig. 8). Many species have ascospores that are bilater­
turn up often produced a large crop, usually spreading ally tlattened but measurements given in the key below
to the blotting paper ". were taken across the widest view (i.e. face view). A
Cribb (1999a and 1999b) recorded Chaetomium few of the Australian collections showed small varia­
murorum and C. homopilatum from Queensland. A tions to the given descriptions. These are listed below:
perusal of the literature indicates that to attempt to
identify species without also culturing them (and ide­ Chaetomium convolutum: One collection did not show
ally seeing sorne type material where it exists) is the petaloid perithecial wall character but it was of
fraught with pitfalls. However, the limited time in angular texture. Von Arx et al. (1986) state that inter­
which to complete this study precluded culturing aIl mediates between C. convolutum and C. bostrychodes
the species keyed and illustrated herein. If a particular are common (the latter also having a perithecial wall of
species grew easily (and agar was at hand), culturing angular texture).
was attempted. Morphological characters that are
useful for identification purposes include morphology Chaetomium indicum: The Australian material showed
of the perithecial hairs (particularly the terminal hairs); sorne straight terminal hairs in addition to branched
perithecial wall structure, and ascospore morphology. ones, which, according to von Arx et al. (1986), would
When available, details of cultural differences also add make the species C. funicola. However, their photo­
further useful characters. Various authors have fa­ graphs of both C. funicola and C. indicum (Plates 26
voured particular features above others although very and 29), show the presence of unbranched hairs.
few of the morphological characters are totally stable
or mutually exclusive thus making key construction Chaetomium piluliferum: This species had a purple
based entirely on morphological features difficult. The perithecial wall in one specimen which von Arx et al.
following key is based upon that of existing literature recorded as characteristic of C. murorum. The Austra­
coupled with my own more limited observations. lian material of C. murorum did not show this feature.
1 found that a minimum of three slides were re­ However, the Australian material did produce the
quired to verify an identification: 1) of a young Botryotrichium anamorph in culture.
perithecium in order to ascertain the arrangement of
the young ascospores within the asci, 2) a whole mount Chaetomium succineum: ln culture this species pro­
of mature intact perithecia in order to ascertain their duced distinctive yeIJow exudates on commeal agar
overall characteristics, 3) a very tlattened and dissected although von Arx et al. ( page 52, 1986) state that this
specimen in order to note the perithecial wall structure species is "without coloured exudates ".
and fine details of the perithecial hairs. Ali specimens
were first mounted in a drop of alcohol to rid the Sixteen species of Chaetomium have been identified
specimen of air bubbles after which the slide was during this project. AIl of these have been previously
flooded with a drop of distiJJed water. In general the described. According to von Arx et al. (1986), regard
publication by von Arx et al. (1986) was used to arrive the following species groups as closely related:
at the identifications, although further confirmation
was sought in the older publications having more 1. C. convolutum, C. bostrychodes, and C. robustum
detailed descriptions and line drawings. Von Arx et al. 2. C. globosum and C. subaffine
distinguish between textura intricata (characteristic of 3. C. piluliferum, C. murorum, and C. succineum
C. globosum & C. affine) and textura epidermoidea 4. C. subspirale and C. ampullare
(characteristic of C. murorum). However, in practice 1

34
 Terminal hai",; general shape Tenninal hairs: arrangement Pe.idial wall structurre

Angular

Entangled Petaloid

Slraight
Sinuous
Intricatafepidermoidea

r
Twotypes Spreading

1(
Haïr ornamentation Ascospore shape Hair ends

.],,~,~
A= Straight
B~ Hooked

C= Circinate
Smooth

Ovoid
,
:;;;.;....~,
'
ABC
.
Ellipscldal

Spiny

Warty
-
Trianqular
~
Lunate
,
'. Lose colis
J Even spiral
" Uneven spiral

Fig. 8. Glossary to terms used in the key below.

Key to the species of Chaetomium

1. Ascospores distinctly triangular in one view 8-10 x 4-5 um; perithecial wall
petaloid; terminal hairs upright, straight, smooth or warted
................................................................................ Chaetomium trigonosporum (Marchal) Chivers (Fig. 53)
1. Ascospores not distinctly triangular, combination of perithecial wall
characters and terminal hair morphology not as above 2

2. Ascospores limoniform, ovoid, lunate, or irregularly shaped, 14-16 x 8-10 um;

hairs entangled, spiny, evenly scattered over the commonly cleistothecial,
sometimes partially submerged, ascocarp Chaetomium irregulare Sorgel (Fig. 53)
2. Ascospores of more uniform shape within the species; perithecial hairs c1early
divided into a terminal cluster and smaller lateral hairs; ascocarps ostiolate 3

3. Terminal hairs of distinctly 2 kinds: 1) fine, entangled, branched and spiny,

forming a ruff around the apex of the perithecium, 2) robust, erect, sinuous,
warty hairs emergening from the ruff; perithecial wall cells angular; ascospores
ellipsoidal, 10-11 x 6-7 um Chaetomium cuniculorum Fuckel (Fig. 54)
3. Terminal hairs ofmainly of 1 type or if2 types present, not sharply distinguished;
combination of perithecial wall and ascospore characters not as above 4

4. Many terminal hairs dichotomously branched, sorne straight, ail warty; perithecial
wall cells angular covered by branched hyphae; ascospores broadly ellipsoidal,
thick-walled, 5-7 x 4-5 um Chaetomium indicum Corda (Fig. 54)
4. Terminal hairs not dichotomously branched; combination ofperithecial wall and
ascospore characters not as above 5

5. Ascospores> 12 um long 6
5. Ascospores < 12 um long 8

35
6. Terminal hairs densely entangled, unbranched, distinctly and evenly spiralled,
spiny; perithecial wall ceIls angular; ascospores ellipsoidal, 14-16 x 7-9 um
.......................................................................................................... Chaetomium succineum Ames (Fig. 59)
6. Terminal hairs arcuate, sinuous or spreading; perithecial wall of textura
intricataJepidennoidea 7

7. Terminal hairs arcuate or sinuous, warty or smooth, tips hooked or straight;

ascospores ellipsoidal, 14-15 x 8 urn Chaetomium murorum Corda (Fig. 58)
7. Terminal hairs spreading, smooth, tips hooked, lateral hairs warty; ascospores
ellipsoidal, 3-15 x 8-10 um Chaetomium piluliferum J. Daniels (Fig. 58)

8. Ascospores uniseriate, ovoid, 9-10 x 7-8 urn; terminal hairs entangled, unevenly
spiralled, warty; perithecial wall cells angular Chaetomlum brasiliense Batista & Pontual (Fig. 56)
8. Ascospores multiseriate, not ovoid 9

9. Ascospores lunate in face view, 9 x 5 um; terminal hairs arcuate, spiny, tips
circinate or hooked or unevenly spiral; perithecial wall ceIls angular
.............................................................................................................. Chaetomium cupreum Ames (Fig. 56)
9. Ascospores broadly ellipsoidal to limoniform in face view 10

]O. Ascospores 9-10 x 7-9 um; perithecial wall of textura intricataJepidermoidea ]]

10. Ascospores smaller; perithecial wall cells angular or petaloid 12

11. Terminal hairs densely entangled, unbranched, mostly sinuous with sorne irregular
spirals, warty; ascospores 9-10 x 7-9 um Chaetomium globosum Kunze (Fig. 57)
1l . Terminal hairs upright, unbranched, sinuous, with few lose coils and irregular
spirals, warty; ascospores 9-] 0 x 8-9 um Chaetomium subaffine Sergeje va (Fig. 57)

]2. Terminal hairs unbranched, distinctly and irregularly long-spiralled ]3

12. Terminal hairs with sorne irregular branching and short irregular spirals 14

13. Perithecial wall cells in a petaloid pattern; terminal hairs profusely and irregularly
spiralled, spiny; ascospores 6-7 x 5-6 um Chaetomium convolutum Chivers (Fig. 59)
13. Perithecial wall cells angular; terminal hairs weakly and unevenly spiralled ending
in twisted hyaline woolly threads, spiny; ascospores 6-7 x 5 um ... Chaetomium subspirale Chivers (Fig. 55)

]4. Terminal hairs upright, mostly sinuous, a few loose coils or uneven spirals, warty;
perithecial wall cells angular or elongated/angular; ascospores 6 x 4-5 um
........................................................................................................ Chaetomium ampulfare Chivers (Fig. 55)
14. Terminal hairs entangled, sinuous, with copious uneven spirals, warty; ascospores
larger 15

15. Perithecia > 1mm high; perithecial wall cells angular with elongatedlangular
portions; ascospores 7-8 x 6-7 um Chaetomium robustum Ames (Fig. 60)
15. Perithecia <Imm high, perithecial wall ceIls angular; ascospores 7-8 x 5--6 um
.................................................................................................. ......Chaetomium bostrychodes Zopf. (Fig. 60)

Family SORDARIACEAE

Sordaria Ces. & De Not.

The genus Sordaria is more or 1ess an exclusively produce asci each with a clear apical ring. In ail spe­
coprophi1ous genus of pyrenomycetes, and the type cies known to date the asci each contain eight uniseri­
species, Sordaria fimicola (Rob.) Ces. & De Not., is ate, single celled, more or less ellipsoidal to globose,
one of the most frequent1y encountered ascomycetes dark ascospores, each of which is fumished with a
upon dung. The genus is characterized by producing single germ pore. The germ pore normally is situated
many small (ca 0.5 mm diam) smooth, black perithecia at the end of the ascospore nearest the ascus stipe, but
in swards across the substrate. Perithecia may be sub­ the asci quickly swell in water mounts and the asco­
merged, semi-immersed or superficially produced. In spores easily reorientate themselves and the viewer is
sorne collections there is a dense white tomentum frequently presented with polar views of ascospores or
clothing the lower portions of the perithecia (e.g. as in ascospores that are angled within the ascus. A gelati­
collection A5 of Sordaria humana). Ali known species nous sheath is usually present around each ripe asco­

36
 spore in fresh specimens and this often swells in water recognized sorne 16 species. Guarro & von Arx (1987)
mounts. In dried material, or in lactophenol mounts the give a key to 14 species, and Kirk et al. (2001) state
sheath may be absent. The genus Sordaria is as that the genus comprises ten species. In 1987 von Arx
Lundqvist states: "very homogeneous and rather uni­ et al. established the genus Asordaria for those species
form" (1972, p. 299). A number of mycologists have lacking a gelatinous sheath and they reclassified seven
contributed information on the genus (see for example species of Sordaria on the strength ofthis feature. This
Olive & Fantini 1961, Fields & Maniotis 1963, Fields view, however, has not been universally accepted
& Grear 1966, Fields 1968, Cailleux 1971, Barrasa et amongst mycologists (including myself). Many of the
al. (1986), Guarro & von Arx (1987), Watanabe species of Sordaria described in the literature have
(1989), Khan & Krug (1989b) and Robertson et al. overlapping ascospore sizes and in aIl other respects
1998). A complete review of the history of Sordaria seem remarkably similar.
spp. can be found in Lundqvist (1972) who at that time

Ê 14
I S

. . .. . •
.:!:
'5 -
:2 13
it ...------s. alcina
e ~
&.1
III
o
u
1• ••
• • •
~ 11

10

1 1 25 1 1 30 1
Ascospore length (l'm) •

Fig. 9. Ascospore measurements of Australian collections of Sordaria alcina. Boxed areas denote size range for published
species as indicated. • Collections from Eastern Grey Kangeroo, • Collections from Quokka dung. For further information
see text.

A number of species grow with comparative ease in 63 E, it will be seen that the unnamed species has
culture and this, coupled with the fact that the genus ascospores that are more acute at their ends but the
as a whole exhibits both homothallic and heterothallic differences are slight. They provide a good example
species, has resulted in much interest by geneticists of what Hawksworth (2001) refers to as cryptic spe­
and teachers alike. Homothallic species include S. cies, i.e. species that can very easily be overlooked
fimicola, S. superba and S. macrospora. Heterothallic since they are so difficult (sorne might say impossi­
species include: S. alcina N. Lundq., S. brevicollis ble) to distinguish on morphological grounds. How­
Olive & Fant., S. heterothallis Fields & Maniotis, S. ever, culturally they show a number of clear differ­
sclerogenia Fields & Grear, S. thermophila Fields, S. ences not the least of which is the fact that they will
tomento-alba Cailleux, and Sordaria sp. nov. (iso­ not cross with one another. These details will be
lated in New Zealand by D. Mahoney). discussed in a future paper. Meanwhile, for those
Each mating strain of these heterothallic species identifying material from the field, S. brevicollis
produces microconidia which presumably act as appears early during the incubation phase and the
spermatia (Guarro & von Arx 1987). During this perithecia are more superficial on the dung, the un­
study 1 have identified seven species of Sordaria from named Sordaria species occurs after the dung has
Australia, three of which have proven to be hetero­ been incubating for 2-3 weeks and the perithecia are
thallic. Ail of the research conceming these hetero­ more submerged. Dade also saw this new species
thallic Sordaria species has been undertaken by D. (always on possum dung) and he sent a sample ofit to
Mahoney. The unnamed species first identified by Cain (see his slide V328c) who identified it as
him from New Zealand collections is morphologically Sordaria brevicollis, without performing any crossing
very close to Sordaria brevicollis Olive & Fant. experiments.
However, our isolates will not mate with the type The third heterothallic species found during this
culture of S. brevicollis. In Australia we have found it study was S. alcina N. Lundq. Lundqvist (1972, p.
on koala and possum dung and in New Zealand the 328) states that Mirza (1963) mentioned in his Ph.D.
same species is found exclusively on possum dung thesis that Canadian specimens of S. alcina produced
Sordaria brevicollis is also found in Australia (thus spermatia. D. Mahoney has now cultured both mating
far only on cassowary dung). These two morphologi­ strains of the Australian S. alcina from kangaroo dung
cally close species are illustrated together in order to collected in Victoria (dung collection A33). Later 1
express their similarity (Fig. 63). By examining Fig. identified what 1 considered to be S. baltica N. Lundq.
37
from quokka dung collected in Western Australia within the S. superba/macrospora size range. Closer
(dung collection A255), according to the original investigation by means of single-ascospore cultures
description and illustrations. Samples of the asco­ may reveal hidden or cryptic species within the S.
spores from these two collections are plotted in the superba/macrospora assemblage. But on CUITent
scatter diagram (Fig. 9). Single-ascospore isolations (morphological) evidence, 1 cannot satisfactorily
of the A255 specimen were found to cross with com­ separate these two species in these Australian collec­
patible mating strains of the A33 (s. alcina) speci­ tions (Fig. 10).
men. 1thus concluded that Sordaria alcina N. Lundq. To date 31 collections of dung have yielded
and S. baltica N. Lundq. are the same species. Sordaria fimicola Ces. & De Not. Ascospores of a
Of the homothallic species of Sordaria, 1 have sample of these are displayed in the scatter diagram
identified sorne 24 records that straddle the sizes (Fig. 10). It can be seen that their size range falls
given for S. superba De Not. and S. macrospora between the ranges given for both S. fimicola and S.
Auersw. as described by Lundqvist (1972). The asco­ lappae Poteb. described by Lundqvist (1972).
spore size range of a representative sample of these Dade also identified Sordaria fimicola (siide VI 87f),
24 collections is displayed in the scatter diagram in S. macrospora (slides VI03c, V445c & d, and
Fig. 10. Lundqvist recognized that sorne mycologists V977b), S. arctica (slide V206e) and S. alcina (the
may choose to unite these two species although he latter he named as S. spermatiophora J.H. Mirza &
preferred to consider them as distinct in order to Cain).
"catch attention" to the problems surrounding their Other records of Sordaria species collected in
circumscription. However, by uniting them 1 do not Australia include S. lappae from dung collected in
wish to convey that there is no alternative interpreta­ Queensland (Jahn 2000), Sordaria fimicola recorded
tion of this data. The heterothallic species Sordaria by Cribb (1988, 1995), S. goundaënsis Caill. (Cribb
heterothallis Fields & Maniotis was described as 1992), and S. macrospora (Tibell 11205-g) loaned to
having ascospores 30-32 x 18-19 urn, which is also me by Lundqvist.

20
0

0
0
0 0 -, i
Ct,
1

_0 000 1
S mecrospora 0. _---J
18 0
S scperba­ ~o·
cr 00
o
• o
~ 16

~
S ïappee
~-9 . 0 00
l ~o
~14
8 ••'4--;;0 f o
: o
••1 '1
12 . ··1
.~~
.1
,

S fimICola ~~._..I._-----
j
1~
"';'~ 1 1 20 1 --r-~-'--- 1 1 ' 28 I-··T--~I 31 1 1 34r-T36T
Ascoapore lenath luml ...

Fig. 10. Ascospore measurements of Australian collections of Sordaria fimicola (e) and S. macrospora (0). Boxed areas
denote size range for published species as indicated. For further information see text.

Key to the species of Sordaria

1. Ascospores globose, subglobose, or obovoid 2

1. Ascospores ellipsoidal 3

2. Ascospores globose, 15-18 x 14-16 um Sordaria arctica Cain (Fig. 62)

2. Ascospores subglobose to obovoid, 20-26 x 15-19flm
................................................................................................ Sordaria humana (Fuckel) G. Winter (Fig. 62)

3. Ascospores < 20 um long 4

3. Ascospores mostly >20 um 5

4. Ascospores 14-16 x 8-11 um acute at the apices Sordaria n. sp. (Fig. 63)
4. Ascospores 13-18 x 9-11 um rounded at the upper end, slightly acute at the germ
pore end Sordaria brevicollis Olive & Fant. (Fig. 63)

38
5. Ascospores 25-34(-36) x (14-)15-18(-20) um Sordaria macrospora Auersw. (Fig. 61)
5. Ascospores mostly < 14 um diam, ascospores appearing narrow in relation to their width 6

6. Ascospores (19-)20-25 x 11-13 um Sordaria fimicola Ces. & De Not (Fig. 61)
6. Ascospores (19-)20-31 x 10-12 um Sorâaria alcina N. Lundq. (Fig. 62)

Note: A culture of S. arctica is deposited at Landcare (lCMP 14389). Cultures of Sordaria n. sp. are deposited at CBS (CBS
112039, 112040, 122040). 1mistakenly included this taxon together with S. fimicola (Bell 1983).

Family CONIOCHAETACEAE

Coniochaeta (Sace.) Cooke

Coniochaeta is a cosmopolitan genus of terricolous, both their anamorph and teleomorph on agar (Hawk­
lignicolous or coprophilous pyrenomycetes, having sworth & Yip 1981). For further discussion of the
small (ca 300-500 um diam), semi-immersed, usually relationships between Lecythophora and terricolous
gregarious, dark perithecia bearing stiff dark bristles species of Coniochaeta see Weber 1 & II (2002). How­
clustered around the neck and sometimes also scattered ever, coprophilous species generally only form their
on the venter. The coprophilous species usually only Lecythophora anamorph in culture (D. Mahoney pers.
appear on dung which has been incubating for a long corn.).
time. The peridial structure is variable within the ge­
nus. Coniochaeta scatigena has a peridium of textura
L C sceeçene ---- - - .
epidermoidea while the peridia of C. leucoplaca and C.
discospora are predominantly of textura angularis
often with patches, which are cephalothecoid (areolate)
in texture. However, in a few collections of these latter
le ..
two species the peridial structure appears diffuse and
almost epidermoid with irregular thickenings. 1 do not .
AlcOipore
1· .....

::~.:-::
. ...,~.
liIlgth
know the significance (if any) of these differing (pm)
a
o

peridial structures. Jensen (1985) describes C. disco­

spora as having "a peridium of textura angularis with
many concave angles that give the peridium a jig-saw .,
puzzle like appearance" and his photograph shows
clearly a textura epidermoidea although he does not
cali it by that term. When flattened, the perithecial Fig. Il. Ascospore measurements of Australian collections
necks exhibit rows of cells altemating with highly of Coniochaeta leucoplaca (e) and C. discospora (0).
melanised strips. The genus is characterized by asci Boxed areas denote size range for published species as
with apical rings and dark discoid ascospores each indicated. For further information see text.
with a germination slit along the narrow side. An
evanescent hyaline sheath surrounds the ascospores. Coniochaeta velutina (Fuckel) Munk is a species
Generally the perithecia are extremely fertile and originally described as lignicolous with ascospores 6-8
contain many asci but sorne species have a tendency to x 4-6 x 3--4 um (Munk 1957). Coniochaeta leuco­
produce asci with odd numbers of ascospores within placa (Berk. & Ravenel) Cain is described as a copro­
them, these being usually larger than those from (nor­ philous species whose ascospores are within the range
mally) 8-spored asci. However, even within 8-spored 7-9 x 6-8 x 5-6 um (Cain 1934). Hawksworth & Yip
asci there may be quite a considerable variability in (1981) and Checa et al. (1988) considered the differ­
size between the spores within the one ascus. ences in ascospore sizes between these two species to
About 30 species were known in the early 1980's be insignificant and considered C. leucoplaca Cain
(Mahoney & LaFavre 1981), and a few more have synonymous with the earlier named C. velutina. Nei­
been added since that date (see for example Hawk­ ther paper indicates that the authors ever examined
sworth & Yip 1981, Richardson 1998a, and Romero et type material of either C. velutina or C. leucoplaca.
al. 1999). Keys to the species can be found in Cain However, examination of 29 collections examined
(1934), Munk (1957), Hawksworth & Yip (1981), during this study did yield two Coniochaeta species
Checa et al. (1988), Richardson & Watling (1997) and which could be separated upon ascospore dimensions,
Romero et al. (1999). Fine illustrations of two copro­ one of which (comprising seven collections), exhibited
philous species of Coniochaeta can be found in Koba­ spore measurements which straddle the given ranges of
yasi et al. (1969). A number of species have been both C. velutina and C. leucoplaca. The other (19
grown in culture and sorne may be induced to form collections) exhibited ascospores approximating the C.

39
discospora size range (scatter diagram Fig. Il). Taking To date 1 have identified three species of Conio­
into account the afore-mentioned cultural differences chaeta during the course of this CUITent study. Dade
between terricolous and coprophilous species, I remain also found three species of Coniochaeta and left c1ear
yet to be convinced that the lignicolous C. velutina and notes and slides. In addition to C. leucoplaca (slides
C. leucoplaca are synonymous. Similarly I consider V979a, and V147f) and C. discospora (slide V938e),
that further studies are required to ascertain whether or he also identified C. hansenii (Oudem.) Cain which I
not the coprophilous C. discospora (Auersw.) Cain is have not yet found on Australian material (slide
synonymous with the lignicolous C. ligniaria (Grev.) V555f). Consequently I have included his drawings of
Massee, as stated by Hawksworth & Vip (1981) and this species alongside my own of the others. Cribb
Checa et al. (1988). (1995) recorded Coniochaeta velutina from macropod
dung collected in Queensland.

Key to the species of Coniochaeta

1. Asci 128-spored; ascospores broadly ellipsoidal to subcircular in face view,

slightly flattened in side view, 7-9 x 4-7 x 4 um Contochaeta hansenii (Oudern.) Cain (Fig. 65)
1. Asci 8-spored; ascospores broadly ellipsoidal to subcircular in face view
flattened in side view 2

2. Ascospores < 16 x 12 um in face view 3

2. Ascospores (17-)20-25 x 14-17 x 8-10 um Conlochaeta scatigena (Berk. & Broome) Cain (Fig. 65)

3. Ascospores 6-10 x 5-8 x 3-5 um Coniochaeta leucoplaca (Berk. & Ravenel) Cain (Fig. 64)
3. Ascospores (9-)10-15 x 6-12 x 4--6 um Coniochaeta discospora (Auersw.) Cain (Fig. 64)

Family LASIOSPHAERIACEAE

Cercophora Fuckel emend. N. Lundq.

Cercophora is a genus of the Lasiosphaeriaceae many into a dark cell and a hyaline pedicel. In this genus
of the species of which are widespread on woody features of the perithecia provide the most useful char­
substrates (Hilber & Hilber 1979). Other species are acteristics by which to distinguish the species. Those
coprophilous. For descriptions of coprophilous species species that have been cultured, frequently exhibit a
in particular see Lundqvist (1972) and Udagawa & Cladorrhinum anamorph. Five species of Cercophora
Muroi (1979). One of the characteristics of the genus is have been identified during the course of this study
the morphological similarity of the ascospores between two ofwhich are new.
species and the tendency of the ascospores to remain in
a hyaline state only belatedly becoming differentiated

Ascus ring l!"-­

~
Apical cauda

'--0\
On
.11
Subapical
1 globulus
_ Darkcell

.(

œphalothecoid perithecial wall

(sometimes called: areolate)

Hyaline sigmoid
ascospore

Fig. 12. Glossary to terms used in key (page 41).

40
Key to the species of Cercophora

1. Perithecial wall cephalothecoid and sparsely hairy, neck massive and clearly
ridged; asci with clear apical ring and smooth subapical globules; ascospores
seen only in their hyaline, sigmoid state 62-75 um long, 25-30 um wide at
the upper end, 11-13 um at the lower end Cercophora californlca (Plowr.) N. Lundq. (Fig. 87)
1. Perithecial wall not cephalothecoid; combination of characters not as above 2

2. Perithecial wall smooth 3

2. Perithecial wall distinctly hairy 4

3. No subapical globulus; young hyaline ascospores cylindrical, straight, and aligned

in a central cluster in the ascus; ascospores with straight pedicels of variable lengths,
dark cell ellipsoidal, symmetrical or asymmetrical with a truncate base,
26-32 x15-18 um, pedicels 43-58 x 6--7 um Cercophora recta n. sp. (Fig. 89)
3. Subapical globulus minutely spiny; young hyaline ascospores sigmoid, entwined,
biseriate within ascus; ascospore dark cell ellipsoidal with truncate base and
conical apex, l4~15 x 9-10 um, pedicel distinctly bent below, 25-34 x 5 um
...................................................................................... Cercophora sordarioldes (Speg.) N. Lundq. (Fig. 89)

4. Perithecial venter clothed in a dense brown tomentum, neck with short inflated
cells; asci with apical ring and minutely spiny subapical globulus; ascospore
dark cell ellipsoidal, more or less symmetrical, with truncate base and conical apex
19-20 x 9-10 um, pedicel slightly bent below 40-50 x 4 um Cercophora mirabilis Fuckel (Fig. 88)
4. Perithecial venter clothed in a dense white tomentum, emergent dark necks with
few short, hyaline-tipped hairs, necks becoming su1cate with age; asci with apical
ring; ascospore dark cell ellipsoidal, symmetrical with truncate base,
27-32 x 15-16 um, pedicel bent below, 45-55 x 7-8 urn Cercophora grandiuscula n. sp. (Fig. 88)

Note: for the descriptions ofthe Cercophora recta and C. grandiuscula see page 59.

Podospora Ces.

For sorne time there was confusion surrounding the Species of the genus may be found in both temperate
legitimacy of this generic name and explanations con­ and tropical regions worldwide on various dung sub­
ceming the complexity of the situation can be found in strates. Currently 78 species have been named (Kirk et
Lundqvist (1972). The matter was resolved when the al. 2001). On morphological grounds alone, it would
genus Podospora was made legitimate citing P. fimicola seem that the genus is probably polyphyletic and a
sensu Ces. (in Rabenhorst 1865) as the type species by number of authors (see for example Lundqvist 1972,
Donk (1964). Barrasa & Solans 1989) separate species having inflated
Podospora is a genus of the Lasiosphaeriaceae in agglutinated perithecial hairs (together with certain
which the majority of described species are coprophi­ other characteristics), into the genus Schizothecium.
lous. It is one of the most frequently encountered genera However, as long as vast tracts of the world remain
on herbivore dung in both Australia and New Zealand. relatively unexplored by mycologists, it would seem
A number of species are found in soil and an interesting premature to invest an inordinate amount of resources
discussion of these species together with an explanation on arranging and rearranging those few examples that
accounting for the morphological differences between wehave.
the dung inhabitants and the soil species is to be found Analogous to the situation in other pyrenomycetes,
in a paper by Stchigel et al. (2002). There exists an attention has been drawn to the problem surrounding the
extensive list of publications conceming the genus continued separation of the ostiolate genus Podospora
Podospora ranging from papers on genetics, papers Ces. and its c1eistothecial counterparts Zopjiel/a G.
dealing with country-specifie species lists, monographie Winter and Tripterospora Cain (e.g. Malloch & Cain
and semi-monographic accounts. Sorne of the taxo­ 1971 b, Lundqvist 1972, von Arx 1973, Barr 1990).
nomic papers that have been consulted for this research Guarro et al. (1991) united Tripterospora with the older
include the following: Bell & Mahoney (1995), Cailleux name Zopjiel/a and provided a key to the species use­
(1969), Cain 1962, Doveri et al. (1998 and 2000a), fully including in it those species of (mainly soil-borne)
Furuya & Udagawa (1972), Krug & Khan (1989), Podospora that most closely resemble species of Zop­
Lundqvist (1970, 1972, 1973), Lundqvist, Bell, Ma­ jiel/a.
honey & Lorenzo (1999), Mirza & Cain (1969), Sharma Podospora is characterized by small (ca 0.5-1 mm
(1976), Stratton (1921), and Wong (1992,1994,2000). diam), dark perithecia, which are superficial, partially

41
 submerged or totally submerged in the dung. The water mounts carefully infiltrated with Indian ink are
perithecia may be smooth or variously adorned with necessary for identifying species of this genus. In spite
hairs or bristles. In sorne species the extent of perithe­ of these obstacles, the genus is accessable to the begin­
cial hair or bristle development may be variably ex­ ner since there are a number of excellent weil illustrated
pressed and like other pyrenomycetes the necks may monographs or serni-monographic works on the genus
lengthen considerably with age. Very few (? any) spe­ (see for example Lundqvist 1972, Mirza & Cain 1969).
cies can be identified by perithecial morphology alone, This was not the situation when Dade was trying to
although the experienced operator may gain a certain identify his Australian collections of Podospora in the
guarded confidence for recognizing certain species by mid-196ü's.
this means. However, it is not uncornmon for as many Dade recognized sorne 19 species of Podospora.
as three or four different species to grow within milli­ There are three boxes of his slides of this genus at the
metres of each, other causing much confusion. The most herbarium (DAR) at Orange. In one box he left a note
characteristic feature of the genus is the morphology of "Several are wrongly identified & some dubious ­
the ascospores, which at maturity are composed of a serious revision necessary". In addition to the slides, he
dark cell attached to a hyaline cell (pedicel), which is left thorough notes and illustrations of his observations
usual1y short-lived. Mucilaginous evanescent append­ and corresponded with Cain and others who were work­
ages (caudae) of various shapes and sizes adorn both the ing on the genus at that time. He sent duplicates of his
dark cell and the pedicel and these are usually necessary slides to Gunnell at Kew. She in turn incorporated sorne
items for identification to the species. The caudae usu­ of his slides into the IMI collections. Dade drew up two
ally dry up on discharged ascospores and are most charts listing the species of Cain 1934 and 1962 (Chart
frequently absent from herbarium specimens. Even 1) and the additional species listed in Mirza & Cain
when fresh, the caudae of sorne species are notoriously 1969 (Chart 2). In 1982 1 studied a number of his Po­
difficult to observe and several slides in different dospora slides and Table 3 represents my observations
mounting media have to be employed in order to ascer­ on Dade's collections:
tain identification. Thus Congo red, Cotton blue and

Table 3. Reidentification of Podospora slides from Dade's collections.

Dade's identification Collection num bers studied in 1982 Current identification

Podospora anserina Vl19g(l), V182e, V590d, V594b Podospora anserina
Podospora austraIis V192h, V257f, V268a, V970d Podospora austraIis
Podospora coronifèra V182j Podospora decipiens
Podospora curvula V399c, V397g, V543a, V586c, V799c Podospora conica
Podospora decipiens VI03e, Vl27b, V182j, V192j, V485b, V513g, V548c, V770f Podospora decipiens
Podospora fimicola V257j, V301j, V609d, V636d Podospora fimiseda
Podospora perplexans Vl27j, V130e, V 187d, V573c Podospora perplexans
Podospora prethopodaIis V205j Podospora excentrica
Podospora pyriformis V441f Podospora pyriformis
Podospora tetraspora V244a, V528a, V718d Podosporateifaspora
Podospora tax. sp. 1 None seen ? Podspora vesticola
Podospora tax.sp. Il V713f Podospora excentrica
Podospora tax. sp. III V229c, V525c, V605f, V953f, V954h, V968c Podospora curvicolla
Podospora tax. sp. IV None seen ?
Podospora tax. sp. V V540g ? (immature)
Podospora tax. sp. VI None seen ?
Podospora tax. sp. VII V959j Podospora austrohemisphaerica
Podospora tax. sp. VIII Vl19k Arnium arizonense
Podospora tax. sp. IX V119j, V964h Podospora pleiosEora

The find of Podospora pyriformis (represented by slide Zealand, 1 have recorded P. pyriformis from both the
V441f) by Dade is particularly interesting. During this North and South Islands and the more southern Enderby
current Australian study, 1 have never found P. pyri­ Island (part of the Auckland Island group). As stated also
formis, which is considered to be more or less confined to by Lundqvist, P. pyriformis develops on dung which has
temperate regions with few subtropical records (see been incubated for four or more weeks.
Lundqvist 1972, p. 161). Dade's single sample of this ln addition to Dade's observations, there are other
species was obtained from cattle dung collected in the records of Podospora species from Australia. Cribb
Baw Baw Mountains, Gippsland, Victoria, "above the (1989, 1998) recorded the following species: P. anserina.
snow Une", which would corroborate Lundqvist's asser­ P. australis, P. absimilis, P. argentiniensis, P. decipiens,
tions. Since Dade left good drawings of this species, they and P. prethopodalis. Lundqvist forwarded five herbar­
have been included in this publication. In temperate New ium specimens of Podospora that were collected in

42
Australia which 1 consider are aIl Podospora curvuloides Amium. Similarly, P. petrogale could be mistaken for a
(Tibell 11285-q, Tibell 11381-d, TibelllI94-f, Tibell species of Strattonia if old discharged ascospores were
12074-g & Tibell 12094-k). Jahn (2000) recorded the observed (at which point the pedicel would have disap­
following species from Australia: P. anserina, P. aus­ peared). Consequently, this species also appears in the
tralis, P. bicolour, P. communis, P. curvicolla, P. de­ key to Strattonia.
cipiens, P. fimiseda, P. perplexans and P. prethopodalis. A single species of Apiosordaria is also included in
Thus far 1 have recorded 35 species of Podospora the key below. This is purely for convenience, since it too
from Australian dung collections. P. australis and P. is characterised by having ascospores composed of a dark
gigantea have also been included in the key to the genus cells and a hyaline pedicel. Apiosordaria species are
Arnium since their caudae are practically non-existent more commonly found in soil (Krug et al. 1983). Stchigel
and thus they could easily be thought of as belonging to et. al. (2000) provide a key to soil-bome species.

Perithecial adornment ....

( Agglutinated hairs/bristles Non-agglutinated hairslbristles

Tubercles

Supernumerary caudae

Hyaline tipped hair

/) '-~,I/
, 1

"
",li

;;:
i"
Single rigid hair

~",'il~
;';1
".,

Fig. 13. Glossary to terms used in the key below.

Key to the species of Podospora

1. Asci 4-spored; dark cell of ascospores covered with coarse warts and/or spines
which may also partially cover the hyaline pedicel, dark cell ellipsoidallconical
with truncate base, 18-20 x 14-17 um, pedicel conicallasymmetrical, 7-10 um
..................................................................... Aplosordaria verruculosa (O. Jensen) Arx & W.Gams (Fig. 82)
1. Dark cell ofascospores smooth (or very minutely) horizontally striate, size and/or
shape not as above 2

2. Perithecia adomed with tufts of inflated agglutinated hairs 3

2. Perithecia bare, or if adomed, hairslbristles composed of non-inflated cells 8

3. Asci 4-spored; dark ce11 of ascospore ellipsoidal, with a smail truncate base,
22-24 x 14-15 um, pedicel 6 x 2 um; a single cauda at each end ofthe
ascospore Podospora tetraspora (G. Winter) Cain (Fig. 75)
3. Asci 8-spored 4

43
4. Dark cell of ascospore ellipsoidal, (33-)37-52 x 19-27 um, sometimes with
minute horizontal striations; pedicel5-10 x 2~3 um; single apical and basal
caudae; perithecium frequently reclining, neck covered with minute agglutinated
hairs resembling velvet pile Podospora curvuloides Cain (Fig. 78)
4. Dark cell of ascospore smaller; other features not as above 5

5. Dark cell of ascospore broadly ellipsoidal with truncate base, 35-37 x 25-28 um;
pedicel5-10 x 2-3 um, single apical and basal caudae Podospora glutinans (Cain) Cain (Fig. 76)
5. Dark cell of ascospore smaller 6

6. Dark cell of ascospore ellipsoidal with a truncate base, 25-28 x 13-16 um;
pedicel6-7 x 2-3 um, single apical and basal caudae ..
........................................................................ Podospora conica (Fuckel) A. Bell & D.P. Mahoney (Fig. 78)
6. Dark cell of ascospore mostly < 25 um long 7

7. Dark cell ofascospore ellipsoid with a truncate base, 17-19 x 10-13 um,
pedicel 8 x 2 um, single apical and basal caudae
............................................................... Podospora vesticola (Berk. & Broome) J.H. Mirza & Cain (Fig. 76)
7. Dark cell of ascospore ellipsoidal with a truncate base, (19-) 21-25 x 15-17 um pedicel 7-8 x 2 um,
single apical and basal caudae Podospora miniglutinans J.H. Mirza & Cain (Fig. 77)

8. Exposed portions of perithecia without adomment of hairs or bristles 12

8. Exposed portions of perithecia adomed with non-intlated hairs or bristles,
which may be agglutinated, and/or with minute tubercles 9

9. Perithecia more or less superficial, or with neck only protruding, exposed

portions evenly scattered with short, blunt, hyaline-tipped hairs 10
9. Perithecial adomment not as above 17

10. Perithecia more or less submerged, neck only protruding; asci 4-spored; dark cell
ellipsoidal, 53--60 x 29-32 um; pedicel absent or as minute apiculus at the base of
the dark cell Podospora australis (Speg.) Niessl (Fig. 83)
10. Perithecia superficial or only partially submerged; asci 8-spored 11

11. Perithecia more or less superficial, often clustered and reclining; asci 8-spored;
dark cell ellipsoidal, 40-50 x 19-22 um; pedicel cylindrical 35-40 x 4--6 um
............................................................................................ .........Podosporaperplexans (Cain) Cain (Fig. 75)
11. Perithecia more or less superficial mostly upright; asci 8-spored; dark cell
ellipsoidal with a truncate base 58-64 x 27-36 um; pedicel 53-55 x 5 um,
swelling at free end Podospora fimiseda (Ces. & De Not.) Niessl (Fig. 74)

12. Apical and basal cauda of ascospores absent; dark cell ellipsoidal,
(45-)50~58(- 60) x (24-)26-30(~32) um, enveloped in a mucilaginous jacket;
pedicel short-lived, 10-11 x 2 um, straight or curved, protruding through jacket,
.............................................................................................................. Podospora petrogale A. Bell (Fig. 90)
12. Apical and basal caudae present 13

13. Dark cell ofascospore ellipsoidal, 120-132 x 40--60 um; pedicel non-existent or
as a minute apiculus at the base of the dark cell; single large striated cauda at each
end of the dark cell Podospora gigantea J.H. Mirza & Cain (Fig. 90)
13. Ascospores much smaller; pedicellarger; caudae not as above 14

14. Apical and basal caudae only present 15

14. Many supemumerary caudae also present besides the apical and basal ones;
dark cell ellipsoidal with truncate base, 50--61 x 25-28 um; pedicel 26-32 x 5--6
um; apical and basal caudae in Iyre-shaped tuft Podospora pectinata N. Lundq. (Fig. 68)

15. Ascospores with a single apical and basal cauda 16

15. Ascospores with 3 apical and 3 basal caudae; dark cell ellipsoidal with truncate
base, 33-40 x 20-22 um; pedicel 31-34 x 5--6 um Podospora communis (Speg.) Niessl (Fig. 72)

44
 16. Dark cell ofascospore ovoidlellipsoidal52-74 x 32-37 um; pedicel19-30 x 11­
20 urn; single large apical and basal fugaceous caudae swelling enormously in
water Podospora intestinacea N. Lundq. (Fig. 69)

16. Dark cell of ascospore ovoid/ellipsoidal with truncate base, 30-60 x 20--40 um;
pedicel straight or curved, 28-60 x 8-12 um; caudae not noticeably swelling in
water Podospora pyriformis (A. Bayer) Cain (Fig. 80)

17. Perithecial adomment in the form of short tubercles only 18

17. Perithecial adomment consisting of agglutinated or clustered stiff or flexuous dark
or light hairs/bristles 21

18. Apical and basal caudae only present; dark cell ovoid/ellipsoidal with truncate
base, 29-35 x 16-20 um; pedicel straight 44-72 x 11-13 um, sornetimes with
transverse septum Podospora dolichopodalis J.H. Mirza & Cain (Fig. 74)
18. Supemumerary caudae present in addition to lyre-shaped apical caudae 19

19. Asci 8-spored 20

19. Asci 16-spored; dark cell ovoid/ellipsoidal sornetimes with truncate base,
33--40 x 20-25 urn; pedicel cylindrical often flattened and twisted,
50-64 x 5-7 urn Podospora pleiospora (G. Winter) Niessl (Fig. 72)

20. Dark cell of ascospore ovoid/ellipsoidal, 26-31 x 13-17 um; pedicel cylindrical
often flattened and twisted, 30 x 5-7 um a single basal cauda also present
........................................................................ Podospora argentiniensis (Speg.) J.H. Mirza & Cain (Fig. 73)
20. Dark cell of ascospore ovoid/ellipsoidal, 36--42 x 19-24 urn; pedicel often
flattened and twisted, 46-63 x 6 um, no basal cauda
............................................................................ Podospora decipiens (G. Winter ex Fuckel) Niessl (Fig. 73)

21. Asci 4-spored; dark cell ovoid/ellipsoidal with a truncate base, 36--45 x 20-23 um;
pedicel straight, cylindrical, 21-29 x 4-5 um Podospora anserina (Ces. in Rabenh.) Niessl (Fig. 70)
21. Asci >4-spored 22

22. Asci 8-spored 23

22. Asci >8-spored 33

23. Dark cell of ascospore distinctly flattened on long side; ascospores with a single
apical and basal cauda 24
23. Dark cell of ascospore symmetrical about its long axis 27

24. Perithecial neck hairs flexuous, yellowish/hyaline 25

24. Perithecial neck hairs robust, dark, sorne agglutinated 26

25. Size of dark cell of ascospore very variable, even within the same ascus,
20-27 x 15-17 um; pedicel conical, 8-10 x 4-6 um Podospora ignota n. sp. (Fig. 82)
25. Size of dark cell of ascospore variable, even within same ascus,
19-26 x 12-17 um; sometimes I-septate; pedicel cylindrical, straight or slightly
curved 9-12 x 3--4 urn Podospora cf. didyma J.H. Mirza & Cain (Fig. 81)

26. Dark cell of ascospore 34-38 x 17-20 um; pedicel stout, straight or curved,
13-16 x 4-7 urn; caudae frequently fugacious, lobed, enveloping the ascospore
...................................................................... ..................................Podospora e.xcentrica N. Lundq. (Fig. 70)

26. Dark cell of ascospore 30--40 x 19-23 um; pedicel slender, straight,
19-24 x 4-5 um; single apical and basal caudae and sorne supemumerary caudae
............................................................................................................... Podosporafabiformis n. sp. (Fig. 79)

27. Ascospores uniseriate, with minute pedicels, ca 5-7 x 2-3 um; dark cell of ascospore
ellipsoidal, 34--40 x 21-25 um; caudae absent; dark cells surrounded by
frothy bubbles Podospora n. sp. (Fig. 79)
27. Ascospores biseriate; pedicels larger 28

45
28. Pedicels straight, 23-30 x 5 um with distinct central swelling (ca 9 um); dark
cell ovoid!ellipsoidal with truncate base, 29-40 x 15-22 um; single apical and
basal cauda Podospora prethopodalis Cain (Fig. 69)
28. Pedicels without distinct central swelling 29

29. Dark cell of ascospore > 35 um long; supernumerary caudae present 32

29. Dark cell of ascospore < 35 um long; supernumerary caudae present or absent 30

30. Ascospores with single whip-like apical and basal caudae 31

30. Ascospores with inflated, irregularly lobed caudae (visible in Indian ink mounts);
dark cell ellipsoidal with truncate base, 24-30 x 14-18 um, pedicels straight or
curved, 17-18 x 7 um Podospora bicornisN. Lundq. (Fig. 80)

31. Dark cell of ascospore variable, even within the same ascus, ellipsoidal with
truncate base, 24-30 x 14-17 um; pedicel delicate, cylindrical, 18-30 x 5-7 um
........................................................................................... Podospora hyalopilosa ? (Stratton) Cain (Fig. 81)
31. Dark cell of ascospore even-sized within the same ascus; dark cells ellipsoidal
with truncate base, 22-26 x 10-14 um, pedicel cylindrical, straight or slightly
curved, 10-11 x 2 um Podospora ellisiana (O. Griff.) J.H. Mirza & Cain (Fig. 66)

32. Dark cell of ascospore ellipsoidal, 38-45 x 22-25 urn; pedicel straight or curved,
27-30 x 5-9 um; 3-4 apical caudae, 2~3 basal caudae plus lobed supernumerary
caudae Podospora dactylina N. Lundq. (Fig. 71)
32. Ascospore dark cell ellipsoid 49--64 x 26-31 um; pedicel straight or curved,
49-70 x 10~-15 um; 3-4 apical caudae, 3-4 basal caudae plus a c1uster
ofsupernumerary caudae Podospora austrohemisphaerica N. Lundq. (Fig. 71)

33. Ascospores with 2 apical caudae; dark cell 19-24 x 12-15 um; pedicel 10 x 5 um
.............................................................................................................. Podospora bijida N. Lundq. (Fig. 67)
33. Ascospores with single apical and basal caudae 34

34. Dark cell of ascospore 22-25 x 13-16 um; pedicel of irregular width,
21-24 x 4-5 um Podospora ampullacea Cailleux (Fig. 68)
34. Ascospore dark cell < 20 um long 35

35. Dark cell ofascospore 16-20 x 10-13 um; pedicel 10-11 x 2-3 um
................................................................................................. Podospora setosa (G. Winter) Niessl (Fig. 67)
35. Dark cell ofascospore 14-15 x 7-8 um; pedicel 5 x 2-3 um
.......................................................................................... Podospora curvicolla (G. Winter) Niessl (Fig. 66)

Note: A culture of Apiosordaria verrucolosa has been deposited at CSS (CBS 110042). Type material of Podospora petro­
gale is deposited at Melbourne (MEL 2000062227) and a culture at ATCC (MYA 1392). Stratton (1921) and Mirza &
Cain (1969) described P. hyalopilosa as having four apical caudae. The present collection is not c1early 4-caudate at the
spore apex.
For the descriptions of the new taxa Podospora ignota and P. fabiformis see page 60.

Arnium Nitschke ex G. Winter emend. N. Lundq.

The genus Arnium is characterized principally by its cia are never produced in great abundance and conse­
ascospores composed of single dark cells provided quently are not recorded as frequently as are species of
with one or two inconspicuous germ pores. The spores Podospora. Keys to the known species of Arnium may
lack pedicels but do possess mucilaginous appendages be found in Lundqvist (1972) and Krug & Cain (1972).
usually in the form of single caudae, one at each pole. Additional species have been added by Cain & Mirza
Sorne species have asci provided with apical rings. The (1972), Furyua & Udagawa (1976) and Jeng & Krug
imrnersed or semi-immersed dark perithecia (ca 0.5 (1977a).
mm diam) may be adomed with conspicuous aggluti­ Considering Australian records of the genus,
nated or nonagglutinated hairs or bristles or the necks Dade's "Podospora taxonomie species VIIF' (specimen
may bear short inconspicuous tubercles. By them­ V119k) is Arnium arizonense (see Fig. 83). Cribb
selves, the morphology of the perithecia is of little help (1995) recorded Podospora absimilis Cain from
for the purposes of identification. Sorne Arnium spe­ Queensland (Arnium absimile (Cain) J.c. Krug &
cies are terricolous, others coprophilous. The perithe- Cain) and Jahn (2000) recorded Amium maerotheea
46
from dung collected in Queensland. To date 1 have Podospora gigantea and P. australis may be con­
recorded seven species of Arnium from Australia and fused with the genus Arnium due to the minute pedicel
Lundqvist sent a further new species to me from mate­ exhibited by these two species. For this reason they
rial collected in Tasmania. This will be formally de­ also appear in the key below.
scribed in a later publication.

Key to the species of Arnium

(G1ossary as for Podospora, Fig. 13)

1. Asci 4-spored 2
1. Asci > 4-spored 3

2. Protruding portion of perithecia adomed with short, single, hyaline-tipped hairs;

ascospore dark cell ellipsoidal, symmetrical, 53-60 x 29-32/lm; with a minute
apiculus (pedicel) at the base Podospora australis (Speg.) Niessl (Fig. 83)

2. Protruding portion of perithecia adomed by conspicuous agglutinated and/or single

bristles; ascospore dark cell ellipsoidal, symmetrical or slightly flattened on one
side, 50-53 x 27-30 um, without apiculus at base
....................................................................... Arnium arizonense (D.Griff.) N. Lundq. & J.C. Krug (Fig. 83)

3. Asci 8-spored 4
3. Asci multispored (at least 128); ascospores ovoid, with 1 germ pore,
17-22 x 10-12 um, surrounded by an irregular evanescent mucilaginous jacket
......................................................................................... Arnium absimile (Cain) le. Krug & Cain (Fig. 87)

4. Ascospore dark cell ellipsoidal, symmetrical 120-132 x 40-60 um, with a minute
apiculus (pedicel) at the base Podospora gigantea J.H. Mirza & Cain (Fig. 90)
4. Ascospores smaller, without apiculus 5

5. Ascospores with 2 strap-shaped caudae 6

5. Ascospores surrounded by a fringed mucilaginous jacket; spores ellipsoidal,
39-44 x 25-26 um; perithecia with a conspicuous tuft of agglutinated bristles
.......................................................................... Arnium macrotheca (Crouan & Crouan) N. Lundq. (Fig. 84)

6. Ascospores uniseriate, ellipsoidal, symmetrical, 42-47 x 23-30 um with 1 germ

pore Arnium olerum (Fr.) N. Lundq. & J.e. Krug (Fig. 85)
6. Ascospores biseriate; other features not as above 7

7. Perithecia adomed with a few short, non-agglutinated bristles; ascus with an apical
ring; ascospores ellipsoidal, symmetrical or slightly asymmetrical,
35-40 x 22-27 um Arnium caballinum N. Lundq. (Fig. 85)
7. Perithecia without bristles; ascospores not as above 8

8. Ascospores ellipsoidal or ovoid, sometimes slightly flattened on one long side,

52-59 x 22-28 um; caudae striate and wider than the dark cell at the base
.................................................................................... Arnium cirriferum (Speg.) J.e. Krug & Cain (Fig. 86)
8. Ascospores ellipsoidal, symmetrical; caudae narrower than dark cell at base 9

9. Ascospores 42-48 x 22-27 um; caudae with central channel .Arnium mendax N. Lundq. (Fig. 86)
9. Ascospores 48-57 x 21-27 um; no channel in caudae Arnium n. sp. (Fig. 86)

Strattonia Cif. emend. N. Lundq.

The genus Strattonia erected by Ciferri was emended mucilaginous sheath that swells considerably in water.
and circumscribed by Lundqvist (1972). It encom­ Sorne morphological characters of the genus indicate
passes both terricolous and coprophilous pyrenomy­ that it may stand equidistant between Sordaria and
cetes, characterized by having dark single-celled asco­ Podospora. For example the ring at the tip of the ascus
spores that have tiny isodiametric or s1ightly elongated and the encompassing ascospore sheath is reminiscent
pedicels at their bases. The dark cell is surrounded by a of Sordaria and the tiny pedice1 is a characteristic of

47
Podospora. Not aIl mycologists agree with the separa­ duced a Phialophora anamorph (Fig. 92 G), and S.
tion of Strattonia from Podospora, and the merit of the insignis produced a few aleuriospores together with its
separation becomes aIl the more questionable as fungi teleomorph. The Australian material of S. insignis had
are found which encompass features of both genera glabrous perithecia but in aIl other respects it matched
(e.g. Podospora petrogale). Lundqvist (1972) stated the published description of this species. In culture,
that the genus comprises sorne seven described species however, the perithecia of S. insignis are densely hairy.
together with four to five undescribed ones. In the S. zopfii was described as having a yellow tomentum
same publication he constructed a key to five Nordic on the perithecial venters (Lundqvist 1972), but this
species of the genus. Since then Barassa et al. (1985) feature was not especially noticeable in the Australian
have added a new species, Strattonia dissimilis. There collections, although in aIl other respects they matched
are relatively few records of Strattonia. The coprophi­ the given description of that species. Herbarium sam­
lous species prefer to fruit on very old dung, and usu­ pIes of a fourth (undescribed) species have been kindly
al1y only a few perithecia are produced at any one loaned to me by Lundqvist (Tibell 12û74-b and Tibell
time. 1 have found three species of Strattonia from 12û84-a). This species will be fully described in a
Australia, two of which 1 have found but once, the future paper.
other twice. In culture S. oblecythiformis n. sp., pro­

<, <,

'\
!
/ \
/

/
/
<;

Isodiametric Oblecythiform

Fig. 14. Glossary to terms used in the key below.

Key to Strattonia

] . Pedicel distinctly longer than wide 2

1. Pedicel more or less isodiametric 3

2. Dark cell of ascospore ellipsoidal, symmetrical or tlattened on one side, with

truncate base, 35-45 x 2]~25 um, pedicel straight somewhat oblecythiform,
7-15 x 3-7!lm; ascus ring indistinct Strattonia oblecythiformis n. sp. (Fig. 92)
2. Dark cell ofascospore ellipsoidal, symmetrical, (45-)50-58(-60) x
(24-)26-30(- 32) um; pedicel cylindrica1, straight or at an angle
to the dark cell, evanescent, ] 0-11 x 2 um Podospora petrogale A. Bell (Fig. 90)

3. Perithecial necks distinctly hairy; ascospore dark cell ellipsoidal,

symmetricalor slightly asymrnetrical, with truncate base, 63-77 x 31-42 um;
pedicel more or less isodiametric, 6.5-7 x 5.5 um Strattonia grandis n.sp. (Fig. 91)
3. Perithecial necks not hairy; ascospore dark cell < 50 um long 4

4. Dark cell of ascospore ellipsoidal, symmetrical, with truncate base,

35-42 x 20-27 um; pedicel isodiametric, 5-6 x 4 um Strattonia zopfiiN. Lundq. (Fig. 91)
4. Dark cell of ascospore ellipsoidal, symmetrical with truncate base,
45-50(-53) x 25-29(-35) um; pedicel isodiametric, 5 x 5 um
......................................................................................Strattonia insignis (E.C. Hansen) N. Lundq. (Fig. 92)

48
Zygopleurage Boedijn Fimetariella N. Lundq.

This is a very distinctive coprophilous genus who se The genus Fimetariella was established by Lundqvist
affiliations (based upon morphology), probably lie (1964a) with Sordaria rabenhorstii Niessl (in Ra­
close to Podospora and Cercophora within the benh.1872) as the type species. Krug (1995) discussed
Lasiosphaeriaceae. To date three species have been taxonomie aspects of the genus and provided a key to
described, aIl showing a strong preference for growing the eight known species. 1 have observed a hitherto
on cow dung (Lundqvist 1969). During this study, only undescribed species of Fimetariella. This may be the
one species Zygopleurage zygospora (Speg.) Boedijn same fun gus reported and illustrated by Larsen (1971),
has been found (a1so recorded on cow dung). The dark who stated that it was the same as Sordaria minima
cells measured ca 30 x 15 um and in respects matching Sace. & Speg. sensu Massee & Salmon. She was of the
the descriptions ofthis species (Fig. 93 A-E). opinion that the fungus was misplaced within
Sordaria.
Because of the smail size of the ascospores (7-9 x 4-5
Zygospermella Cain um), 1 cali it Fimetariella minuta n. sp. (Fig. 94 A-C).
1 found it difficult to ascertain the existence of "minor
A coprophilous genus compnsmg two species, pores" mentioned as being characteristic of the genus
(Lundqvist 1969). The genus is characterised by hav­ by both Lundqvist and Krug, although 1 have observed
ing two-celled, dark ascospores, constricted at the minute granules (?) in sorne ascospores at the pole
central septum, each cell furnished with a single polar opposite the gerrn pore. 1 have also located the same
gerrn pore. A single-strap shaped channelled cau da species on rabbit dung collected in the Orongorongo
covers each gerrn pore. Zygospermella insignis (Mou­ Valley in New Zealand.
ton) Cain has been recorded on cattle dung during this
study. The Australian material has ascospores within
the range: 60-69 x 14-16 um and in aIl respects Order XYLARIALES
matches the given description of this species (Fig. 93
F-L). Phomatospora Sace.

Phomatospora is a genus usuafly found on decaying

Bombardioidea e. Moreau ex N. Lundq. vegetation and currently considered to belong to the
Xylariales (Kirk et al. 2001) and a number of species
A small genus of sorne four species, aIl of which are have been shown to have a Spororthrix anamorph
coprophilous (Krug & Scott 1994). The dark perithecia (Rappaz 1992). Two species, Phomatospora minutis­
are approx. 1 mm high with a multi layered coriacious sima (Crouan & Crouan) N. Lundq. and P. coprophila
peridium. They appear singly or in small clusters on M.J. Richardson are coprophilous, but the anamorphs
the dung surface. A full description of the genus to­ ofthese remain unknown. To date 1 have found only P.
gether with historical data concerning its taxonorny is minutissima on three dung samples (cattle 2, swamp
to be found in Lundqvist (1972) and Krug & Scott wallaby 1). The tiny (ca 170 um diam) perithecia occur
(1994). Lundqvist considered the perithecial wall to be in swards, usually with the venters of the perithecia
in part stromatic and an archaic feature, while Krug & submerged, (although on one collection the perithecia
Scott consider it to be nonstromatic, its thickness being were superficial). The cylindrical asci have a tiny
due to adaptation to its xeric environment. apical ring and the eight ascospores are uniseriate. The
1 have not found this genus in Australia, but Dade left ascospores are hyaline, bluntly ellipsoidal, with a tiny
excellent records of a 4-spored species which he re­ guttule at each end. Ascospores sizes were within the
ferred to it by its old name Fimetariella maxima (Ni­ range 5-6 x 2-3 urn (Fig. 51 E-H).
essl) Cain and elsewhere as Bombardiodea. In 1982 1 Dade also found Phomatospora minutissima on wom­
studied his slide V532g and the current name for this bat and rabbit dung but wrote in his Rough Case Notes
fungus is Bombardiodea stercoris (De.: Fr.) N. Lundq. "1 don 't know il". Elsewhere he queried "? Allan­
Dade gives the ascospores measurements as (32-)39­ tosphaeriaceae/?". In 1982 1 studied two of his slides,
42 x (15-) 17-22 um. His illustrations are reproduced V101j (lMI 101018j) and V548g). At that time 1 also
in Fig. 94 O-G. did not know this fungus and left a note to this effect.
Krug (pers. comm.) has subsequently recorded B. Richardson (pers. comm.) also identified P. minutis­
anartia J.e. Krug & Scott from Australia. sima from cattle dung which he collected in Australia
in 1999. Jahn (2000) recorded P. coprophila on wal­
laby dung collected from Queensland by T.R. Loh­
meyer.

49
Family: XVLARIACEAE nately remained sterile. Attempts were made to induce
them to form their teleomorphs in the laboratory by
Hypocopra (Fr.) J. Kickx, Podosordaria Ellis & Holw. employing the technique described by Webster &
and Poronia Willd. Weber (2000) without success. Ali six species found
during this current study have asci which are fumished
These members of the Xylariaceae all have perithecia with prominent rings which stain a bright blue in Mel­
embedded in a stroma, asci with a prominent ring zer' s reagent.
which stains in Melzer' s reagent, and dark, single­ One of the first keys to species of Hypocopra is that
celled more or less ellipsoidal ascospores provided of Griffiths (1901). The most recent is that by Krug &
with a longitudinal germ slit. A smail evanescent Cain (1974b) and the same authors published a key to
hyaline basal cell may be present in Hypocopra spe­ Podosordaria species in the same year (1974a). Hyde
cies. The ascospores of ail three genera are surrounded et al. (1996) described a new species P. australiensis,
by an evanescent gelatinous sheath. Poronia was dis­ from wallaby dung.
tinguished from the other two genera by having 1 find the synoptic key to Hypocopra in Krug &
perithecia and embedded in a flattened stromatic dise Cain (1974b) confusing to use. Although one may
which may or may not be subtended by a long (Le. at theoretically enter the key using a number of features
least lem) stalk emerging from the dung surface. In (e.g. stroma colour, texture, position in relation to the
Podosordaria the protruding perithecia are typically substrate, etc.), in practise each identification has
c1ustered on a semicircular to subglobose stalked dise ultimately to rely upon ascospore dimensions, which in
(although a few species are described as having a this key are drawn extremely cIosely with many over­
sessile stroma). In Hypocopra the stroma is always lapping ranges between the species. For these reasons,
sessile and situated just below the dung surface. In 1 have been unable to identify sorne of the species to
practice it is difficult to distinguish between sessile my satisfaction. Sorne of the Australian finds may be
Podosordaria and Hypocopra species. A Lindquistia new, but without consulting type material, 1 am unable
anamorph was first associated with Poronia by the to ascertain this point.
brothers Tulasne (volume Il, 1863). Later Po­ Sorne authors have described various species of
dosordaria was defined as having a Lindquistia ana­ Poronia, P. punctata being one of the best-known
morph (Rogers & Lœssoe 1992), although Rogers et species (Dennis 1978). Poronia erici Lohmeyer &
al. (1998) now relegate species shown to possess a Benkert was described from Australian material by
Lindquistia anamorph to Poronia, based upon their these authors in 1988, the specifie name honouring the
further examination of the type species of Po­ late mycologist Erich lahn. Other authors dealing with
dosordaria. The number of genera involved here re­ this genus include Lim (1968) and long & Rogers
mains speculative. Here 1 have confined Poronia to (1969).
species with a flat-topped, sta1ked or sessile stroma in Dade recorded Poronia punctata which he obtained as
which the perithecia are embedded. a "dried specimen" presumably collected in an already
Ali three of these genera tend to occur on old dung mature state in the field), by Crichton (sIide V426).
or after the dung has been incubated for a long time (at Another sIide (V143A) is recorded as the same spe­
least six weeks), none of them are particularly com­ cies. Dade (surprisingly) did not leave any drawings of
mon on dung as compared with other taxa such as this species. Cribb (1988) recorded P. punctata from
Podospora or Sporormiella. Although a number of dung in Queensland, but his illustration suggests that
species of Podosordaria and Poronia developed their the species he saw was P. erici. In another article
stromata on incubated dung during this study, only (1996a), Cribb recorded Xylaria brasiliensis from
three produced their teleomorphs, the others unfortu­ Queensland.

Key to the species of Hypocopra, Podosordaria and Poronia

1. Peritheciaborne aloft on tall (ca 1-2 cm) branched or unbranched, dark stromata;
ascospores more or less uniseriate, more or less symrnetrical, and ellipsoidal 2
1. Stromata sessile, appearingeither as a ca 2 mm diam pale dise, or dark and
partially hidden withinthe dung; ascospores more or less symmetrical or distinctly flattened on one side ..... 3

2. Peritheciaembedded in flattenedtops of branched stromata; ascospores 21-30 x

3-17 um Poronia oedipus (Mont.) Mont. (Fig. 97)
2. Peritheciaembedded along the sides ofunbranched stromata which have whitish
conidial heads; ascospores 24-29 x 14-16 um Podosordaria sp. (Fig. 96)

3. Pale (3A3), sessile dise in which the perithecia are embedded; ascospores more or
less symmetrical, 26-30 x 14-18 um Poronia erlci Lohmeyer & Benkert (Fig. 97)
3. Peritheciaemerging from dark, more or less subterranian stromata 4
50
4. Ascospores > 30 um long 5
4. Ascospores 20-30 x 13-16!!m Hypocopra equorum? (Fuckel) G. Winter (Fig. 95)

5. Ascospores conspicuously flattened on one side, ellipsoidal in other view

8--42 x 13-14 um Hypocopra sp. (Fig. 95)
5. Ascospores symmetrical or slightly flattened on one side, 40--45 x 21-25Jlm,
germ slit extending the whole lengthof the ascospore
.................................................................Podosordaria violacea (Ellis & Everh.) J.C. Krug & Cain (Fig. 96)

Family PHAEOTRICHACEAE

Trichodelitschia Munk

Trichodelitschia Munk is a small bitunicate genus outline of residual cytoplasrn is a matter of conjecture.
comprising three coprophilous species. It is character­ The genus is placed in the family Phaeotrichaceae a
ized by small (ca 300-500 um diam), gregarious, family comprising both c1eistothecial and perithecial
superficial or serni-irnrnersed, black perithecia. Fre­ genera, and the genus Phaeotrichium is the c1eistothe­
quently the perithecia are reclining rather than upright cial analogue of Trichodelitschia.
and in Trichodelitschia adelphica the perithecial neck Dade also found species of Trichodelitischia and left
is sometimes poorly defined and the ascomata superfi­ slides and iIIustrated notes on his observations. He
cially resemble cleistothecia of genera like Zopfiella or identified his specimens as T. bisporula but recognized
Kernia. Copious stiff, dark, non-septate bristles adom that there were "a smaller and a larger strain" in his
the perithecial necks and are sometimes also unevenly collections. In 1982 1 studied sorne of his slides and
scattered over the venters. The asci are bitunicate and found that V132e, V267b and V618f were T. munkii
fumished with an apical ring. The ascospores are while 1 recorded V173e as T. bisporula at that time.
uniseriately arranged, each comprising two dark cells However, the sizes of the ascospores which 1 measured
each with a prominent polar germ pore. The exact border on those described for T. adelphica (26-31 x 9­
nature of the inner ascus wall remains elusive. Barr 10 um) and hence 1 may have misidentified this slide
(1990) stated that there is no inner ascus wall but at that time. The publications by Lundqvist (1964b)
instead the terminal ascospore upon dehiscence carries and Luck-Allen (1970) have been used to identify the
aloft the ascus ring. In water mounts 1 have observed Australian collections.
the faint outline of something resembling an inner
wall; whether this is structurally a wall or rnerely the

Key to the species of Trichodelitschia

1. Ascospores 29~34 x 13-15 urn; sorneperithecia reclining with ill

definednecks; bristles scattered over the venter and/or concentrated
around the ostiole Trichodelitschla adelphica Luck-Allen (Fig. 99)
1.Ascospores smaller 2

2. Ascospores 22-25 x 8~9 um Trlchodelitschla bisporula (Crouan) Munk (Fig. 98)

2. Ascospores 16-20 x 7-8 um Trichodelitschia munkii N. Lundq. (Fig. 98)

Order DOTHIDIALES

Family SPORORMIACEAE

Delitschia Auersw. and Semidelitschia Cain & Luck-Allen

Delitschia was established by Auerswald (1866), with widely distributed around the world. The history of the
Delitschia didyma Auersw. as the type species. Since genus together with an extensive key to the coprophi­
that date, sorne species have been added to the genus lous species can be found in Luck-Allen & Cain
(Cain 1934, Breton & Faurel 1970, Furuya & Udagawa (1972). Lignicolous species are keyed out in Hyde &
1976, Jeng et al. 1977, Hyde & Steinke 1996, Richard­ Steinke (1996).
son 1998b). Species of Delitschia are coprophilous or Delitschia is characterized by having clearly bituni­
Iignicolous, tropical or temperate, and appear to be cate asci and dark, ellipsoidal, mainly two-celled,
51
 smooth ascospores, fumished with longitudinal, pro duce pycnidial anamorphs. Ali of thern have 8­
straight or sinuous germ slits. Wh en fresh, the asco­ spored asci. None of the multispored species have yet
spores are surrounded by a transparent sheath. The been detected in Australia. This is somewhat unex­
ascus tip of many species appears to have a ring and it pected since the rnultispored species seem to be found
is particularly clear in D. vulgaris. The asci are ac­ predorninantly in tropical or wanner c1imates (Jeng et
companied by copious branched paraphyses that do not al. 1977c). Besides the other papers mentioned, the
appear to have free ends. The perithecia are usually papers by Cain (1934) and Luck-Allen & Cain (1975)
subrnerged in the substrate with only the necks pro­ have been the most useful for identification. As the
truding. Several of the species have roughened perithe­ authors state, sorne of the characters exhibited by the
cial necks or necks fumished with hairs, although the genus prove to be rather unreliable for identification
latter may disappear with age. Sorne perithecial necks purposes. For exarnple, the presence of hairs on the
have a distinct greenish tinge when viewed under perithecial necks is seemingly a constant feature in
transmitted light (e.g. D. winteriï, and the perithecial sorne species and variable in others. For this reason,
necks of many species elongate with age. species D. winteri and D. pachylospora are difficult to
Dade did not illustrate any species of Delitschia, distinguish since their ascospore sizes are very similar.
although he did leave sorne slides of various species, Similarly, D. furfuracea, D. tomentosa, and D. pata­
sorne of which 1 saw in 1982. Dade recorded slide gonica are hard to separate. The arrangement of asco­
VI02h (duplicates IMII 0109h, and DAR42848h), and spores within the asci is also an unreliable character
V274c (duplicate IMIl13409c) as Delitschia winteri. since the spores can be uniseriate at first but quickly
Dade's VI73d (duplicates IMII 05542d and DAR assume a biseriate arrangement when irnrnersed in
42909d), not identified to species, is Delitischia mar­ water on a si ide. Hence I have only used this character
chalii, a species that I have not found rnyself on Aus­ in the key for species where it appears to be stable.
tralian material. The genus Semidelitschia was established by Cain
Specimens of Australian dung fungi held at Upp­ & Luck-Allen (1969) to accommodate pyrenomycetes
sala forwarded to me by Lundqvist are: Tibell l2074e which possess bitunicate asci containing dark single­
and Tibell 11381 h (both D. winterîï, and Tibell 11194a celled ascospores, which otherwise resemble features
which I have identified as Delitschia gigaspora. Cribb of the genus Delitschia, with S. agasmatica as type
(1994) recorded D. vulgaris from dung collected in species. Mirza & Khan described a further species S.
Queensland. tetraspora in 1979. During this study a third species, S.
Ten species of Delitschia have been identified nannostellata, was found (Bell & Mahoney 2001 ).
during this study, two of which are in culture and

Key to the species of Delitschia and Semidelitschia

1. Perithecia dark, semi-immersed; ascospores uniseriate, ellipsoidal,

single-celled, with a longitudinal germ slit, 34-42 x 19-22 um
................................................................... Semidelitschia nannoste/lata A. Bell & D.P. Mahoney (Fig. 100)
1. Perithecia dark, semi-immersed; ascospores uniseriate or biseriate; two-celled,
ellipsoidal, each with a longitudinal, straight, or sinuous germ slit (Delitschia spp.) 2

2. Septa of ascospores very oblique and deeply constricted such that the
two cells of the ascospore appear to be adjacent, part-spores
25-26 x 15-16 um Delitschla didyma Auersw. (Fig. 101)
2. Septa of ascospores transverse or slightly oblique, constricted or
non-constricted, cells separable or non-separable 3

3. Ascospores bi-to multiseriate, straight or slightly curved, ce1ls

separable, 27-30 x 4---6 um Delitscùia leptospora Oudem. (Fig. 105)
3. Ascospores uniseriate or biseriate, and much larger 4

4. Ascospores ellipsoidal, equilateral or inequilateral, with conical,

rounded or hemispherical ends, septa transverse, cells readily separable
at maturity; germ.slits fulllength of the cells, sometimes extending at
right angles along the central septum; ascospores
39-54 x 19-22 um Delitschia variispora (?) Luck-Allen & Cain (Fig. 105)
4. Ascospores not as above 5

52
5. Most ascospores > 60 um long 6
5. Most ascospores < 60 um long 9

6. Ascospores ellipsoidal, ends rounded to conical, septa non-constricted

or slightly constricted; cells with minute pores in addition to germ slits;
ascospores 65-76 x 30-34 um Delitsehia sp. (Fig. 102)
6. Ascospores only with longitudinal germ slits 7

7. Ascospore septum transverse or oblique, conspicuously constricted;

cells separating at maturity, ascospores 66--76 x 27-32 um
........................................................................................ Delitschla ehodoeola Luck-Allen & Cain (Fig. 101)
7. Ascospore septa transverse or slightly oblique, non-constricted;
cells not separating at maturity 8

8. Ascospores ellipsoidal, equilateral or slightly inequilateral,

82-90 x 26-28 um, with conical to rounded ends; septa transverse,
non-constricted Delitsehia gigaspora Cain (Fig. 103)
8. Ascospores ellipsoidal, equilateral, 63-70 x 23-27 um, conical
to rounded ends, septa transverse to slightly oblique, non-constricted
or slight1yconstricted; cells not separable Delitsehia wlnterl PhiIl. & Plowr. (Cain) (Fig. 102)

9. Ascospores ellipsoidal, equilateral or slight1y inequilateral,

cells non-separable, 34-41 x 13-16 um; septa transverse;
asci with conspicuous apical ring Delitsehia vulgaris D. Griff. (Fig. 104)
9. Ascospores wider 10

10. Perithecia hairy; ascospores ellipsoidal, equilateral to slightly

inequilateral, minutely punctate, 47-59 x 20-24 um
...................................................... ..................................Delitsehia intonsa (?) Luck-Allen & Cain (Fig. 103)
10. Perithecia smooth; ascospores smooth, of different size II

II. Ascospores ellipsoidal, symmetrical, 49--65 x 22-25 um; septa

transverse, non- constricted or slightly constricted Delitschla furfuracea Niessl (Fig. 104)
II. Ascospores ellipsoida1, sometimes equilateral, more often
inequilateral, 45-53 x 22-25 urn; septa transverse or slightly
oblique, non-constricted or slightly constricted Delitschia excentriea Griff. (Fig. 100)

Note: Luck-Allen & Cain (1975) described the germ slit of Delitschia variispora as longitudinal, otherwise their description
fits the specimen here. The same authors made no mention of D. intonsa having minuately punctate walls to the ascospores, a
feature which was noticed in the Australian collection.

Sporormia sensu S. Ahmed & Cain and Sporormiella (Ellis & Everh.) S. Ahmed & Cain

Although they are most probably unrelated, these two both genera. The differences between them are sum­
genera are often considered together since the major marized in the key.
key available for identifying the species deals with

Key to Sporormia and Sporormiella

1. Ascomata ca 150 urn diam, not clearly ostio1ate; ascospores clustered
together centrally within ascus; ascospores 16-celled, 50-58 x 5 um,
each ascospore surrounded by a sheath which is extended to form prominent
lobed hyaline appendages at the ends ofthe ascospores Sporormia fimetaria (De Not.) De Not. (Fig. 106)
1. Ascomata larger, with clearly defined ostiole; ascospores uniseriate
or biseriate in the ascus Sporormie/la spp. see key below

Richardson & Watling (1982) described the ascospores with ascospores of 37--40 x 3.5--4 um considering at
of St fimetaria as 37.5--42 x 3-3.8 um, while Ahmed & that time that this cou Id represent a new species since
Cain (1972) describe them as larger, 50-57 x 3-3.5 the measurements were smaller than those given by
um. In 1983 1 recorded S. fimetaria from New Zealand Ahmed & Cain. In a later paper Richardson (l998a)

53
 clearly described the confusion that surrounds the within the same family or genus (e.g. Malloch & Cain
specifie epithets Sporormia fimetaria (De Not.) and S. 1971b, von Arx 1973). Thus von Arx (1973) suggested
fimetaria sensu Ahmed & Cain. He also indicated that that the ostiolate genus Sporormiella should be united
the original description by De Notaris in 1849 gave with the non-ostiolate genus Preussia. His main argu­
neither ascospore measurements nor a scale with the ment hinged upon the fact that sorne fungi ascribed to
illustration (the latter being at odds with the descrip­ pyrenomycete genera may be non-ostiolate in culture.
tion). Dissing (1992) described and i1lustrated S. He also stated that poorly preserved and non­
fimetaria (De Not.) De Not. from several collections representative herbarium material is no substitute for
made in Greenland giving ascospores as 40-55 x 3-3.5 working with living fungi, preferably in pure culture,
um, thus intermediate between the measurements where ail stages of development can be studied under
given by Ahmed & Cain and those given by Richard­ defined, controlled conditions. This argument presup­
son & Watling (1982). Dissing also illustrated the poses that ail these pyrenomycetes can be induced to
large single hyaline ascospore appendages described as grow and fruit in culture, which is not yet the case for
"pointed". He assumed that the ascospores were most many coprophiles. It also presupposes that morphology
likely discharged as a whole bundle in the manner of is stable in cultured fungi, although they frequently
Sacco bolus species. Lundqvist (pers. comm.) described produce abnormal ascomata with multiple ostioles,
a collection in which the appendages were "spear­ submerged reclining perithecia, partially fertile
like". However, one of the most outstanding features perithecia, etc. Furthermore it may be difficult to acer­
ofthe Australian material is the presence ofhuge lobed tain whether a particular ascocarp is ostiolate or not,
ascospore appendages extending from both ends of particularly in Sporormia where they are very small
each ascospore (Fig. 106 F and G). The very pale and collected so infrequently. During the thirty years
purple-brown ascomata of the Australian collection are since the publications of Malloch & Cain and von Arx,
so small that 1 have been unable to ascertain whether a number ofmycologists have taken up their views and
or not they are cleistothecia or if in fact a tiny ostiole is transferred species of Sporormiella to the earlier genus
present. In water the ascomata immediately burst Preussia (see Guarro et al. 1997, Valldosera & Guarro
t1inging the asci out. Discharged ascospores readily 1990, and Abdulla et al. 1999. Other mycologists have
separate from one another in water mounts and are chosen to uphold the older scheme of recognizing the
held in the meniscus suspended by their enormous ostiole as an important distinguishing feature of
appendages. pyrenomycete genera. Most species of Sporormiella
have not been obtained in culture but do form clear
unambiguous ostioles to their ascomata on the natural
Sporormiella (Ellis & Everh.) S.I. Ahmed & Cain dung. Therefore 1 choose to recognize Sporormiella for
the species, reserving the name Preussia for clearly
Sporormiella includes sorne of the most commonly non-ostiolate counterparts. Von Arx (1973) also rec­
encountered fungi found on dung, S. minima being the ognized that coprophilous species of Sporormiella are
most frequently recorded species. Sorne species de­ principally ostiolate and soil-borne species of that
velop early on the incubated dung fruiting (typically) genus tend to be non-ostiolate, and he convincingly
within 2 weeks, others occur on old dung or dung suggested that selection pressures have caused this
which has been incubated for a long time (after 1-2 difference, insofar as soil and root-inhabiting species
months). There are little or no distinguishing features have no void in which to eject ascospores. In any case,
in the perithecia of most species except for the fact that should a particular species be proven to form both
those having large ascospores tend to have larger ostiolate and non-ostiolate ascomata it seems a simple
perithecia. The perithecial necks of most species elon­ enough task to key them out under either generic
gate and/or branch with age, the elongated portion name. There is another practical reason for maintaining
frequently being semi-transparent. Ali species have the genus Sporormiella. The genus is in dire need for
clearly bitunicate asci, but details of the ascus tip vary revision. Until this occurs, simply transferring ail
between the species. All species have transversely existing putative species of Sporormiella to Preussia
septate multicellular ascospores, usually four-celled. will only add further confusion.
The ascospores of many species have a tendency to The most complete key to the species is Ahmed &
separate at maturity. In all species the ascospores are Cairi's Revision of the genera Sporormia &
encased in a transparent sheath that disappears with Sporormiella (1972), in which the authors describe
age. The morphological variation within this large sorne 66 species from Argentina, Canada, Mexico and
genus suggests that further evidence may prove the United States. Morphological features which they
Sporormiella as it currently stands is polyphyletic. use to delineate the species include: smooth vs. hairy
In the early 1970's sorne mycologists considered perithecia, the nature of the ascus stalk, the number of
that the variable presence or absence of an ostiole in cells per ascospore and the shape and position of the
soil-bome and coprophilous pyrenomycetes should germ slit. In addition, Furuya and Udagawa (1972) is
either not be used to separate genera or that non­ helpful for identification purposes. However, identifi­
ostiolate and perithecial genera should be placed cation of the most commonly encountered species of
54
 Sporormiella found in Australia and New Zealand to those with button-like stipes), e.g. S. dubia S.I.
proved to be a challenge because few of the stated Ahmed & Cain, S. grandispora (Speg.) S.1. Ahmed &
characters are clearly distinctive. For example, 1 find Cain and S. megalospora (Auersw.) S.1. Ahmed &
(as Ahmed & Cain themselves intimate) that there is Cain, or the series: S. isomera S.I. Ahmed & Cain, S.
an overlap between ail their described species: leporina (Niessl) S.I. Ahmed & Cain and S. tetramera
Sporormiella minima - S. minimoides - S. australis ­ S.I. Ahmed & Cain, aIl of which (based on ascospore
S. intermedia - S. teretispora - S. eylindrospora dimensions), form a series of closely related taxa. In
through to S. borealis, with no clear separation in any addition, the Australian material of S. insignis strad­
of the morphological features. Booth (1982) was also dIes two species, S. insignis and S. splendens of Ah­
unable to separate S. australis, S. intermedia and S. med and Cain, having the ascospore width of S. in­
similis and he assigned collections of Arctic species to signis and the ascospore length of S. spendens.
S. intermedia because "this name predates those ofthe Dade left a comprehensive set of notes regarding
other eonsidered taxa". On present evidence, it is only his observations of Sporormia and Sporormiella. He
a matter of opinion as to where the specifie divisions also corresponded with Cain prior to the publication of
may lie. If all these species were combined, one would Ahmed & Cain's 1972 revision of the genera. Dade
be left with a single species whose ascospore sizes initially referred to aIl these species under the older
range would be 27-90(-100) x 4-16(-19) um! Far name of Sporormia but changed this in later notes.
broader than even the most ardent "Iumper" would When Ahmed & Cain's monograph was published,
contemplate! Evidence from sources other than anat­ Dade drew up a table of ail the species, illustrating the
orny and morphology may elucidate the situation. Ms. various diagnostic features to aid identification. He
Asa Nyberg is currently undertaking research using also sent various specimens to Cain for his opinions.
DNA sequencing techniques (Lundqvist pers. comm.) Dade certainly recognized S. minima but he too re­
on this genus. In the meantime one is left with the very corded difficulties in separating collections that fell
practical problem of leaving sorne guidelines to those into the minima/australis/intermedia taxa and commu­
wishing to identify their collections. For these reasons nicated the problems to Gunnell (Nov. 1964). He could
1 have chosen to recognize S. minima, extending its recognize S. minima with ascospores 28-35(-38) x 4­
circumscription to include S. minimoides Ahmed & 5 um listing slide nos. V128e(1), V214b* and V251 b
Cain. 1 have found collections that fit the given de­ as examples. Later (1966), he found other collections
scriptions of S. australis (Speg.) Ahmed & Cain, S. (V442f, V444e*, and V586IIe), which he also identi­
borealis (1. Egel.) J.C. Krug and to combine specimens fied as S. minima, and sent portions ofV442fto Cain).
which straddle S. intermedia, S. teretispora and S. At one point during his research Dade did identify
eylindrospora under the single (temporary) name of S. sorne of his specimens as S. australis (V 132d* and
intermedia sensu lato. For this publication S. interme­ V442h*). Other collections he found more difficult to
dia is more broadly defined: ascospores (50-) 55-75 x place. His collections VI28e(2), VI7Ih*, VI74f*,
(9-)10-13(-14) urn and the germ slits variously orien­ V182p, V191j, V256e*, and V258e were intermediate
tated from parallel (with a central kink), to diagonal. between S. minima and S. australis. To summarize,
The problem is made even more confused because Dade found from his own observations that there was a
Ahmed & Cain (1972) did not investigate Des­ size continuum in S. minimalS. australislS. intermedia.
mazière's type of Sphaeria intermedia held in Paris, About a year later (June 1965), Dade described his
but they studied secondary collections, and the correct specimen V251b (which he hitherto had called S.
name may in fact be Sporormiella ovina (Roberge ex minima), as an unknown species (t'Taxonomie speeies
Desm.) (Lundqvist, pers. comm.). Sorne seven collec­ F'). He described the asci as having "short sinuous
tions encountered had ascospores that were larger than stipes" and the ascospores as "tapering a little at eaeh
S. intermedia as defined above, and may match S. end", which would not place this specimen with S.
borealis (1. Egeland) J.C. Krug with ascospores minima as delineated by Ahmed & Cain. A pencil note
(73-)75-90(-100) x (11-)13-16(-19) um. These written by Dade indicated that he re-identified V251b
collections frequently show asci containing irregular as S. leporina. He sent sorne of a matching specimen
numbers (Jess than 8) of ascospores per ascus, a factor (V322a) to Cain who stated that it was "similar to S.
that may bedevil correct ascospore measurements. isomera but has narrower ascospores", This observa­
Scatter diagrams have been used to iIIustrate the range tion vindicates my view that S. isomera and S. leporina
of ascospore sizes in these particularly problematical may be synonyms. The fact that Dade originally con­
species (Fig. 106). It should be stressed that for this sidered specimen V251b to be S. minima also iIIus­
particular piece of research 1 have not seen any of the trates the difficulty that sometimes exists in separating
type material to which Ahmed & Cain refer, nor have 1 species with short-stipitate asci (S. isomera) from those
seen many collections outside those made in Australia with button-like stipes (S. minima), which is a neces­
or New Zealand. sity at one point in the Ahmed & Cain key.
A similar problem may be encountered when trying Dade grouped his problematical Sporormiella
to separate the species having stipitate asci (as opposed species which straddled the taxa S. australis - S. in­
termedia as "Taxonomie sp. II", which he described as
55
having ascospores 36-47 x 7.5-8 urn (specimens V988c), which he tentatively named as: S. lata (?),
V313IIb*, V329d, and V442h). He sent a portion of having ascospores 60 x 15 um, but he did not leave
the latter specimen of this group to Cain. Cain replied any illustration of this specimen and 1 did not locate
that S. australis had a "more oblique germ slit and that any slide of it in 1982.
he would like to see more material before arriving at a ln addition to the siides left by Dade, 1 received
decision". Later Dade added more specimens, which several collections of Australian Ascomycetes by
fell into this problematical group (V450a, V6051ik, Lundqvist that are held at the herbarium in Uppsala. Of
V616j and V998e). these collections, the following identifications of
Dade confirmed his specimen V 1190* as Sporormiella spp. have been made: S. borealis (Jonsell
Sporormiella megalospora. His "Taxonomie speeies 5247-b and Tibell 11194-g), S. eymatomera (Tibell
V" may be what 1 am identifying here as S. borealis. 122094-q), and S. insignis (Tibell 11258-n).
The material exhibiting 8-celled ascospores (Dade's Cribb, (1988, 1992, 1994, 1998), identified the
V209c, V230b, V373c and V405e; ail samples on following species of Sporormiella from a variety of
possum dung) was confirmed as S. bipartis by Cain. dung substrates collected in Queensland: S. australis,
Dade left excellent drawings ofthis species and since 1 S. dakotensis, S. leporina, S. nigropurpurea, S. min­
have not yet found this species in the Australian mate­ ima. S. bipartis/ontariensis ? Jahn (2000) also identi­
rial myself, 1 include his drawings here (Fig. 113 F ­ fied S. minima from wallaby dung collected in Austra­
1). lia by Lohmeyer.
1could find no references to any Taxonomie speeies Ali these species of Sporormiella are illustrated
III or IVofthis genus in Dade's notes. There is a small here with the same magnification to allow for easier
reference to a Taxonomie speeies VI (specimen comparisons.

Asci

1 /~~
t /-
Perithecia

L
Hairy Smooth

r - A

l •- •
Asospores
• III
9
Cylindrical Tapered

1 ••••
Button stipe

Parallel Oblique Diagonal Transverse

Germ. slits

l • Il Kinked Right·angled

Fig. 15. Glossary to terms used in the key below.

Elongated stipe

56
Key to the species of Sporormiella

1. Exposed portions ofperithecia densely and conspicuously hairy 2

1. Exposed portions of perithecia smooth 3

2. Perithecial neck hairs brown, sparsely branched; asci long-stipitate; ascospores readily
separable into l-celled units; gennslits parallel; ascospores 40--46 x 6-7 um
..................................................................Sporormiel/a chaetomoides (Griff.) S.I. Ahmed & Cain (Fig. 115)
2. Perithecial neck hairs hyaline, not branching; asci long-stipitate; ascospores
readily separable into 1-celled units; germ slits parallel; ascospores 20-25 x 3--4 um
.......................................................................................................... Sporormiel/a albolanata n.sp. (Fig. 115)

3. Ascospores 4-celled 4
3. Ascospores > 4-celled 15

4. Asci (prior to discharge) contracted abruptly below to fonn a button-like stipe; ascospores cylindrical 5
4. Asci (prior to discharge) with short or elongated stipes; ascospores tapered or cylindrical 8

5. Majority of ascospores < 50 um long 6

5. Majority ofascospores > 50 um long 7

6. Ascospores readily separable into 1- or 2-celled units with parallel, oblique, diagonal
and/or kinked germ slits; ascospores 27-38 x 4-7 um ..
.......................................................................Sporormiel/a minima (Auersw.) S.I. Ahmed & Cain (Fig. 107)
6. Ascospores readily separable into 1-celled units, with parallel, oblique, diagonal
and/or kinked germ slits; ascospores 35--48 x 7-8 um .
..........................................................................Sporormiel/a australis (Speg.) S.l. Ahmed & Cain (Fig. 107)

7. Ascospores readily separable into 1-celled units, with oblique, diagonal,

and/or kinked germ slits; ascospores (50-)55-75 x (9-)10-13(-14) um
............................................................................... ...................Sporormiel/a intermedia sensu lato (Fig. 108)
7. Most cells readily separable into l-celled units, with parallel, oblique, diagonal and/or
kinked germ slits; ascospores (73-)75-90(-100) x (11-)13-16(-19) um
...................................................................................Sporormiel/a borealis (1. Egeland) J.c. Krug (Fig. 108)

8. Ascospores < 50 um 9
8. Ascospores > 50 um Il

9. Ascospores cylindrical; germ slits parallel, oblique and/or kinked, 27-32 x 6-8 um
..................................................................................... Sporormiel/a subtllls S.I. Ahmed & Cain (Fig. 109E)
9. Ascospores tapered at both ends; germ slits parallel, oblique, diagonal and/or kinked 10

10. Ali ascospore cells more or less same size: ascospores 26-34 x 5-8 um
..................................................................Sporormiel/a cymatomera S.I. Ahmed & Cain (Fig. 109 A-D) 10.
10. Ascospores with elongated end-cells; ascospores (27-)30--41 x 6-8 urn
............................................................................. Sporormiella tetramera S.l. Ahmed & Cain (Fig. 109 G-J)

11. Ascospore cells each with several parallel, oblique or diagonal germ slits;
ascospores 68-98 x 20-24 um Sporormiel/a euryospora (?) S.I. Ahmed & Cain (Fig. 112)
11. Ascospore cells each with one germ slit.. 12

12. Ascospores mostly < 80 um long 13

12. Ascospores mostly > 80 um long 14

13. Ascospores 50-56 x 10-13 um; germ slits oblique, diagonal, or right-angled
.................................................................... Sporormiel/a grandispora (Speg.) S.I. Ahmed & Cain (Fig. 110)
13. Ascospores 70-85 x 15-17 um; germ slits oblique, diagonal, transverse, or
right-angled Sporormiel/a megalospora (Auersw.) S.l. Ahmed & Cain (Fig. 110)

14. Ascospores 82-93 x 13-16 um Sporormiel/a longisporopsis S.l. Ahmed & Cain (Fig. 111)
14. Ascospores 120-130 x 19-241lm Sporormiel/a ovina (Desm.) S.I. Ahmed & Cain (Fig. 111)

57
15. Ascospores 8-11 celled, tapered at both ends; uppermost ascospore in
the ascus with one very enlarged cell (approx. 22 pm wide); ascospores 125-140 x 16-17 um
............................................................Sporormiella herculea (Ellis & Everh.) S.I. Ahmed & Cain (Fig. 114)
15. Ascospores 8-celled; all cells of ascospores more or less of same size 16

16. Ascospores 44-60 pm long 17

16. Ascospores mostly > 60 pm long 18

17. Ascospores 44-60 x 5-7 um; cells readily separable; germ slits diagonal
............................................................................. Sporormiella bipartis (Cain) S.I. Ahmed & Cain (Fig. 113)
17. Ascospores 50-60 x 9-10 um; cells inseparable; germ slits oblique to diagonal
......................................................................................Sporormiella octonalis S.I. Ahmed & Cain (Fig. 113)

18. Ascospores 78-94 x 12-14 um, cells separable; germ slits transverse in
mid-cells parallel in end-cells Sporormiella platymera S.l. Ahmed & Cain (Fig. 114)
18 Ascospores 164-180 x 19-21 um, cel1s separable; germ slits oblique
to diagonal Sporormiella insignis S.I. Ahmed & Cain (Fig. 112)

Note: Dade's slides marked with * (page 55 & 56) are slides which 1 examined in 1982. At that time 1 erroneously label1ed
his slide VII90 as Sporormiella cylindrospora.

Glossary

Acerose Clavate Doliiform Ellipsoid Fusiform

Oblate Ovoid Obovoid Pyriform Obpyriform Saccate

• •• • •• ••
.. • • ••
• •
••• .•
• •
L _ _ _ _·

Acicular Subulate Uncinate Granulate Punctate Reticulate

Fig. 16. General glossary of various shapes and omamentations.

58
Appendix 1 Characteristics on dung: Perithecia scattered or aggre­
gated, ovate, about 0.75 mm diam, immersed in sub­
strate with only the dark necks protruding. A dense
Ascobolus dadei A. Bell & D.P. Mahoney, n. sp. white tomentum composed of narrow (circa 4 um
wide), thick-walled branched hyphae clothing the
Apothecia aggregata, sessilia, semi-immersa vel immersa, doIi­ venter (Fig. 88 G). Neck clothed in short straight blunt
iformia, ca 0.5 mm diam, albida vcl ochracea: paraphyses hyaline,
septatae. Asci clavati, operculati ca 260 x 55 um, 8-spori.
hairs, about 15-20 um long and 4-5 um wide. Neck
Ascosporae biseriatae, ellipsoideae, (35-)40-50(-54) x (17-)18­ hairs brownish at the base, hyaline at the tip, with a
22 urn (ornamento incluso). Episporium carmineo-roseum, slightly irregular surface (Fig. 88 H). Neck becoming
crassum, verruculosum, saepe pustula terminali manifesta, ridged upon drying. Ascus mass a very pale pink (12
aliquando fissuratum. Fungus fimicola. A2). Paraphyses slender, 1.5-2 urn wide, appearing
free-ended. Asci cylindrical, about 350-400 x 25 um,
Characteristics on dung: Apothecia scattered, sessile, with a conspicuous apical ring (about 5 urn diam), 8­
usually semi-immersed occasionally almost completely
spored and becoming costate when empty (Fig. 88 J).
immersed, doliform, dull whitish to pale ochre- yellow
Ascospores biseriate, rod-shaped and straight when
(5B4), about 0.5 mm diam. Commonly sorne dung
young, becoming clavate, sigmoid and initially hya­
debris scattered over the sides of the apothecia (Fig. line. At maturity transversely septate to form a dark,
19A). Ususally < 10 ripe asci protruding at any one
ellipsoidal, slightly beaked upper cell, 27-32 x 15-16
time. Apothecial wall of textura angularis (Fig. 19C).
urn with a truncate base and terminal germ pore and a
Paraphyses hyaline, septate. Asci c1avate, about 260 x
lower sigmoid or slightly bent hyaline pedicel, 45-55
55 urn, operculate, 8-spored (Fig. 19C). Ascospores
x 4 urn, A small gelatinous cauda present at each end
biseriate, ellipsoid, rose in colour at maturity (12A5-6),
of the ascospores. Many ascospores remaining in a
(35-)40-50(-54) x (17-)18-22 um, epispore omamen­
hyaline or fuscus, non-septate state (Fig. 88 K). Asco­
tion of darker rose, composed of coarse, coalescing
spores capable of germination in ail states of develop­
warts, which may be enlarged to form distinctive
ment producing germination hyphae from both the
blisters at the poles. Sorne ascospores having paler
apical and pedicel ends of the spores (Fig. 88 M).
fissures besides the coarse warts. A crescent of a hya­
Anamorph Cladorrhinum sp., exhibiting the same pink
line gelatinous substance sometimes attached to one or
colouration as the centrum contents (Fig. 88 L).
both sides of the ripe ascospores (Fig. 19B).
Frequently found on dung of marsupial mammals,
Found on three collections of wombat dung collected
particularly kangaroo, wallaby and wombat. During
from Tasmania.
this current study, the species has been recorded 14
times from dung collected in Queensland, Tasmania Etymology: grandiusculus = 'rather large " referring to the
and Victoria. ascospores which are larger th an those encountered in most
coprophilous Cercophora species.
Etymology: dadei named after Dade who tirst recorded this
species. Holotype: On wombat dung collected on E. shore of
Bathurst harbour, Tasmania, 43° 21' 46" S,145° 13' 30" E,
h
Holotype: On wombat dung collected at Spain Bay Port coll. Janet Fenton, 1i Apr. 1998. PDD 73858.
Davey, Tasmania,. 43° 22' S, 145' 57' 47" E, coll. Janet
Fenton, Il th Apr. 1998. PDD 76460.
Other specimens: AUSTRALIA: (DAR) Orange, NSW, Cercophora recta A. Bell & D.P. Mahoney, n. sp.
slides made by H.A. Dade: V194f, V396f, V537d, V827b,
and V878a. SWEDEN: (UPS) Tibe1l11205-b (slide). Perithecia disseminata, ca 0.5 mm diam, ventre subterraneo, colla
emergente, superficie velutina. Peridium textura prismatica. Asci
clavati, ca 250 x 50 um, S-spori, anulo apicali indistinto praediti.
Cercophora grandiuscula A. Bell & D.P. Mahoney, n. Ascosporae juvenes bacillares, fasciculum centrale in asco
formantes. maturae uniseptatae, cellula superiore brunnea,
sp.
ellipsoidea, basi truncata 26-32 x 15-18 um, poro germinali apicali
praeditae. Pedicellus rictus 43-58 x 6-7 um. Cauda gelatinosa
Perithecia disseminata vel aggregata, ventre immerso, ovoideo, ca attenuata, ascosporae utrinque affixa. Fungus fimicola.
0.75 mm diam, tomento albo ubique obtecta colla conico excepto.
Collum emersum hirsutum, atrurn, siccum corrugatum. Centrum
perithecii dilute roseum. Paraphyses graciles, septatae, 1.5-2 um Characteristics on dung: Perithecia scattered, about
latae, liberae. Asci cylindracei, ca 350-400 x 25 um, anulo apicali 0.5 mm diam, immersed in the substrate, with the dark,
praediti, 8-spori. Ascosporae biseriatae, maturae uniseptatae, slightly roughened neck protruding (Fig. 89 G). Neck
cellula superiore brunnea, ellipsoidea, basi truncata, apice conico surface covered by dark, rounded papillae resembling
vel umbonato, 27-32 x 15-16 um, poro germinali apicali.
velvet pile (Fig. 89 H). Venter wall of textura pris­
Pedicellus sigrnoideus, hyalinus, 45-55 x 4 urn. Cauda gelatinosa
attenuata, in praeditae ascosporae affixa. Fungus fimicola. Status matica. Asci long-stalked about 250 x 50 urn, with an
conidialis Cladorrhinum. inconspicuous apical ring (Fig. 89 L), 8-spored. Asco­
spores rod-shaped wh en young and arranged in a
central bundle within the asci, becoming multiseriate
59
when mature (Fig. 89 K). At maturity ascospores
transversely septate to form a dark, ellipsoidal sym­ Characteristics on dung: Apothecia scattered, sessile,
metrical or slightly asymmetrical upper cell, 26-32 x pulvinate about 0.5-0.75 mm diam, whitish to pale
] 5-] 8 um, with an apical germ pore and a lower ochre (5B2,) beset with a dense vestiture of conspicu­
straight pedicel of variable length, about 43-58 x 6- 7 ous, hyaline, sparingly septate, smooth, thick-walled
um. A transparent single cauda situated at both ends of bristles, which are straight and pointed at their tips, up
the spore (Fig. 89 M). to 900 um long and somewhat inflated (about 5-8 urn)
Found on two collections of dung collected from at their bases (Fig. 31A and C). Rows of ripe asco­
the Kimberley region, Western Australia. spores can be seen protruding above the hymenium in
ripe apothecia (Fig. 3] A). Paraphyses slender, hya­
Etymology: rectus = 'straight", referring to the straight line, septate with free ends (Fig. 31 E). Asci cylindri­
pedicel of this species of Cercophora. cal, about ] 00 x ] 0 um, with indi stinct apical appara­
tus, 8-spored (Fig. 3] D). Ascospores uniseriate, hya­
Holotype: On dung of northern brush tail possum (?), col­ line, globose, 9-]] urn diam, with a prominently and
lected in Barnett river gorge, 16 S, ]26 10 E, Kimberley
03] 0

evenly reticulate hyaline episporium, particularly

region, Western Australia, coll. Pat Grey, 31 st Aug. 2000.
visible in lactic acid cotton blue (Fig. 3] D and F).
POO 73885.
One collection found on koala dung, collected from
the Koala Park, Sydney, New South Wales, ] 98].
Fimetariella minuta A. Bell & D.P. Mahoney, n. sp.
Etymology: sphaera = 'sphere', 'sporus ' = 'spore', referring to the
spherical ascospores.
Perithecia disseminata, late ovoidea, semiirnmersa, ca 150 um Holotype: On koala dung collected at Koala Park, Sydney, NSW,
diam. Collo conico pilis unicellularibus induto. Venter coll. A. Bell. 25'h Aug. 1981. PDD 75305.
perlucidulus. textura globulosa/angulari. Asci clavati, breves, ca
50-55 um longi, aqua inflata, 8-spori. Ascosporae primum
uniseriatae, deinde aqua biseriatae, unicellulares, hyalinae vel
dilute brunneae. ovoideae, symmetricae vel asymrnetricae, 8-9 x Podosporafabiformis A. Bell & D.P. Mahoney, n. sp.
4-5 um, poro germinali juxta extremum praeditae. Fungus fimi­
cola. Perithecia disseminata, serm-immersa vel irnmersa, globosa vel
ovoidea, ca 0.75 mm diam, perlucida, textura angulari. collo brevi
Characteristics on dung: Perithecia clustered, broadly hirsuto. Setae brunneae, septatae, ca 200-300 x 3- 4 um
Paraphyses filiformes. ventricosae. Asci clavati, 8-spori, ca 200 x
ovoid, minute, about 150 urn diam, partially sub­
45 um. annulo apicali parvo praediti. Ascosporae biseriatae,
merged in the substrate, the conical ostiolar region uniseptatae, cellula superiore brunnea, ellipsoidea vel deltoidea,
beset with short, one-celied hairs (Fig. 94 A). Perithe­ basi truncata, poro gerrninali apicali 30-40 x 19-23 um. Pedicellus
cial venter semitransparent and composed of textura hyalinus, cylindraceus, rectus, 19-24 x 4-5 urn. Cauda gelatinosa
globulosa/angularis. Paraphyses not in evidence. Asci attenuata in extremis ascosporae affixa. Duo caudae laterales
short-clavate, about 50-55 um long, rapidly swelling praesentes. Fungus fimicola.
in water, with no discernable apical apparatus, 8­
spored (Fig. 94 B). Ascospores initial1y uniseriate, but Characteristics on dung: Perithecia scattered semi­
rapidly assuming a biseriate arrangement in water, immersed or almost completely immersed in the sub­
one-celled, hyaline or a pale brown, ovoid, symmetri­ strate. Venters hairy, about 0.75 mm diam, of textura
cal or slightly asymmetrical, 8-9 x 4-5 um, with a
angularis, semi-transparent, with ripe asci visible
inside (Fig. 79 A). Perithecial necks short, dark,
germ pore at the narrower end and often small granular
clothed in clusters of stiff, dark, thick-walled, septate
cell inclusions at the opposite ends (Fig. 94 C).
bristles, ca 200-300 x 3- 4 um (Fig. 79 B). Old
Etymology: minutus = 'minute', referring to the tiny asco­ perithecia, with elongate necks and bristles almost
spores. disappearing. Paraphyses/interascal filaments com­
posed of columns of cells ofvarying width (Fig. 79 D).
Holotype: On rabbit dung collected from Macquarie Island, Asci clavate, about 200 x 45 um, with a small apical
54.5 S, 158.95 E, coll. S. Stephenson ]995, POO 76475. ring, 8-spored (Fig. 79 D). Ascospores biseriate, the
dark cell ellipsoid in one view, flattened or somewhat
D-shaped in another, with a narrow truncate base and a
Mycoarctium sphaerosporum A. Bell & D.P. Ma­ small apical germ pore, 30--40 x 19-23 urn. Pedicel
honey, n. sp. hyaline, straight, ]9-24 x 4-5 urn (Fig. 79 E). The
apical and basal caudae usually persistent. Sometimes
Apothecia disseminata, sessilia, pulvinata, ca 0.5--0.75 mm diam,
albida vel dilute ochracea, dense hirsuta. Setae hyalinae, raro
two lateral caudae arising from the top of the pedicel
septatae, laevigatae, crassitunicatae, rectae, basi inflatae, ad apicem visible (Fig. 79 E).
acuminatae, ca 900 x 5-8 um. Paraphyses graciles. hyalinae, Found twice during the course of this study on
septata, summae liberae. Asci cylindracei, ca 100 x 10 urn, pademelon (or wallaby?) dung from Tasmania and
apparatu apicali arnbiguo, 8-spori. Ascosporae hyalinae, globosae, quokka dung collected in Western Australia . Po­
9-11 um diam, regulariter reticulatae. Fungus fimicola.
dospora fabiformis somewhat resembles the descrip­
60
tions of P. horridula (Sace.) Dennis & S. Francis, a attenuatae 20-25 x 3--4 um, fissura germinali longitudinali
soil species (Francis & Sparrow 1984 and Guarro et al. praeditae. Fungus fimicola.
1991). Characteristics on dung: Perithecia scattered, im­
mersed in dung, venters dark, globose ca 200 um diam
Etymology: fabae = 'broad bean', referring to the shape of the dark with necks protruding, the latter becoming elongated
cell of the ascospores. with age (Fig. 115 F). Dark necks clothed with a dis­
tinctive woolly covering of radiating, white, un­
Holotype: On dung of pademelon or wallaby (?), collected at Bond
Bay, Tasmania, 43" 15' 14" S, 145" 54' 04" E, coll. Janet Fenton, branched hairs, which are about 200 urn long and 1-2
Il th Apr.1998. POO 76455. A culture of this species has been urn wide (Fig. 115 G). Venter of textura prismatica
deposited with the Centraalbureau voor Schimmelcultures (CBS with thickened patches in the neck region (Fig. 115 G).
112043). Paraphyses slender ca 2-3 um wide of indeterminate
length. Asci bitunicate, slender cylindrical with elon­
Podospora ignota A. Bell & D.P. Mahoney n. sp.
gated stipe, ca 120 urn long prior to dehiscence, 8­
Perithecia disserninata, ventre immerso, subgloboso. ca 0.75 mm spored (Fig. 115 H). Ascospores biseriate, 4-celled,
diam. Collo emerso, atrato, tomento luteo. Paraphyses filiforme, cylindrical with tapered ends, 20-25 x 3-4 um, each
ramosa, ca 3 um latae. Asci cylindracei, ca 250 x 25 um, 8-spori. cell with a more or less longitudinal germ slit (Fig. 115
Ascosporae uniseriatae, uniseptatae, cellula superiore brunnea, 1); ascospore cells tending to split apart upon maturity.
magnitudine variabiles, ellipsoideae, asyrnmetricae, basi truncatae,
This species has been seen twice on the dung ofwalla­
poro germinali apicali praeditae, 20-27 x 15-17 um, Pedicellus
hyalinus, conicus, 8-10 x 4-6 um. Cauda gelatinosa attenuata roo, coIlected in Western Australia and the Northern
basilaris, cauda superior canaliculata. Fungus fimicola. Territories.

Characteristics on dung: Perithecia scattered, im­ Etymology: albus = 'white', lanatus = 'woolly'; refer­
mersed in substrate, venter about 0.75 mm diam, semi­ ring to the distinctive white woolly coating to the
transparent, ripe asci visible within, only the cylindri­ perithecial necks.
cal neck emerging (Fig. 82 A and B); neck dark,
clothed in a fine weft of agglutinated, septate, yellow­ Holotype: On dung of wallaroo collected in the Keep
ish (4A5) hairs. Paraphyses filiform, branched, about 3 River National Park, Northern Territories, coll. Pat
urn wide seemingly not free ending. Asci cylindrical, Grey, 15th Aug. 2000. PDD 73888.
about 250 x 25 urn, 8-spored, no apical apparatus
recognisable, becoming costate when empty (Fig. 82
D). Ascospores uniseriate, dark cell ellipsoidal, asym­ Strattonia grandis A. Bell & N. Lundqvist, n. sp.
metrical, with a truncate base and a small apical germ
pore, 20-27 x 15-17 urn. Dark cell of ascospores Perithecia immersa vel semi-immersa, singula vel bina,
ovoidea, ca 0.75 diam, 1.5 mm alta, collo et ventre pilis
frequently of different sizes even within the same 8­
griseis obtecto. Asci cylindracei, ca 560 x 45-57 um, annulo
spored ascus (Fig. 82 D); pedicel hyaline, conical, 8­
apicali non manifesto, 8-spori. Ascosporae uniseriatae,
10 x 4-6 urn. A single lash-like cauda at either end of uniseptatae, cellula superiore brunnea, ellipsoidea, 63-77 x
the ascospores, the upper one being channeled, the 31-42 um, plus minusve symmetrica, basi truncata, apice
lower one enclosing the pedicel (Fig. 82 E). conico, poro germinali api cali praedita; pedicellus hyalinus,
This species has been seen only once during this 6.5-7 x 5.5 um. Tota spora tunica gelatinosa crassa in aqua
current study. intlata. Fungus fimicola.

Etymology: ignotus = 'unknown', This specimen has been Characteristics on dung: Perithecia developed singly or
puzzling since in a number of respects it resembles Po­ in pairs, dark, ovoid, venters totally or partially sub­
dospora cf didyma (compare Fig. 81 G·-M with Fig. 82 A­ merged in the substrate, about 0.75 diam, 1-1.5 mm high.
E). Emergent necks short, dark, cy1indrical. A tomentum of
grey hyphae sometimes present on the necks and/or the
Holotype: On dung of wallaby (?), Gregory Park, Arkaroola,
South Australia, 30 19' S, 139 20' E, 22 nd Aug. 1999, coll.
0 0 upper venters (Fig. 91 H). Asci cylindrical, about 560 x
Pat Grey, POO 76448. 45-57 um with no visible apical apparatus, 8-spored (Fig.
91 1). Ascospores uniseriate, the dark upper cell ellipsoid,
symmetrica1 or slightly asymmetrica1, with a truncate
Sporormiella albolanata A. Bell & D.P. Mahoney n. base, 63-77 x 31-42 um, pedicel more or less isodiamet­
sp. rie, 6.5-7 x 5.5 urn. Ascospores enveloped by a hyaline
ge1atinous sheath (Fig. 91 J).
Perithecia disserninata, ventre immerso, globoso ca 200 um diam. Materia1 of this species was kind1y loaned to me by
Collo maturitate emerso. Capillis ca 200 x 1-2 um, lanuginoso. Nils Lundqvist who sent me two specimens for observa­
Peridium ventris textura prismatica, partim incrassatum.
tion (Tibell12074-b and Tibell 12084-a).
Paraphyses ca 2-3 urn lata, longitudine indeterminata. Asci
bitunicati, cylindracei, attenuati, ca 120 um longi, 8-spori.
Ascosporae biseriatae, brunneae, ellipsoideae, triseptatae, utrinque Etymology: grandis = 'large', referring to the large ascospores.

61
Holotype: On kangaroo dung (Macropus sp.), 18 km NW of 92 1and K). Ascospores uniseriate, at maturity composed
0
Adarninaby, NSW, uppermost part of Alum Creek, 35 53' S, of an ellipsoidal symmetrical or slightly asymmetrical
E, ca 1260m. ait. 5th Apr. 1981, coll. LeifTibell, UPS
055'
148 dark cell with a truncate base and a smail apical germ
12074-b.
pore, 35-45 x 21-25 um and a small hyaline (occasion­
ally pale brown), oblecythiform pedicel, 7-15 x 3-7 um,
Ascospores surrounded by a hyaline gelatinous sheath
Strattonia oblecythiformis A. Bell & D.P. Mahoney n.
which swells in water (Fig. 92 J & L). Usually the asco­
sp.
spores are arranged with their dark cell uppermost within
Perithecia immersa vel semi-immersa, singula vel bina, ovoidea, ca
the ascus, but occasionally this arrangement may be
0.75 mm diam, collo emerso, breve, verruculoso protrudente; peridium reversed (Fig. 92 K). Ascospores germinating from the
ventris textura prismatica, delicatula Paraphyses hyalinae, rarnosae, dark cell via the germ pore (Fig. 92 H), the resulting
septatae, ca 4-5 um latae. Asci cylindracei, ca 360 x 40 um, annulo mycelium producing an anamorph which forms dense
apicali indistincto, 8-spori. Ascosporae uniseriatae. uniseptatae, cellula floccose c1usters of densely septate hyphae, each cell of
superiore brunnea, ellipsoidea vel ovoidea, plus minusve aequilateralis,
basi truncata, apice conico, poro germinali apicali praedita, 35--45 x
which produces an active phialide (Fig. 92 G). Conidia in
21-25 um; pedicellus hyalinus vel luteobrunneaus, oblecythiformis, 7­ mass pale yellow (3A3-4). Sizes of individual conidia
15 x 3-7 um. Tota spora tunica gelatinosa crassa, aqua inflata within the range 2.5-3 x 2-2.5 um,
circumdata. Fungus fimicola This is a slow-growing species; the perithecia took 81
days to develop after initial incubation.
Characteristics on dung: Perithecia developed singly or
in pairs, about 0.75 mm diam, dark, submerged in the Etymology: ob = 'reversed , 'lecythiform' = 'bottie-shaped',
substrate, with short, blunt-domed, slightly verrucose referring to the pedicel which is shaped somewhat like an
necks protruding (Fig. 92 F). Venter wall of textura invertedstoppered bottle.
prismatica, delicate with substrate adhering to the sur­
face. Paraphyses hyaline, branched, septate, about 4-5 Holotype: On macropod dung collected at White Mountain
National Park, Queensland, coll. Kevin Teys, Apr. 2001. PDD
um wide. Asci cylindrical, about 360 x 40 um, with
75154. A culture is deposited at the Centraalbureau voor
scarcely visible apical ring about 5 urn diam, 8-spored,
Schimmelcutures (CSS 110350).
appearing costate after discharge of the ascospores (Fig.

62
Appendix2 staining paraphyses and hyaline ascospore omamenta­
tion.
Formulae of stains used for microscopie examination
Melzer's reagent (formula taken from Dennis, 1968)
Congo red Potassium iodide 1.5 g, iodine 0.5 g, distilled water 20
Congo red 0.2 g, a1cohol (92%) 6.6 ml, distilled water ml, chloral hydrate (poisonous) 20 ml.
100 ml. This stain is used principally to test for amyloid reac­
This mounting medium renders the walls of fungi tions in fungal structures. A positive reaction is manifest
clearer. It should be used as a tempororary mounting by a blue colouration. The entire asci of sorne species of
medium only. 1 have found it useful for observing the ascomycetes may become blue in Melzer's (e.g. Iodo­
caudae ofcertain species of Podospora. phanus), while in other species (e.g. Hypocopra, Peziza,
Poronia) only the tip of the ascus is affected. Cytoplas­
Indian ink mie structures stain reddish brown.
Diluted Indian ink (see Lundqvist 1972) is used for ren­
dering visible the caudae of sorne Podospora spp. (e.g. P. Shear's mounting medium
dactylina & P. bicornis). Patience has to be exercised 2% Potassium acetate 300 ml, glycerine 120 ml, alcohol
with this technique and the procedure often has to be (95%) 180 ml.
repeated several times before getting a satisfactory prepa­ Mix and filter the ingredients. 1 find this superior to
ration (see Bell 1983). Specimens are first mounted in clear Lactophenol for making semi permanent slides
water after which a drop of very dilute ink is placed to (provided the edges of the coverslips are sealed).
one side of the coverslip. The ink is drawn under the
coverslip by means of a piece of tissue or blotting paper Use of nail varnish as a slide sealant
applied to the opposite edge. By this means the ink is ln recent years there has been an increase in the number
drawn in and around the specimen and will condense of commercial cosmetic preparations available for fin­
around the (previously invisible) caudae. Such ink gemail decoration. One of the disadvantages of nail
mounts are generally temporary, although on occasions polish as a slide sealant has been that it is difficult to affix
the caudae remained visible even when the slide is irri­ to a slightly greasy surface and it chips with time and the
gated with Shears mounting fluid and sealed with nail siides dry out. 1 have found that a primary application of
polish. a layer of commercial nail base coat (specifically de­
signed to adhere to slightly greasy surfaces such as finger
Lactophenol cotton blue nails), over which one or two thin coats of standard nail
Phenol 20 g, lactic acid 16 ml, glycerol31 ml, cotton blue polish are painted is a great improvement. Slides made
0.05 g, distilled water 20 ml. this way have remained (thus far) sealed with their
This is a useful stain for protoplasmic structures, but it mounting fluid intact.
leaves the fungal cell walls unstained. It is useful for

63
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69
OL
 A ::, /

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20~l m

')

1mm

Fig. 17. Asco bolus amoe nus. A- C. A. Two apothecia. B. Mature asco spore . C. Asc i, ascospores and paraph yses. Asco bolus
immersu s. D-F. D. Immature & mature apothecia (one subm erged). E. Mature asco spores showin g variations in shape. F.
Mature ascospore .
71
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.

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Fig. 18. Ascobolus quezelii. A-O. A. Mature apo thecium. B. Tip of ascu s showing operculum. C. Mature ascus with
ascospores and par aphyses . D. Matur e ascospore. Ascobolus bistisii. E- H. E. Mature apothecium . F. Mature asco spores. G.
Outline showing genera l shape of matur e asci. 1. Base of ascu s and its attachment to lining cell. H, Mature ascospore.

72
 A

1mm

D
,

1mm

Fig. 19. Ascobolus dadei. A-C. A. Mature apothecia one submerged. B. Two mature ascospores illustrating range of size &
epi spore omamentation. C. Apothecium, asci & paraphyses. Ascobolus stictoideus. D-F. D. Immature & mature apothecia
(one submerged). E. Two mature ascospores. F. Top of mature ascus showing operculum, ascospores & paraphyses.

73
 A

1mm
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B

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Fig. 20. Ascobolus hawaiiensis. A-C. A. Two matu re apo thec ia. B. Mature ascospores showi ng epispore ornamentation
(bro wn asco spore show ing the co lour change as the y age) . C. Mature as cus containing ascospores & acc ompany ing
paraphyses. Ascobolus brassi cae. D-F. D. Mature asci show ing uniseriate arrangement of asco pores together with
paraphyses. E. Detail of mat ure asc us & ascospores. F. Mature ascos pore .

74
 A

III
(1-­ Il
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1

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Fig. 21. Ascob o/us crenulatus. A-O . A. Immature & mature apothecia. B. Stipitate apothecium which is sometimes seen. C.
Mature ascospores show ing a range of sizes and epispore patterns. D. Mature asci showing uniseriate & biser iate arrangement
of ascospores and branched paraphyses . Ascobo/us michaudii. E-H. E. Immature & mature apothecia one iIJ ustrating the
stipitate cond ition. F. Mature ascospores. G. Mature ascu s & tips of paraphyses. H. Silhouette of mature ascus .

75
 A

,,
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Fig. 22. Ascobo/us a/bidus . A-E. A. Three mature apothecia & detail of peridium . B. Mature ascopores. C. Older ascospore
sectional view to show thick wall. D. Mature ascus and branched paraphyses. E. Two mature asci containing ascospores .
Ascobo/us albidus "British Forrn" . F-1. F. Two mature apothecia. G. Portion of peridium to illustrate patchy dark
pigmentation . H. Three mature ascospores to show range of epispore pigmentation . 1. Mature asci & paraphyses.

76
 A - apothecia may reach 5mm

Scatter diagram illustrating the range of ascospore sizes of 5 species

of Ascobolus.
Sketches represent apothecia & ascospores (=Iatter drawn to scale)

20

d~ ,d",

1 1 1 1 l ,---'------'--'-----'--,---.,' I-,- r..--r-' -.-----.-.--r-.­

, 1 j 1

20 25 30 35 40
Ascospore lenglh ÜJm) ~

Fig. 23. Ascobolus fur furaceus . A-B. A. Diagrammatic representation of matur e apothec ia (materia l too old to illustrate
actual specimen). B. Two ascospores, one mature the other old showing peelin g of epispore. Below : scatter diagram to
illustrate range of asc os pore sizes & othe r morphological feature s of 5 species of Ascobolus.

77
 in
A ~\K)\ I( Ji

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Fig. 24. Thecotheus crustaceus. A-E. A, Almost mature apothecium. B, Mature apothecium showing protruding asci. C.
Tops of two asci showing opercula. D, Mature asci showing uniseriate arrangement of ascospores, E. Mature ascospores with
nuclei one drawing in sectional view to show thick wall. Thecotheus pelletieri. F-L F. Immature & mature apothecia. G.
Silhouette of mature apothecium. H. Mature ascospore showing nucleus, thick wall & hyaline sheath. I. Top of mature ascus
showing line ofweakness delineating the operculum & sorne of the ascospores.

78
 c )
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Fig. 25. Saccob olus verrucisp orus. A-E. A. Group of mature and immature apothecia. B. Scatter diagram illustrating the
range of ascospore sizes. C. Two mature asci containing ascospores & accompanying paraphyses. D. Ascus top showing
operculum. E. Discharged older ascospores showing disintegration of episporium. Sacco bolus versicolor. F-K. D. Mature
apothecia from several views. G. & J. Cluster of ripe ascospores. H. Opercula of asci. l. Mature ascus together with
ascospores and paraphyses. K. Mature ascus.
79
 ---7
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Fig. 26. Saccobolus geminatus A-C. A. Matur e apotheciurn . B, Pair of ascospores. C. Asci, ascospores & paraph yses.
Saccobolus portoricensis D-F. D , Mature apothecim . E, C luster of 8 mature ascospores. F. Mature & immature asci,
ascospores and paraphyses.

80
 A

1
1
(
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Fig. 27. Saccobolus minimus. A-C. A. Group of apothecia . B. Cluster of 8 ripe asco spores showing pigment change wirh
ageing. C. Mature asc i & ascospores with paraphyses, "Queens/and variant " D. (inset): asci , asco spore s & paraphyses (for
further discus sion see text). Saccobolus citrinus. E-G . E. Two mature apothecia . F. C/uster of mature ascospores. G. Asci,
ascospores & paraphyses.

81
A
~

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...,

1O/-!m
1mm
B

lûum
Fig. 28. Saccobolus in/es/ans. A-C. A. Gro up of apo thecia. B. Matur e asc us, asco spores & paraphyses. C. C luster of mature
ascospores.

82
 A

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Fig. 29. Lasiobolus ciliatus. A-G. A. & B. Mature apothec ia showing a range of colour & sizes. C. Silh ouett e of apothecial
bristles. D. Bases & tips of apothecial bristles. E. Ripe asci showin g uniseriate & biseriate ar rangement of ascos pores. F.
Tops ofbranched paraphy ses. G. Mature ascospores show ing luteu s co louration, nu cJei and tran sparent jacket.

83
 ,.

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Fig. 30. Lasiobo/us microsporus. A-E. A. Two mature apothecia. B. Details of bases & tips of apothecial bristle s. C.
Paraph yses. D. Mature ascus showing uniseriate arra ngeme nt of ascospores and an operculum. E. Mature ascospores one
show ing nucleu s. Lasiobo/us diversisporus. F-I. F. Mature apoth ecium . G. Base of apothecial bristle. H. Ascus showing
uniser iate arran gement of ascospores. 1. Mature ascospores two showing nucl ei.
84
 /
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Fig. 31. Mycoarctium sphaerosporum. A- F. A. Mature apothecium. B. Scatter diagram to illustrate size range ofascospores.
C. Detail of apothecial bristles. D. Mature ascus showing uniseriate arrangement of ascospores. E. Free-ended paraphyses. F.
Mature ascospore. Lasiobolus trichoboloides. G-J. G. Two mature apothecia. H. Detail of apothec ial bristles. I. Branched &
unbranched paraphyses. J. Mature ascus, paraphyses, ascospores & operculum.

85
 l
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Fig . 32 . Orbilia inflatula. A- O. A. Apothec ia. B. Paraph yses show ing incrusted apices . C. Mature ascu s. O. Ascospore s.
Orbilia alnea. E-J . E. Apothec ia. F. Co nidia l state on agar. G. Paraphyses showing pigrnent ed inflated apices. H. Matu re
ascu s. 1. T ip of discha rged ascus. J. Matu re ascospores.

86
 1mm

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r
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Fig. 33. lodophanus carneus. A-O. A. Group of mature apothecia. B. Mature ascospores showing nuclei. C. Immature &
mature asci & paraphyses. O. Oischarged asci showing opercula. Jodophanus verrucosporus. E-H. E. Group of mature
apothecia. f. Mature ascospore showing nucleus. G. & H. Asci containing mature ascospores, paraphyses and discharged asci
showing opercula.
87
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Fig. 34. Peziza vesiculosa. A-E. A. Young apothec ium. B. Opercula showing distribution of blue reaction in Melzer' s
reagent. C. Paraphyses. D. Mature ascus showing uniseriate ascospores. E. Mature ascospores one showing nucleus. Peziza
p leurota. F-K. F. Mature apotheciurn. G. Ageing apothecium. H. Operculum showing distribution of blue reaction in
Melzer' s reagent. 1. Paraphyses. J. Ascus containing ascospores. K. Mature ascospores one showing the nucleus.
88
 r,

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Fig. 35. Cheily menia theleboloides. A-F. A. Details of apothecia. 8 . Colour chip showing typical colour of mature apothecia.
C. Details of apothecial bristles. D. Mature ascus & uniseriate ascospores. E. Paraphyses. F. Mature ascospores. Cheily menia
rarip ila. G-L. G. Mature apothecium . /-1 . Colour chip showing typical coour of mature apothecia. T. Paraphyses. 1. Apothecial
bristles. K. Mature ascus. L. Mature ascospores. Ali illustrations taken fro m Dade 's notes.
89
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Fig. 36. Cheilymenia coprinaria. A-F. A. Mature apothec ium. B. Co lour chip showing ty pica l co Jour of mature apothec ia. C.
Apothec ial bristles. D. Tip s of paraph yses. E. Mature ascos pores. F. Asc us with ascos pores . Cheilymenia stercorea . G-L. G.
Mature apothecia . H. Detail of apothec ia l margin. 1. Co lour chip show ing typi cal co Jour of matu re apothec ia. J. Apoth ecial
bristles. K. T ips of paraph yses. L. Mature asc i & ascospores. Ali illustrations takenfrom Dade 's notes.

90
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Fig. 37. Ch eilvinenia granulata. A-E A , Mature apot hecium B, Colom chip showi ng typical co lour 0"m ,H U': upothcc ia. l'
Mature ascus & ascospor es. D, Mature asco spores. L. T ips of paraphy ses, Ai l illustrationstakcn [icnn !JIlIie ', 11 11 /<\ ,
(' ()j7I'1J/ il S lu/eus, F- I I. F, Mature aporheciurn . G. Mature ascus & ernpty ascus show ing operculu m 8:. p a ra p h:- s l' ~ I l , vl aturc
élSCOS p O IT S with de Bary bubb les.
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li 1
10 1 131 1 15 1 1 1
: ~sco~pore I.~ngth (IJm) ~

Fig. 38. Copra/us cf disculus. A-C. A. & B. Groups of asci , ascospores & paraphyses illustra ting range of variation. C.
Mat ure apothec ia. Below: Sca tter diagram of asco spore sizes of 18 co J1ections of Copr a/us cf disculus. Boxed areas denote
size range for published spec ies as ind icated . Forfurther explanation see text.

92
 A

1mm

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1mm

Fig. 39. COpro/IIS baeosporus. A-C. A. Mature apothecium. B. Ascospores showing de Bary bubbles. C. Group of mature
asci, ascospores & paraphyses. Cop ra /lis ochrace us. D-H . D. Mature apothecia. E. Mature ascospores one showing de Bary
bubble & lateral gelatinous substance the other drawn in sectional view. F. Mature ascus & paraphyses. G. Opercula of asci.
H. Detail of ascus base.

93
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F

1mm

10flm
H

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10f lm

Fig. 40. Copra/us sexdecimsp orus. A- E. A. Matu re apothecia (inset drawings fro m Dade). B. Sca tter dia gram showing
distrib ution of ascospores sizes. C. Matu re ascu s. ascos pores & paraph yses. D. Ope rculum of d ischarged ascus. E. Two
Matu re ascos pores one w ith de Bary bubble. Copr a/us winteri. F- f. F. Mature apot hecia . G. Matur e asc us & parap hyses. H.
Opercu lum of disc harged asc us. I. Ma ture ascos pores one with de Bary bubbl e.

94
 c~
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a

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Fig. 41. Ascozonus woolhopensis. A-D. A. Profile views of apothecia. B. Details of mature asci. C. Tip of discharged ascus
& ascospores. D. Tip of undischarged mature ascus. Pulvinula cinnabarina. E-H. E. & G. Stages in the development of
apothecia. F. Tip ofasci. H. Mature asci, ascospores & paraphyses. Ali illustrations takenfrom Dade's notes.

95
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Fig. 42. Selinia pulchra. A-D. A. Perithecium embedded in str oma on dung surface . B. Mature ascus & ascospores. D.
Immature ascospore (lower) and two mature ascospore s showing eac h containing 2 nu clei and thin gelatinou s coating around
ascospores. Pseudombroph ila theioleuca. E-G . E. Development of apothecia. F. Sket ches of various stages in apothecial
development. G. Mature asci & ascospores. li/us/rati ons E - G takenfrom Dade 's no/es.

96
 B

G
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200pm
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Fig. 43. Thelebolus polysporus . A-E. A . Cluster of 3 mature apothecia on dung surface. B. Immature apothecium . C. Mature
apothecium. D. Mature ascospores. E. Mature asci and paraphyses . Trichobolus zukalii. F- J. F. Mature apothecium on dun g
surface. G. Slightly imm ature apothecium showing large ascu s tip emergent from apothecium . H. Apothecial bristle s. 1.
Squash of mature apothec ium showing many mature and immature asc i. J. Mature asco spores two showing de Baty bubbles.
Illustrations H & 1 taken fr om Dade 's notes.
97
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Fig. 44. Thelebolus microsporus . A-F, A. Apothecia in various sta ge s of development (Dod e 's drawin gs) . B. Immature &
mature apothecia on dung surface. C. Detail of apothecia margin. D. Mature asci, ascospores & paraphyses ( V409a &
uncoloured paraphyses fr om Dade 's notes) . E. Mature ascospores. F. Mature asco spores (from Dade 's notes). Thelebolus
stercoreus. G-L. G., H. & l. stages in the development of apothecia. 1. One huge ascus present in the apothecium. K. Det ails
of the operculum at ascu s tip . L. Mature asco spores.

98
 A
o
v
c
e ~
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Fig. 45. Pseudoarachniotus roseus. A-C. A. Gymnothecia. B. Asci, ascospores & portion s of gy mnotheci um. C. Matu re
ascospores. Gymnoascus reesii. D-F. D. Gymnothecium. E. Secti onal view of gymnothecium hyphae . F. Mature asci &
ascos pores viewed throu gh gy mnothec ium hyph ae , the latter with hooked appendages.

99
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Fig. 46. Lasiobolidium orbiculoides. A-E. A. Cleistothecium. B. Peridium cells of cIeistothecium. C. Cleistothecial hairs, one
in sectional view to show septa and thick walls. D. Portion of mature ascus showing mature ascospores. E. Mature ascospores
one with de Bary bubble. Kernia nitida. F-l. F. Cleistothecia with appendages. G. Detail of cIeistothecial wall. H. Details of
two types of appendages on cIeisthothecia. l. Mature ascus & ascospores.
100
 E

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Fig. 47. Pleuroa scus nicholsonii. A-E. A. Cleistothecia. B. Peridium cells of cleistotheciu m. C. View of cleistothecium at
greater magnific ation than in A. D. Coiled cleistoth ecial appendage . E. Mature ascospores & asci. Nigrosabulum globosum.
F-J . F. Cleistothecia as they appear on dung surface. G. Peridium cells of cleistothecium . H. Acremonium anamorph From
culture . 1. Mature ascospores. J. Asci in various stages of maturity .

101
 A

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Fig. 48. Preussia funiculata. A-O. A. Cleistothecium. B. Peridial cells of cleistothecium. C. Mature ascus & ascospores. D.
Mature ascospores. Preussia cylindrica. E-H. E. Cleistothecium. F. Peridium cells of cleistothecium. G. Asci & ascospores,
the latter splitting into individual cells within the ascus. H. Released portions of ascospores.

102
 B

Fig. 49. Zopjiella erostrata. A-D. A. Cleistothecia. B. Portion of peridium with one attached appendage. C. Mature
ascospores. D. Ascus containing mature ascospores.

103
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Fig. 50. Pyxidiophora arvensis. A. Details of perithecium, asci & ascospores. Pyxidiophora. spp. B-E. B. Perithecia &
ascospores. C. Ascospores D, & E, Details of perithecia & ascospores of V776 b & 790f. Ali illustrations taken from Dade's
notes.

104
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Fig. 51. Ophiostoma n. sp. A-D. A. Perithecia. B. Detail of perithecial neck apex showing discharge of asci. C. Ascospores.
D. Ascus containing ascospores. Phomatospora minutissima. E-H. E. Young & old perithecia. F. Detail of peridial wall. G.
Ascus tops showing apical ring & ascospores. H. Mature ascospores. Sphaeronaemella fimicola. I-K. 1. Perithecia. J.
Perithecia at higher magnification. K. Fimbriate tip to perithecium & discharged ascospores. Illustrations 1 -K taken from
Dade 's notes.

105
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Fig. 52. Petriella sordida. A-F. A. Perithecia seated on a web of hyphae. B. Graphium anamorph from culture. C. Peridium
celis ofperithecium. D. Young ascus. E. Mature ascospores. F. Germinating ascospore. Melanospora brevirostris. G-L. G.
Perithecia seated on a web of hyphae. H. Two asci. 4 & 8 spored. 1. Details of rostrum. J., K., L. Samples of ascospores from
3 different collections.

106
 A

1mm

10IJm

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F

Fig. 53. Chaetomium trigonosporum. A-D. A. Perithecium. B. Mature ascospores. C. Details ofperithecial hairs. D. Petaloid
peridium of perithecium. Chaetomium irregulare. E-H. E. Subterranean, partially immersed & superficial cleistothecia. F.
Angular peridial wall cells. G. Ascus & mature free ascospores. H. Cleistothecial hairs.
Arrows indicate germ. pores.

107
 0.

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Fig.54. Chaetomium indicum. A-E. A. Perithecium. B. Mature ascospores. C. Asci in various stages of development. D.
Terminal hairs ofperithecium. E. Peridial wall cells. Chaetomium cuniculorum. F-K. F. Perithecium. G. Details of entangled
perithecial hairs. H. Mature ascospores. I. Angular peridial cells. 1. Details of upright terminal hairs. K. Ascus containing
immature ascospores. Arrows indicate germ. pores.
108
 F
A

1mm

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1mm

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Fig. 55. Chaetomium subspirale. A-f. A. Perithecium. B. Immature ascus. C. Mature ascospores. D. Angular peridium cells.
E. Anamorph in culture. F. Details of terminal hairs. Chaetomium ampullare. G-K. G. Perithecium. H. Mature ascospores. 1.
& 1. Terminal hairs. K. Immature ascus. Arrows indicate germ. pores.

109
 o

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10IJm 10IJm

Fig. 56. Chaetomium cupreum. A-E, A. Perithecium. 8. Mature ascospores. C. Angular peridial wall cells. D. Terminal hairs.
E. Asci. Chaetomium brasiliense. F-J. F. Perithecium. G. Mature ascospores. H. Angular peridium cells. 1. Asci in various
stages ofmaturity. 1. Terminal hairs. Arrows indicate germ, pores.

110
 E

1mm M
1
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D
10l..lm

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F
Fig. 57. Chaetomium glob osum. A-F. A. Perithecium. B. Mature ascospores. C. Anamorph from culture. D. Asci in various
stages of maturity. E. Terminal hairs. F. lntricata/epiderrnoidea peridium cells. Chaetomium subaffine. G-K. G. Perithecium.
H. Mature ascospores. 1. Ascus. 1. lntricatalepidermoidea peridium & base of a terminal hair. K. Terminal hairs. Arrows
indicate germ . pores.

111
 1mm

,~
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..

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10~m

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E

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1mm

10~m
10~m

H
Fig. 58. Chaetomium piluliferum. A-E. A. Perithecium. B. Mature ascospores. C. Ascus. D. Smooth circinate terminal hairs
& warty lateral hair. E. IntricataJepidermoidea peridium cells. Chaetomium murorum. F-J. F. Perithecium. G. Mature
ascospores. H. Smooth terminal hair. 1. Asci. J. Spiny terminal hairs. Arrows indicate germ. pores.

112
 D

10J.lm

1mm

10J.lm

10J.lm

E 1mm

Fig. 59. Chaetomium convolutum. A-D. A. Perithecium. B. Mature ascospores. C. Unevenly spiral, spiny, terminal hairs. D.
Petaloid peridium cells. Chaetomium succineum. E-1. E. Perithecium. F. Angular peridial cells. G. Mature ascospores. H.
Evenly spiral terminal hairs. 1. Ascus containing immature ascospores. Arrows indicate germ. pores.

113
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Fig. 60. Chaetomium robustum. A-D. A. Perithecium. B. Mature ascospores. C. Angular & elongate peridial wall cells. D.
Irregularly branched teminal hairs. Chaetomium bostrychodes. E-H. E. Perithecia. F. Mature ascospores. G. Asci in various
stages of development. H. Terminal hairs & angular peridium cell. Arrows indicate germ. pores.

114
 1mm

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Fig. 61. Sordaria jimicola. A-O. A. Perithecia. B. Ascus tips showing apical rings. C. Ascus containing mature ascospores
and interascal filaments. D. Mature ascospores. Sordaria macrospora. E-H. E. Mature perithecia. f. Old perithecia with
elongated necks. G. Mature ascus. H. Mature ascospores.

115
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Fig. 62. Sordaria alcina. A-C. A. Mature perithecia. B. Mature ascus & interascal filaments. C. Mature ascospores. Sordaria
H 'il' "-' ",/

10J.lm
­

arctica. D-G. D. Perithecia. E. Silhouette of mature ascus. F. Mature ascus with ascospores. G. Mature ascospores. Sordaria
humana. H. (inset) Mature ascospores.

116
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Fig. 63. Sordaria brevicollis. A-D. A. Perithecia. B. Mature asci. C. Mature ascospores. D. Scatter diagram showing size
distribution of mature ascospores. Sordaria n. sp. F-I. F. Perithecia. G. Mature ascus H. Mature ascospores. 1. Scatter
diagram showing size distribution of mature ascospores. E. Mature ascospores of both species side by side for comparisons.
For further information see text.

117
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Fig. 64. Coniochaeta leucoplaca. A-E. A. Perithecium. B. Peridial structure of upper perithecial wall close to neck. C.
Perithecial bristles. D. Mature ascus. E. Mature ascospores. Coniochaeta discospora. F-I. F. Mature perithecia. G. üld
perithecia showing elongated necks. H. Mature ascus. 1. Mature ascospores.

118
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Fig. 65. Coniochaeta scatigena. A-E. A. Perithecia. B. Epidennoid peridium & bristles. C. Mature ascus. O. Tip of mature
ascus. E. Mature ascospores. Coniochaeta hansenii. F-G. F. Mature asci & ascospores. G. Perithecium. Illustrations F & G
from Dade 's notes.

119
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Fig. 66. Podospora ellisiana. A-F. A. Perithecium. B. Perithecial hairs. C. Silhouette of mature ascus. D. Mature asci. E.
Ascus tip showing minute ring. F. Immature & mature ascospores. Podospora curvicolla. G-K. G. Perithecia. H. Perithecisl
bristles. 1. Peridium. J. Mature & immature ascospores. K. Mature ascus.

120
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Fig. 67. Podospora setosa. A-G. A. & B. Perithecia. C. Peridium of perithecial neck. D. Immature ascospores. E. Mature
ascospores. F. Ascus tips, one showing apical ring. G. Mature ascus & interascal filaments. Podospora bifida. H-K. H.
Perithecia. 1. Mature ascospores. J. Mature ascus. K. Silhouette of mature ascus.

121
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Fig. 68. Podospora ampullaceae. A-F. A. Perithecia. B. Perithecial hairs. C. Silhouette of mature ascus. D. Three views of
ascus tip with ring. E. Mature ascus. F. Immature & mature ascospores. Podospora pectinata. G-K. G. Perithecia. H. Hyphae
covering venter. 1. Mature ascus. J. Detail of ascus tip with small ring. K. Mature ascospores.

122
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Fig. 69. Podospora intestinacea. A-E. A. Perithecium. B. Silhouette of mature ascus. C. Mature ascus containing ascospores
of varying sizes. D. Immature ascospore showing nucleus. E. Mature ascospores. Podospora prethopodalis. F-K. F.
Perithecia. G. Conidia in culture. H. Perithecial bristles. 1. Mature ascus swel1ing in water. J. Mature ascospores. K. Immature
ascospores.
123
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Fig.70. Podospora anserina. A-E. A. & B. Perithecia. C. Silhouette of mature ascus. D. Mature ascus & interascal filaments.
E. Mature ascospores. Podospora excentrica. F-J. F. Perithecia. G. Perithecial bristles. H. Mature ascus. 1. Immature
ascospore. J. Mature ascospores.

124
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Fig. 71. Podospora dactylina. A-G. A. Perithecia. B. & C. Hairs & tubercles on perithecial neck. D. Mature ascospore &
ascus. E. Immature ascospore. F. Ascus tip. G. Mature ascospores viewed from various angles. Podospora
austrohemisphaerica. H-K. H. Perithecia. J. Silhouette of mature ascus. J. Mature ascus. K. Mature ascospores.
125
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Fig. 72. Podospora pleiospora. A-E. A. Perithecium. B. Tubercles on perithecial neck. C. Mature ascus. D. Empty (costat).
E. Ascus. E. Mature ascospores. Podospora communis. f-I. F. Perithecia. G. Perithecial hairs. H. Ascus tip & mature ascus.
1. Mature ascospores.

126
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Fig. 73. Podospora argentiniensis. A-D. A. Perithecium. B. Neck tubercles. C. Immature ascus. D. Mature ascospores.
Podospora decipiens. E-H. E. Perithecium. F. Neck tubercles. G. Mature perithecium with apical ring. H. Mature ascospores.

127
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Fig. 74. Podospora fimiseda. A-O. A. Perithecia. B. Silhouette of mature ascus. C. Ascus with immature ascospores & apical
ring. D. Mature ascospores. Podospora dolichopodalis. E-H. E. Perithecium. F. Neck tubercles. G. Mature ascus swelling in
water. H. Mature ascospores.

128
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Fig. 75. Podospora perplexans. A-E. A. Perithecia. B. Hyaline tipped perithecial hairs. C. Mature ascus. D. Tip of immature
ascus showing tiny apical ring. E. Mature ascospores. Podospora tetraspora. F-J. F. Perithecia. G. Inflated & agglutinated
perithecial hairs. H. Phialophora anamorph in culture. 1. Mature ascus & interascal filaments. J. Mature ascospores.

129
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Fig. 76. Podospora glutinans. A-E. A. Perithecia. B. Inflated perithecial hairs. C. Germinating ascospore. D. Mature ascus &
interascal filaments. E. Mature ascospores. Podospora vesticola. F-J. F. Perithecia. G. Inflated perithecial hairs. H. Mature
asci(one swollen in water) & interascal filaments. 1. Phialophora anamorph. J. Mature ascospores.

130
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Fig. 77. Podospora miniglutinans. A-E. A. Perithecia. B. Intlated perithecial hairs. C. Silhouette of mature ascus. D. Mature
ascus and ascospores. E. Mature ascospores.

131
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Fig. 78. Podospora curvuloides. A-F. A. & B. Perithecia (often reclining). C. Inflated neck hairs. D. Silhouette of mature
ascus. E. Mature asci. F. Ascospores, two showing fine horizontal striations (visible in slightly immature ascospores).
Podospora conica. G-K. G. Perithecia one showing positive phototropism when located on side of dung bail. H.
Agglutinated inflated perithecial hairs. I. Silhouette ofmature ascus. J. Mature ascus. K. Mature ascospores.

132
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Fig. 79. Podospora fabiformis. A-E. A. Perithecia. B. Perithecial hairs. C. Silhouette of mature perithecium. D. Mature
perithecium & interascal filaments. E. Mature ascospores. Podospora n. sp. F-J. F. Mature & old perithecia (latter with
longer neck). G. Perithecial hairs. H. Silhouette of mature ascus. 1. Immature & mature asci with interascal filaments. 1.
Mature & immature ascospores.
133
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Fig. 80. Podospora pyriformis. A-O. A. Perithecium. B. Ascospore drawn with full complement of caudae. C. Asci &
ascospores. O. Mature ascospores (Except for B. al! drawing from Dade 's notes). Podospora bicornis. E-H. E. Perithecia.
older ones with elongated necks. F. Perithecial hairs. G. Mature ascus. H. Mature ascospores.

134
 1mm

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Fig. 81. Podospora hyalopilosa ? A-F. A. Perithecia. B. Peridial wall. C. Phialophora anamorph in culture. D. Perithecial
hairs. E. Immature & mature asci. F. Mature ascospores. Podospora cf didyma. G-M. G. Perithecia with luteus hyphae. H.
Details of luteus hyphae. 1. Tubercles around perithecial neck. 1. Germinating ascospore. K. Silhouette of mature ascus. L.
Mature asci with ascospores of varying sizes, empty costate ascus & paraphyses. M. Mature ascospores.

135
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Fig. 82. Podospora ignota. A-E. A. & B. Perithecia with luteus neck hairs. B. Neck hairs. D. Empty costate ascus & mature
ascus with ascospores of varying sizes inside. E. Mature ascospores. Apiosordaria verruculosa. F-I. F. Perithecium. G.
Anamorph in culture. H. Empty costate ascus & mature ascus with ascospores. 1. Various stages in the development of
ascospores. Arrows indicate germ. pores & minute hyaline appendages.

136
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f 1

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il
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lCJ
Fig. 83. Podospora australis. A-F. A. Perithecia & perithecial neck hair. B. Empty asci showing fissurate dehiscence. C. Tip
of intact ascus. D. Mature ascus & free ended paraphyses. E. Silhouette of mature ascus. F. Immature & mature ascospores.
Arnium arizonense. G-K. G. Perithecia with agglutinated bristles & mat of white hyphae. H. Mature ascospores. 1. Mature
ascus. 1. Immature & mature ascus tips with small rings. K. Mature ascospore.

137
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Fig. 84. Arnium caballinum. A-D. A. Perithecia. B. Ascus tips showing rings. C. Mature ascus. D. Various views of mature
ascospores. Arnium macrotheca. E-I. E. Perithecia. F. Agglutinated perithecial bristIes. G. Silhouette of mature ascus. H.
Mature ascus. 1. Mature ascospores.

138
 / , "'

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1mm

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Fig. 85. Arnium olerum. A-f, A. Perithecium. B. Tip of ascus showing clear ring. C. Mature ascus. D. Silhouette of mature
ascus. E. Claddorhinum anamorph from culture. F, Mature ascospores. Arnium cirriferum. G-K. G. Perithecium. H.
Silhouette of mature ascus. 1. Two mature asci, one showing aberrant ascospores. J. Tip of young ascus. K. Mature
ascospores.

139
 ""~-'\

A'
1mm
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c

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E 10~m
10~m

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Fig. 86. Arnium mendax. A-C. A. Perithecium. B. Tubercles on perithecial neck. C. Mature ascospores Old specimen- asci
ï

not seen). Arnium n. sp. (Tibell 11045 B). D-H. D. Perithecium. E. Neck of perithecium showing pigmented furrows &
tubercles at base of neck. F. Mature ascus. G. Various asci. mature & those which have discharged their ascospores. H.
Mature ascospores. For further discussion ofthis species see text.

140
 A

1mm

10IJm
B

Fig 87. Arnium absimile. A-G. A. Perithecium. B. Perithecial hairs. C. Silhouette of mature ascus. D. Empty costate ascus. E.
Portions of young & mature asci. F. Immature ascospores. G. Mature ascospores. Cercophora californica. H-L. H. Side &
polar view of perithecium. 1. Cephalothecoid peridium. J. Hairs from venter of perithecium. K. Portions of two asci, one
containing mature ascospores with subapical globulus, the other an empty ascus. L. Ascospores in various stages of
development.
141
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Fig. 88. Cercophora mirabilis. A-F. A. Perithecia. B. Tuberc1es on neck. C. Silhouette of mature ascus. D. Mature ascus with
subapical globulus. E. Ascospores in various stages of develoment. F. Ascus tip showing ring & subapical globulus.
Cercophora grandiuscula. G-N. G. Various views of perithecia covered with distinct white hyphae. H. Detail of neck
bristles. I. Dried perithecium showing sulcate neck with short bristles. J. Asci, ascospores & free ended paraphyses showing
apical rings & costate nature of discharged asci. K. Mature ascospore one which is completely pigmented. L. Claddorhinum
anamorph from culture. M. Germinating ascospore. N. Immature ascospore.

142
 /1~
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\ i

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\
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K ~
J
Fig. 89. Cercophora sordarioides. A-F. A. Perithecia. B. Peridium wall. C. Germinating ascospores & portion of ascus tip.
D. Phialophora anamorph from culture. E. Various stages in the maturity of asci & ascospores. F. Ascospores showing
various states of pigmentation & development. Cercophora recta. G-M. G. Perithecia. H. Verrucose tubercles from neck. I.
Peridium cells. J. Silhouette of mature ascus. K. Ascus showing centrally positioned immature ascospores. L. Tip of ascus
showing tiny ring. M. Mature ascospores.

143
 A 1mm

~\
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10l..lm
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Fig. 90. Podospora gigantea. A-F. A, Perithecia. B. Discharged fissurate ascus. C. Silhouette of mature ascus, D. Two
immature & one mature ascospores. E. Tmmature & mature ascospores. F. Enlarged mature ascospore without pigment to
show apical germ pore & reduced basal "pedicel", Podospora petrogale. G-K. G. Perithecia. H. Silhouette of mature ascus. L
Portion of mature ascus to show ascospore uniseriate arrangement. J. Mature ascospores. K. Phialophora anamorph from
culture.
144
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Fig. 91. Strattonia zopfii. A-G. A. Perithecium. B. Discharged asci showing apical portions which split around the edges of
the apical ring. C. Mature ascospores. D. Ascus tip ofundischarged ascus. E. Immature ascospore. F. Mature ascus showing
uniseriate arrangement of ascospores. G. Silhouette of mature ascus. Strattonia grandis. H-J. H. Perithecia with hairs
developed on necks and/or exposed upper venters. 1. Portion of mature ascus showing uniseriate arrangement of ascospores.
J. Mature ascospores.
145
 """,­

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Fig. 92. Strattonia insignis. A-E. A. Perithecia. B. Silhouette of mature ascus. C. Germinating ascospore. D. Top of mature
ascus showing uniseriate arrangement of ascospores & paraphyses. E. Immature & mature ascospores. Strattonia
oblecythiformis. F-L. F. Perithecia. G. Anamorph in culture. H. Germinating ascospores. I. Empty costate ascus &
paraphyses. J. Mature ascospores. K. Tip of mature ascus showing apical ring. L. Two immature ascospores.

146
 tr>
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Fig. 93. Zygopleurage zygospora. A-E. A. Perithecia. B. Immature ascospore, C. Mature ascus. D. Ascus with immature
ascospores. E. Mature ascospore. Zygospermella insignis. F-L. F. Perithecia. G. Silhouette of mature ascus. H. Mature ascus.
1. Germinating ascospore J. Tip of ascus showing small ring. K. Interascal filaments. L. Immature & mature ascospores.

147
 + 200~m

ft
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Fig. 94. Fimetariella minuta. A-C. A. Perithecium. B. Mature asci showing swelling in water. C. Mature ascospores.
Bombardiodea stercoris. D-G. D. Details ofperithecial developrnent. E. Bases of mature ascospores showing germ pores. F.
Mature ascospores. G. Mature asci. Illustrations D-G from Dade 's notes.

148
 A

2mm

10~m

10~m

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l '
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1

2mm

H
G

Fig. 95. Hypocopra equorum ? A-E. A. Perithecia & surface stroma. B. Detail of interwoven hyphae of stroma. C. Free­
ended paraphyses & portion of mature ascus with large apical ring. D. Ascospores in various stages of development sorne
with small hyaline basal cell. E. (inset) collection A 321 identical in other respects but ascospores slightly longer & more
apiculate. Hypocopra sp. F-H. F. Perithecia in dung substrate. G. Tip ofmature ascus. H. Mature ascospores.

149
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Fig. 96. Podosordaria sp. A-O. A. Stroma. B. Lindquistia anamorph on stroma tip. C. Details of mature ascus & paraphyses.
D. Mature ascospores. Podosordaria violacea. E-1. E. Perithecia in dung substrate showing pale crenulate flange around
ostiole. F. Mature asacus. G. Tip of ascus. H. Various views of mature ascospores. 1. Base of ascus.

150
 A

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1mm

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Fig. 97. Poronia erici. A-O. A. Peritheeia embedded in prominent stroma. B. Seetional view through stroma & detail of
peritheeium. C. Mature aseus. O. Mature aseospores. Poronia oedipus? E-G. E. Stroma peritheeia embedded in apical dises.
F. Mature aseus showing uniseriately arranged aseospores. G. Mature aseospores.
151
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Fig. 98. Trichodelitschia munkii. A-D. A. Perithecia. B. Enlarged view of perithecia. C. Details of bitunicate ascus with
apical ring, bases of asci & interascal filaments. D. Details of mature ascospores, showing one germinating through ascus
wall. Trichodelitschia bisporula. E-G. E. Perithecia, one recumbent & resembling a cleistothecium. F. Two mature asci, one
discharging ascospores. G. Mature ascospore.
152
 c

A
10IJm

1mm

A
D a~ ••
T. munkii T. bisporula
.,
T. adelphica

Fig. 99. Trichodelitschia adelphica. A-C. A. Perithecia, two recumbent resembling cleistothecia. B. Top of undischarged
mature ascus. C. Mature ascospore. D. Mature ascospores of three species of Trichodelitschia side by side for easy
comparison.

153
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Fig. 100. Semidelitschia nannostellata. A-F. A. Perithecium. B. Details of hairs on neck. C. Mature ascospores with central
nucleus. E. Silhouette ofburst bitunicate ascus showing subapical rupture of outer ascus wall. F. Mature ascus & paraphyses.
Delitschia excentrica. G-1. G. Perithecia one with hairs, the other glabrous. H. Mature ascus showing uniseriate arrangement
of ascospores. r. Unruptured (upper) & ruptured (lower) asci with paraphyses. 1. Mature ascospores.

154
 1mm

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10~m;

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H

+
+
10~m

Fig. 101. Delitschia didyma. A-D. A. Perithecium. B. Mature unruptured & mature ruptured asci. C. Branched paraphyses.
D. Mature ascospores. Delitschia chodocola. E-H. E. Mature perithecium. F. Mature asci. G. Mature & germinating
ascospores. H. Portion ofpycnidial state in culture.

155
 é.llll
A" 1mm

.l 1 li 1 ·1 10IJm
Ê34 o 0
2:
.&:
lj
Cb 00
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l!
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65' '70' 75
Ascospore length (IIm) ~
B

0­ 1mm

20IJm

Fig. 102. Delitschia sp. "AI63". A-e. A. Perithecium. B. Scatter diagram showing range of ascospore sizes. C. Details of
mature ascospores showing pores & wrap around germination slits. (For further discussion of this species see text).
Delitschia winteri. D-F. D. Perithecium. E. Mature ascus. F. Mature ascospores.

156
 A 1mm
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Fig. 103. Delitschia intonsa. A-O. A. Perithecia, B. Details ofunruptured & ruptured ascus apices. C. Mature ruptured ascus
& branched paraphyses. D. Mature ascospores with fine punctations. Delitschia gigaspora. E-H. E. Perithecia, hairy &
hairless. F. Tip of mature unruptured ascus. G. Mature ascus & branched paraphyses. H. Mature ascospores sorne with
slightly punctate walls (=arrow) & pale slightly immature ascospore showing two nuclei in each cell.
157
 c-~,

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1mm
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1mm /
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1

10~m
20~m
Fig. 104. Delitschia vulgaris. A-F. A. Two perithecia, old one with elongated neck. B. Ascus tips one unruptured other
ruptured. C. Portion of pycnidium in culture. D. Mature asci, one unruptured, the other ruptured together with branched
paraphyses. E. Silhouettes of mature asci. F. Mature ascospores two germinating. Delitschiafurfuracea. G-K. G. Submerged
perithecia. H. Tips ofunruptured & ruptured asci. 1. Silhouette ofmature ascus. J. Mature ascus. K. Mature ascospores.

158
 E
A

1mm

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B

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1mm

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Fig. 105. Delitschia variispora. A-F. A. Perithecia. B. Detail of peridium. C. Silhouette of mature ascus. D. Mature ascus
showing subapical rupture of outer ascus wall & base of ascus. E. Tip of unruptured mature ascus. F. Mature ascospores.
Delitschia leptospora. G-I. G. Perithecium. H. Ruptured mature ascus & branched paraphyses. I. Mature ascospores.

159
 -,

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Fig. 106. Sporormiafimetaria. A-G. A. Ascoma in situ, one discharging asci in water. R Sectional view through ascoma, C.
Details ofperidium. D. Ruptured mature ascus. E. Unruptured mature ascus, empty outer ascus wall & branched paraphyses.
F. Bundle of 8 mature ascospores. G. Single mature ascospore. Below Scatter diagram illustrating the ascospore dimensions
of Sporormiella intermedia sensu lato & Sporormiella borealis. (For further explanation see text).

160
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Sporormiella australis. D-H. D. Perithecia, enlarged one showing hyaline neck. E. Peridium cells. F. Mature ascospores. G.
Silhouette of unruptured ascus with button stipe. H. Two mature asci one showing subapical rupture & ascus base.

161
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the tips. C. Peridium cells. D. Mature ascospores. E. Silhouette of unruptured ascus with button stipe. F. Mature unruptured
ascus. Sporormiella borealis. G-J. G. Old perithecia with elongated necks. H. Younger perithecium with short neck. 1. Three
Mature asci containing 8, 4 or 1 ascospores. 1. Mature ascospores.

162
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ascospores. Sporormiella subtilis. E. Three Mature ascospores. F. Scatter diagram illustrating the overlap in ascospore
dimensions of S. cymatomera (x) & S. subtilis ('). Sporormiella tetramera. G-J. G. Perithecia. H. Silhouette of ascus. 1.
Details of mature asci, ruptured & unruptured. J. Mature ascospores.

163
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ascospores. Sporormiella megalospora. E-I. E. Perithecia. F. Peridium. G. Mature unruptured ascus. H. Tip of mature ascus
1iberating part -spores. \. Mature ascospores.

164
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branched paraphyses. D. Tip & base of ascus . E. Mature ascospores. Sporormiella ovina. F-J. F. Perithecium. G. Peridium.
H. Tip of Mature ascus. 1. Ascus bases showing star like attachments to lining ofperithecium. J. Mature ascospores.

165
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paraphyses. D. Mature ascospore. Sporormiella euryospora ? E-H. E. Perithecia, mature & old with elongated bifid neck. F.
Peridium. G. Mature ruptured ascus. H. Tip of mature ascus & mature ascospores.

166
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E. Mature ascospores. Sporormiella bipartis. F-I. F. Perithecia. G. Mature ascospore drawn to same scale as S. octonalis. H.
Mature ascospores. I. Mature ascus.( Illustrations F. H. & l from Dade 's notes).

167
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ascus. D. Portions of mature asci. E. Mature ascospores. Sporonniella herculea. F-H. F. Perithecia older one with elongated
neck. G. Mature ascus. H. Portion of mature ascus showing upper two ascospores.

168
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Ruptured mature ascus & paraphyses. E. Mature ascospores. Sporormiella albolanata. F-l. F. Perithecia, older ones with
elongated necks, to right one enlarged perithecium. G. Detail of perithecial hairs & peridial cells. H. Asci, unruptured &
ruptured with paraphyses. 1. Mature ascospores.

169
INDEX OF FUNGAL NAMES (page numbers in bold refer to illustrations)

Acremonium, 101 - cunicu1orum, 35, 108

Apiosordaria verrucu1osa, 43, 136 - cupreum, 36, 110
Amium, 42, 43, 46, 47, 65-67, 69,137-141 - fimico1a, 33
- absimi1e, 46, 47,141 - funico1a, 34
- arizonense, 42, 46, 47,137 - globosum, 8, 33, 36, 111
- caballinum, 47, 138 - homopi1atum, 34
- cirriferum, 47, 139 - indicum, 34, 35, 108
- macrotheca, 46, 47, 138 - irregu1are, 28, 29, 33, 35, 107
- mendax, 47, 140 - murorum, 6, 34, 36, 65, 112
- n. sp., 47, 140 - pi1u1iferum, 34, 36, 112
- olerum, 47, 139 - robustum, 36, 114
Ascobo1us, 6, 8, 9, 11, 13-15, 17,59,64-66,68,69, - subaffine, 36, 111
71-77 - subspira1e, 36, 109
- a1bidus, 14, 76 - succineum, 34, 36, 113
- americanus, 13 - trigonosporum, 35, 107
- amoenus, 6, 13, 15, 71 Chei1ymenia, 6, 8,21-23,64-66,68,69,89-91
- archeri, 13 - coprinaria, 6,22,23,90
- bistisii, 15, 72 - coprogena, 23
- brassicae, 15, 74 - granu1ata, 23, 91
- crenu1atus, 6, 14, 75 - pallida, 22, 64
- dadei, 8, 13, 15,59, 73 - pulcherrima, 22
- furfuraceus, 6, 13, 14, 77 - raripi1a, 6, 23, 89
- hawaiiensis, 6, 13, 15, 74 - stercorea, 6, 21-23, 90
- immersus, 6, 9,13-15, 71 - the1ebo1oides, 22, 23, 89
- michaudii, 14, 75 C1adorrhinum, 40, 59, 139, 142
- queze1ii, 14, 72 Coniochaeta, 6, 39,40, 65-69, 118, 119
- scategenus, 13 - discospora, 6,40,118
- stictoideus, 15, 73 - hansenii, 6,40,119
Ascodesmis, 17,68 -leucop1aca, 6, 39,40, 118
Ascophanus, 6,20,21,24,26,66,68, 69 - scatigena, 39,40, 119
- argenteus, 6, 24 - ve1utina, 39,40
- microsporus, 6, 26 Coprotus, 8,21,24,64,66,91-94
- subfuscus, 26 - baeosporus, 24
Ascozonus, 6, 8,25,26,95 - breviascus, 24
- woo1hopensis, 6, 8, 25, 26, 95 - cf. discu1us, 24
- dup1us, 24
Bombardioidea, 49, 67, 148 - 1acteus, 24
- anartia, 49 - 1eucopocillum, 24
- stercoris, 49, 148 - 1uteus, 24, 91
Botryotrichium, 34 . - niveus, 24
- ochraceus, 24, 93
Cercophora, 9, 31,40,41,49, 59, 60, 66, 141-143 - sexdecimsporus, 24, 94
- ca1ifomica, 41,141 - winteri, 24, 94
- grandiuscu1a, 41,59,142
- mirabilis, 41, 142 De1itschia, 6, 9, 51-53, 64, 66--68,154-159
- recta, 41,59,143 - chodoco1a, 53, 155
- sordarioides, 41, 143 - didyma, 51, 52, 155
Chaetomium, 6, 8, 9,28,29, 33-36, 64, 65, 67-69, - excentrica, 53, 154
107-114 - furfuracea, 53, 158
- ampullare, 36, 109 - gigaspora, 52, 53, 157
- bostrychodes, 36, 114 - intonsa, 53, 157
- brasiliense, 36, 110 - 1eptospora, 52, 159
- caprinum, 33 - sp., 53, 156
- coch1iodes, 33 - variispora, 52, 53, 159
- convo1utum, 34, 36, 113 - vulgaris, 53, 158
170
 -winter~6,52,53,156 - minutissima, 49, 105
P1euroascus nicho1sonii, 28, 29, 101
Fimaria, 25, 64, 68 Podosordaria, 50, 51, 66-69, 150
Fimetariella, 6, 49, 60, 67, 148 - sp., 50, 150
- maxima, 6, 49 - vio1acea, 51, 150
- minuta, 49, 60, 148 Podospora,6,8,31,41-50,60,61,63-65,67,69,
120-137, 144
Graphium, 31, 106 - ampullacea, 46,122
Gymnoascus, 6, 28, 29, 99 - ampullaceae, 122
- reessii, 28, 29 - anserina, 6, 42, 45, 124
- argentiniensis, 45, 127
Hypocopra, 50,51,63,67, 149 - austra1is, 6, 31,42,44,47, 137
- equorum?, 51 - austrohemisphaerica, 42, 46, 125
- sp., 51, 149 - bico1our, 43
- bicornis, 46, 134
Iodophanus,8,9,20,63,66, 69,87 - bifida, 46, 121
- carneus, 8, 9, 20, 87 - cf. didyma, 45
- granulipo1aris, 20 - communis, 44,126
- verrucosporus, 20, 87 - conica, 42, 44, 132
- coronifera, 42
Kernia nitida, 29, 100 - curvicolla, 42, 46, 120
- curvu1oides, 43,44, 132
Lasiobolidium orbicu1oides, 29, 100 - dactylina, 46, 125
Lasiobolus, 3, 6,11,17-19,21,26,64,69,83-85 - decipiens, 6, 42, 45, 127
- ciliatus, 3, 18, 19, 83 - dolichopodalis, 45, 128
- cunicu1i, 18 - ellisiana, 46, 120
- diversisporus, 19,84 - excentrica, 42, 45, 124
- intennedius, 18 - fabiformis, 45, 60, 61, 133
- microsporus, 19,84 - fimiseda, 6, 42, 43, 44, 128
- pi1osus, 6, 18 - gigante, 44, 47, 144
- trichoboloides, 19, 85 - gigantea, 44, 47, 147
Lecythophora, 39, 69 - glutinans, 44, 130
Leucoscypha, 6,25 - hya1opi1osa ?, 46, 135
Lindquistia, 50, 68, 150 -ignot~45,46,61,136
- intestinacea, 45, 123
Me1anospora, 6, 32, 65, 106 - miniglutinans, 44, 133
- brevirostris, 32, 33, 106 - n. sp., 45, 133
- damnosa, 6, 32 - pectinata, 44, 122
- raffae1ea, 32, 66 - perp1exans, 6, 42, 44, 129
- zamiae, 6, 32 - petrogale, 8,44,46,48,64, 144
Mycoarctium, 9, 26, 60, 66,85 - p1eiospora, 6, 42, 45, 126
- ci1iatum, 26 - prethopoda1is, 8,42,46, 123
- sphaerosporum, 26, 60, 85 - pyriformis, 6,42,45, 134
- setosa, 46, 121
Nigrosabu1um globosum, 6, 28, 29, 101 - tetraspora, 6, 8,42, 43, 64, 129
- vestico1a, 6, 44, 64, 130
Ophiostoma, 31, 32, 105 Poronia, 6,50,63, 66,67, 151
Orbi1ia, 9,11,19,20,26,66-69,86 - erici, 50, 67, 151
- a1nea, 20, 86 - oedipus, 50, 66, 151
- inflatu1a, 20, 86 - punctata, 6, 50
Preussia, 28, 29, 54, 65, 66, 69, 102
Petriella, 31,32, 64, 66, 106 - cylindrica, 29, 102
- sordida, 31, 106 - funicu1ata, 29, 102
Peziza, 6,21,63,66,88 Pseudoarachniotus roseus, 29, 99
- p1eurota, 21, 88 Pseudombrophi1a, 25, 64, 96
- vesicu1osa, 21, 88 - theio1euca, 25, 96
Phia1ophora, 48, 129, 130, 135, 143, 144 - virginea, 25
Phomatospora, 49, 68, 105 Pu1vinula, 6, 25, 95

171
 - cinnabarina, 6, 25, 95 - chaetomoides, 57, 169
- subpro1ata, 25 - cymatomera, 57, 163
Pyxidiophora, 6, 31, 67, 104 - dakotensis, 56
- arvensis, 31, 104 - euryospora ?, 57, 166
Rhinocladiella, 17 - grandispora, 57, 164
Ryparobius, 26 - hercu1ea, 58, 168
- insignis, 6, 58, 166
Saccobo1us, 6, 8, Il,15-17,54,64,65,67,79-82 - intermedia, 6, 57, 160, 162
- beckii, 15, 16 - leporina, 55, 56
- citrinus, 17,81 - longisporopsis, 57, 165
- depauperatus, 16 - megalospora, 6, 56, 57, 164
- geminatus, 17, 80 - minima, 6, 55, 57, 161
- g1aber, 15, 16 - nigropurpurea, 56
- g1obu1iferellus, 15, 16 - octonalis, 58, 167
- infestans, 16, 82 - ovina, 55, 57, 165
- ? minimus, 81 - platymera, 58, 168
- minimus, 6, 17, 81 - subtilis, 57
- obscurus, 16 - tetramera, 57, 163
- portoricensis, 17, 80 Spororthrix,49
- reticu1atus, 15 Strattonia, 9,43,47,48,61,62,64,145,146
- thaxteri, 15 - grandis, 48, 61, 145
- verrucisporus, 17, 79 - insignis, 48, 146
- versicolor, 6, 17, 79 - oblecythiforrnis, 48, 62, 146
Scedosporium, 31 - zopfii, 48, 145
Schizothecium, 41, 64 Thecotheus, 17,64-69,78
Sclinia, 33, 66, 68, 96 - crustaceus, 17, 78
- africana, 33 - pelletieri, 17, 68, 78
- pulchra, 33, 68,96 Thelebo1us, 6, Il,26-29,65,66,69,97,98
Semidelitschia, 8, 51, 52,64,65,68, 154 - microsporus, 26, 27, 65, 98
- agasmatica, 52 - polysporus, 27, 97
- nannostellata, 52, 64, 154 - stercoreus, 6, 26, 27, 98
- tetraspora, 52 Trichobolus,6, Il,26,27,67,97
Sordaria,6, 9,36-39,47,49,64-66,68,69, 115-117 - pilosus, 27
- alcina, 37-39, 116 - sphaerosporus, 27
- arctica, 6, 38, 116 - zukalii, 6, 27, 97
- brevicollis, 37, 38, 68, 117 Trichodelitschia, 6, 28, 51, 67, 152, 153
- fimicola, 6, 9, 36-39, 115 - adelphica, 51, 153
- goundaënsis, 38 - bisporula, 6, 51, 152
- heterothallis, 37 - munkii, 51, 152
- humana, 36, 38, 116
-lappae,38 Xylaria brasiliensis, 50
- macrospora, 6, 37-39, 115
- rabenhorstii, 49 Zopfiella erostrata, 29, 103
- sclerogenia, 37 Zygopleurage, 49,67, 147
- spermatiophora, 38 - zygospora, 49, 147
- superba, 37, 38 Zygosperrnella, 49, 67, 147
- thermophila, 37 - insignis, 49, 147
- tomento-alba, 37
Sphaeronaemella, 6, 31,65,69, 105
- firnicola, 6, 31, 65, 69, 105
Sporormia, 53-55, 64, 65, 160
-fimetaria, 53, 54, 65,160
Sporormiella, 6, 8, 9, 28,50,53-58,61,64,69, 160­
169
- albolanata, 57, 61, 169
- australis, 6, 57, 161
- bipartis, 58, 167
- borealis, 57, 160, 162

172