The Botanical Review 1970 v36

THE BOTANICAL REVIEW
VOL. 36 APRIL-JUNE, 1970 No. 2
CURRENT T R E N D S IN T H E C L A S S I F I C A T I O N
OF DISCOMYCETES 1
JAMES W. KIMBROUGH
Department of Botany
University of Florida
Gainesville
A HISTORICAL REVIEW ................................................................................ 92

INFLUENCE OF APOTHECIAL STRUCTURE ON THE CLASSIFICATION
O F D I S C O M Y C E T E S ............................................................................................ 104
D e v e l o p m e n t o f t h e A s c o g e n o u s S y s t e m ................................................................ 104
E x c i p u l u m .................................................................................................................. 112
Asci .......................................................................................................................... 113
A s c o s p o r e s ........................................................................................................ 117
P a r a p h y s e s ........................................................................................................... 120
Apothecial Shape .......................................................................................... 120
U l t r a s t r u c t u r e of D i s c o m y c e t e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
THE ORDERS AND FAMILIES OF DISCOMYCETES ...................................... 121
T h e H e l o t i a l e s ........................................................................................................... 123
A s c o c o r t i c i a c e a e ................................................................................................... 123
Geoglossaceae ............................................................................................... 123
Sclerotiniaceae ............................................................................................. 125
Caliciaceae ............................................................................................ 125
H e l o t i a c e a e ............................................................................................................. 126
Hyaloscyphaceae ...................................................................................... 128
Orbiliaceae ............................................................................................. 129
D e r m a t e a c e a e ...................................................................................................... 129
Hemiphacidiaceae ................................................................................ 130
The Phacidiales ......................................................................................... 130
Hypodermataceae ....................................................................... 131
Rhytismaceae .......................................................................................... 131
Phacidiaceae ................................................................................................. 132
The Ostropales ............................................................................................... 132
The Cyttariales .................................................................................................. 133
T h e Pezizales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
The suborder Sarcoscyphineae ................................................................... 134
S a r c o s c y p h a c e a e ...................................................................................................... 135
T h e s u b o r d e r Pezizineae ................................................................................... 135
M o r c h e l l a c e a e ................................................................................................. 137
H e l v e l l a c e a e ..................................................................................................... 137
Otidiaceae ................................................................................................................. 139
A l e u r i a c e a e .................................................................................................... 140
Pezizaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
T h e l e b o l a c e a e .................................................................................................... 142
A s c o b o l a c e a e ................................................................................................... 143
P y r o n e m a t a c e a e ............................................................................................... 144
R e l a t i o n s h i p to t h e T u b e r a l e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Synoptical Key .......................................................................................... 146
LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
W l a . A g r i . E x p . Sta. J o u r . Series # 3 6 0 1 .
91
92 THE BOTANICAL REVIEW
A H I S T O R I C A L REVIEW
The term discomycete is one coined in the early days of mycology.
The name persists in mycological literature, even though its meaning
has changed somewhat as our knowledge of the fungi has increased.
Generally, it may be said that the discomycetes are those ascomycetes
which possess an apothecium. In the apothecium, the asci are arranged
in tufts or typically palisade layers, or hymenium, almost always lying
between paraphyses, and the ascocarp is an open structure, exposing the
hymenium at various stages of development. The apothecium may be
variously shaped, though typically it is saucer- or cup-shaped, giving to
this group the common name, cup-fungi.
Pliny (23-79 A.D.), a Roman naturalist and scholar, mentioned nearly
a thousand plants in his Historia Naturalis. Among his writings one
finds, perhaps, the first mention of a discomycete in the following quote:
"Belonging to the mushroom kind, also there is a species known to the
Greeks by the name 'Pezica' which grows without root or stalk." Dillenius
(1719) left little doubt in his description that Peziza applied to the cup-
shaped groups of fungi. No species were mentioned, but in the catalogue
proper, which is arranged by months in which the plants appear, several
Pezizas are listed. Linnaeus (1753) in Species Plantarum adopted the
genus Peziza essentially in the same sense of Dillenius. Linnaeus treated
all the fungi under ten genera, two of which, Elvela and Peziza, comprised
the discomycetes recorded at that date. He recognized Elvela with two
species and Peziza with eight. Persoon (1801) was one of the first to
attempt a systematic classification of the discomycetes. They were placed
in his second class, Gymnocarpi, which included 6 orders. Order 5, the
Hymenothecii, contained most of the fleshy fungi. Among the 6 sub-
groups of Hymenothecii, 8 discomycete genera are found, Gleoglossum,
in the Clavaeformes; Spatularia, Leotia, Helvella, Morchella, Peziza,
Ascobolus, and Helotium in the Helvelloidei. By this time almost 200
species of discomycetes were recognized.
The next significant step in the systematic arrangement of discomy-
cetes is that of Fries (1822). The Friesian arrangement will be recognized
as an important step in the development of the classification of this
group. Fries (1821) recognized 4 broad classes of fungi, the Coniomycetes,
Hyphomycetes, Gasteromycetes, and Hymenomycetes. With the exception
of Thelebolus, found in the Gasteromycetes, the remaining discomycetes
were placed in the Hymenomycetes. Four orders of Hymenomycetes
treated were the Sclerotiacei, Tremellini, Uterini, and Hymenini. The
cupulate discomycetes were found in the Uterini and the stipitate forms
among the genera of Hymenini. In 1822, Fries established the order
Elvellaceae which was more or less equivalent to the Helvelloidei of
Persoon (1801) and Hymenomycete-Uterini of his 1821 classification. The
Elvellaceae contains two suborders, the Mitrati founded to include the
genera with mitrate or clavate apothecia, while the cup-shaped forms
made up the Cupulati. The genera of clavate forms included Mo~'chella,
Helvella, Verpa, Leotia, Vibrissea, and Rhizina; distinquished largely
KIMBROUGH: CLASSIFICATION OF DISCOMYCETES 93
on the configuration of the hymenium. The Cupulati was divided into

three families: the Peziaeae, in which he placed Peziza, PatelIaria,
/lscobolus, and Bulgaria; the Dermeae, containing Ditiola, Tyrnpanis,
Cenangiurn, and Stictis; and the Heteroclitae, with Solenia and Cyphella.
These divisions were made primarily on shape and texture of apothecia.
Although the Friesian system was founded initially on that of Persoon,
it is very significant that the establishment of subgeneric groups led to
a still further narrowing of generic limits. Fries (1822) divided Peziza
into three series, which in turn were divided into twelve tribes. Many
of these tribes were elevated to generic rank by later workers. Another
important fact is that Fries established ordinal, subordinal, and familial
taxa, which did not exist in systems of classifications proposed earlier.
The Friesian system of classification remained in use for almost 60 years.
In 1849 Fries modified his previous system with the establishment of a
family Discomycetes which included six "orders": Helvellaceae, Bul-
gariaceae, Dermateae, Patellariaceae, Phacidiaceae, and Sticteae. He
leaned strongly on the consistency of apothecia in establishing these taxa
and acknowledged the difficulty in using microscopic characters to base
natural genera.
Even though the Friesian system was followed by a number of my-
cologists (Nylander, 1869; Fuckel, 1869; Karsten, 1871; Cooke, 1879), a
number of significant changes occurred in the mid-19th century. Most
work done before this time had emphasized the external or gross mor-
phology of fungi. Textbooks such as DeBary's (1866) Morphology and
Physiology of the Fungi, Lichens and Myxornycetes placed great emphasis
on structural detail and to the development of fungi. More and more
students were encouraged to look at the microscopic features of fungi,
and this, of course, led to the type of investigations on which sounder
systems of classification could be based. De Notaris (1864) catalogued
26 genera of discomycetes without following any systematic arrangement.
It is important, however, that microscopic characters such as form and
color of asci, paraphyses, and spores as well as cellular nature of the
excipulum were used. As time passed more emphasis was placed on
microscopic measurements and, to a limited extent, microchemical reac-
tions of certain elements of the apothecium. Nylander (1869) was one
of the first to stress the iodine reaction in asci. With the accumulation
of both microscopic and microchemical data many genera, especially
large ones like Peziza, were broken into many subunits. This is especially
evident in Karsten's (1869) Monographia Pezizarura fennicarurn in which
Peziza is divided into 25 subgenera. Some of the more important sub-
genera were Aleuria with 57 species; Phialea, 41 spp.; Mollisia, 15 spp.;
Trochila, 12 spp.; Patellaria, 9 spp.; Orbilia, 8 spp.; Lachnetla, 24 spp.;
and Lachnea, 20 spp. Fuckel (1869) followed the familial arrangement
of Fries, yet his system displayed a further narrowing of generic limits.
Eighty genera of discomycetes were recognized, 32 of which were placed
in the Pezizei, 18 in the Phacidiacei, 11 in the Patellariacei, 7 in the
Helvellacei, and 6 each in the Stictei and Bulgariacei. The trend toward
94 THE BOTANICALREVIEW
use of microscopic features in delimiting species is evident in Fuckel's
work. The excellent descriptive and illustrative work of Tulasne and
Tutasne (1865) was a landmark at the time. This work greatly influenced
the later works of Karsten and others. Karsten (1871) recognized the
order Discomycetes and placed within it three families, the Helvellaceae,
Pezizaceae, and Phacidiaceae. He was one of the first to designate sub-
families, relying heavily on the cellular nature of sterile elements of the
apothecium. The HeIvellaceae contained three subfamilies based both
on texture and shape of apothecia, and shape of ascospores. The Peziza-
ceae contained seven subfamilies distinguished generally by a similar set
of characters. Karsten included 53 genera, but in this respect he was
more conservative than Fuckel.
Although some of the more important systems of classification were
made by Fries, Karsten, and Fuckel, a number of other workers made
significant contributions to the systematics of discomycetes. Among these,
the work of Crouan and Crouan (1857) is important in that they were
the first to illustrate and describe the operculate dehiscence of asci. The
significance of this discovery was not fully appreciated until the later
emphasis by Boudier (1879). Cooke (1879) published plates and descrip-
tions of 406 species of primarily operculate discomycetes. The value of
this publication lies in our ability to still use his illustrations and
measurements to identify specimens, and also the fact that most of his
drawings were based upon authentic herbarium collections. In 1884,
Saccardo used the novel approach of arranging discomycetes according
to the size, shape, color, and septation of ascospores. He adopted an
order Discomycetes which included eight families. Saccardo tended to
be more conservative than Karsten or Fuckel, having rather large genera
with a large number of subgenera. This system, modified by adding
the Gymnoascaceae, was adopted by Phillips (1887) in his British Dis-
comycetes. Here, many of the Fuckel and Karsten genera are lowered
to subgeneric rank.
Perhaps one of the most significant publications of the late 1800's
was Boudier's (1885) new natural classification of discomycete genera
based primarily on the presence or absence of an operculum. In Boudier's
system, external characters were still regarded as of primary importance,
but a number of microscopic characters were introduced, such as the
amyloidity of asci, number of oil drops of spores and others. He intro-
duced two large sections, the operculate and inoperculate discomycetes.
The operculates were initially divided into the tribes Mitres (stipitate),
Cupules (sessile, mostly cupulate), and Lenticules (sessile, becoming hemi-
spheric, convex). Seven operculate families were recognized, distinguished
both on external and microscopic characters. The inoperculates were
divided into three tribes: Clavules, Carnoses, and Cyathules, and a total
of 7 families. This important division was not accepted by Boudier's
contemporaries such as Quelet, 1886; Phillips, 1887; Saccardo, 1889-1931;
Rehm, 1887-1896; Cooke, 1892; Schroeter, 1893; Massee, 1895; Lindau,
1897; Clements, 1909; and was later rejected by Clements and Shear,
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added. This immense work included 213 genera with more than 3,500
species. It was quite obvious that spore length-width ratio, color, and
septation frequently changed with age. This, in essence, caused this prac-
tical system to soon loose its influence on the classification of this group.
Saccardo erected two divisions: (1) those with a receptacle, and (2) those
without. These divisions were then divided into families, and the families
further divided into the hyalosporae, phaeosporae, hyalodidymae, phrag-
mosporae, dictyosporae, and scolecosporae. This system greatly influenced
the classification used by Cooke (1892) and Clements and Shear (1931).
As the influence of Saccardo's system of classification was being felt
in many areas, so was that of some prominent German mycologists. One
of the most important of these was Rehm's (1887-1896) massive study
of the discomycetes for Rabenhorst's Kryptogamen-Flora. The order
Discomycetes was divided into two divisions, the Pezizaceae and Hel-
vellaceae. The division Pezizaceae was broken into five suborders, the
Phacidiaceae, Stictideae, Tryblidieae, Dermatiaceae, and Pezizeae. The
most striking innovation in this work was the inclusion of many lichens
among the first three or four suborders. Rehm followed Karsten in using
characters of the sterile portions of the apothecium in making sub-
divisions. The large suborder Pezizeae included four families, the Mol-
lisiaceae, Helotiaceae, Eupezizeae, and Ascobolaceae. The generic limits
were a combination of both those of Fuckel and Saccardo. For many years,
this was the most practicable textbook for the identification of the
discomycetes. This work was somewhat discredited because the author
failed to recognize the operculate and inoperculate series.
Schroeter (1893) followed Rehm for the most part, but differed from
him in including the Taphrinineae and Hysteriineae under the dis-
comycetes. A number of genera were shifted around, such as certain
members of Rehm's Dermatiaceae were placed in the Pezizaceae. Schroe-
ter in 1894, initiated the writing of the discomycetes for Engler and
Prantl. Unfortunately, this work was cut short by death when it was
just begun. It was continued along similar lines and brought to com-
pletion by Lindau (1897).
Durand (1900) presented a classification of the fleshy Pezizineae. This
included what was equivalent to Rehm's suborder Pezizeae, i.e. most of
the cup-shaped discomycetes which are fleshy or waxy. Durand treated
four families, the Pezizaceae, Ascobolaceae, Helotiaceae, and Mollisiaceae.
The first two families mostly fleshly or rarely leathery, the last two, waxy,
fleshy-waxy or membranous.
Much fundamental work on the taxonomy of the cup-fungi was done
by von H6hnel (1903-1918), and although he ignored a systematic treat-
ment, his work was extremely useful for those seeking to establish a
natural arrangement. One of the greatest assets of this work was the
large number of authentic specimens described or recorded.
Boudier (1907) expanded his 1885 system of classification to include
other subdivisions and 7 operculate families and 12 inoperculate. In the
earlier paper, there were subfamily "groups" which, with some rearrange-
KIMBROUGH~ C L A S S I F I C A T I O N OF DISCOMYCETES 99
ments, were here designated as tribes. Twelve operculate tribes, contain-
ing 72 genera were recognized. Boudier used not only external features
of apothecia, but also cytochemical and microscopic features in delimiting
tribes. As with the operculates, the inoperculates were divided into two
broad divisions, the marginates with 63 genera and the immarginates 102
genera. One of the greatest criticisms of the Boudier system is that many
of his taxonomic decisions were based on descriptions rather than the
examination of authentic material. This is readily apparent when one
examines a number of species in many of the genera.
The scope of the discomycete orders has varied considerably from one
author to another. G~iumann (1926), Seaver (1928), Clements and Shear
(1931), and Bessey (1950) would accept the whole of the fleshy cup fungi
in the Pezizales, but the last two authors recognized the significance of
the ascus character in the classification of these fungi. Seaver (1928)
accepted the difference between operculate and inoperculate discomycetes,
but otherwise developed his own, practical system. Earlier authors such
as Rehm (1887-1896), Schroeter (1893), and others recognized the order
Helvellales to accommodate the larger and distinctly stipitate species
without giving any consideration as to whether the asci were operculate
or inoperculate. Seaver (1928) recognized two families of the section
operculates, the Pezizaceae and Elvelaceae, and three, the Geoglossaceae,
Helotiaceae, and Cenangiaceae, in the section inoperculates. The large
families, Pezizaceae and Helotiaceae were further subdivided into tribes
(Seaver, 1928 and 1951). He relied heavily on ascospores, external features,
and substrate to distinguish 8 tribes of the Pezizaceae. Many of these
tribes corresponded closely to families recognized by Boudier (1907).
Similar features were used in distinguishing 5 tribes of the Helotiaceae.
Nannfeldt (1932) pointed out the weakness of this scheme of classification,
which is largely due to the fact that Seaver founded his system on sets of
convenient, but wholly unnatural, genera.
Nannfeldt's (1932) outstanding work laid the basis for a critical re-
evaluation of many genera, together with a revision of their nomenclature
during the 1950's and 1960's. Three main groups of higher Ascomycetes
were proposed, the Plectoascales, Ascoloculares, and Ascohymeniales. The
main subdivisions of these groups were based on the development of the
ascus itself and of the fructifications within or upon which they are
assembled. The discomycetes were considered in the Ascohymeniales,
along with the Pyrenomycetes and others. In these the ascogonium, and
antheridium, if it exists, lie free or enclosed in various amounts of sterile
tissues. The ascogonium and resulting ascogenous hyphae in the pyre-
nomycetes and many discomycetes soon become enclosed and a hymenium
is formed from parallel hyphae (future paraphyses) among which asci
arise. Nannfeldt separated under the Ascohymeniales a number of nat-
urally related groups in which, however, the boundaries in general be-
tween the discomycetes and pyrenomycetes do not overlap. The discomy-
cetes seemed to form, in contrast to the pyrenomycetes, a proportionally
homologous group. Since the fruit body exposed the hymenium directly
to the air and they do not attain as many highly differentiated forms,
the discomycetes were considered the most primitive. He recognized
nine orders of Ascohymeniales, 4 of which, the Pezizales, Ostropales,
Helotiales, and Lecanorales comprised the discomycetes.
The Ostropales with the single family Ostropaceae, was proposed by
Nannfeldt (1932) for certain genera of Helotiales characterized by long-
cylindric asci with greatly thickened apices and very long, filiform
ascospores. He included the genera Vibrissea, Apostemidium, Ostropa,
Robergea, Schizoxylon, and Stictis. There is currently considerable
controversy over the validity of the Ostropales. Six families of Helotiales
were recognized; each distinguished by a set of characteristics of spores,
asci, excipular and other sterile structures, and parasitic nature. These
are the Dermateaceae, Phacidiaceae, Orbiliaceae, Hyaloscyphaceae, Helo-
tiaceae, and Geoglossaceae. He relied heavily on the nature of excipular
structure, as correlated with ascal and spore features, to further divide
the Dermataceae, Hyaloscyphaceae, and Helotiaceae into subfamilies. In
delimiting the genera of Hyaloscyphaceae, Nannfeldt accepts most of the
conclusions arrived at by von H6hnel in his exhaustive studies of type
material. The family is divided into two subfamilies, Lachneae and
Hyaloscypheae, mainly on the basis of apothecial size, with a third rather
ill-defined group, Arachnopezizeae, segregated on the basis of being seated
on a subicilum. Detailed structure of hairs was used to distinguish 13
genera. This treatment was followed, with modifications, by Dennis
(1949). Nine subfamilies of Helotiaceae were recognized by Nannfeldt,
comprising a total of 30 genera. The Dermateaceae was likewise divided
into 9 subfamilies with 30 genera. Again, size and texture of apothecia,
and detailed structural features of the excipulum were used in distinguish-
ing genera of these families.
During the late 1940's and early 1950's a re-evaluation of many dis-
comycete genera began. These re-evaluations all may be said to have
their basis in Nannfeldt's critical studies, especially those of the inoper-
culates (1932) and operculates (1949). Outstanding among these studies
were those on ascus structure by Chadefaud (1940a, 1940b, 1942, 1943,
1944, 1946a, 1946b, 1949), those on ascospore, ascal, and apothecial
anatomy by Le Gal (1942, 1946a, 1946b, 1947, 1949, 1953a, 1953b, 1957,
1963), and those predominantly on nomenclature by Korf (1953, 1954,
1957a, 1959, 1960a, 1963b, and Korf and Rogers, 1967).
Chadefaud's (1940a) study of ascus structure in Bulgaria inquinans
Pers., and subsequently followed by his studies of many other groups, has
stimulated a great interest in ascus structure as a means of showing
natural relationships. Although exterior features of asci had been de-
scribed by various workers, Chadefaud was first to describe the internal
apical apparatus of operculate and inoperculate asci in detail. One of
the first major discoveries from these studies was that the genera of
Sarcoscyphaceae and others appeared to have an apical apparatus inter-
mediate between the operculate and inoperculates. Le Gal (1946b)
suggested the taxon "Subopercules" for groups with this ascus type.
KIMBROUGH: C L A S S I F I C A T I O N O F DISCOMYCETES 101
Nannfeldt (1949), following Le Gal, recognized two families of Pezizales,
the Sarcoscyphaceae and Pezizaceae. That ascus structure still plays an
important role in classification is evidenced by recent work of Kimbrough
(1966a, 1966b) and Kimbrough and Korf (1967) who leaned heavily on
ascal characters in delimiting genera of the Thelebolaceae. This work
also points out possible relationships of operculates to inoperculates.
Le Gal (1947) gave extremely valuable information for taxonomic and
phylogenetic evaluations in her monumental study of the formation of
spore walls and ornaments in the operculate discomycetes. She believed
that spore character should take precedence of all other characters. The
system of classification proposed by Le Gal in this study included the
families Morchellaceae, Helvellaceae, Aleuriaceae (= Pezizaceae), Hu-
mariaceae, Ascobolaceae, and Sarcoscyphaceae. This familial arrange-
ment has been followed by most current mycologists (Dennis, 1960; Moser,
1963; G~iumann, 1964; Rifai, 1968; Dennis, 1968; Eckblad, 1968). Le Gal
pointed out that the ascospores of operculate discomycetes were always
unicellular, often ornamented and larger than those of the inoperculate
group, which were smooth walled, always small, one to many celled and
often with a very high length-width ratio. Unfortunately, Le Gal pro-
posed two new ordinal names, Homospermales and Heterospermales, to
replace the classical orders Pezizales and Helotiales.
Beginning with new rules of typification as they affect Sarcoscypha
and Felutaria, Korf (1953) has contributed greatly to the clarification of
nomenclature of both operculate and inoperculate discomycetes (cited
above), as well as other groups of ascomycetes. Korf has emphasized that
few nomenclatural acts have greater potential consequences than the
designation of types, that typification of genera frequently involves wide-
spread name changes, but when wisely applied may achieve stability in
nomenclature, and that implications of typification are felt at all tax-
onomic levels. Korf's (1954) revision of the classification of operculate
discomycetes had a great influence on subsequent family delimitation in
this group. He proposed 3 families, the Cyttariaceae, Pezizaceae, and
Sarcoscyphaceae. The large family Pezizaceae encompassed 11 tribes,
many of which were the basis for families erected by subsequent workers
(Rifai, 1968; Eckblad, 1968).
Of particular significance on the taxonomy and phylogeny of dis-
comycetes has been the interesting studies of Berthet (1961, 1963, 1964a,
1964b) on the number of nuclei in various apothecial structures, the
presence or absence of imperfect stages, developmental aspects such as
crozier formation, and other cytological and cultural aspects.
The rejuvination of interest in the discomycetes resulted in the pub-
lication of numerous studies of both major and minor groups. Outstand-
ing among those of major groups was Dennis' (1960) British cup fungi
and their allies. Even though he treated the discomycetes in fair detail,
nearly hal{ of the volume is actually devoted to the plectomycetes and
pyrenomycetes. This book was recognized at that time as the only reason-
ably complete synthesis of modern thought on ascomycete classification.
Dennis relied heavily on Nannfeldt for an over-all system for major

grouping in the inoperculates; on Boudier and Le Gal for concepts in
the operculates; and on Luttrell (1951), Munk (1957), yon Arx and
Miiller (1954) for recent thought in the pyrenomycetes. Keys for all
genera were provided, but unfortunately keys to species were not given;
but instead descriptions and illustrations of the most common British
species were included.
Some of the more important regional treatments of discomycetes, or
major groups of discomycetes, are those of Grelet (1942-1959) on the
discomycetes of France; Dennis (1949), the British Hyaloscyphaceae;
Ramsbottom and Balfour-Brown (1951), for the British Isles; Dennis
(1954a, 1954b), operculate and inoperculate discomycetes of South Amer-
ica; Le Gal (1953b, 1961), discomycetes in the Crouan Herbarium; Gilkey
(1954), Tuberales of North America; Mains (1954-1956), North American
Geoglossaceae; Dennis (1956, 1962a), British Helotiaceae; Rawlings (1956),
Australian Cyttariaceae; Eckblad (1957), the Sarcoscyphaceae of Norway;
Dennis (1958), Australian Helotiales and Ostropales; Le Gal (1959), dis-
comycetes of the Belgium Congo; Korf (1958, 1959), on Japanese discomy-
cetes; Batra (1961a, 1961b), Batra and Batra (1963), Indian discomycete
flora; Gamundi (1956, 1957, 1959, 1960, 1964), discomycetes of Argentina;
Eckblad (1963), Geoglossaceae of Norway; Korf (1963), status and scope of
discomycetes flora of Asia; Denison (1963), preliminary study of Costa
Rican Pezizales; Denison (1965, 1967, 1969), Central American Pezizales;
Maas Geesteranus (1965), Geoglossaceae of India and adjacent countries;
Thind et al. (1959-1967), on operculate and inoperculates of India;
Naumov (1964), discomycetes of the Leningrad district; Rifai (1968), the
Australasian Pezizales.
The latter reference (Rifai, 1968), along with Dennis' (1968) revised
and enlarged edition of "British Ascomycetes" and Eckblad's (1968) re-
evaluation of the taxonomy, phylogeny and nomenclature of operculate
discomycetes, has greatly expanded our knowledge and understanding of
the Pezizales. Rifai (1968) adopted, in general, the system of classification
proposed by Le Gal (1947). The suggestions made by Nannfeldt (1949),
Korf (1954), and Berthet (1964b) to give equal rank to the Sarcoscyphaceae
and Pezizales were incorporated into Rifai's scheme by recognizing two
suborders, the Sarcoscyphineae and the Pezizineae. Eight families were
treated, the Sarcoscyphaceae, Morchellaceae, Helvellaceae, Humariaceae,
Pyronemataceae, Thelebolaceae, Pezizaceae, and Ascobolaceae.
Rifai (1968), following G~iumann (1964), recognized the monotypic
family Pyronemataceae and elevated to family rank the Thelebolaceae.
He considered the Humariaceae the largest and probably the most dif-
ficult family of the Pezizales. It was divided into four tribes, the Otideae,
Lachneae, Ciliarieae, and Aleurieae. These corresponded closely to a
number of tribes formerly recognized by Korf (1954). Clear, easy to follow
keys for most taxa were included by Rifai, and excellent illustrations of
most all apothecial characters were provided.
Another of the most significant contributions was Eckblad's (1968)
KIMBROUGH: C L A S S I F I C A T I O N OF DISCOMYCETES 103
reevaluation of the taxonomy, phylogeny and nomenclature of the genera
of operculate discomycetes. All genera or generic names once associated
with the operculate discomycetes are treated, their taxonomic and phy-
logenetic position is discussed on the basis of a study of the type species
or other species, and their nomenclatural status is ascertained. The single
order Pezizales is divided into the following families: Thelebolaceae,
Ascobolaceae, Pyronemataceae, Pezizaceae, Otideaceae, Helvellaceae, Mor-
chellaceae, Rhizinaceae, and Sarcoscyphaceae. The phylogeny of the
operculate and inoperculate discomycetes is discussed and the Thele-
bolaceae suggested as their common origin. The helvelloid families and
the Sarcoscyphaceae are regarded as derived.
In what likely will become one of the most widely accepted treatments
of the ascomycetes, Dennis' (1968) British Ascomycetes, includes the follow-
ing orders of discomycetes: Pezizales, Helotiales, Phacidiales, Lecanorales,
and Ostropales. For the Pezizales he follows Le Gal (1947), with the
addition of the Thelebolaceae. The idea of giving the Pezizaceae and
Sarcoscyphaceae equal rank is also expressed here. The large inoperculate
order Helotiales consists of eight families, the Ascocorticiaceae, Geoglos-
saceae, Sclerotiniaceae, Caliciaceae, Helotiaceae, Hyaloscyphaceae, Orbi-
liaceae, and Dermateaceae. The basis of much of his system of inoper-
culate classification was derived from Nannfeldt (1932), influenced greatly
by Whetzel (1945), Seaver (1951), and others. The Phacidiales have been
variously distributed in most of the systems of classification. Many
modern authors have followed Nannfeldt (1932), who treated them as a
family of the Helotiales. Dennis, supported by the study of Terrier (1942)
considers them as an order with three families, the Hypodermataceae,
Cryptomycetaceae, and Phacidiaceae. Dennis treated a heterogeneous
group of fungi among the Lecanorales. Most were lichenized. However,
of those accepted, the Patellariaceae show affinities to the bitunicate
ascomycetes and the Lecideaceae to the unitunicates. The Ostropales,
with the single family Ostropaceae, is based also on the same genera as
accepted by Nannfeldt.
The recent report by Burdsall (1965) on the puffing of ascospores and
the presence of opercula in the asci of Geopora cooperi Harkness adds
support to the views that together with the Pezizales the Tuberales, or
at least many of its genera, belong to the Operculatae. The distinction
between the Pezizales and the Tuberales is not always clear. Generally
the epigeous habit and the presence of forcible discharge of spores in
the Pezizales are contrasted to the hypogeous habit and lack of a mech-
anism of forcible spore discharge in the Tuberales. The subsequent
transfer (Burdsall, 1968) of Sepultaria and Pseudhydnotrya to the genus
Geopora shows that perhaps the affinities are much closer than most
mycologists have realized. To him the division of the Tuberales based on
gross morphology seemed extremely artificial. He relied heavily on the
basic structure and cytochemical reactions of spores, asci, and exipula
to show relationships.
The interesting work of Arpin (1968), on the carotenoides of disco-
mycetes, compliments beautifully the cytological studies by Berthet. This

study has already added much color to the numerous gray areas of dis-
comycete classification.
There have been many recent detailed studies of the discomycetes
which have so far not been included in a complete system. Those that
are considered most significant, especially monographic treatments of
genera, will be cited in the discussion of their respective families later
in this paper.
T H E INFLUENCE OF A P O T H E C I A L S T R U C T U R E ON
T H E CLASSIFICATION OF DISCOMYCETES
T o fully appreciate and understand the variations in apothecia among
the various groups of discomycetes one must first understand their
developmental patterns and the basic tissues involved. The arrangement
of the different parts of the apothecium varies, but most apothecia consist
of a hymenium, hypothecium, medullary or ental excipulum, and ectal
excipulum. The types and amount of cells or tissues in each region may
be extremely variable from one taxon to another. Thus, the forms and
shapes assumed by the ascocarp of discomycetes are highly diverse. This
can be attributed to not only the ratio of hymenial to excipular growth,
but also to a great extent the time at which hymenial tissues are formed.
The cytological aspects of the mycelial system in the discomycetes
has been thoroughly summarized by Berthet (1964b). He reported that
the mycelia of the Pezizales (including the suborder Sarcoscyphineae and
Pezizineae) were coenocytic; coenocytic mycelia have also been observed
in many species of the inoperculate families Sclerotiniaceae and Geoglos-
saceae, but the other families of the Helotiales usually have mycelia with
uninucleate cells. The studies conducted by Berthet (1961, 1963, 1964a,
1964b) have greatly improved our understanding of the phylogenetic
relationships between members of the discomycetes as a whole.
Development of the Ascogenous System. Very few attempts have been
made to correlate the types of reproductive structures and the mechanisms
of sexual reproduction with classification of the discomycetes. Principal
features and patterns of variation in this critical phase of ascocarp develop-
ment could be extremely useful in demonstrating natural relationships.
As a point of departure we might consider the types of ascogonia of
various groups of discomycetes. Among the inoperculate discomycetes sex
organs were observed by Tulasne and Tulasne as early as 1865. Spermatia
or conidiola were believed to be sperms functioning like the spermatia
of lichens. Drayton (19~4) proved that microconidia, which were found
in many species of Sclerotinia, stimulated the production of apothecia
when placed on certain structures which develop on another thallus of
the same fungus. He termed this type of reproduction "spermatization."
The receptive bodies containing deeply stained, coiled ascogonial hyphae
on the stromatic tissue are the only structures capable of fertilization by
the microconidia. In Coccomyces heimalis Higg. the elongated ascogo-
nium extends up to the coiled acervuli differentiating microconidia.
KINIBROUGH: C L A S S I F I C A T I O N OF DISCOMYCETES 105
Pseudopeziza differentiates its microconidia at lower temperatures than
macroconidia (Blodgett, 1936). Griffiths (1959) showed that apothecia
of Gloeotinia temulenta (Prill. and Delacr.) Wilson and Gray develop
from ascogonial stromata which have been fertilized by microconidia from
a compatible strain. The work of Drayton (1934) and Groves and Drayton
(1939) aroused considerable interest in the sexual behavior of the Sclero-
tiniaceae (Drayton and Groves, 1943; Groves and Loveland, 1953; Groves
and Bowerman, 1955; Groves and Elliott, 1961; Elliott, 1962, 1964a, 1964b,
1967).
Much less cytological information is known about most other families
of the Helotiales. The monotypic family Ascocorticiaceae, considered by
Dennis (1968) and others in the Helotiales, has recently received attention
by Oberwinkler, et al. (1967). Although they provided little cytological
information, it is evident from their illustrations that the asci developed
from a crozier system and that paraphyses are also produced by the same
mycelial system.
Most of our knowledge of the development of the Geoglossaceae was
contributed by Duff (1922). A variety of ascogonial coils were observed
in early stages of development, some long, cylindric, multiseptate coils
as in the Humariaceae, others reduced as a single cell with trichogyne
as in Pyronema. In Leotia lubrica (Scop.) Per. a large branching cell,
presumably an ascogonium, occurs at the base of the very young ascocarp
and appears to give rise to ascogenous hyphae. The ascogenous and sterile
tissues grow simultaneously until the mechanism of ascal development
is triggered, at which time a rapid expansion of hymenial growth occurs.
Details of apothecial development of species of Mitula, Microglossum,
Geoglossum, and Trichoglossum were well illustrated and explained by
Corner (1930a). In each there is a primordial corticated shaft of hyphae,
at the distal end of which the hymenium forms on cessation of upward
growth. Gymnocarpic development was noted in all but Mitula.
Members of the Caliciaceae may be distinguished from other Helo-
tiales by the evanescent character of their asci, which break down and
completely disappear, leaving a globose head of loose dry ascospores held
together by undulating hyphal threads. According to Funk (1963), fruit-
ing of Caliciopsis pinea Peck begins with the aggregation of hyphae
beneath the periderm of infected bark to form a flattened stroma. Sper-
mogonia are formed within the enlarging stroma. As the ascocarp begins
to lengthen, certain filaments in the central core thicken and become
dark-staining. These coils of cells making up the ascogonium send out
trichogynes. After fertilization, cells of the coil bud out side branches
that give rise to asci.
Quite a number of Helotiaceae have been cultured but with little or
no success in inducing apothecia to form in culture: Pycnopeziza (White
and Whetzel, 1938), Tryblidiella (Voorhees, 1939), Retinocyclus (Groves
and Wells, 1956), Godronia (Groves, 1965), Vibrissea (Sfinchez and Korf,
1966), Discocainia (Reid and Funk, 1966). Bell~mere (1958) observed
ascogonia and the development of ascogenous hyphae in Cyathicula.
Here, ascogonia developed early and the ascogenous hyphae proliferated

as the stalk grew upward. Ontogenetic studies have been made on mem-
bers of the tribes Ombrophiloideae and Leotioideae. Biffin (1901) studied
the earlier developmental stages of Bulgaria inquinans Ft. in culture on
wooden blocks. Howarth and Chippindale (1931) studied the gross mor-
phological changes during development of .4scocoryne (Coryne) and the
origin of the ascogenous system.
Among the Hyaloscyphaceae, cultural studies were performed on a
number of species of Dasyscyphus by Hahn and Ayers (1934). Only
imperfect stages were formed. According to Bayliss-Elliott (1927) Arach-
nopeziza has an angiocarpic development. Korf (1951) cultured a number
of species of Arachnopeziza but did not succeed in inducing conidial or
ascal stages.
Cultural studies of the Dermateaceae have been made on Dermea
(Groves, 1946), and Tyrnpanis (Groves, 1952). Cultures were obtained
from both ascospores and conidia of Pezizula (Groves, 1939) and of Duran-
diella (Groves, 1954) but cytological aspects of apothecial production
were not recorded. Thompson (1939) was able to produce apothecia of
Neofabraea in culture but he also did not indicate the type of ascogonia
or mechanism of fertilization.
There have been a number of significant studies on the development
of ascocarps of the Phacidiales. According to Killian and Likhite (1924)
long chains of approximately 40 ascogonial cells develop from one or
more enlarged cells (ascogonial mother cells) in the immature hystero-
thecial stroma of Lophodermiurn hysteroides (Pers.) Sacc. Jones (1935)
discussed two types of ascocarpic development, one in which the ascocarp
evolves directly from the spermogonial stage and the other in which
ascocarps occur on areas of the needle devoid of spermogonia. The best
illustrated and most comprehensive investigation of ascocarpic centrum
ontogeny of species of Hypodermataceae is that of Gordon (1966, 1968).
Two basic types of centrum ontogeny were described in which ascal
initiation within Type I centrum (Gordon, 1966) occurs in the basal cells
of the pseudoparaphyses and involves anastomoses, while ascal initiation
within Type II occurs in cells of a plectenchymatous centrum, with no
visible anastomosing in the ascocarp. The Type III centrum (Gordon,
1968) has a layer of hyaline cells in the primordium that is meristematic
and gives rise to all subsequent structures of the centrum.
Wehmeyer (1966) reported that fern fronds infected with Cryptomy-
cina produced pycnidia and ascostromata in which the archicarps and
emergent trichogynes are formed in a central meristem but later degen-
erate. Ascogeneous hyphae arise from intact or fragmented strips of the
central "meristem." Asci develop from these strips without producing
croziers. Ascocarps mature on overwintering fronds and mature ascospores
are produced in early spring. Killian (1918) contrasted the elongate
ascogonia of Cryptomyces with those of other discomycetes. Helical
ascogonia with trichogynes were reported in Coccomyces (Higgins, 1914)
and Phacidium (Satina, 1921). Studies of ascal evolution in Rhytisma
were reported by Aragno (1967). Ascogonia elongate while in the interior
of the plectenchymous stroma numerous apothecial cavities develop.
Septa between ascogonial cells break down and pairs of nuclei migrate
into the ascogenous hyphae.
The ontogeny of the apothecia of Pyronema has been studied perhaps
more than that of any other operculate discomycete. Copulation of the
antheridium and trichogyne was seen by DeBary (1863), but the sexual
nature was not established. The Tulasne brothers (1865) initially found
no such copulation, but later (1866) reported it. Kihlman (1883) reported
a connecting pore between the antheridium and the trichogyne but did
not consider them functional reproductive structures. Others (Harper,
1900; Claussen, 1912; and Gwynne-Vaughan and Williamson, 1931)
believed that male nuclei passed through the trichogyne into the ascogo-
nium, but Dangeard (1907) and Moreau and Moreau (1930) felt that
no fertilization occurred. Harper (1900) and Gwynne-Vaughan and
Williamson (1931) reported brachymeiosis, with nuclear fusion occurring
in the ascogonium followed by a second fusion in the young ascus. Hirsch
(1950) and McIntosh (1954) presented cytological evidence to refute the
theory of double fusion. Moore (1963) showed that a single hyphal strand
produces a cluster of antheridia and ascogonia in P. domesticum (Sow.
ex Gray) Sacc. A sterile system of hyphae arises from the stalk cells
supporting the ascogonia and antheridia. All elements proliferate to
give apothecial expansion. Corner (1930b) placed Pyronema among
those with a gymnocarpic development.
Because of the lack of an excipulum, species of Ascodesmis seem to be
very suitable for cytological work. Van Tieghem (1876) and Zukal (1886)
did not observe sexual stages. Dangeard (1903, 19'07) first saw pairs of
coiled hyphae growing out of short side branches but claimed that there
was no fusion between these structures, the only sexual fusion of nuclei
taking place in the young asci. Claussen (1905) and Swingle (1934), how-
ever, observed nuclear fusion both between the coiled hyphal pairs and
in the young asci. Since all cells are multinucleate and the coiled hyphal
pairs occur in dense clusters in which cell walls seem to disintegrate, this
problem is yet to be solved (Obrist, 1961). Only scant mycelium remains
associated with the asci, placing these among species with a gymnocarpic
development (Corner, 1930b). A vast amount of work has been published
on other members of the Ascobolaceae. Van Brummelen (1967), in his
monograph of Ascobolus and Saccobolus, summarized many of the cyto-
logical studies performed on these genera. The ease of handling and
availability of specimens during most seasons of the year has resulted in
large amounts of cytological evidence. He lists the earlier work of Dodge
(1912), Lohwag (1927), Kniep (1928), G/iumann and Dodge (1928), Moreau
and Moreau (1930), Gwynne-Vaughan and Williamson (1932), Drayton
(1932), Dodge (1936), Dodge and Seaver (1946), Whitehouse (1949),
Langeron (1952), and Olive (1954). Modern research mentioned was that
of Bistis (1956a, 1956b, 1957), Bistis and Raper (1963), Esser and Kuenen
(1965), and Esser (1967). In both genera the ascogonium is usually a few
108 T H E BOTANICAL REVIEW
to many-celled coil. An antheridial element may be present or absent.

G~iumann (1964) states that in this group plasmogamy leading to fertiliza-
tion may occur through (1) spermatization, (2) parthenogamy, (3) autog-
amy, (4) somatogamy, or (5) apomixis.
There appears to be a variation in the type of ascocarp development
after fertilization. Corner (1930b) listed species of Ascobolus in each of
the hemiangiocarpic, angiocarpic, and gymnocarpic groups. Van Brum-
melen (1967) uses two sets of terms in describing the development of
apothecia. Chronological phases were distinguished as (1) archihymenial
phase: before the initials of the hymenium are formed; (2) prohymenial
phase: paraphyses are present but no croziers are as yet formed; (3)
mesohymenial phase: the hymenium is in the process of developing but
no mature asci are present; (4) telohymenial phase: mature asci are
present and ascospores are normally discharged; and (5) posthymenial
phase: the hymenium becomes overripe or obsolete and decomposes. With
regard to the hymenial development, two main types of ascomata were
distinguished: (1) cleistohymenial ascoma: the hymenium is enclosed, at
least through early development. This phase may be further divided into
pro-meso telohymenial phases. (2) Gymnohymenial ascoma: the hy-
menium is exposed from the first until the maturation of asci; with two
subgroups, paragymnohymenial in which the ascogonium is over-arched
by unfused hyphae, and eugymnohymenial in which the hymenium is
exposed early but is soon covered with an excipulum or sheath (Fig. 2).
Among the Thelebolaceae, ascocarp initiation has been studied (Dan-
geard, 1907) in Coprotus (Ascophanus) ochraceus (Cr. and Cr.) Kimbr.
On a single hypha there arises a small group of up to 15 stipitate, swollen
unicellular ascogonia, each of which is provided with an occasionally
branched trichogyne. Antheridia are not formed, the female nuclei pair
within one ascogonial cell and migrate into the ascogenous hyphae. A
similar finding was made by Delitsch (1926) in Lasiobolus pulcherrimus
(Cr.) Schroet., in which species nuclear pairing takes place in only one
cell of the ascogonium. In Thelebolus (Ramlow, 1906) and TrichoboIus
(Kimbrough, 1966a), one of the ascogonial cells becomes the privileged
cell, i,e. bears a single pair of compatible nuclei. Karyogamy occurs and
this cell becomes the ascus mother cell. Meiosis, followed by a series of
mitoses, results in an ascus with several thousand spores. A number of
species of Thelebolus (Ryparobius sensu Boudier) have been studied in
culture (Barker, 1904; Dangeard, 1907; Ramlow, 1915). All asci of the
ascocarp are initiated from a single ascogonial coil, and the number of
asci appear to be nutritionally controlled. Bergman and Shanor (1957)
showed a similar pattern for Thelebolus psychrophila (Berg. and Shanor)
Eckbl. According to Corner (1903b) species of Thelebolus (including
Ryparobius) have an angiocarpic development. This was confirmed for
Trichobolus, a setose segregate of Thelebolus, by Kimbrough (1966a).
Apothecial ontogeny in the Pezizaceae has been studied for a number
of species. In Iodophanus carneus (Pers. ex Fr.) Korf, each ascogonial coil
has a privileged cell from which the ascogenous hyphae arise (Cutting,
KIMBROUGH" C L A S S I F I C A T I O N O F D1SCOMYCETES 109
1909). Branching of stalk cells and adjacent mycelium soon enclose the
young ascogenous system. Corner (1930b) placed it among those species
with an angiocarpic development. In Van Brummelen's (1967) scheme
it would have a cleistohymenial ascoma opening during the telohymenial
phase. An Oedocephalurn conidial stage has been reported for species
of Iodophanus by Korf (1958) and Gamundi and Ranalli (1964). Overton
(1906) found that apothecia of Thecotheus pelletieri (Cr. and Cr.) Boud.
developed from several ascogonial coils. The ascogenous hyphae arose
from any or all cells of the ascogonium, and asci arose from subterminal,
binucleate cells. Peziza spp. have been cultured by Dodge (1937), Fergus
(1960), Berthet (1964a), Webster et al. (1964), and Paden (1967). Fraser
and Welsford (19.08) observed sexual reproduction in which ascogenous
hyphae were derived from somatic cells. Similar results were reported by
Schultz (1927) for Peziza adae Sadd. and by Gwynne-Vaughan (1922) for
P. tectoria Cke. All produced an Oedocephalurn, Ostracoderma, or
Rhino trichum-like conidial stage. Wolf (1958) described another Ostra-
coderma as Rhinotrichum trachycarpa, the conidial stage of Plicaria
trachycarpa (Curr.) Boud.
A number of the Humariaceae (sensu Le Gal) have been studied in
culture. Dodge (1922) found a Botrytis-like conidial stage in a species
of Lachnea (Trichophaea). Sexual structures were not reported. Fraser
(1913) reported that the archicarps of Trichophaea abundans Boud. con-
sisted of a long, eight- to nine-celled trichogyne, and projecting beyond
the hyphal tangle, a three- to four-celled, helical ascogonium with a multi-
cellular stipe. Webster et al. (1964) induced a Botrytis stage on cultures.
Whitney and Parmeter (1964) were able to produce apothecia of T. bullata
Kan. by overlaying cultures of Rhizoctonia hiemalis Sak. and Voar. with
sterile sand and placing outside for several weeks. Apothecial ontogeny
was not followed. An unidentified conidial stage was reported for T.
confusa (Cke.) Berthet (Berthet, 1966). In Leucoscypha rutilans (Fr.)
Sckbl. the apothecium arises as a dense weft of tangled filaments, which
for a time differ from one another only in the relatively thick walls of
the outer hyphae, and the richer protoplasmic content of the inner
(Gwynne-Vaughan, 1922). Nuclei of two sizes were seen later in the central
part of the mass. Sexual organs are completely lacking and fertilization
is accomplished through fusion of nuclei of somatic hyphae. Anthracobia
melaloma (Albt. and Schw. ex Pers.) Boud. is a heterothallic species in
which archicarps are produced by both mating types when they are grown
separately (Rosinski, 1956). Antheridia cannot be distinguished except
when they are in contact with a trichogyne. Plasmogamy follows the
coiling of the tips of the trichogyne around an antheridial hypha. Asco-
genous hyphae develop from 2, 3, or 4 ascogonial cells. The hyphae giving
rise to the sterile portion of the apothecium develop from the stalk cells
of the archicarp. Similar findings were reported for A. nitida Boud. by
Durand (1968). A cytological study of ascus development in A. melaloma
was made by Olive (1950). The results contradicted those of Gwynne-
Vaughan in showing that there is no brachymeiotic division in the ascus.
There were four pairs of chromosomes in the primary nucleus of the

ascus, and four chromosomes pass to each spindle pole in all three nuclear
divisions. Corner (1930b) places A. melaloma with the hemiangiocarpic
forms. Cheilymenia stercorea (Wigg. ex Gray) Boud. produces pyriform
ascogonia with a long multicelled trichogyne as a branch which is sep-
arated from the main hyphae by several stipe cells (Fraser, 1907; Fraser
and Brooks, 1909). Antheridia and trichogynes appeared nonfunctional;
ascogonial nuclei, after a single division, pair autogamously and form
ascogenous hyphae. In Cheilymenia theleboloides (Alb. and Schw. ex
Pers.) Boud. there is a well-marked ascogonial region of one or more cells
(Gwynne-Vaughan, 1922). Corner (1930b) cited C. stercorea among the
operculates with hemiangiocarpic apothecia.
The archicarp of Tricharia cretea (Cke.) Boud. consists of a long,
branched, multicellular trichogyne, an ascogonial region of three to four
coenocytic cells, and a multicellular stalk (Gwynne-Vaughan, 1922). No
antheridia were observed. In the central part of the archicarp the trans-
verse septa are completely broken down, so that a very wide passage is
formed; and nuclei may pass readily from cell to cell. All cells give rise
to ascogenous hyphae. In Scutellinia scutellata (L. ex. Ft.) Lamb. the
archicarp consists of seven to nine cells, the subterminal of which enlarges
to form the ascogonium (Brown, 1911). The nuclei in this cell divide,
and show five short, stout chromosomes. Nuclear fusion was not observed,
only pairs of nuclei in the ascogonium. Large ascogenous hyphae de-
velop, undergo septation, and branch freely. Much to the disturbance of
Gwynne-Vaughan (1922), Brown inferred the occurrence of a single fusion
in the ascus of S. scutellata, followed by a single reduction. In Coprobia
granulata (Qu4l.) Boud. the archicarp develops as a side branch from an
ordinary hypha. The apical cell of this branch increases in size and
becomes spherical, forming the ascogonium (Blackman and Fraser, 1906).
This was one of the first reports of brachymeiosis in this group of fungi.
There were no signs of antheridia or trichogynes. Vegetative cells grow
up and invest the archicarp, forming a close pseudoparenchymatous
sheath in which the ascogenous hyphae ramify. Fusion was reported in
both the ascogonium and ascus mother cell. Corner (1930b) cited C.
granulata among those Pezizales with an angiocarpic development. Mar-
celleina atroviolacea (Del. ex de Sey.) Brumm., = Lamprospora planchonis
(Dun.) Seav., produced variable colonies when grown in culture from
germinating ascospores (Hanlin, 1965). Numerous attempts were made
to induce the formation of apothecia and conidia by growing the fungus
on various media and under different temperatures and light conditions,
but sexual or asexual reproduction was not observed. Reports of conidial
stages in cultures of Lamprospora (Wolf, 1958; Hanlin, 1965) have been
shown to be of species currently placed in the Pezizaceae under Marcel-
leina or Plicaria.
In d leuria aurantia (Pers. ex Hook.) Fckl., Fraser and Welsford (1908)
noted that during the early stages of development a large cell, no doubt
part of an archicarp, was present. Berthet (1964a) was unable to find an
KIMBROUGH: C L A S S I F I C A T I O N OF DISCOMYCETES lll
imperfect stage for this species. Ascogonia of Sphaerospora minuta Cain
appear as side branches on the aerial mycelium when the culture is two
or three days old. Sometimes they become spiraled rather regularly with
up to six coils, the lower one being much broader (Cain and Hastings,
1956). Transverse septa are formed at irregular intervals and branches
develop from the basal cells of the coil. These grow upward to soon
enclose the ascogonial coil. The branches enclosing the ascogonium
develop shorter cells to form a pseudoparenchyma which develops into
the excipulum that continues to enclose the hymenium until late in its
maturation. Conidia are produced in Botrytis-like clusters on long dicho-
tomously branched conidiophores. As was pointed out by Cain and
Hastings (1956) these conidial stages are not congeneric with Botrytis
cinerea Per. ex Fr., the type species of the genus. A similar conidial stage
is found in Sphaerosporella brunnea (Alb. and Schw. ex Fr.) Svrfek and
Kub.( Rifai, 19'68) and S. hinnulea (Berk and Br.) Rifai (Wolf, 1963).
Both are morphologically similar to that produced by Trichophaea
abundans (Karst.) Boud. In Inermisia fusispora (Berk.) Rifai (= Humaria
sensu Sacc.) there is a well marked ascogonial region of one or more
large cells (Gwynne-Vaughan, 1922).
Within the Helvellaceae development has been studied in species of
Helvella, where apothecia arise as tufts of branching, septate hyphae,
without morphologically distinguishable ascogonia (Gwynne-Vaughan,
1922). Carruthers (1911) studied nuclear fusion in young ascogenous
hyphae of H. crispa Scop. ex Fr. and found four chromosomes. In young
apothecia of H. elastica Bull. small, thick hyphal tangles form, gradually
differentiate into stipe and pileus (McCubbin, 1910). In the Helvellaceae
the number of nuclei in the ascospores is invariably four (Berthet, 1964a).
For the Morchellaceae, Greis (19'40) showed somatic copulation in Mor-
chella elata Fr. and Morchella conica Pers. Berthet (1964a) was unable
to find conidial stages in 4 species of Morchella. In the Morchellaceae
the number of nuclei in spores is variable but very high, not less than
15 and up to 60. Five species representing the genera Discoitis, Mitro-
phora, Morchella, and Verpa were studied by Berthet (1964b).
In Rhizina undulata Ft. (Rhizinaceae, Eckblad, 1968; Helvellaceae,
Rifai, 1968) a long multicellular archicarp similar to those of the Asco-
bolaceae were found by Fitzpatrick (1918). He regarded the terminal
cells as a trichogyne but there was no evidence that antheridia existed
or that fertilization ever occurred. They were multinucleate and appeared
to arise from vegetative hyphae. The central cells eventually gave rise
to ascogenous hyphae. He observed paired nuclei in the ascogonial cells
and in the ascogenous hyphae. He inferred from this that nuclear fusion
did not occur until later in the young asci.
A number of the Sarcoscyphaceae have been successfully grown in
culture. Of those, only 3, Desmazierella acicola Lib., Sarcoscypha coccinea
(Jacq. ex Gray) Lamb., and Urnula craterium (Schw.) Fr. have been
induced to sporulate. Gremmen (1949), Hughes (1951), and Berthet
(1964b) obtained Verticicladiura trifidum Pr. from ascospore cultures of
D. acicola Lib. Alexopoulos and Butler (1949) and Rosinski (1953) dis-
covered a conidial stage for S. coccinea but did not suggest a name for it.
Mycelial cultures of U. craterium originating from ascospores were planted
on decaying branches of oak that had been sterilized. After approximately
6 months, black heaps of conidia which appeared to be Strumella cory-
noidea Sacc. were formed (Wolf, 1959). The relationship between
Strumella and Urnula on oaks were suggested earlier by Davidson (1950).
Berthet (1964a, 1964b) was unable to demonstrate conidial stages for
species of Phillipsia, Pithya, Plectania, Pseudoplectania, and Sarcosoma.
It seems that no one has been able to induce apothecia in culture or
study the ontogeny of the ascogenous system in these groups.
Several attempts have been made to utilize the results on sexuality
for taxonomic purposes, but so far with small success. G/iumann (1964)
summarized this area by stating that within one and the same family one
can usually find the whole range of different types of sexuality rep-
resented, from functional ascogonia and antheridia through absence of,
or nonfunctional antheridia, and eventually to the absence of both
antheridia and ascogonia.
Spermatization appears to be more frequent among inoperculates, hav-
ing been reported in families of both the Helotiales and the Phacidiales.
That this type of fertilization is not restricted to the inoperculates is
evident from studies of sexuality of the Ascobolaceae. It would not be
surprising to find that both microconidial (spermatia) and macroconidial
forms reported for other Pezizales may act in fertilization. As was pointed
out by Eckblad (1968), there is a tendency for somatogamy to be more
common among the "helvelloid" families of the Pezizales. Little tax-
onomic information can be drawn from the shape and size of ascogonia
and antheridia. There appears to be extreme variations, even among
species of the same genus. The presence or absence of antheridia is also
an inconstant feature of species of a genus. Berthet (1964b) summarized
some of the present knowledge of the occurrence of homothallism and
heterothallism among the discomycetes and concluded that as far as can
be judged from meager data available, the distribution of the two types
of sexual reproduction is of no taxonomic importance. Olive (1954)
pointed out that many investigations indicate that the sexual process in
the fungi is much more labile and variable than was presumed by early
investigators. Heterokaryosis, heterozygosity, and segregation may occur
freely in homothallic as well as heterothallic fungi.
Excipulum. Aside from the ascogenous hyphae and resulting hyme-
nium, the remainder, and sterile part of the apothecium, is termed
excipulum. The ectal excipulum forms the major part of the margin and
outer layers, while within these layers is the medullary or ental excipulum.
Corner (1930a, 1930b) described in detail three patterns of growth of the
excipulum in ratio to hymenial growth. Van Brummelen (1967) proposed
a scheme of seven developmental types, one of which may be characteristic
of any specific discomycete. St/irback (1895) developed a terminology for
describing the tissues of the sterile parts of the apothecium. This system
was modified by Korf (1951) and has been illustrated by Korf (1958),
Van Brummelen (1967), and Eckblad (1968). Not every type of tissue
occurring within the operculate and inoperculate discomycetes is covered
by these terms. However, deviations rarely occur that would require
detailed descriptions beyond those of the 6 basic types.
The types of excipulum have been widely used in discomycete clas-
sification. Nannfeldt (1932) relied heavily on the nature of excipular
cells in delimiting the Orbiliaceae, Dermateaceae, and Hyaloscyphaceae.
He also recognized nine subfamilies of the Helotiaceae in which the
excipulum was one of the most distinguishing features of most of the
taxa. Specialists in the Pezizales have tended to place much more emphasis
on asci and ascospores, with excipular characters playing a secondary role
in the delimiting of taxa at the family level or above. These features
have been used to a greater extent at the generic level, especially in
delimiting certain genera of the Sarcoscyphaceae (especially Sarcoscypha,
Aurophora, and Phillipsia), and the Humariaceae (especially Octospora,
Inermisia, Pulvinula, and Lamprospora). A large number of discomycetes
possess apothecia clothed with hairs. Eckblad (1968) defined hairs as
modified hyphae differing from ordinary hyphae in one or more of the
following ways: either in form or by having thicker walls, or by being
colored, or by being of conspicuous length. He recognized the following
types of hairs in the Pezizales: (1) short, blunt, colored, hyphoid, often
fascicled hairs (characteristic of Anthracobia and Melastiza); (2) hyaline,
septate, flexuous, hyphoid hairs (as in Leucoscypha and Sarcoscypha);
(3) colored, flexuous, septate, hyphoid hairs (as in Sepultaria, Pseudo-
plectania, and Urnula); (4) straight, hyaline, acute, non-septate bristly
hairs (Lasiobolus); (5) straight, colored, acute, septate, bristly hairs (Des-
mazierella, Sphaerosporella, Humaria, and Tricharina); (6) straight,
colored, acute, septate, bristly hairs with forked or rooting base (Cheily-
menia, Scutellinia); (7) colored, acute, septate, bristly hairs with a stellate
apical part (Cheilymenia). Many of these types of hairs may be found
among the approximately twenty genera of Hyaloscyphaceae (Dennis,
1949). His tribe Lachneae contained genera almost invariably clothed
with rather long, septate, rough hairs, while those of the tribes Hya-
loscypheae and Arachnopezizae were clothed with hairs of diverse shapes.
Berthet (1964b) studied the number of nuclei in mature spores, para-
physes and mycelium of a large number of discomycetes. Some of his
findings will be discussed under the respective taxa in part III.
Asci. Asci of the discomycetes exhibit a great variation. The majority
of genera and species have asci that are cylindrical, straight, narrowed
at the base, rounded at the top, and with a hyaline, unitunicate wall.
Truly, there are many species of Phacidiales, and perhaps Helotiales and
Pezizales which, when examined critically, will prove to have asci with
two or more layers. Several authors have indicated that the so-called
unitunicate wall is actually two layered (G~iumann, 1964; Kimbrough,
1966b; Van Brummelen, 1967), but the wall is never bitunicate in the
sense that the two layers separate at spore discharge. It appears that in
A B C
O E F
FIG. 1.
l
D i a g r a m m a t i c d r a w i n g s of the apical structure of asci. A. A n 8-spored species of
Lasiobolus s h o w i n g a sharply i n d e n t e d area at o p e r c u l u m . B. A m u l t i s p o r e d
Lasiobolus with a t h i n n e r walled area at t h e p o i n t of dehiscence. C. A n ascus
of Trichobolus s h o w i n g a m u c h b r o a d e r t h i n area at t h e apex. D. A n ascus of
a species of Thelebolus with a swelling in t h e o u t e r wall layer. E. A n ascus of
Ascozonus w i t h a n extremely thin-walled, n i p p l e d tip. F. A y o u n g ascus of
Caccobius s h o w i n g a b r o a d core in t h e apex. G. A m a t u r e ascus of caccobius
with t h i n area of dehiscence. H. A y o u n g ascus of Cyttaria gunnii in Meltzer's
reagent. I. A y o u n g ascus of C. gunnii in lactophenol-cotton blue. J. A y o u n g
ascus of C. gunnii in Congo red with 2 percent K O H s h o w i n g apical a p p a r a t u s .
K. Apical a p p a r a t u s of C. gunnii in Congo red with 2 percent K O H .
the coprophilous genera Thelebolus, Trichobolus, Coprotus, and perhaps
Lasiobolus, in which there are both 8-spored and multispored species,
there have been a number of modifications in the ascus to accompany
the enormous number of spores produced. Aside from having fewer asci
per ascocarp, the multispored species have additional ascal wall layers.
In Trichobolus zukalii (Heimerl) Kimbr. and Thelebolus stercoreus Tode
ex Fr. as many as five layers can be distinquished with proper staining
(Kimbrough, 1966a, 1966b).
Preliminary evidence indicates that the shape of the ascus, influenced
by the number of asci per apothecium and the number of spores per ascus,
may, along with the basic wall structure, determine whether the apical
apparatus functions as an operculum or an irregular tear. T h e 8-spored
members of Lasiobolus and Coprotus have a sharply indented area in the
inner wall layer marking the point of the operculum (Fig. 1A). In multi-
spored members of these genera, L. caini Kimbr. and C. rhyparobioides
(Heimerl) Kimbr., in which the ascus apex is much broader, the indented
area is much broader (Fig. 1B) and the inner wall layer is much thicker
around the sides and base. In Trichobolus zukaIii, with a single, large,
ovoid to pyriform ascus, the ascus apex is broadly rounded above and
all of the ascus wall layers become abruptly thinner near the apex. T h e
weakness to allow spore liberation is spread over a larger area. Thus,
only on rare occasions will the apex function as an operculum but most
often as an irregular tear. T h e lateral walls are more than 5 /~ thick
and appear stratified, while in the apex they are slightly more than 2/~.
Further modifications can be noted in the related genus Thelebolus in
which T. microsporus (Berk. and Br.) Kimbr. (in Kobayasi, 1967), an
8-spored species, the ascus wall is clearly observed to be two layered when
stained in Congo red or acid fuchsin. The outer layer has a slight inward
thickening near the apex. In T. crustaceus (Fckl.) Kimbr. (in Kobayasi,
1967), a 64-spored member, the outer wall thickening is more pronounced
and a ring is often evident, especially when stained with Congo red (Fig.
1D). In T. stercoreus Tode ex Fr., a uniascal species with more than 2,000
spores, the apical ring is very apparent without staining. T h e increase
in spore number per ascus in these taxa is also accompanied by an increase
in ascal wall thickness. The inner wall, as in Trichobolus, appears to be
sublayered. In all species of Thelebolus observed, the wall layers of the
ascus apex are much thinner than those below the ring. With this par-
ticular apical apparatus, dehiscence is usually irregular, but occasionally
in 8-spored forms as an operculum.
Further modifications may be seen in the asci of Ascozonus in which
the apical ring is more pronounced than in Thelebolus. It is similar to
the latter genus in that the ring is due to a sharp inward thickening of
the outer wall layer. Unlike Thelebolus, however, the outer wall layer
remains relatively thick almost to the tip of the ascus, at which point,
over a nippled tip, it is almost indistinguishable (Fig. 1E). In Caccobius
minusculus Kimbr. the ascus wall is of uniform thickness, with a 15-20/~
plug in the apex. Although the young asci of Caccobius are similar to
those of Thelebolus (Fig. 1F), in the process of maturation an obvious

core or plug of a different chemical nature develops in the apex. At
maturity spores are liberated through a broad irregular tear in the ascus
apex. As was pointed out by Eckblad (1968), these members of the
Thelebolaceae may be the living representatives of the common origin of
the discomycetes. Truly, Caccobius, Thelebolus, Trichobolus, Ascozonus
(non-operculate forms) are related to Lasiobolus and Coprotus (oper-
culates). Hopefully, we can see that minor changes in wall configuration,
plus broadening and stretching of the apex with increased spores, may
alter the manner of dehiscence even though their basic structure is quite
similar.
For the major groups of operculate or inoperculate discomycetes, the
most comprehensive and stimulating work on ascus structure has been
that of Chadefaud (1940-1969) and Le Gal (1946a, 1946b). Details of
the apical structure have been studied for the following genera of
operculate discomycetes: Peziza, Bulgaria (Chadefaud, 1940a, 1940b),
Ascobolus, Lachnea, Coprobia, Ciliaria, Sepultaria, Helvella, Pulvinia,
Humaria (Chadefaud, 1942), Aleuria, Phaedropezia, Trichophaea (Le
Gal, 1953a), Pustularia (Chadefaud, 1943), Morchella (Chadefaud, 1949);
the suboperculates: Leotia (Chadefaud, 1944), Sarcoscypha (Chadefaud,
1946; Le Gal, 1966), Pseudoplectania, Melascypha, Pithya, Urnula, Sar-
cosoma, Wynnea, Phillipsia (Le Gal, 1946b); and the inoperculates:
Microglossum, Mniaecia, Mollisia, Dasyscyphus (Chadefaud, 1942), Rut-
stroemia, Pachydisca, Helotium, and Hyaloscypha (Le Gal, 1953a). Ascus
structure of the Ostropales was studied by Bellbmere (1960). Chadefaud
(1942) divided the asci of the discomycetes into two broad groups: (1)
Those with a broad apical pad with a well formed, deep and narrow pore,
surrounded by an apical ring, and (2) those with a thin, flat pad with
the pores or depressions reduced to a dent from which the lining or
sheath forms. Among those with a long, slender pore surrounded by a
ring, he separated those with an amyloid ring (inoperculate discomycetes
and certain groups of Sphaeriales) from those with a non-amyloid ring
(other Pyrenomycetes). The operculates were placed among the latter,
and were distinguished from other members of this group in having a
long, funnelform strand that penetrates more or less deeply into the ascus
region occupied by ascospores and attaches to the apical apparatus.
In Chadefaud's (1967) discussion of the phylogenetic origin and evolu-
tion of the ascomycetes from lichen fungi, he placed great emphasis on
the structure of asci. Details of ascal anatomy were given for approxi-
mately 20 lichen genera. He concluded that the discolichens were of
polyphyletic origin, having evolved from very peculiar ancestral types.
He considered the Graphidales to be more primitive than the Lecanorales.
He suggested also that at least part of the fungi of the pyrenolichens
belong perhaps to the same group of discomycetes, despite other aspects
of their fructifications.
Another aspect of discomycete taxonomy was introduced by Chadefaud
(1969) in his remarks on the walls, apical structure, and nutritive reserves
of asci. Chadefaud concluded that due to parallel evolution the same
types of apical structures may exist independently in various groups of
ascomycetes. It was difficult when using a single set of characters to
distinguish inoperculate asci from those of the ascohymenial pyrenomy-
cetes. Correlation of several characteristics was necessary to separate the
groups. He found that the nutritive reserves of the epiplasm and the
ascospores, especially the lipid-glycogene ratio, appeared more or less
characteristic in each group. In the discomycetes as well as the pyrenomy-
cetes, the melanospore type contained proportionately more reserves than
the allantospore types.
Ascospores. One of the best accounts of the influence of ascospore
characteristics on the classification of discomycetes is that of Eckblad
(1968). The shape, color, surface structure, contents, and nuclear con-
dition of ascospores have been of immense importance in the taxonomy
of discomycetes. Eckblad's comparison of the ascospores of operculate
and inoperculate discomycetes are summarized on the following points:
(1) Cell number: spores of operculates are permanently one-celled; inoper-
culates, one, two to many-celled; (2) Form: extremely variable but spores
of operculates appear bipolar-symmetrical; those of the inoperculate have
the distinct feature of being more or less unequally sided, while acicular
or filiform spores do not occur in the operculates; (3) Size: spores of
operculates are generally larger than those of inoperculates, mainly 10 /~
or longer, whereas those of the latter are usually less than 15/~; (4) Color:
outside of the Ascobolaceae, pigmented spores are rare in the operculates,
while in the inoperculates a number of genera with pigmented spores
occur; (5) Contents: oil droplets appear to be of more taxonomic value
in the operculates, while the occurrence of a gaseous phase within asco-
spores (DeBary bubbles) have had a limited use in the delimiting of
genera; (6) Ornamentation: Le Gal's (1947) detailed study of formation
and chemical nature of spore ornamentation is summarized. Five types
were briefly discussed; false ornamentation, ornaments of vacuolar origin,
those of vacuolar and perisporic origin, simple non-callose pectic, and
simple callose pectic ornaments. Eckblad concluded that in no case is
it obvious that a genus should be separated on spore form alone, nor
on the presence or absence of ornaments. However, the type of ornamen-
tation appears extremely significant at the generic and perhaps family
level.
Berthet (1964b) was able to differentiate four main groups of species
by the number of nuclei per ascospore. (1) Spores of the Sarcoscyphaceae
become plurinucleate, the number ranges from 2 to 4 in Pithya, to about
25 in Urnula craterium. (2) The ascospores of Helvellaceae are always
tetranucleate and it appeared that this unique feature is associated with
other features of taxonomic significance such as habit, habitat, and
pigmentation of the apothecia. (3) Nuclei in spores of the Morchellaceae
ranged from 15 to 60 in the 5 species studied. (4) With minor exceptions,
the remaining operculate and inoperculate discomycetes possess uni-
nucleate ascospores. Binucleate spores were reported for species of Scutel-
FIc. 2. Scheme of the developmental types in Ascobolaceae. The gametophytic system

in black, the sporophytic system darker (after Brummelen, 1967).
linia, and both binucleate and plurinucleate spores in members of the

Sclerotiniaceae.
Paraphyses. Paraphyses belong to the monokaryotic hyphal system
and, according to Corner (1930b) through monopodial branching grow
upward from the hypothecium. They are extremely variable in shape,
but usually filiform to clavate, sometimes lanceshaped, pyriform curved,
hooked or branched, and most always septate. Shape, size, and color of
paraphyses have been extremely useful in distinguishing taxa, especially
at the species level. A particular feature of paraphyses is often a constant
feature of certain genera, i.e. extremely clavate in Orbilia, uncinate in
Pulvinia, broadly lanceotate in most of Dasyscyphus, and greening in
iodine in several genera of the Aleuriaceae sensu Arpin. Hymenial color
is almost always located in the paraphyses.
In the area of pigmentation, Arpin (1968) studied the pigment system
of almost 100 species of discomycetes. The presence or absence, and kinds,
of carotenoids in 67 members of the Pezizales and 19 of the Helotiales
was studied, followed by a discussion of their biochemistry and the influ-
ence on taxonomy and phylogeny of the discomycetes. From the tax-
onomic point of view, the study of carotenoids allowed him to erect a
new family, the Aleuriaceae, corresponding to the Ciliarieae and Huma-
rieae sensu Le Gal. Pigment analysis also justified the tribes Urnuleae
and Sarcoscypheae of the Sarcoscyphaceae. Carotenoids were not found
in the tribe Urnuleae, nor in the Morchellaceae, Helvellaceae, Pezizaceae,
and Otideaceae. Arpin considered the species more evolved chemically
as the carotenoids which characterize it are more oxydized. Among the
Hyaloscyphaceae, fl-carotene seemed to be responsible for coloration.
Carotenoids were also found in members of the Helotiaceae and Geoglos-
saceae, but not in the Sclerotiniaceae.
Apothecial Shape. The shape of apothecia has frequently been used
in the past as the sole basis for dividing families of discomycetes. Tradi-
tionally the Geoglossaceae were distinguished from other inoperculates
by being stipitate instead of pezizoid. The Helvellaceae and Morchel-
laceae were similarly segregated from pezizoid operculates. Earlier treat-
ments of the Geoglossaceae had placed prime importance on the gross shape
of the ascocarp. The pileate genera were brought together by Durand
(1908) and Mains (1956b) as the tribe Cudonieae, and by Imai (1941) as
the subfamily Cudonioideae. Imai (1956) placed some of these (Leotia
and Neocudoniella) in the subfamily Leotioideae. Korf (1958) found,
however, that only the stipitate-pileate character of the ascocarp clearly
distinguished the Leotioideae from the Ombrophiloideae (Helotiaceae)
and proposed transferring the subfamily Leotioideae to the Helotiaceae.
It appears difficult to draw family lines in these groups between pileate
and cupulate forms.
Nannfeldt (1932) found a similar situation at the generic level in the
Helvellaceae. Nannfeldt (1937) united a number of helvelloid and pezi-
zoid genera under Helvella, using anatomical characters to delimit genera.
Korf (1956) demonstrated that the genera Daleomyces, Durandiomyces,
and Napomyces were based on sparassoid forms of the same species. Burd-
sall (1968) showed that Sepultaria (Pezizales) and Geopora (Tuberales)
were anatomically similar, differing essentially only in superficial mor-
phology. Eckblad (1968) concluded that in the Pezizales the form of
ascocarp appears to have a very low taxonomic value. On the generic
level it should be used with great care, and then only when the difference
in form is correlated with differences in other characters. I feel these
same conclusions could be applied to the Helotiales, Phacidiales, and
Ostropales.
Ultrastructure of Discomycetes. The fine structure of various aspects
of several discomycetes has been described. These include the type of
mycelial septation, crozier formation, development of young asci, meiosis
in the ascus, delimitation of ascospores, and many other extremely interest-
ing aspects of ascosporogenesis. Comprehensive reviews of the literature
on fine structure of discomycetes may be found in Reeves (1967) and
Carroll (1969). Presently, little or no emphasis has been given to uhra-
structure in solving taxonomic problems. Many questions on the structure
and development of discomycetes that have been raised in light microscope
studies may now be answered. These may aid us greatly in showing
natural relationships between taxa at both higher and lower levels. T h e
use of freeze etch or scanning microscopes to show details of spore orna-
mentations may be shown to be useful in resolving problems at the species
or subspecies level. To arrive at a truly natural system of classification
we must know not only the structure and function of various organelles,
but the origin of such structures. This can best be done through ultra-
structure research.
T H E ORDERS AND FAMILIES OF DISCOMYCETES

T h e number of orders and families that are recognized in the dis-
comycetes differs from one system of classification to another. The fam-
ilies delimited by Boudier (1885, 1907), based mainly on gross morphology
of apothecia, were soon considered to be unnatural. However, this system
has formed the basis for most of the modern systems. Tables 1, 2, and 3
(taken from Boudier) illustrate the major categories of classification for
the inoperculate and operculate divisions down to genera. Table 5 briefly
outlines the orders and families of inoperculate discomycetes recognized
by some of the more prominent workers since Boudier (Nannfeldt, 1932;
Seaver, 1951; Dennis, 1968; and G~iumann, 1964). It is clear that those
of G~iumann (1964) and Dennis (1968), although founded on the superb
work of Nannfeldt (1932), reflects much of the recent work on these taxa.
In the discussion of families, I shall follow the framework of Dennis (1968)
in recognizing the inoperculate orders Helotiales, Ostropales, and Phacid-
iales. In Dennis' list of families, the Hemiphacidiaceae of Korf (1962a)
shall be added to the Helotiales. T h e Cyttariaceae is treated in a separate
order intermediate to the Helotiaceae and Sarcoscyphaceae.
To quote Korf (1969): " T h e year 1968 must surely go down in the
history of systematics of the discomycetes as one of the most significant
TABLE IV
FAMILIESOF OPERCULATEDISCOMYCETESRECOGNIZEDFROM 1885 TO THE pRESENT
Boudier (1885) Seaver(1928) Clements & Shear (1931) Le Gal (1947)

Morchellaceae Pezizaceae Pezizales (17 families) Operculates
Helvellaceae (Tribes) Morchellaceae
Pezizaceae Sphaerosporeae Operculates Helvellaceae
Humariaceae Ascoboleae Pezizaceae Aleuriaceae
Ascobolaceae Aleurieae Ascobolaceae Humariaceae
Pyronemaceae Humarieae Helvellaceae Ascobolaceae
Exoascaceae Lachneae
Otideae Inoperculates 8~Lichens Suboperculates
Sarcoscypheae (14 f a m i l i e s ) Sarcoscyphaceae
Elvelaceae
Nannfeldt (1949)
Pezizaceae
Sarcoscyphaceae
Korf (1954) Dennis (1968) Rifai (1968) Eckblad (1968)
Cyttariaceae Morchellaceae Sarcoscyphineae Sarcoscyphaceae
Sarcoscyphaceae Helvellaceae Sarcoscyphaceae Sarcoscypheae
Pezizaceae Pezizaceae Urnuleae Urnuleae
(Tribes) Humariaceae Sarcoscypheae Rhizinaceae
Pezizeae A s c o b o l a c e a e Pezizineae Morchellaceae
Ascoboleae 8arcoscyphaceae M o r c h e l l a c e a e Helvellaceae
Discineae Thelebolaceae Helvellaceae Otideaceae
Rhizineae Humariaceae Pezizaceae
Psilopezieae Otideae Pyronemataceae
Morchelleae Lachneae Ascobolaceae
Gyromitreae Ciliarieae Thelebolaceae
Acetabuleae : Aleurieae
Ciliarieae Pezizaceae
Octosporeae Ascobolaceae
Pyronemeae Pyronemataceae
Thelebolaceae
ever, with the appearance of three m a j o r works dealing primarily with

Operculate Discomycetes and the appearance of the second edition of
British C u p Fungi ( now more aptly titled)." T h e three m a j o r works in
reference were those of Rifai (1968) on the Australasian Pezizales, Eckblad
(1968) on the taxonomy, phylogeny, and nomenclature of the Pezizales,
and Arpin's (1968) chemotaxonomic essay on the carotinoides of Dis-
comycetes. These treatments reflect not only a surprising increase in the
n u m b e r of families of Pezizales, but a substantial reclassification of genera
within these families. T a b l e 4 briefly outlines the operculate families
recognized by Boudier (1907) and those of subsequent workers. I n the
discussion of families, the system proposed by Rifai (1968) will be followed,
with modifications reflecting the works of Eckblad (1968) and A r p i n
(1968). T h e r e appears to be a current consensus that all operculate dis-
comycetes be placed in the order Pezizales, although, the work of Burdsall
(1966, 1968) clearly points to a close relationship of the Tuberales.
KIMBROUGH" C L A S S I F I C A T I O N O F DISCOMYCETES 123
The Helotiales. The order Helotiales is accepted essentially in the
sense of Nannfeldt (1932), except that his Phacidiaceae is treated as a
separate order, and his subfamily Ciborioideae of the Helotiaceae is
essentially equivalent to the Sclerotiniaceae. The order is characterized
by ascocarps in which the hymenium is of indefinite extent, as in the
Ascocorticeaceae, or is borne in a superficial apothecium in which the
asci are inoperculate, and the ascospores are ellipsoid to needle-shaped.
Surprisingly, very little work has been done on the sexuality and develop-
ment of members of the Helotiales, considering the very large number of
species involved. Dennis (1968) listed 8 families, Caliciaceae, Ascocorticia-
ceae, Geoglossaceae, Orbiliaceae, Dermateaceae, Hyaloscyphaceae, Sclero-
tiniaceae, and Helotiaceae. To this we are adding the Hemiphacidaceae
of Korf (1962).
Ascocorticiaceae. The Ascocorticiaceae is recognized for the single
genus Ascocorticium, which forms definite mats of mycelium bearing asci
over the surface, and recalling various species of the basidiomycete genus
Corticium. There may even be clamp connections present, or as has been
suggested, proliferating ascogenous hyphae. The gray or pink colored
fructifications form a thin coating of indeterminate growth over the
surface of bark or bare wood of dead trees. The hypothecium consists
of 4-6 layers of close interwoven hyphae on which numerous ellipsoid,
8-spored, inoperculate asci are borne. Early reports questioned the pres-
ence of paraphyses. However, Oberwinkler et al. (1967) shows them clearly
for A. anomalum (Ell. and Ever.) Earle. Nannfeldt (1932) suggested that
the genus may be related to the Protomycetaceae, however, the presence
of paraphyses and the peculiar ascus apical structure (G~iumann, 1964)
makes this unlikely.
Geoglossaceae. The erect, clavate sporocarps of this group gives them
the common name of "earth tongues," and led to their being placed
among the Helvellaceae before the importance of ascus dehiscence was
known. The ascocarp is termed an apothecium even though it is in no
way the shape of a cup. One must review Corner's (1930a-1930b) work
to fully appreciate the variations that might exist within an apothecium.
The pileus-like cap bears the hymenium over most of the surface and
often far down the stipe. The asci are inoperculate and have been shown
in Microglossum (Chadefaud, 1942) to have a long slender pore sur-
rounded by an amyloid ring. Ascospores are elongate to filiform, uni-
cellular or with numerous transverse septa, and hyaline or colored. Eleven
genera and 41 spp. were recognized by Durand (1908), who published
additional notes later (Durand, 1921). Other important monographs are
those of Imai (1940) and Nannfeldt (1942). Contributions to our knowl-
edge of North American members of this family are those of Mains (1954,
1955, 1956a, 1956b); for other geographical areas there are those for
Estonia (Raitviir, 1961), Norway (Eckblad, 1963), Finland (Hakelier,
1964), India (Maas Geesteranus, 1965), Switzerland (Rahm, 1966), and
Nova Scotia (Grund and Harrison, 1967). Of the eleven genera first
recognized by Durand (1908), only 8 were considered by Dennis (1968).
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The genera Apostemidium and Vibrissea, treated by Durand and others
in this family, are now considered to belong to the Ostropales, and species
of Gloeoglossum are considered with Geoglossum (Nannfeldt, 1932).
Arpin (1968) considered the Geoglossaceae to have a heterogeneous pig-
ment system, capable of producing both carotenoids and water soluble
forms. The taxonomic position of Leotia has been in question. It has
generally been treated in the Geoglossaceae (Durand, 1908; Nannfeldt,
1932; Imai, 1956; Mains, 1956b) because of its pileate ascocarp. Imai and
Korf (Korf, 1958) considered Leotia and Neobulgaria similar and removed
the subfamily Leotioideae from the Geoglossaceae, placing it in the
Helotiaceae.
Sclerotiniaceae. The Sclerotiniaceae is often united with the Helo-
tiaceae, from which it may be distinguished by the parasitic habit of
most of its members and by having apothecia arising either directly upon
or in association with sclerotia or from blackened stromata. Apothecia
are mostly larger than in other Helotiales. They are usually soft-fleshed,
long-stalked, cup-shaped, funnel-formed, or verpoid, and are typically
yellowish-brown. The asci are inoperculate, broadly ellipsoid, slightly
thickened at the apex and blue in iodine. The ascospores are unicellular,
hyaline or brown (in some, spores are hyaline when shot but turn brown
before germination), and mostly obliquely uniseriate in the ascus. Sper-
matia or microconidia are formed by most species. This group was treated
by Nannfeldt (1932) as a subfamily (Ciborioideae) of the Helotiaceae,
but Whetzel (1945) recognized it as a separate family characterized by
the features described above. He noted, however, that a stroma or sper-
matia may be absent in some, and a few species are saprophytic. Whetzel
recognized 15 genera, delimited primarily on the structure of the sclerotia
or stroma on which they were formed and on characters of the imperfect
states. Function of the spermatial stage was summarized in Part II of
this paper. More than % of the genera of this family were monographed
by Whetzel or his students: Sclerotinia, Septotinia, Streptotinia, Ver-
patinia, Coprotinia, Martinia, Lambertella, and Seaverina (Whetzel, 1926,
1928, 1937, 1942, 1943, 1944, 1945, 1946), Ovulinia (Weiss, 1940), Rut-
stroemia (White, 1941), and Monilinia (Honey, 1928). Additional tax-
onomic studies were made on Chlorociboria (Ramamurthi, Korf, and
Batra, 1958), Ciborinia (Batra and Korf, 1959; Batra, 1960), and Cibo-
riopsis (Dennis, 1962a). The position of Rutstroemia was questioned by
Le Gal (1953a) who discovered a suboperculum in R. nummiformis (Pat.)
Le Gal. She suggested that possibly Rutstroemia and Helotium may have
arisen from the same ancestral stock and that members of the tribe
Sarcoscypheae may have arisen from the former genus. However, the
suggested relationship here appeared, from the chemotaxonomic view,
difficult to conceive (Arpin, 1968).
Caliciaceae. The family Caliciaceae Fr. has been variously classified
in the orders Caliciales (Fink, 1915; Funk, 1963), Coryneliales (Fitzpatrick
and Korf, 1959), Helotiales (Dennis, 1960, 1968), and Sphaeriales (G/iu-
mann, 1964 as Coryneliaceae), depending on the interpretation of the
126 TIlE BOTANICAL REVIEW
writers and the relative importance given to various structures. Fitzpatrick

(1920) was the first to make a critical taxonomic study of the group. He
placed the Coryneliaceae in the Perisporales but later (Fitzpatrick, 1942)
transferred them to the Dothideales. Luttrell (1955) pointed out that
the Coryneliaceae must be excluded from the Dothideales of the Loculoas-
comycetes on the basis of their unitunicate asci. Arnaud (1930) reviewed
Fitzpatrick's monograph and was convinced that they were discomycetes.
After a thorough study of Caliciopsis, Funk (1963) concluded that the
Caliciaceae and Coryneliaceae were related. Both families were placed in
the Caliciales which includes both lichen and non-lichen genera. The
group is characterized by perithecoid or apothecoid-like bodies that are
clavate, turbinate, barrel-shaped, slender flask-shaped, and sometimes
elongate or stipitate. The pluriascal cavities are initially enclosed. The
asci are typically broadly ovate, with extremely long stalks, and occur in
fascicles. Ascospores in some genera are of unusual form and structure.
At maturation the locules of asci dehisce in a definite fashion and open
widely to expose the asci. Other forms remain essentially closed to appear
perithecoid or pseudothecoid. They are readily distinguished from other
groups by the evanescent character of their asci, which completely dis-
appear, leaving globose heads of loose dry ascospores held in a mazaedium.
Most species are associated with unicellular green algae and conven-
tionally classed among the lichens.
Helotiaceae. According to Dennis (1968), the Helotiaceae is the largest
and most heterogeneous family of the Helotiales. It is characterized by
parallel hyphal structure of the outer layers of the excipulum (a textura
intricata to textura prismatica) or by a flesh of woven hyphae covered by
a comparatively thin area of short, cuboid, or globose cells. The apothecia
are typically bright-colored or darker, soft, fleshy to leathery. The asci
are usually 8-spored, but in some genera numerous secondary, rod-shaped
spores occur. They are inoperculate, with a long, slender pore surrounded
by an amyloid ring. Nannfeldt (1932) recognized nine subfamilies, genera
of two of which have currently been placed in other families. Dennis
(1956) accepted these subfamilies with minor changes. With further
modifications, Dennis (1968) continues to recognize 7 of the subfamilies
as tribes of the Helotiaceae. Dennis has shifted more emphasis to ascus
and apothecial characters than those of substrate and color.
The Ombrophiloideae have very soft apothecia, either completely
gelatinous or with only thin non-gelatinized tissues. One of the most
significant taxonomic treatments of members of this group is Groves'
(1952) study of Tympanis. Other studies include those on Sphagnicola
(Dennis and Korf, 1958) and Coryne (Christiansen, 1963; Groves and
Wilson, 1967). Moore (1965), in a study of gelatinous fungi, concluded
that the presence or absence of gel alone is not sufficient to delimit this
tribe.
In the Helotioideae the apothecia are light colored, usually yellow or
brownish, fleshy, and often long stalked. The medullary excipulum is a
rather loose textura intricata and the ectal area of a textura oblita. Dennis
KIMBROUGH." CLASSIFICATION OF DISCOMYCETES 127
(1968) recognized 18 genera, many of which still appear rather ill defined.
Papers that have contributed greatly to an understanding of the taxonomy
of this group are those on Cudoniella (Mains, 1965a), Helotium (White,
1942a, 1942b, 1943, 1944; Mains, 1956a), Hymenoscyphus (Dennis, 1964),
Chloroscypha (Dennis, 1954a; Kobayasai, 1965), Belonioscypha (Grem-
men, 1950).
The tribe Encoelioideae is characterized by the relatively large,
clustered apothecia that are usually erumpent from bark. The ectal
excipulum is composed of a textura angularis to textura globulosa, whose
ceils are generally brown-walled, giving the apothecium a scruffy appear-
ance. This tribe consists essentially of those genera previous placed in the
Cenangiaceae. Dennis (1956) considered 7 genera in this subfamily, but
in a later treatment (Dennis, 1968), only 5 genera are considered. Missing
from the latter list were the genera Holwaya, Cenangiopsis, Encoeliella,
and Midotiopsis, but with the addition of Cenangium and Praeangella.
Le Gal (1953a) demonstrated that Midotiopsis barnbusicola Henn. had
suboperculate asci and for this reason she placed this genus in the Sar-
coscyphaceae. Rifai (1968) feels that M. bambusicola may fill the gap
between the Sarcoscyphineae and the Helotiaceae. The typification of
Velutaria (Korf, 1953) resulted in transferring those species to Velutarina
Korf. Although Holywaya has been considered in this tribe (Dennis,
1956), the work of Moore (1965) suggests that it should be placed within
the Ombrophiloideae, although Durand (1901), in his monograph of the
genus, placed it in the Patellariaceae.
The tribe Scleroderridoideae superficially resembles the Encoelioideae
but lacks the powdery outer surface. The apothecia are cartilaginous,
erumpent, inverted-conical, closed when dry, black or dark colored, and
often crowded. The ectal excipulum is of a textura prismatica and the
outermost cells often run out into cylindrical hairs. The asci are clavate
with the pore staining blue in iodine. The ascospores are hyaline, elon-
gate or needle-shaped, and 1-multi-septate. Dennis (19'56) recognized a
single genus in Britain, Scleroderris, but later stated (Dennis, 1968) that
two British genera are usually recognized, Scleroderris and Crumenula.
However, Groves (1965) synonymized the two under the older name
Godronia. On a different basis, i.e. the iodine reaction of ascal pores,
Dennis (1968) distinguished the genera Godronia, positive in iodine, from
Ascocalyx with a negative reaction. Six conidial and one microconidial
stages were associated with Godronia (Groves, 1965). Gremmen (1963)
also studied Godronia associated with conidial fungi. Perhaps A tropellis
and Discocainia also belong in this tribe (Reid and Funk, 1966).
In the tribe Durelloideae (Dennis, 1956), the apothecia are superficial,
sessile, saucer-shaped, small, dark brown or black, when moist with a flat,
soft-fleshy hymenium, when dry, cartilaginous and wrinkled. The ex-
cipulum is of thick-walled, cartilaginous hyphae lying parallel and
directed upwards and outwards. Asci are broadly clavate, seldom with
a positive iodine reaction. The spores are elongate, hyaline, simple or
septate, and the paraphyses are rather firm, thickened and often brownish
at the tip. A Cystotricha conidial state occurs. Dennis (1956, 19'68) listed
the single genus Durella.
The Heterosphaerioideae are characterized by apothecia that are car-
tilaginous, superficial or erumpent, sessile, flattened, brown to black,
closed and shrunken when dry, and opened and saucer-shaped when moist.
The ectal excipulum is formed from rectangular cells with thick car-
tilaginous walls. The outermost cells are of a textura globulosa, individ-
ual cells almost isodiametric; with dark brown walls. The ascal pore blues
in iodine. The spores are hyaline and simple or septate. The paraphyses
are hyaline, cylindric or slightly lanceolate. They have been associated
with the conidial stages Pseudodiscosia and Heteropatella. Only one
genus, Heterosphaeria, was recognized in this tribe (Dennis, 1956). How-
ever, in his later treatment Dennis (1968) listed three somewhat doubtfully
related genera, superficially similar in their sessile, cup-shaped, blackish
apothecia with toothed margins. He added Triblidium and Pseudo-
graphis to this tribe.
The tribe Polydesmioideae was proposed by Dennis (1968) to accom-
modate a small but distinctive genus which did not seem to fit elsewhere
in the system. It had light-colored, pruinose apothecia in which the
hymenia were composed of relatively large, non-amyloid asci, containing
large, hyaline, non-septate ascospores, each with two or four oil drops.
The paraphyses were slender and highly branched toward their apices.
It included only the genus Polydesmia.
Nannfeldt (1932) and Dennis (1956) had recognized, in addition, two
other tribes, the Phialeoideae and Trichoscyphelloideae. The genera
originally placed in the Phialeoideae, Phialia, Chlorociboria (= Chloro-
splenium sensu Korf, 1959), CyathicuIa, Calycella, and Belonioscypha,
were transferred to the Helotioideae by Dennis (1968). In a separate
paper, Dennis (1949) treated Nannfeldt's subfamily Trichoscyphelloideae
as an addendum to his British Hyaloscyphaceae. This subfamily contains
two genera with hairy apothecia, Trichoscyphella and Lachnellula,
normally treated with other hairy Helotiaceae, i.e. Hyaloscyphaceae
genera, but which differ in their tissue structure, and in possessing co-
nidial states. They are characterized by apothecia that are light in color,
fleshy, and short to long-stalked. The medullary excipulum is of a loose
textura intricata and the ectal excipulum of a textura oblita, composed
of cells with thicker, somewhat cartilaginous walls, which grow out to
form long, cylindrical hairs. The asci are club-shaped to almost cylindrical
and with an amyloid pore. The spores are spherical or elongate, non-
septate and hyaline. Korf (1962b), after examining an exceptionally rare
yet highly characteristic fungus referred to the genus Lachnellula, sug-
gested that from the nature of the excipulum and other features, perhaps
another genus is needed. Seventeen species were included in a taxonomic
study of the genus by Dharne (1965). Dennis (1968) placed Lachnellula,
including species previously in Trichoscyphella, in the Hyaloscyphaceae.
HyaIoscyphaceae. The Hyaloscyphaceae was erected by Nannfeldt
(1932) for the "hairy Helotiaceae" of earlier authors. He divided the
family into three tribes, Lachneae, Hyaloscypheae, and Arachnopezizeae.
They were characterized as saprophytes with apothecia superficial, or
rarely erumpent, often very small, short- or long-stalked, light-colored,
and with soft flesh. The excipulum is of a texture prismatica, with very
thin-walled, hyaline or occasionally colored cells. The apothecial margins
are clothed with distinct, often rough, hairs. The asci are medium to
large, club-shaped, usually amyloid. Ascospores are elongate to almost
needle-shaped, hyaline, simple or septate. The paraphyses are filiform
to lanceolate, hyaline or with light-colored contents. The species accepted
by Nannfeldt are distributed among 13 genera, distinguished largely by
the detailed structure of the hairs. A good treatment of British species of
the Hyaloscyphaceae is that of Dennis (1949). He adopted Nannfeldt's
classification, with the addition of a new genus Hyalotricha for species
somewhat intermediate in character between Unguicularia and Hya-
lopeziza. Members of the tribe Lachneae are relatively large, and usually
stalked. It consists of two genera, Dascycyphus (Lachnum sensu Nann-
feldt, 1932 or Lachnella sensu Seaver, 1951) and Lachnaster. The Hya-
loscypheae have minute apothecia, and are very poorly known. Nannfeldt
recognized 8 genera. The Arachnopezizeae differs from the other tribes
in possessing apothecia borne on a subiculum. It encompassed Arach-
nopeziza, Arachnoscypha, and Eriopeziza. However, Korf (1951), in a
monograph of the Arachnopezizeae, followed Boudier (1907) in placing
Arachnoscypha in synonymy with Arachnopeziza. Korf also rejected
ChrysothaUus and ArachnopezizeUa as members of the tribe but, following
Dennis (1949), included the genus Tapesina. Dennis (1963) published a
lengthy distribution of fungi previously ascribed to the Hyaloscyphaceae.
Otani (1967) has recently published on a number of Japanese Hyaloscy-
phaceae.
Orbiliaceae. The Orbiliaceae is the smallest family of the Helotiales.
Its apothecia are small, subsessile, mostly on wood or bark, occasionally
on woodier herbaceous stems. They are light-colored, having a waxy
consistency and often translucent when fresh. The excipulum is a textura
angularis to globulosa, with thin-walled cells. Their non-amyloid asci
are exceptionally small, often with a characteristically forked base and
truncate tip. The ascospores are usually small, bacilloid to elliptic,
hyaline, and non-septate. There are three genera, Patinella with a dark
excipulum and dark paraphyses tips; Hyalinia with a waxy-translucent
appearance, toothed margin and cylindric paraphyses; and Orbilia with
an even margin and abruptly swollen paraphyses. Svrfiek (1954) provides
keys and descriptions for 16 species of Orbilia; Seaver (1951) treats 20
North American species, including species placed in Hyalinia by others.
Dermateaceae. The Dermateaceae comprises the species usually
brought together as the "Mollisiaceae" of many treatments. Here are
assembled a large number of genera which have in common a usually
soft flesh composed of mostly thin-walled cells of a textura globulosa.
Nannfeldt (1932) divided the family into 9 subfamilies. The number of
species is large, and the taxonomy very confused. There is no good
monographic treatment of the family. Dennis (1968) treats more than

30 genera, many of which were included also by Seaver (1951) under the
tribe Molliseae. There have been several studies on the "Mollisiaceous"
fungi, most important of which are those of Gremmen (1954, 1955, 1956a,
1956b, 1957) and Le Gal and Mangenot (1956, 1958, 1960, 1961, 1966).
Groves has perhaps contributed greatest to our knowledge of this group
through his thorough studies of the following genera: Pezicula (1939),
Dermea (1946), Dermatea (1940), Tympanis (1952), DurandielIa (1954),
and Pragmopora (1967). Other studies of taxonomic significance are those
on Pyrenopeziza (Hurter, 1958; Gremmen, 1952, 1958), Pseudopeziza
(Schiiepp, 1959), Belonidium (Dennis, 1962b), Drepanopeziza (Rimpau,
1962), and Mollisea (Dennis, 1950).
Hemiphacidiaceae. The Hemiphacidiaceae is the most recent family
proposed for the Helotiales. Korf (1962a) erected the family for a number
of inoperculate discomycetes which cause needle-blights or snow-blights
of conifers, and form orange to yellow-brown, occasionally greenish-brown,
fleshy apothecia within or beneath the host epidermis or hypodermis.
The apothecia become erumpent either by an irregular tear or circum-
scissle scale. The ascocarps are simple, with a poorly differentiated basal
stratum giving rise to asci and paraphyses, with scarcely any marginal
excipular tissue and usually lacking any covering layer of fungus growth
(except in Lophophacidium and Naemacyclus). Most of the species con-
sidered in this family were at one time or another considered in Phacidium
of the Phacidiales. They could be distinguished from the latter in the
absence of the outer layers of the ascocarp made up of dark-walled cells
or hyphae, giving the phacidaceous fruit body a black appearance to the
naked eye. The iodine reaction of asci was used in distinguishing two
tribes, the Hemiphacideae (iodine negative) and the Sarcotrochileae
(iodine positive). Korf provided a key to 5 families of ascomycetes com-
monly associated with needle blight, the Hysteriaceae, Lophiostomataceae,
Phacidiaceae, Hypodermataceae, and Hemiphacidiaceae. Keys were also
provided for the three tribes and eight genera accepted in this family.
These genera included Naemacyclus, Didymascella, Sarcotrochila, Rhab-
docline, Fabrella, Lophophacidium, Hemiphacidium, and Gremmenia.
The last 2 genera were described as new. Reid and Cain (1963) added
the new genus Kor[ia to this list. In an addendum, Korf (1962a) removed
Lophophacidium from the family since it had been shown to possess a
dark-walled upper stratum characteristic of the Phacidiales. Korf's (1962a)
criteria for erecting Gremmenia were: (1) lack of an overlying stromal
layer, (2) the reduction in spores per ascus, and (3) non-amyloid asci.
Petrak (1962) and Reid and Pirozynski (1968) considered these criteria
of doubtful value in delimiting Gremmenia and Phacidium. Reid and
Cain (1962a, 1962b) have contributed greatly to our understanding of the
Hemiphacidiaceae and other needle-blight fungi.
The Phacldiales. Several genera of the families Dermateaceae and
Hemiphacidiaceae of the Helotiales appear related to members of the
Phacidiales. The Phacidiales may be distinguished from both of these
families in having the outer layers of the ascocarp made up of dark-walled
cells or hyphae, giving the fruitbody a black appearance to the naked eye
(Korf, 1962a). Many have recognized this group by apothecia which
developed within a stroma that typically ruptures along one or more
longitudinal or radial lines. Each ascocarp often has more than one
hymenial layer. The order as interpreted by Dennis (1968) is a hetero-
geneous group which contains both ascohymenial and ascolocular forms.
Many authors have followed Nannfeldt (1932), who treated the taxon as
a family of the Helotiales (Bessey, 1950; Wolf and Wolf, 1947). A number
of other workers (Terrier, 1942; Fitzpatrick and Korf, 1959; G~iumann,
1964; Dennis, 1968) recognized it as a separate order. Terrier (1942) was
led to believe that the Phacidiaceae sensu Nannfeldt consisted of both
ascohymenial and ascolocular forms. He erected two families, the Hypo-
dermataceae and Rhytismaceae within the order Hypodermatales, which
contained only ascohymenial forms. The Phacidiales sensu Terrier had
only ascolocular forms. However, according to Korf (in Fitzpatrick and
Korf, 1959), he and Nannfeldt are in agreement that Terrier's conclusion
is invalid. In the group which the latter author terms ascolocular the
ascus pore blues in iodine; no ascolocular forms are known with asci
having pores. The group also has true paraphyses, which are not found
in ascolocular forms. Fitzpatrick and Korf (1959) followed Terrier in
recognizing the Hypodermateaceae, Rhytismaceae, and Phacidiaceae. Von
Arx and Miiller (1954) divided the order into three families, the Cryp-
tomycetaceae, Hypodermataceae, and Phacidiaceae. Dennis (1968), how-
ever, states that two natural families can be segregated, the Hypoder-
mataceae and the Cryptomycetaceae, but there remains a residue of genera
keyed together under the Phacidiaceae. The Rhytismaceae of earlier
workers was abandoned and its species placed in the Hypodermataceae.
Hypoderrnataceae. In the Hypodermataceae the stroma develops but
a single apothecium, simple in structure, with dark basal and covering
layers. There is sometimes a preformed line of dehiscence of the ascocarp,
which opens along a line at maturity, except in Coccomyces,with a radiate
pattern. Terrier (1942) provided a key to species, while a monograph of
conifer-inhabiting species was contributed by Darker (1932). A compre-
hensive taxonomic revision of species is that of Tehon (1935). Darker
(1963) has continued to add to our knowledge of this family, the latest
item (Darker, 1967) being a revision of the genera of the Hypodermata-
ceae. With the more common genera of earlier treatments, i.e. Lopho-
dermium, Hypoderma, Elytroderma, and Coccomyces,he includes Virgilla,
Isthmiella, Lirula, Davisomycella, Ploioderma, Soleela, and Meloderma.
Ouellette and Magasi (1966) added Lophomerum. Cryptomcyes and
Potebniamyces were the only genera placed in the Cryptomycetaceae by
Dennis (1968). Centrum studies of Gordon (1966, 1968) suggest that the
subgelatinous to fleshy stroma characteristics of the Cryptomycetaceae
is not unlike those of the Hypodermataceae.
Rhytismaceae. The pluriapothecial stoma of the Rhytismaceae easily
distinguishes this family from most other families of the order. Separate
apothecia within the common stroma are usually delimited by dark zones
of tissue. The upper and lower layers of stroma are of dark tissue, while
the internal layers are hyaline or white. The asci are cylindric, with a
thin, non-amyloid pore. Terrier (1942) recognized two genera, Rhytisma,
the cause of "tar spot" of a number of plants, and Nymanomyces, orig-
inally a monotypic genus with ovoid brown spores occurring on ,4cer in
Java (Hennings, 1900). Cash in Chardon (1946) described a NTmanomyces
species with brown, ellipsoid spores on Xolisma sp. from the Dominican
Republic.
Phacidiaceae. Terrier (1942) placed in this family those Phacidiales
with amyloid asci, and with a stromatal structure of vertical, rather than
radiating hyphae. He included the genera Phacidium, Phacidiostroma,
Macroderma, and with some hesitance, Myxophacidium and Myxopha-
cidiella. Genera and species are described by Nannfeldt (1932) and
Terrier (1942). Dennis (1968), who includes 12 genera here, doubtlessly
has taxa that belong to the Helotiales and Hysteriales. Of these, Korf
(1962a) earlier transferred Naemacyclus, Didymascella, and Rhabdocline
to the Hemiphacidiaceae. Lophophacidium, originally placed in the
Hemiphacidiaceae (Korf, 1962a), on re-examination was retained in the
Phacidiaceae. Reid and Pirozynski (1966) erected the genus Ceratopha-
cidium for a species of Pezizella that seemed to fall between the Hemi-
phacidiaceae and Phacidiaceae. Since the ascal pore turns blue in iodine,
it appears to lean more toward the Phacidiaceae. Although the genus
Therrya has been placed in this family (Dennis, 1968), Reid and Cain
(1961) feel that it can not be placed here.
The Ostropales. The Ostropales was erected by Nannfeldt (1932) for
a group of genera with asci and ascospores reminiscent of the Clavicipi-
tales, and with rather distinctive tissues in the ectal and medullary
excipula. The asci are long-cylindrical, with a minute pore, and amyloid.
The highly diagnostic ascospores are filiform, multiseptate and hyaline,
tapering towards both extremeties. The idea that this order is related to
the Clavicipitales is suggested by Dennis (1968) in his placing the two
side by side in his latest treatment of British Ascomycetes. G~iumann
(1964) considered the Ostropaceae as a second family of the Clavicipitales.
Gross morphology of the ascocarps is extremely variable, clavate to
stipitate in Vibrissea, discoid in Stictis, to almost perithecoid in Ostropa
and Robergea. This great variation lead earlier workers to place the
pileate forms in the Geoglossaceae (Schroeter, 1897; Mains, 1956b), the
sessile forms in the Mollisioideae, Stictidieae, etc. (Lindau, 1897; Rehm,
1912; Mains, 1956a; and Gremmen, 1958). Aside from the genera above,
Nannfeldt included Apostemidium, Acrospermum, and Schizoxylon in
the order. The studies of Bell~mere (1960) on the apical apparatus in
asci of the Ostropales have raised some doubt as to the homogeneity of
the order. His studies have shown two types of apical apparati within
the group. From this, he divided the order into the "pseudo-Ostropales"
with an apical structure as in Vibrissea and ,4postemidium, and the "eu-
Ostropales" with a structure as in Schizoxylon and Stictis. Without
personal observations, he also considered Ostropa, Robergea, and perhaps
Nanostictis to belong to the latter type. Correlation of ascal structure
and apothecial anatomy led Bell~mere to suggest that the Ostropales be
restricted to those members with the "eu-Ostropales" type, and that the
remaining genera might best remain in the Geoglossaceae. A great deal
of study is needed to clear the controversity which has arisen over the
validity of this order, and to its relationship to other groups. Of help
will be the study of Graddon (1965) on the lignicolous species of Apos-
temidium, and those of S~tnchez and Korf (1966) and S~tnchez (1967) on
the genus Vibrissea. S~inchez and Korf discuss a number of names that
have been applied to the aquatic to semiaquatic discomycetes referred to
the Ostropales. These included Leptosporium, Apostemium, Aposte-
rnidium, Gorgoniceps, Ophiogloea, and Vibrissea. They concluded that
all but Gorgoniceps were synonyms of Vibrissea, and that Gorgoniceps
rightfully belonged in the Helotiales where it had been placed by most
modern authors. S~inchez (1967) provides keys and descriptions to the
species of the sections Microstemium and Apostemium of the genus
Vibrissea.
The Cyttariales. There is still considerable controversity as to the
position of the curious genus Cyttaria. The apothecia are numerous,
imbedded in a common, gelatinous to fleshy stroma produced externally
on the twigs of Notho]agus. The ascocarp appears somewhat "morchel-
loid" but the tissue is solid rather than hollow. The asci have been
considered inoperculate by some (Santesson, 1945; White, 1954; Chade-
faud, 1960), operculate by others (Nannfeldt, 1932; Korf, 1954; Batra and
Batra, 1963). White (19'54) showed the asci of C. gunnii Berk. to be
narrowly cylindric and with a broad, refractive ring in the apex. Korf's
(in Fitzpatrick and Korf, 1959) studies of herbarium material of Cyttaria
showed the ascus apex very much like that in the Sarcoscyphaceae, with
a definite operculum. Moreover, the ring at the ascus apex turns blue
with iodine, and resembled some species of Peziza. Personal observations
of C. gunnii from Nothofagus menziesii preserved for several years in
formalin-acetic acid-alcohol substantiate the findings of Korf. However,
I was unable to detect an amyloid reaction in asci preserved in this
manner. The apical ring stained readily in Congo red in material that
was allowed to soak in distilled water over night. A small plug or core
is present in the apex of young asci. Ascal apices are unequally rounded
as in the Sarcoscyphaceae. As asci mature the plug of material appears
to broaden and is pushed to one side of the ascus, or is divided unevenly
with the larger portion remaining toward the apex (Fig. 1H-K) and the
small portion on the opposite side and farther down the ascus. A single
pore at the base of asci suggests also that they may not have developed
through crozier formation. Illustrations of Le Gal (1946b, 1953a) suggest
that this may be true for the Sarcoscyphaceae. Broadly ellipsoid to ovoid
ascospores is a feature more common to the operculate than to the
inoperculate discomycetes. So far as I am aware, the tissue structure of
the excipulum is unlike that of any of the Pezizales. The smalHumened
cells are extremely thick-walled and with a unique corrugated outer wall
sheath. The orientation of hyphae and the pigmentation of those toward
the exterior is suggestive of that found in the tribes Scleroderridoideae
and Heterosphaerioideae of the Helotiaceae. Strangely enough, some of
those genera are also parasitic, with erumpent stroma, and with amyloid
asci. The nature of their ascal apical structure has not been studied, but
would perhaps be worthwhile in attempting to establish relationships.
The studies of Rifai (1968) were inconclusive, but because of their peculiar
ascocarp anatomy and the unique association with Nothofagus, he sug-
gested that we follow Luttrell (1951) and establish an order Cyttariales
for this genus.
Bessey (1950) indicated that two genera of inoperculate discomycetes
from Sumatra described by Boedijn (1934) were perhaps a transition to
the Cyttariaceae. These were Jacobsonia on wood and Myriodiscus on
bamboo. Palm (1932) described Cyttariella as a new genus for the
pycnidial stage of Cyttaria. The Australian species of Cyttaria were
revised by Rawlings (1956).
The Pezizales. Boudier (1885) divided the cup fungi into major
groups, the Operculatae and the Inoperculatae. Although this system
was not accepted initially, most mycologists now agree that this subdivision
is fully justified. Other characters of ascospores, excipula, and to some
extent cytological and cytochemical features (discussed in part II) are
associated with the manner of ascus dehiscence. The number of families
recognized in the Pezizales varies considerably from one author to another
(Table 4). The number of families that have been recognized, and that
should be recognized, has been discussed thoroughly by both Rifai (1968)
and Eckblad (1968). Although the family names proposed by Boudier
(1885) still exists for taxa of the Pezizales (except Exoascaceae), they are
considered unnatural in their original sense. The distinction of sub-
operculate from operculate asci (Le Gal, 1946a), and weakening influence
of gross morphology of apothecia on classification, brought about more
natural systems of classifications. The suggestion made by Nannfeldt
(1949), Korf (1954), and Berthet (1964a) to give equal rank to the Sar-
coscyphaceae and the rest of the Pezizales was incorporated by Rifai (1968)
who recognized two suborders, the Sarcoscyphineae and the Pezizineae.
Arpin (1968) felt that this subdivision was justified based on pigment
analysis also.
THE SUBOROEaSAaCOSCYrmNEAE. Rifai (1968) characterized this sub-
order as follows: Asci suboperculate, thick walled, long, cylindrical above,
usually much narrower, flexuous, trailing hypha-like base, of "aporhy-
nique" type (Berthet, 1964a); ascospores hyaline, always guttulate, pluri-
nucleate, often asymmetrical, smooth or covered by transverse or longi-
tudinal striations or ridges unaffected by cotton-blue in lactic acid (except
in Galiella and Neournula, the ascospores of which are covered with
callose-pectic warts); paraphyses delicate to slender, sometimes interspersed
with distinctive hymenial hairs; medullary excipulum always prosenchy-
matous, sometimes embedded in a gelatinous matrix; apothecia tough,
corky, leathery, subgelatinous but never truly fleshy, lignicolous or rarely
terricolous. He included the single family Sarcoscyphaceae.
Sarcoscyphaceae. Based upon pigmentation of apothecia, the Sar-
coscyphaceae has been divided into two tribes, the Urnuleae with dark
apothecia and the Sarcoscypheae with bright colored apothecia (Le Gal,
1947; Nannfeldt, 1949; Dennis, 1960, 1968; Rifai, 1968; Eckblad, 1968;
Arpin, 1968). This subdivision is further supported by the cytological
studies of Berthet (1964b) in which paraphyses of the Urnuleae were
uninucleate, the Sarcoscypheae, muhinucleate. Some of the genera pres-
ently considered in this family were placed in the tribes Ciliarides and
Peziz6es of the Pezizaceae by Boudier (1907). These included the genera
Sarcoscypha, Urnula, Sarcosoma, Melascypha, and Pseudoplectania. Le
Gal (1946b) considered these plus Cookeina, Phillipsia, Pithya, and Wyn-
nea as members of the Sarcoscyphaceae. In 1953a, she added Plectania
and Midotiopsis. A number of other genera have been subsequently
added: Desmazierella (Nannfeldt, 1949); Galiella (Korf, 1957b); Geodina
(Denison, 1965); Chorioactis, Nannfeldtiella, Wolfina, Phaedropezia, and
Pseudopithyella (Eckblad, 1968); and Boedijinopeziza and Aurophora
(Rifai, 1968). Even though this family has received considerable attention
in recent years, there are still many areas of confusion. Korf (1953, 1957b)
has discussed the confusion of the application of the generic name Plec-
tania, and Rifai (1968) pointed out a similar problem in Sarcoscypha.
The disposition of a number of species is still in question. Ito and Imai
(1937) made Peziza institia Berk. and Curt. the type of Boedijnopeziza
(accepted by Rifai, 1968); however, Le Gal (1958) and Eckblad (1968)
retained it in Cookeina. Peziza dochmia Berk. and Curt., the type of
Aurophora Rifai, is retained in Phillipsia by Le Gal (1953b) and Eckblad
(1968). Although Phaedropezia has suboperculate asci, Rifai (1968) in-
cluded it in the Pezizineae because the other characters were markedly
different from other Sarcoscyphaceae. He suggested that another family
be proposed to accommodate Phaedropezia in the Pezizales. Urnula
geaster Peck was placed in a new genus Chorioactis Kupfer (1902) because
of differences in cellular nature of the excipulum from that of U. crater-
ium. This was strongly opposed by Heald and Wolf (1910) who were
followed by Seaver (1928) and others. On the basis of differences in
excipular structure and tomentum in connection with habit, Chorioactis
was accepted as clearly distinct from Urnula by Eckblad (1968). Rifai
(1968) provides keys and descriptions to seven Australasian genera, Dennis
(1968) includes seven genera in the British Ascomycetes, while Eckblad
(1968) treats eight genera in the Sarcoscypheae and ten genera in the
Urnuleae. Denison (1967, 1969) has provided some interesting studies on
Phillipsia, Geodina, and Cookeina and other Sarcoscyphaceae from
Central America. Paden and Tylutki (1968) described the new genus
Neournula from Idaho with one species, N. nordmanensis. In a later
paper (Paden and Tylutki, 1969), new species of Sarcosoma and Plectania
are described from the same area.
Tn~ SVBORDEkPEmZlNEAE. Rifai (1968) described the Pezizineae as
follows: Asci operculate, or very rarely suboperculate, thin walled,

cylindrical, subclavate, clavate or saccate, base not hypha-like, or "pleuro-
rhynique" or "aporhynque" type; ascospores hyaline to dark colored, with
or without oil globules, plurinucleate, tetranucleate or mostly uninucleate,
mostly symmetrical, smooth or covered by various kinds of ornaments of
callose-pectic material; paraphyses slender to stout, rarely delicate;
medullary excipulum pseudoparenchymatous or prosenchymatous but
never embedded in a gelatinous matrix; apothecia fleshy to finn fleshy,
brittle, mostly terricolous but sometimes also on dung or other vegetable
remains or lignicolous. Rifai included seven families in this suborder
(Table 4), basing the system mainly on that of Le Gal (1947) and with
strong support provided by Berthet's (19.61-1964) work. The scope of
the Pezizineae was identical with that of the family Pezizaceae as con-
ceived by Nannfeldt (1937, 1949) and Korf (1954). Korf, however, divided
the family into eleven tribes, four of which correspond to the Morchel-
laceae, Pyronemataceae, Pezizaceae, and Ascobolaceae that were recognized
by Rifai. He suggested that the tribes Discineae, Gyromitreae, and
Acetabuleae (in part), and possibly the Rhizineae also, should be in a
taxon equal in rank to the Morchellaceae and Pezizaceae. Berthet's
(1963, 1964a) cytological work indicated their affinities to the Helvella-
ceae. Eckblad (1968) proposed the family Rhizinaceae for essentially the
same groups.
The Humariaceae, recognized by Boudier (1885) and a number of
other mycologists (Table 4), is the largest and most difficult family of
the Pezizales. Currently, there is considerable controversity as to the limits
of this taxon. Rifai (1968) conceives it as a broad family, related to the
Pyronemataceae on one hand and the Helvellaceae on the other. He
proposed four tribes, the Otideae, with "helvelloid tendencies," the
Lachneae, the Ciliarieae, and the Aleurieae. Dennis (1968) recognized
only the tribes Lachneae, Ciliarieae, and Aleurieae in the Humariaceae,
placing the Otideae as a second tribe of the Pezizaceae. Eckblad (1968)
placed most of the Otideae in the new family Otidiaceae; those of the
tribes Aleurieae, Lachneae, and Ciliarieae were mostly transferred to
the Pyronemaceae Corda emend Eckblad. Rifai (1968), however, limits
the Pyronemataceae to the single genus Pyronema. Arpin (1968) discussed
both the taxonomic and nomenclatural problem of the Humariaceae and
proposed in a restricted sense to replace it with the new family Aleuriaceae
Arpin. Based on a detailed analysis of pigments, he proposed the Aleu-
riaceae for taxa previously placed in the tribes Aleurieae and Ciliarieae.
The tribes Otideae and Lachneae would become part of the Otideaceae.
Arpin would conserve the Pyronemataceae sensu Rifai distinct from the
Aleuriaceae. In the discussion of families of Pezizales, I shall follow
Arpin in treating "humariaceous" genera as members of the Otidiaceae
and Aleuriaceae.
The Pezizaceae is restricted to include those genera with amyloid asci
and many species of which produce in culture Oedocephalum or Ostraco-
derma conidial stages. The Ascobolaceae is also restricted to those Pezi-
zales in which the ascospores have purple to dark brown epispores in
which the pigments originate from extra sporal or vacuolar material
(Le Gal, 1947). She pointed out that the pigmented species of the Ascobo-
laceae form a homogeneous and natural entity which is related to the
Pezizaceae. The Pseudoascoboleae have traditionally been included in
this family. Van Brummelen (1967), one of the latest to do so, divided
the Ascobolaceae into three tribes, the Ascoboloideae, the Ascodesmidoi-
deae, and the Theleboloideae. Dennis (1968), Rifai (1968), and Eckblad
(1968) chose to follow Kimbrough and Korf (1967) in placing the Pseu-
doascobolaceae outside of the Ascobolaceae. They chose to elevate the
Theleboleae to family Thelebolaceae, even though Dennis used it in a
very restricted sense. Thus by basing a system on the work of Berthet
(1961-1964), Rifai (1968), and Eckblad (1968), the following families may
be recognized in the suborder Pezizineae: the Morchellaceae, Helvella-
ceae, Otidiaceae, Aleuriaceae, Pezizaceae, Ascobolaceae, Thelebolaceae,
and Pyronemataceae.
Morchellaceae. Apothecia of the Morchellaceae are large and vary
greatly in shape from sponge-like in Morchella, campanulate in Verpa,
to sometimes pezizoid and substipitate in Disciotis. The medullary ex-
cipulum is of a textura intricata and the ectal excipulum is of a textura
angularis to textura prismatica. The asci are cylindrical, non-amyloid,
not protruding at maturity, with an "aporhynque" type base, usually
8-spored, but sometimes with 2, 4, or 6 spores. The most outstanding
feature of the family is in the ascospores which are ellipsoid, smooth,
usually hyaline. Mature ascospores do not contain oil globules but during
their development their polar regions are crowned with numerous ex-
ternal guttules. Berthet (1963, 1964a) has shown that the spores of this
family are invariably plurinucleate. The paraphyses are straight, septate
and pale brown. The current works of Rifai, Eckblad, and Dennis agree
in the inclusion of Morchella, Verpa, and Disciotis. Dennis also includes
Mitrophora and Ptychoverpa, both of which were placed in synonymy
by Eckblad. Rifai (1968) discusses the history of the Morchellaceae and
the confusion that has existed between stipitate and sessile forms. The
genus Mitrophora differs from Morchella in having a smaller receptacle
with its lower half free from the stalk (Boudier, 1907; Le Gal, 1947;
Dennis, 1960, 1968). This small difference has led many to merge it with
Morchella (Saccardo, 1889; Seaver, 1928; Imai, 1954; Moser, 1951; Groves
and Hoare, 1953; Eckblad, 1968; and Rifai, 1968). As is noted by Rifai
(1968), the number of species that can be recognized in Morchella is
uncertain and varies from three (Groves and Hoare, 1953), six (Seaver,
1928), to 32 (Boudier, 1907). Wojewoda (1966) listed 6 species as occurring
in south Poland. Heim (1966) described two new species of MorcheIla
from the south Pacific. gebek (1966) provided keys to 11 taxa of Mor-
chella, 2 of Mitrophora, and 2 of Verpa found in Czechoslovakia. Rifai
suggested the possibility of Morchella having many "micro-species" and
recommended the "species aggregate" concept.
Helvellaceae. There is currently a wide difference in opinion as to
the limits of the Helvellaceae. Superficial similarities between members

of the present family and the Morchellaceae have led many to unite them
(Scarer, 1928; Bessey, 1950; Imai, 1954; Alexopoulos, 1962; Benedix, 1962;
Batra and Batra, 1963). Many would exclude the cupulate or sessile
genera, but cytological evidence (Berthet, 1964b) indicates that they
belong with the Helvellaceae. He would include the genera Discina,
Rhizina, and Neogyromitra together with Helvella, Wynnella, and
Gyromitra in the Helvellaceae because they always have tetranucleate
ascospores. It appears that this unique spore character is associated with
other features of taxonomic significance such as the habit, habitat, pig-
mentation of apothecia, guttulation of ascospores, and the internal struc-
ture of the apothecium. Eckblad (1968) conceives the family to correspond
to the tribe Acetabuleae Nannf. and the Helvelleae Dissing (1966a).
However, he refused to include Gyromitra, Discina, and their allies merely
on the strength of tetranucleate spores. For these he proposed the family
Rhizinaceae with Discina, Neogyromitra, Maublancomyces, Rhizina,
Gyromitra, and Pseudorhizina. The main criterion on which the Rhizina-
ceae was based was the presence of similar callose-pectic marked spores.
Rifai (1968) states that the presence of such spore markings cannot be
regarded as an obstacle for placing a genus in the Helvellaceae because
the spores of H. macropus, which is an unquestioned member of the
Helvellaceae, are also provided with minute callose-pectic warts. Similar
markings are present on Underwoodia (Rifai, 1968), a genus which Eck-
blad states does not differ in any other character than form, and was
therefore included in Helvella. For these reasons I cannot accept the
family Rhizinaceae sensu Eckblad. Eckblad restricted the Helvellaceae
to include only Helvella and Wynnella. The Helvellaceae as conceived
by Berthet (1964b) has been accepted by Dissing (1966a), Dennis (1968),
and Rifai (1968). Nannfeldt (1937) pointed out that the segregation of
several small genera from Helvella by Boudier (1885, 1907) was difficult
to accept because characters such as the shape of the receptacle, the type
and absence or presence of the stipe, were not of important taxonomic
value at the generic level. Thus, he placed in synonymy the genera
Leptopodia, Cyathipodia, Macropodia, and Paxina. Rifai (1968) and
Eckblad (1968) add several other synonyms to this list. Nannfeldt's sug-
gestion of merging these genera was not accepted by a number of workers
(Le Gal, 1947; Dennis, 1960, 1968; Benedix, 1962; and Berthet, 1964b).
Le Gal (1963), however, admitted this view should be accepted. Batra
and Batra (1963), Dissing (1964, 1966a, 1966b), Dissing and Nannfeldt
(1966), and Dissing and Lange (1967) have found the wide generic concept
to be the most natural arrangement. Although the genus Underwoodia
was synonymized by Eckblad (1968), I follow Rifai (1968) who accepts
it as a distinct genus of the Helvellaceae. Korf (1956) considered that
some of the species assigned to Underwoodia were merely sprassoid forms
of species of Peziza, while others were like Helvella in that the sterile
tissues were similar to stipe tissues of the latter genus. Although Nann-
feldt (1937) and Dennis (1960) placed Pustulina in the present family,
Rifai (1968) believes that because of uninucleate ascospores it should
be excluded and referred to the Humariaceae. Dennis (1968) does exclude
it, but places it in the tribe Otideae of the Pezizaceae.
Therefore, the Helvellaceae in the present sense may be said to include
genera with usually more or less stalked ascocarps, varying in form from
cup-shaped to saddle-shaped, often lobed, veined, contorted or intricately
folded. Most are terrestrial, but Rhizina is often associated with sub-
merged wood. The asci are non-amyloid and the ascospores contain
conspicuous oil droplets but without polar granules. The ascospores are
smooth and ellipsoid in some genera, fusiform, apiculate, and minutely
roughened in others.
Otidiaceae. The new family Otidiaceae Eckblad (1968) was diagnosed
as follows: "Apothecia medium-sized, sessile or more often shortly stipi-
tare, externally glabrous or furfuraceous. Excipulum distinctly two-
layered, ental layer of a textura intricata, ectal layer of a textura angularis,
often externally clothed with warts of globose cells or short hyphoid hairs.
Hymenium yellowish, brownish or pale, rarely orange. Asci cylindrical,
eight-spored non-amyloid. Spores ellipsoid, hyaline, smooth or finely
warted, containing two oil drops or none. Paraphyses septate, mostly
straight, rarely apically curved or hooked, hyaline or containing granular
matters." He included five genera: Geopyxis, Otidea, Pustulina, Sower-
byella, and Ascosparassis. The type, Otidea, had previously been placed
in the tribe Peziz~es of the Pezizacfies (Boudier, 1907), the tribe Otideae
of the Aleuriaceae (Le Gal, 1947), the Acetabuleae of the Pezizaceae (Korf,
1954), the Otideae of the Pezizaceae (Dennis, 1968), and the Otideae of
the Humariaceae (Rifai, 1968). Scarer (1928) treated genera of both
Pezizaceae and Sarcoscyphaceae in his tribe Otideae.
The absence of carotenoids prompted Arpin (1968) to emend the
Otidiaceae sensu Eckblad to accommodate the genera Sepultaria, Tri-
charia, Mycolachnea, Pseudombrophila, and Trichophaea. Most of these
had been treated in the Humariaceae, tribe Lachneae (Le Gal, 1947;
Rifai, 1968; Dennis, 1968), the Pezizaceae, tribe Ciliarieae (Korf, 1954),
or as members of the Pyronemaceae (Eckblad, 1968). The presence of
carotenoids in Geopyxis and Sowerbyella prompted their transfer to the
Aleuriaceae (Arpin, 1968). Morphological features suggest that Ja[nea
(Korf, 1960b), Notho]a/nea and Ja[naedelphus (Rifai, 1968), and Sphae-
rosporeIla (Svrfiek and Kubi6ka, 1961) belong here. The genus Marcel-
leina was only provisionally referred to the tribe Otideae by Rifai (1968).
It was similarly treated (as Barlaeina) by Dennis (1968). Eckblad (1968),
however, considered it related to Plicaria, and placed it in the Pezizaceae,
despite the non-amyloid asci. He contended that MarceIleina differs from
Plicaria only in the lack of an amyloid reaction of asci. But in Marcelleina
the disc is purple due to the presence of purple granules around the
paraphyses, the excipulum is essentially hyaline to yellowish, and the
spores are smooth to verrucose and hyaline, while in Plicaria, with amyloid
asci, the excipulum is brownish in section, the apothecia are darker brown,
the ascospores become light yellow or brown and are smooth to variously
ornamented, and the paraphyses usually become encrusted with dark

brown amorphous material. Whether Marcelliana should be included in
the Otidiaceae is still open to question.
The genus Otidea was monographed by Kanouse (1949).
Aleuriaceae. Arpin (1968) proposed the new family Aleuriaceae for
a number of genera previously placed in tribes Aleureae and Ciliareae
of the Humariaceae (Rifai, 1968) and parts of the Pyronemaceae Eckblad
(1968). The diagnosis from Latin is as follows: Apothecia humicolous,
or rarely epixylous, often sessile, rarely subsessile, cup-shaped or lenticular,
and then broadening, glabrous or pilose. Excipulum for the most part
consistently of two layers; interiorly a textura intricata, exteriorly a
textura globulosa or angularis; sometimes a single zone of textura globu-
losa-angularia. Hymenium colored yellow, orange, red, brown, rarely
white or ivory; asci cylindrical, 8-spored, operculate, not at all amyloid;
spores smooth or ornamented; ornaments of callose-pectic nature, varying
greatly, sometimes of complex types; spores uninucleate. Paraphyses
straight or curved upwards, filled with carotenoid pigments, dissolved
in guttules, or as granules or crystals in others. Carotenoid pigments have
either a bicyclic structure (B-carotene) or monocyclic (y-carotene, torulene,
plectaniaxanthine, dehydro-2 plectania-xanthine, aleuriaxanthine).
Arpin was able to divide the Aleuriaceae into three groups on the basis
of carotenoid ratios. Group a contained Coprobia, Cheilymenia, Scutel-
linia, and GeopTxis, all with generally a higher ratio of y- to B-carotene.
Group b contained .41euria, Melastiza, and Octospora, each with approxi-
mately equal concentrations of carotenes. Group c contained Pulvinula,
Anthracobia, Caloscypha, and Sowerbyella in which ratio of /3-carotene
was almost 10--1 over y-carotene. With the exceptions of Geopyxis and
SowerbyeUa, genera of all three groups were placed in the Pyronemaceae
sensu Eckblad (1968). All of group a except Geopyxis were in the tribe
Ciliareae, and group b and group c, the tribe Ciliareae of the Humariaceae
(Rifai, 1968). Of the other genera currently placed in the Humariaceae
or Pyronemaceae, the following would appear from morphological features
to belong in the Aleuriaceae: Fimaria, Leucoscypha, Lamprospora,
Inermisia, Genosperma, and Rhizoblepharis.
Worthwhile taxonomic studies have been published on the following
genera: Lamprospara (Dennis, 1955), Leucoscypha (Le Gal, 1957), Mela-
stiza (Le Gal, 1958; Maas Geesteranus, 1967), Octospora (Svrfiek and
Kubifika, 1963), Cheilymenia (Denison, 1964; Moravec, 1968), and Scutel-
linia (Denison, 1961; Gamundi, 1956; Le Gal, 1962, 1964, 1966).
Pezizaceae. The Pezizaceae has traditionally encompassed essentially
all of the larger, cupulate or sessile, operculate discomycetes (Boudier,
1907; Seaver, 1928; Le Gal, 1947; Nannfeldt, 1949; Korf, 1954). The
number of tribes recognized varied from three (Le Gal) to eleven (Korf).
Current treatments of the Pezizales, however, more or less restrict the
Pezizaceae to correspond to the tribe Pezizeae of Korf (1954), i.e. genera
with amyloid asci, smooth or callose-pectic marked spores, and many with
special conidial stages. Although, Eckblad (1968) would include certain
genera with non-amyloid asci, as did Dennis (1968) who included the tribe
Otideae here, I follow Rifai (1968) who restricts the family to those
amyloid groups. In this sense the family may be diagnosed as follows:
Apothecia from one to two millimeters to more than ten centimeters,
variable in shape, globose to cupulate, discoid, pulvinate, sessile or slightly
stipitate, some becoming sparassoid. Excipulum usually of two layers,
the medullary area variable, hyphoid with globose cells to almost a textura
globulosa, interspersed with hyphae or often with lactiferous hyphae;
the ectal area of a textura angularis to globulosa, and occasionally beaded
rows of cells or parallel hyphae oriented upwards. Hymenium variable
in color from whitish or yellowish, through various shades of violet,
purple, ochraceous to brown, never bright orange or red. Asci cylindrical,
8-spored, operculate, diffusely amyloid their entire length or strongly
blueing only at their apices. Ascospores globose, fusiform to ellipsoid,
hyaline or rarely pale yellow, smooth or with callose-pectic ornaments,
rarely apiculate, uninucleate, with or without oil guttules. Paraphyses
septate, simple or branched below, filiform to slightly clavate, straight or
curved, hyaline or containing various pigments, sometimes encrusted
with amorphous matter.
Traditionally the genera Peziza, Plicaria, Sarcosphaera, and Pachyella
have been placed in this family. Kimbrough and Korf (1967) referred
the coprophilous genera Iodophanus and Thecotheus to this family.
Rifai (1968) believed that the genera Gelatinodiscus, Boudiera and per-
haps Sphaerosoma belonged here also. Dennis (1968) accepted two tribes,
Pezizeae with Plicaria, Sarcosphaera, and Peziza, and Otideae with Otidea,
Barlaeina, Pseudotis, Pustularia, and Sowerbyella. The latter tribe, ex-
cepting Sowerbyella, is currently referred to the Otidiaceae Eckblad
emend Arpin. Eckblad (1968) treats two genera with amyloid asci, Peziza
and Plicaria, and three with non-amyloid, Marcelleina, Psilopezia, and
Discomycetella. Preliminary observations indicate that species of Iner-
misia Rifai and Discomycetella Sanwal are perhaps congeneric. It is sug-
gested above that Inermisia probably belongs to the Aleuriaceae. The
excipular structure and ascospores of Psilopezia are reminiscent to those
of Iodophanus, except that the latter possesses spores without oil guttules.
Spores of both have fine, callose-pectic markings. Eckblad (1968) states
that asci of Psilopezia are non-amyloid. I find, however, that in several
North American collections of P. aquatica (DC.) Rehm and P. babingtonii
(Berk. and Br.) Le Gal the asci are diffusely amyloid their entire length.
The remaining non-amyloid genus, Marcelleina, should perhaps be re-
tained in the Otidiaceae as was suggested by Rifai (1968).
There has been considerable controversy as to the typification and
nomenclature in the Galactinia-Peziza complex. A number of workers
have abandoned the name Peziza and chosen the name Galactinia (Cooke)
Boud. emend. Le Gal, while other authors have been reluctant to drop
the name. The long history of this genus is ably discussed by Rifai (1968).
The genus is cited differently in the most recent treatments of the group:
Peziza St-Amans (Dennis, 1968; Arpin, 1968), Peziza (Dill.) L. ex St-Amans
(Rifai, 1968), and Peziza Dill. ex. Fr. (Eckblad, 1968). Korf (1960a) dis-
cussed a similar confusion in application of the generic name Plicaria.
Korf (1960a), Le Gal (1953a, 1962), Denison (1963), and Berthet (1964a)
feel that Plicaria should be merged with Peziza, while Maas Geesteranus
(1955), Dennis (1960, 1968), Batra (1961a), Batra and Batra (1963), Moser
(1963), Rifai (1968), and Eckblad (1968) treat them as distinct genera.
Rifai (1968) feels that Plicaria represents an intermediate taxon between
Peziza and Boudiera.
There have been but few recent taxonomic treatments of genera
belonging to this family; they are: Galactinia (Le Gal, 1969); Thecotheus
(Kimbrough, 1969a), and Iodophanus (Kimbrough, 1969b).
Thelebolaceae. Most of the taxa represented in this family were placed
originally in the "Ascobolei spurii" (Boudier, 1869), or what later became
known as the Pseudoascoboleae. A historical survey of this group was
discussed by Kimbrough (1966b). In this study it was found that the
genera of the Pseudoascoboleae should be founded on qualitative micro-
scopic and micro-chemical characters, especially those of asci and asco-
spores, rather than on quantitative and superficial evidence such as the
number of asci and ascospores as has been traditionally accepted. Kim-
brough and Korf (1967) placed those genera with hyaline spores, 8- and
multispored, non-amyloid, operculate or irregularily dehiscing asci in
the tribe Theleboleae of the Pezizaceae. Van Brummelen (1967) elevated
this tribe to the subfamily Theleboloideae of the Ascobolaceae, while
both Rifai (1968) and Eckblad (1968) placed them one step higher as
the family Thelebolaceae. Based on the studies of Cooke and Barr (1964),
Dennis (1968) restricted the family to the single genus Thelebolus.
The genera Iodophanus and Thecotheus, each with amyloid asci and
callose-pectic marked spores, were removed from the Pseudoascoboleae
and placed in the tribe Pezizeae of the Pezizaceae (Kimbrough and Korf,
1967). Rifai (1968) supports this transfer and accepts these as genera of
the Pezizaceae. Eckblad (1968), however, retained Thecotheus in the
Thelebolaceae because he felt amyloidy of asci and callose-pectic marked
spores were common features of many operculates, that the excipular
structure of Thecotheus was less complex than those of the Pezizaceae,
and that multispored asci suggested a different phylogenetic level. He
felt that Iodophanus was more related to Pyronema and placed it in the
Pyronemataceae. Dennis (1968) continues to treat both the "theleboloid"
and "pezizoid" genera in the Pseudoascoboloideae of the Ascobolaceae.
I accept the Thelebolaceae to correspond to the tribe Theleboleae
(Kimbrough and Korf, 1967) or Thelebolaceae (Rifai, 1968). Eckblad's
(1968) diagnosis of the family, with minor modifications, can be accepted
as follows: Ascocarps apothecial or cleistothecial, small, sessile, smooth
or furfuraceous to distinctly hairy. Excipulum of one to several layers,
mostly of a textura angularis to globulosa, sometimes almost a textura
epidermoid, ectal and medullary are present in some. Hairs of different
kinds, including straight, acute, thick-walled, colored or hyaline, septate
or non-septate. Asci one, few or many, obovate, clavate to clavate-cylin-
KIMBROUGH: CLASSIFICATION OF DISCOMYCETES ]43
drical, dehiscing by an operculum, bilabiate split, irregular tear, or apical

plug, non-amyloid, mostly protruding above the hymenial level, or ruptur-
ing the cleistothecium apically at maturity, 8 to several thousand spores
per ascus. Ascospores uniseriate, biseriate, or irregular, ellipsoid, ovoid,
naviculate, globose, subglobose or fusoid, smooth, hyaline, often sur-
rounded by a mucilaginous sheath, uninucleate, contents granular, some
with de Bary bubbles, never with oil guttules. Paraphyses highly variable,
simple to very branched, filiform to clavate, straight or curved, rarely
forming an epithecium.
Kimbrough and Korf (1967) accepted the following genera: Ascozonus,
Caccobius, Coprobolus, Coprotus, Lasiobolus, TheleboIus, and Tricho-
bolus. Eckblad (1968) would include Thecotheus and Leporina. Accord-
ing to Eckblad (1968) van Brummelen studied the type of the latter genus
and found it to be conspecific with Ryparobius sexdecirasporus (Cr. and
Cr.) Sacc. This implies that Leporina Vel. is the correct name of Coprotus
Korf and Kimbrough. Personal observations of the type, L. multispor
Vel. confirms van Brummelen's conclusions. Extensive taxonomic studies
on generic and species level are needed in this group, although the work
of Kimbrough (1966a, 1966b) and Kimbrough and Korf (1967) have done
much to clarify the generic concepts.
Ascobolaceae. The Ascobolaceae in the present sense may be diagnosed
as follows: Apothecia superficial to immersed, white, yellowish, greenish
or brownish to purple brown apothecial to cleistothecial, with intergrades
between, mostly sessile, sometimes very shortly stipitate, receptacle
smooth, furfuraceous or villose, without true hairs. Excipulum poorly
developed or absent in some, usually simple, varying from a textura
globulosa, angularis, to epidermoid, medullary and ectal areas not dis-
cernable in most. Asci clavate with a well formed operculum, mostly
8-spored, sometimes less, amyloid or non-amyloid, protruding above the
level of the hymenium at maturity. Spores ellipsoid, fusiform, globose or
subglobose, uninucleate, small or large episporial ornaments which are
mostly colored, purple at first, becoming purple-brown or dark brown.
Ornaments at thin crust, at first continuous, later cracked or discontinu-
ous, an alveolate reticulum or strong spines, big warts or crests. Para-
physes filiform, septate, straight or curved, with hyaline walls, sometimes
colored contents or surrounded with mucilaginous material with or with-
out amorphous colored material.
Following Eckblad (1968), the family as here conceived comprises only
the subfamilies Ascoboloideae and Ascodesmidoideae of van Brummelen
(1967), the first containing dscobolus and Saccobolus, the latter with
Ascodesmis, Bo.udiera and Svr~ekia (the last two added by Eckblad, 1968).
In adding Boudiera, Eckblad (1968) contended that spore sculpturing in
this genus appears to be closer to Ascodesmis than any other taxon except
perhaps the tuberaceous Sphaerosoma. He tentatively placed Svr~ekia
close to Boudiera. Le Gal (1947) excluded the genus Boudiera from the
Ascobolaceae and placed it in the Humariaceae. Rifai (1968) felt that
Boudiera should be classified as a member of the Pezizaceae because the
type species had amyloid asci. Dennis (1968) places it in the tribe Aleu-
rieae of the Humariaceae. The hyaline spored genera traditionally placed
in the family as the subfamily or tribe Pseudoascoboleae or Theleboloi-
deae are treated above as a separate family, the Thelebolaceae.
As is pointed out in part II, the Ascobolaceae have received the
attention of many who are studying development, cytology, and genetics
of fungi. A number of taxonomic studies have been made, foremost
among these is the current world monograph of Ascobolus and Saccobolus
by van Brummelen (1967). Until recently no attempt has been made to
revise the numerous species of Ascobolaceae. Le Gal (1953b, 1961) exam-
ined many original collections in the herbarium of the Crouans, while
van Brummelen (1962) investigated the collections and types in the
herbarium of Spegazzini. Others include a study of the genus Ascodesmis
(Obrist, 1961) and the Ascobolaceae of Argentina (Gamundi and Ranalli,
1966).
Pyronemataceae. Following Rifai (1968) and Arpin (1968), the Pyro-
nemataceae is restricted to a single genus, Pyronema. As such, the family
may be diagnosed as follows: Apothecia minute, red, pink, orange, ivory,
or white, scattered to gregarious, sometimes confluent and forming crusts
of fused apothecia, borne on a more or less prominent subiculum, always
on recently sterilized substrate (particularly fireplaces, charcoal kilns,
steam-sterilized soil, new plaster), of very simple structure, at first conical,
expanding to turbinate, with or without delicate hairs on the ectal
excipulum, arising from one or more pairs of large sex organs supported
by conspicuous stalk cells, each pair consisting of a clavate, unicellular
antheridium and a globose, unicellular ascogonium surmounted by a
tubular, unicellular tricogyne; asci operculate, 8-spored, clavate, not
blueing in iodine, arising from croziers; ascospores uniseriate, hyaline,
elliptic, 1-celled, non-guttulate, smooth or very finely punctate at complete
maturity; paraphyses filiform, seldom branching except at the very base,
sparingly septate, slightly or not enlarged above, about as long as the
asci. Conidia absent. Sclerotia known only in culture of certain species.
Rifai (1968) states that the true relationship of the Pyronemataceae
with others can be established only when the anatomy and the ontogeny
of many brightly pigmented species of the Humariaceae and the Thele-
bolaceae are more fully understood. He suggested that the family should
possibly be regarded only as a tribe of the Humariaceae, but since the
latter appears already polyphyletic the introduction of another aberrant
element there seems to be unwarranted. This family (as Pyronemaceae)
was taken in a very wide sense by Eckblad (1968) who included along
with Pyronema most all of the genera, except the Otideae, that had been
placed by earlier workers in the Humariaceae. Dennis (1968) includes
it in the Pseudoascoboleae of the Ascobolaceae. Arpin (1968) follows
Rifai (1968) and maintains Pyronema in a distinct family because the
precise pigments appear completely different from those of Coprobia and
similar Aleuriaceae to which others contend it is related.
The voluminous studies on this genus have been thoroughly covered
by Boedijn (1940) and Moore (1963). Moore and Korf (1963) stated,
however, that there was no guarantee that all these works were actually
based on species of Pyronema. They include keys and descriptions for
two species.
RELATIONSHIP TO THE TUBERALES. Most modern authors agree that the
Pezizales and Tuberales are closely related and that the latter should be
derived from the former. There is no agreement about where the line
between them should be drawn. Generally the epigeous habit and the
presence of forcible discharge spores in the Pezizales are contrasted to
the hypogeous habit and the lack of a mechanism for forcible spore
discharge in the Tuberales. The simplest forms (Hydnocystis, Hydnotrya)
are cupulate or wholly spherical with a central cavity lined with asci and
paraphyses. The hymenium becomes convoluted into labyrinthine cham-
bers in higher forms (Hydnotrya, Genea). The paraphyses tips often
form an extra layer of tissue over the hymenium (Genea, Petchiomyces),
and in more advanced forms completely fill the chambers with tissue
(Tuber). The work of Fischer (1938) in Engler and Prantl was the
standard treatment for this group for many years. Gilkey (1939), who
treated the North American species, at first placed all the Tuberales in
one family, the Tuberaceae, but later (Gilkey, 1954) she followed Fischer
in breaking the order into three families, still including both Pseudo-
tuberaceae and Eutuberaceae sensu Knapp (1950) in the Tuberaceae.
Malencon (1938) divided the order using the orientation of the ascocarp
and of the hymenium with respect to the soil surface as his criteria, i.e.
the interae, superae, and ubiquarieae. Burdsall (1968) felt that this
division was extremely artificial since it takes into account only the gross
morphology. He felt that the most logical breakdown of families of the
Tuberales was that of Knapp (1950) in which four families, the Hydno-
tryaceae, Geneaceae, Tuberaceae, and Terfeziaceae were recognized. They
all lacked a definite method of ascus dehiscence, presumably because
natural selection for such a mechanism is inoperative in their specialized
habitat. All evidence seems to point to their origin from the Pezizales.
Burdsall's (1965) discovery of spore "puffing" from operculate asci in
Geopora added more weight to this theory and provoked skepticism
about the validity of the present classification of it in the Tuberales. In
a continued study of Geopora and related genera, Burdsall (1968) showed
that Hydnocystis is a monotypic genus in the Geneaceae of the Tuberales.
Sepultaria, of the Pezizales, was shown to be congeneric with the older
genus Geopora (Tuberales by most). Two species of Sepultaria and several
in Hydnocystis were transferred to Geopora, while H. convoluta McA1p.
was transferred to Peziza.
If we go on the above premises that natural selection for opercula
is inoperative, we can see that selection for features associated with ascal
arrangement and spore discharge would be similarly effected. To show
natural relationships within this Pezizales-Tuberales complex, it will
perhaps be necessary to turn to cytological and cytochemical evidence.
Although morphological features may be drastically changed, the nuclear
condition of vegetative and reproductive cells and the basic chemical

nature of such cells may be unaltered. Yet, the dividing line between
the two perhaps will always be open to question.
A Synoptic Key to the Orders and Families of Discomycetes
1. Asci inoperculate, opening by an apical pore; spores mostly ellipsoid, asym-
metrical, sometimes filiform to acicular.
2. Asci cylindrical with a thickened apical cap and thread-like pore; spores
filiform, often segmenting. OSTROPALES.
a. A single family. Ostropaceae.
2. Asci and spores not as above.
3. Hymenium typically formed in an open disc, sometimes naked or in a
stroma, usually superficial or becoming so, without a dark covering
layer. HELOTIALES.
a. Ascocarps apothecoid or perithecoid, turbinate, barrel-shaped to flask-
shaped; asci deliquescing, leaving a dry mass of ascospores. Caliciaceae.
a. Asci and ascocarps not as above.
b. An excipulum absent or poorly defined; palisade of asci of indefi-
nite extent, or stromatic, variously rupturing the host epidermis.
c. Palisade of asci with indeterminant growth, forming irregular
patches on dead wood or bark. ,4scocorticiaceae.
c. Ascocarps simple, with a poorly differentiated basal stratum giv-
ing rise to asci, becoming erumpent by an irregular tear or
circumscissle scale, causing needle blights. Herniphacidiaceae.
b. A well defined ascocarp with excipulum and paraphyses.
d. Apothecia clavate, stipitate, or pileate; spores elongate to fili-
form; paraphyses often brown; mostly soil inhabiting. Geoglossaceae.
d. Apothecia eupulate to lenticular, stalked or sessile.
e. Apothecia superficial, short-stalked, soft-fleshy, waxy, trans-
parent, hyaline to yellow or red, sometimes darker; asci
usually minute. Orbiliaceae.
e. Not with the above combination.
f. Apothecia erumpent or sessile, dark, rarely yellowish to red,
cartilaginous, leathery, or fleshy; excipulum soft, composed
of subglobose cells. Dermateaceae.
I. Excipulum formed of elongated or prismatic cells.
g. Apothecia hairy, mostly bright colored; excipulum of a
textura prismatica to textura angularis. Hyaloscyphaceae.
g. Apothecia smooth or only minutely downy; excipulum
typically of a textura intricata.
h. Apothecia arising from sderotia or stomatised areas
of host; mostly plant parasites. Sclerotiniaceae.
h. Apothecia not arising from sclerotia, most bright-
colored or darker; excipulum usually composed of
parallel hyphae. Helotiaceae.
3. Hymenium formed within a stroma, rupturing to expose the asci; an
upper stromatic layer present. PHACIDIALES.
a. Ascal pore bluing in iodine; ascocarp composed of vertical hyphae,
without radial hyphae; without preformed lines of dehiscence. Phacidiaceae.
a. Ascal pore not bluing in iodine; ascocarp layers of radiating hyphae;
often with preformed lines of dehiscence.
b. Monoapothecial stromata immersed in host tissue, stromata poorly
developed, of two layers. Hypodermataceae.
b. Pluriapothecial stromata superficial to slightly immersed; stromata

well developed. Rhytisrnaceae.
Asci mostly operculate, sometimes with an oblique lid (suboperculate), a
bilabiate split or irregular tear.
4. Numerous apothecia imbedded in a common, gelatinous to fleshy stroma;
growing externally on the twigs of Nothofagus. CYTTARIALES.
a. A single family. Cyttariaceae.
4. Apothecia not imbedded in a common stroma. PEZIZALES.
a. Asci suboperculate, thick-walled, long cylindrical above, narrow, flexuous,
hypha-like at the base; apothecia corky, leathery, subgelatinous but
never truly fleshy; lignicolous or rarely terricolous. Sarcoscyphaceae.
a. Asci operculate or sometimes with bilabiate split or irregular tear; flesh
brittle to firm; occurring on various substrates.
b. Asci not bluing in iodine; ascospores hyaline to pale yellow (if pig-
mented see Ascobolaceae); ectal and medullary excipulum differen-
tiated, the inner area of a texture angularis to textura globulosa.
c. Ascospores tetranucleate or plurinucleate; receptacle broadly effused
or sponge-like, variously shaped, cupulate to stipitate; terricolous.
d. Ascospores tetranucleate, smooth or rarely reticulate, containing
1-3 oil guttules. Helvellaceae.
d. Ascospores plurinucleate, smooth, without oil guttules, but with
numerous small external, polar globules. Morchellaceae.
c. Ascospores uninucleate; apothecia minute to large, lenticular, cupu-
late, sessile or rarely substipitate, smooth or with distinct hairs.
e. Apothecia very small, gregarious, densely crowded to confluent,
orange or pinkish red, lenticular, seated on a well formed
subiculum on recently sterilized substrates. Pyronernataceae.
e. Not with the above characteristics.
f. Apothecia minute to small, scattered to gregarious, mostly
translucent, white, or dull colored; ectal and medullary
excipulum poorly differentiated, varying from textura an-
gularis, epidermoidea, to globulosa; asci one to many per
apothecium, subcylindric to broadly clavate to saccate; as-
cospores 8 to thousands per ascus, uniseriate, biseriate, or
irregularly disposed, hyaline, smooth, without oil droplets,
but with DeBary bubbles in some. Thelebolaceae.
f. Not with the above characteristics.
g. Apothecia small to large, discoid, cupulate or ear-shaped,
dull whitish or brownish, often externally clothed with
tufts of short hyphoid hairs; ascospores mostly smooth
walled, hyaline, guttulate or without; paraphyses septate,
mostly straight, rarely apically curved or hooked, hyaline
or with granular matters, not greening in iodine. Otidiaceae.
g. Apothecia variable in size, mostly sessile, or rarely subsessile,
cup-shaped or lenticular; hymenium colored yellow, orange,
red, brown, rarely white or ivory; spores smooth or orna-
mented; paraphyses straight or curved, filled with carote-
noid pigments. Aleuriaceae.
b. Asci bluing in iodine, if not spores with dark thick walls; excipulum
variable.
h. Ascospores hyaline or rarely brown colored, mostly thin-walled,
smooth or with callose-pectic markings or apiculi. Pezizaceae.
h. Ascospores dark colored and thick-walled, with purple or brown
markings of extra sporal or vacuolar origin. Ascobolaceae.
LITERATURE CITED
ALEXOPOULOS, C . j . 1962. Introductory Mycology. 613 pp. John Wiley & Sons. New
York.
, and E. E. BUTLER. 1949. Conidia like structures in Plectania coccinea.
Mycologia 41: 180-182.
ARAGNO, M. 1967 9 Formation et 6volution de l'asque chez Rhytisma acerinum (Pers.)
Fr. Ber. sehweiz, botan. Ges. 77: 173-186.
ARNAUD, G. 1930. Les Ast~rinees. V. l~tude sur les champignons parasites: Caliciac6s,
Hemisphaeriac6es, etc. Ann. Epiphyt. 16: 235-302.
ARPIN, N. 1968. Les Carotdnoides de Discomyc6tes: Essai Chimiotaxonomique. Thbses
527. Univ. Lyon. 170 pp. Villeurbanne, France 9
Aax, J9 A. YON, and E. M/SLLER. 1954. Die Gattungen de amerosporen Pyrenomyceten.
Beitr/ige zur Kryptofl. Schwiez 11 (1): 1-4349
BARKER, P. T . B . 1904. Further observations on the ascocarp of Rhyparobius. Rept.
Brit. Assoc. Adv. Sci. Cambridge 1904: 825-8269
BATRA, L. R. 1960. T h e species of Ciborinia pathogenic to Salix, Magnolia, and
Quercus. Amer. J. Botany 47: 819-827.
1961a. New species of discomycetes from India. I. Mycologia 52: 524-527.
9 1961b. New species of discomycetes from India. II. Mycologia 52: 665-6679
, and S. W. T. BATRA. 1963. Indian Discomycetes. Univ. Kansas Sci. Bull. 44:
109-256.
, and R. P. KORF. 1959. T h e species of Ciborinia pathogenic to herbaceous
angiosperms. Amer. J. Botany 46: 441-450.
BAYLIss-ELLIOTT, J. S. 1927. Studies in Discomycetes. IV. Trans. Brit. Mycol. SOc.
12: 290-302.
BELL~MERE, A. 1958 9 Quelques observations SUE le dfivelopment de l'apothficie d'un
discomycbte inopercul~ Cyathicula coronata (Bull.) DeNot. Bull. Soc. Mycol.
France 74: 70-93.
1960. SuE les asques des Ostropales. Bull. SOc. Mycol. France 76: 69-82.
BENEmX, E. H. 1962. Zur polyphyletischen Herkunft dee Helvellaceen ss. lat. Zeit.
Pilzk. 27: 93-102.
BERGMAN, P9 and L. SHANOR. 1957. A new species of Streptotheca. Mycologia 49:
879-883.
B~RTHET, P. 1961. Variation de nombre des noyaux darts les articles mycfiliens des
Discomyc~tes, en rapport avee la systfimatique et la phylogfinie. C. R. Acad. Sci.,
Paris 252: 3855-3857.
9 1963. Le hombre des noyaux dans la spore et son int6r~t pour la syst~m-
atique des Discomycbtes opercul6s. C. R. Acad. Sci., Paris 256: 5185-5186.
1964a. Formes conidiennes de divers Discomyc~tes. Bull. Soc. Mycol.
France 80: 125-149.
1964b. Essai biotaxonomique SUE les Discomyc~tes. Th~se, 160 pp., Univ.
Lyon.
1966. L'appareil conidien de Trichophaea confusa (Cooke) Berthet, nov.
comb. Bull. Soc. Mycol. France 82: 473474.
B~ssEv, E. A. 1950. Morphology and taxonomy of fungi. 791 pp., Philadelphia.
BIFFIN, R . H . 1901. On the biology of Bulgaria polymorpha Wett. Ann. Botany 15:
119-134.
Bxs~s, G . N . 1956a. Studies on the genetics of Ascobolus stercorarius (Bull.) Schroet.
Bull. Torrey Botan. Club 83: 35-61.
9 1956b. Sexuality in Ascobolus stercorarius. I. Morphology of the asco-
gonium; plasmogamy; evidence for a sexual hormonal mechanism. Amer. J.
Botany 43: 389-394.
9 1957. Sexuality in Ascobolus stercorarius. II. Preliminary experiments on
various aspects of the sexual process. Amer. J. Botany 44: 436-443.
, and J. R. RAeER. 1963. Heterothallism and seuuality in Ascobolus
stercorarius. Amer. J. Botany 50: 880-891.
BLACKMAN, V. H., and H. C. I. FRASER. 1906. On the sexuality and development of
the ascocarp of Humaria granulata. Proc. Royal Soc. Botany, London 77:
354-368.
BLOt)GEar, E. C. 1936. The anthracnose of currents and gooseberry caused by
Pseudopeziza ribis. Phytopathology 26: 115-152.
Bov_oIJN, K. B. 1934. Two new Malaysian genera of Discomycetes. Bull. Jard.
Botan. Buitenzorg. III. sea'. 13: 478-483.
9 1940. T h e genus Pyronerna in the Netherlands Indies (Fungi, Ascorn.)
Bull. Jard. Botan. Buitenzorg III 16: 272-2759
BOUDIER, J. L9 E. 1869. Mdmoire sur les Ascobolds. Ann. Sci. Nat. Bot. V. 10: 191-2689
1879. On the importance that should be attached to the dehiscence of asci
in the classification of the Discomycetes. Grevillea 8: 4549.
9 1885. Nouvelle classification naturelle des Discomycbtes charnus connus
gdndralement sous le nom de Pezizes. Bull. Soc. Mycol. Ft. 1: 91-120.
9 1907. Histoire et classification des Discomyc~tes d'Europe. I-VII: 1-223.
P. Klincksich, Paris.
BROWN, W . H . 1911. The development of the ascocarp of Lachnea scutellata. Botan.
Gaz. 52: 275-3059
BRUMMELEN, j. VAN. 1962. Studies on Discomycetes 1. Types of species of .4scobolus
and Saecobolus in Spegazzini's herbarium 9 Persoonia 2: 195-1999
1967. A world-monograph of the genera ilscobolus and Saccobolus (Asco-
mycetes, Pezizales). Persoonia (Suppl.) 1: 1-260.
BURaSALL, H. H. 1965. Operculate asci and puffing of ascospores in Geopora
(Tuberales). Mycologia 57: 485-487.
1968. A revision of the genus Hydnocystis (Tuberales) and of the hy-
pogeous species of Geopora (Pezizales). Mycologia 60: 496-525.
CAIN, R. F., and N. A. HASXlNC.S. 1956. Studies of soil fungi. II. A new species of
Sphaerospora with a Botrytis-like stage9 Canad9 J. Botany 34: 360-376.
CARROLL, G . C . 1969. A study of the fine structure of ascosporogenesis in Saccobolus
kerverni. Arch. Mickrobiol. 66: 321-339.
CARRUTHERS, D. 1911. Contributions to the cytology of Helvella crispa. Ann. Botany
25: 243-252.
CHADEFAUO, M. 1940a. Etudes d'asques. I. Les asques et les ascospores de Bulgaria
inquinans. Rev. Mycol. 5: 87-101.
9 1940b. Le reseau ornemantal de spores et l'appareil apical des asques chez
Peziza aurantia. C. R. Acad. Sci., Paris 211: 650-660.
9 1942. Etudes d'asques. II. Structure et anatomie compar~e de l'appareil
apical de asques chez divers Disco- et Pyrdnomyc&es. Rev. Mycol. 7: 57-88.
9 1943. Sur les divers types d'dldments dangeardiens de Ascomyc6tes, et sur
la formation des asclues chez la Pdzize Pustularia catinus. La. Rev. Sci. 81: 77-80.
9 1944. Etudes d'asques. IV. L'asques hdmiopercule dd Leotia lubrica. Rev.
Mycol. 9: 3-13.
9 1946a. Les asques para-operculds et la position systdmatique de la Pdzize
Sarcoscypha coccinea. C. R. Acad. Sci., Paris 222: 753-755.
1946b. Sur les asques ~t nasse apicale. Bull. Sci. Bot. de France. 93: 128-130.
1949. Observations sur les asques de quelques Morchellaceae. Le Botaniste
341 75-85.
9 1960. Les vdgdtaux non vasculaires (Cryptogamie). In Chadefaude, M.,
and L. Emberger. Traitd de Botanique. 1018 pp. Librairie Masson, Paris.
9 1969. Remarques sur les patois, L'appareil apical et les reserves nutritives
de asques. Osterr. Bot. Zeit. 116: 181-202.
- - , M. LETROUIT-GALINOU, and M. JANEX-FAVRE. 1967. Sur l'origine phylo-
gdndtique et l'6volution des Ascomycbtes des Lichens. Soc. Bot. France, Colloque
sur les Lichens. 1967: 79-111.
CHAV,OON, C. E. 1946. Fungi Domingenses novi vel minus cogniti. I. Farlowia 2:
455-473.
Cnl~STIANSEN, M. P. 1963. Danish species of the genus Coryne. Friesia 7: 75-85.
CI~USSEN, P9 1905. Zur Entwicklungsgeschichte der Ascomyceten. Boudiera. Botan.

Ztg. 63: 1-28.
9 1912. Zur Entwicklungsgeschichte der Ascomyceten. Pyronema con[luens.
Zeit. Bot. 4: 1~o4.
CLEMENTS, F . E . 1909. The genera of fungi. 227 pp. Minneapolis.
9 and C. L. SHEAR. 1931. The genera of fungi. 496 pp. New York.
COOKE, J. C., and M. E. BARR. 1964. The taxonomic position of the genus Thelebolus.
Mycologia 56: 763-7699
COOKE, M. C. 1879. Mycographis, seu icones fungorum. 1. Discomycetes. 267 pp.
London.
1892. Handbook of Australian fungi. London.
CORNER, E. j. H. 1930a. Studies in the morphology of discomycetes. III. The
Clavuleae. Trans. Brit. Mycol. Soc. 15: 107-120.
9 1930b. Study in the morphology of Discomycetes. IV. The evolution of
the ascocarp. Trans. Brit. Mycol. Soc. 15: 121-134.
CROUAN, P. L., and H. M. CROUAN. 1857. Note sur quelques Ascobolus nouveaux et
sur une bspece nouvelle de Vibrissea. Ann. Sci. Nat. (Bot.) IV 7: 173-178.
CUTTING, E. M. 1909. On the sexuality and development of the ascocarp in Asco-
phanus carneus Pers. Ann. Botany 23: 339-4169
DANGEARD, P. A. 1903. Sur le genre Ascodesmis. Le Botaniste 9: 33-35.
1907. L'origine du p6rithbce chez les Ascomyc6tes. Deuxi~me Partie.
Recherches sur le developpement du p6rith6ce chez les Ascomyc~tes. Le
Botaniste I0: 1-3859
DARKER, G . D . 1932. The Hypodermataceae of conifers. Contr. Arnold Arbor9 Har-
vard Univ. 1: 1-131.
9 1963. A new phragmosporous genus of the Hypodermataceae. Mycologia
55: 812-818.
9 1967. Revision of the genera of the Hypodermataceae. Canad. J. Botany
45: 1399-1444.
DAVlDSON, R. W. 1950. Urnula craterlum is possibly the perfect stage of Strumella
coryneoideae. Mycologia 42: 735-742.
DEBARv, A. 1863. Uber die Fruchtenwicklung der Ascomyceten. 38 pp. Englemann,
Leipzig.
1866. Morphology und Physiologie der Pilze, Elechten und Myxomyceten.
316 p9 Leipzig.
DELXTSCR, H. 1926. Zur Entwicklungsgeschite der coprophilen Ascomyceten: Lasio-
bolus pulcherrimus Crouan: Humaria anceps Rehm, var. aurantiaca, n. var.; und
Sporormia leporina Niessl. Diss. 40 p. Leipzig.
DENISON, W . C . 1961. Some species of the genus Scutellinia. Mycologia 51: 605-635.
1963. A preliminary study of the operculate cup-fungi of Costa Rica.
Rev. Biol. Trop. 11: 99-129.
1964. The genus Cheilymenia in North America. Mycologia 56: 718-737.
9 1965. Central American Pezizales. I. A new genus of the Saccoscyphaceae.
1967. Central American Pezizales. II. The genus Cookeina. Mycologia
59: 306-317.
1969. Central American Pezizales. III. The genus Phillipsia. Mycologia
61: 289-304.
DENNIS9 R. W. G. 1949. A revision of the British Hyaloscyphaceae with notes on
related European species. Commonw. Mycol. Instn., Mycol. Papers 32: 1-97. Kew.
1950. Karsten's species of Mollisia. Kew Bull. 1950: 171-178.
1954a. Chloroscypha Seaver in Britain. Kew Bull. 1953: 410.
9 1954b. Some inoperculate Discomycetes of tropical America. Kew Bull.
9: 289-348.
9 1955. A note on the spiny-spored species of Lamprospora. Kew Bull. 1955:
571-572.
1956. A revision of the British Helotiaceae in the Herbarium of the
Royal Botanic Gardens, Kew, with notes on related European species9 Mycol.
Papers 62: 1-262.
9 1958. Critical notes on some Australian Helotiales and Ostropales. Kew
Bull 1958: 321-358.
1960. British cup fungi and their allies. An introduction to the Asco-
mycetes. 280 pp. The Ray Soc., London 9
9 1962a. New or interesting British Helotiales. Kew Bull. 16: 317-327.
1962b. A reassessment of Belonidium Mont. 8c Dur. Persoonia 2: 171-1919
1963. A redisposition of some fungi ascribed to the Hyaloscyphaceae. Kew
Bull. 17: 319-379.
1964. Remarks on the genus Hymenoscyphus S. F. Gray, with observations
on sundry species referred by Saccardo and others to the genera Helotium,
Pezizella, or Phialea. Persoonia 3: 29--80.
1968. British Ascomycetes. 455 pp. J. Cramer. Lehre.
, and R. P. KORF. 1958. A Japanese species of Sphagnicola Velenovsky. Kew
Bull. 1958: 321-358.
DE NOTARIS, G. 1864 9 Proposte di alcune rettificazione al profilo dei Discomiceti.
Comm. Soc. critt, ital. l: 357-388.
DnARNE, C. G. 1965. Taxonomic investigations on the discomycetous genus Lach-
nellula Karst. Phytopath. Zeit. 53: 101-144,
DILLEmUS. J. J. 1719. Catalogus plantarum sponte circa Gissam nascentium. Cure
a p p e n d i c e . . . Francofurti ad Moenum.
DIssINe, H. 1964. Studies in artic and subartic Discomycetes I. The genus Helvella.
Botan. Tidskr. 60: 108-128.
9 1966a. The genus Helvella in Europe with special emphasis on the species
found in Norden. Dansk Botan. Ark. 25: 1-172.
9 1966b. A revision of collections of the genus Helvella L. ex St-Amans
emend. Nannf. in the Boudier Herbarium. Rev. Mycol. 31: 189-224.
, and M. LANCE. 1967. Notes on the genus Helvella in North America.
, and J. A. NANNVELDT. 1966. Helvella cupuliformis sp. nov., H. villosa
(Hedw. ex O. Kuntze) comb. nov., H. macropus (Pers. ex Fr.) Karst., and their
allies. Svensk Botan. Tidskr. 60: 325-337.
Do~E, B. O. 1912. Methods of culture and the morphology of the archicarp in
certain species of the Ascobolaceae. Bull, Torrey Botan. Club 39: 139-197.
9 1922. A Lachnea with a botryose conidial stage. Bull. Torrey Botan. Club
49: 301-305.
1936. Facultative and obligate heterothallism in Ascomycetes. Mycologia
281 399-409.
1937. The conidial stage of Peziza pustulata. Mycologia 29: 651-655.
, and F. J. 8eaver. 1946. Species of Ascobolus for genetic study. Mycologia
38: 639-651.
DRAYTON, F . L . 1932. The sexual function of the microconidia in certain discomycetes.
1934. The sexual mechanism of Sclerotinia gladioli. Mycologia 26: 46-72.
- - , and J. W. GROVES. 1943. A new Sclerotinia causing a destructive disease of
bulbs and legumes. Mycologia 35: 517-528.
, and - - . 1952. Stromatinia narcissi, a new sexually dimorphic disco-
mycete. Mycologia 44: 119-140.
DUFF, G. H. 1922. Development of the Geoglossaceae. Botan. Gaz. 74: 264-291.
DURANn, E. j. 1900. The classification of the fleshy Pezizineae with reference to
the structural characters illustrating the bases of their division into families.
Bull. Torrey Botan. Club. 27: 463-495.
1901. Studies in North American discomycetes. I. The genus Holwaya
Sacc. Bull. Torrey Botan. Club 28: 349-355.
1908. T h e Geoglossaceae of North America. Ann. Mycol. 6: 387477.

9 1921. New or noteworthy Geoglossaceae. Mycologia 13: 184-187.
DUaAND, F. 1968. Sur l'ontog6nie du Discomycete opercul6 Antracobia nitida Boudier.
C. R. Acad. Sci., Paris 267: 2316-2318.
ECKBLAO, F.-E. 1957. Norges Sarcoscyphaceer. Blyttia 15: 2-12.
13~-11963"..~v. Contributions to the Geoglossaceae of Norway. Nytt. Mag. Bot. 10:
1968. T h e genera of the Operculate Discomycetes. A reevaluation of their

taxonomy, phylogeny and nomenclature. Nytt. Mag. Bot. 15: 1-191.
ELLIOTr, M9 E. 1962. Streptotinia caulophylli sp. nov. produced in culture. Canad.
J. Botany 40: 1197-1201.
1964a. Sclerotinia artica n. sp. on Carex from Northern Canada. Canad.
J. Botany 42: 1065-1070.
9 1964b. Self fertility in Botryotinia pori. Canad. J. Botany 42: 1393-1395.
9 1967. Rutstroemia cuniculi, a coprophilous species of the Sclerotiniaceae.
Canad. J. Botany 45: 521-524.
ESSER, K. 1967. Die Incompatibilit~it. Handbuch der Pflanzenphysiologie (W. Rhu-
land ed.) Vol. 18.
, and R. KUENEN. 1965. Genetik der Pilze. 497 pp. Springer-Verlag, Berlin.
FERGUS, C . L . 1960. A note on the occurrence of Peziza ostracoderma. Mycologia 52:
959-961.
FINK, B. 1915. T h e ascomycetes of Ohio. I. Ohio Biol. Sur. Bull. 5, Vol. II. 70 pp.
FISCHER, E. 1938. Tuberineae, In Engler, A. u n d K. Prantl., Die natiirlichen
Pflanzenfamilien II, 5b(8): 1-42.
FITZPATRXCK, H. M. 1918 9 Sexuality in Rhizina undulata. Botan. Gaz. 65: 201-226.
1920. Monograph of the Coryneliaceae. Mycologia 12: 206-267.
1942. Revisionary studies in the Coryneliaceae. Mycologia 34: 464-514.
, and R. P. KORF. 1959. Lectures on mycology, Ascomycetes. Plant Pathology
Department, Cornell University, Ithaca, N. Y. (Mimeo).
FRASER, H. C . I . 1907. On the sexuality and development of the ascocarp in Lachnea
stercorea. Ann. Botany 21: 349-360.
1913. T h e development of the ascocarp in Lachnea cretea. Ann. Botany
27: 553-563.
- - , and S. W. S. BROOKS. 1909. Further studies on the cytology of the ascus.
Ann. Botany 23: 537-549.
- - , and E. J. WELSFOag. 1908. Further contributions to the cytology of the
ascomycetes. Ann. Botany 22: 465-477.
From, E.M. 1821. Systema mycologicum. I. Lundae.
1822. Systema mycologicum. 2. Lundae.
1849. Summa vegetabilium Scandinaviae. 259-572.
FUCKEL, L. 1869. Symbolae mycologicae, Beitr~ige zur Kenntniss der rheinischen
Pilze. Jahrb. Nassauischen Ver. Naturk. 23-24: 1-459.
FUNK, A. 1963. Studies in the genus Caliciopsis. Canad. J. Botany 41: 503-543.
GAMUNm. I . J . 1956. E1 genero Scutellinia en Argentina 9 Contr. cient. Fac. Co. exact.
fis. nat. Univ. B. Aires (Bot.) 1: 69-889
1957. El genero Cookeina en la Republica Argentina 9 Boln Soc. argent 9
Bot. 6: 212-2229
1959. Addenda alas especies argentinas de Cookeina Kuntze. Boln Soc. argent.
Bot. 7: 201-204.
1960. Discomycetes operculados de la Argentina, familias Pezizaceae y
Humariaceae. Lilloa 30: 257-338.
1964. "Discomycetes" operculados del Parque Nacional Nahuel Huapi,
Argentina. Darwiniana 12: 568-606.
and M. E. RANALLt. 1964. Estudio sistem~itico y biol6gico de las Asco-
bol~tceas de Argentina 9 I. Nova Hedwigia 7: 517-5339
and . 1966. Estudio sistem~ttico y biol6gico de las Ascobol~tceas de
Argentina. II. Nova Hedwigia 10: 337-3669
KIMBROUGH" CLASSIFICATION OF DISCOMYCETES 153
GAUMANN, E . A . 1926 9 Vergleichende Morphologie der Pilze. 626 pp. Jena.

1964. Die Pilze, Grundzfige ihrer Entwicklungsgeschichte u n d Morphologie.
541 pp. Birkh~iuser Verlag, Basel und Stuttgart.
, and C. W. DODGE. 192'89 Comparative morphology of fungi 9 I-XIV: 1-7019
New York.
GXLKEY, H. M. 1939. Tuberales of North America. Oregon State Monogr. (Stud.
Botany) 1: 1-63.
9 1954. Tuberales. No. Amer. Flora II, 1: 1-29.
GORDON, C. C. 1966. Ascocarp centrum ontogeny of species of Hypodermataceae of
conifers9 Amer. J. Botany 53: 319-327.
9 1968. Ascocarpic centrum ontogeny of species of Hypodermataceae of
conifers. II. Amer. J. Botany 55: 45-529
GRADDON, W . D . 1965. A guide to lignicolous species of Apostemidium. Trans. Brit.
Mycol. Soc. 48: 639-646.
GREIS, H. 1940. Befruchtungsarten bei Morchella. Jahrb. Wissen. Botan. 89: 245-253.
GRELET, L . J . 1942. Les Discomyc&es de France d'apres la Classification de Boudier.
Rev. Mycol. 7: 3-22.
1943. Ibid. 8: 3-25.
1944. Ibid. 9: 14-35, 78--99.
1945. Ibid. 10: 96-116.
1946. Ibid. 11: 89-106.
1947. Ibid. 12: 24-36, 45-689
1948. Ibid. 13: 22-24, 105-134.
1949. Ibid. 14: 26-57.
1950. Ibid. 15: 29-58.
1951. Ibid. 16: 11-26, 80-100.
1953. Ibid. 18: 24-28, 200-220.
1954. Ibid. 19: 138-166.
1956. Ibid. 21: 14-39, 146-173.
1957. Ibid. 22: 26-52, 166-182.
1959. Ibid. 24: 81-92, 234-263.
GRE~MEN, j. 1949. De imperfecte vorm van Desmazierella acicola Lib. Fungus
(Wagen.) 19: 32-35.
1950. Her genus Belonioseypha sensu Rehm en de indentiteit van B.
ciliatospora (Fuckel) Rehm en Helotium scutula (Pers. ex Ft.) Karst. Fungus
(Wagen.) 20: 7-17.
1952. Het genus Pyrenopeziza. Levenswijze en cultuur van P. plataginis.
Fungus (Wagen.) 22: 22-249
9 1954. Taxonomical notes on Mollisiaceous fungi. I. A study on some
Dutch species growing on Rubus stems9 Fungus (Wagen.) 24: 1~/~.
9 1955. Taxonomical notes on Mollisiaceous fungi. II. Some caulicolous
Mollisia species inhabiting various hosts, mainly Compositae. Fungus (Wagen.)
25: 1-12.
9 1956a. Taxonomical notes on Mollisiaceous fungi 9 III. T h e polyphagous
species Mollisia pastinaceae Nannfeldt. Fungus (Wagen.) 26: 28-319
1956b. Taxonomical notes on Mollisiaceous fungi 9 IV. Species inhabiting
prevous year's stems of Epilobium and Ulmaria. Fungus (Wagen.) 26: 32-37.
1957. Taxonomical notes on Mollisiaceous fungi 9 V. On some species
described by Velenovsky. Fungus (Wagen.) 27: 30-33.
1958. Taxonomical notes on Mollisiaceous fungi 9 VI. T h e genus Pyre-
nopeziza Fuckel. Fungus (Wagen.) 28: 37-46.
9 1963. Preliminary notes on Godronia and association with conidial fungi 9
Nova Hedwigia 6: 15-20.
GRIFFITHS, E. 1959. T h e cytology of Gloeotinia temulenta (blind seed disease of Rye-
grass). Trans. Brit. Myeol. Soc. 42: 132-148.
GROVES, J . w . 1939. Some Pezicula species and their conidial stages. Canad. J. Res.
17: 125-143.
9 1940. Some Dermatea species and their conidial stages. Mycologia 32:
736-751.
1946. North American species of Dermea. Mycologia 38: 351-431.
1952. The genus Tympanis. Canad. J. Botany 30: 571-651.
1954. The genus Durandiella. Canad. J. Botany 32: 116-144.
1965. The genus Godronia. Canad, J. Botany 43: t19,6-1276.
1967. The genus Pragmopara. Canad. J. Botany 45: 169-181.
, and C. A. BOWERMAN. 1955. The species of Ciborinia on Populus. Canad.
J. Botany 33: 577-590.
- - , and F, L. DRAYTON, 1939. The perfect stage of Botrytis cinerea. Mycologia
31: 485-489.
, and M. E. ELLIOT;. 1961. Self-fertility in the Sclerotiniaceae. Canad. J.
Botany 39: 215-231.
- - , and S. C. HOARE. 1953. The Helvellaceae of the Ottawa district. Canad.
Fld. Nat. 67: 95-102.
, and C, A. LOVELANV. 1953. T h e connection between Botryotinia fucketiana
and Botrytis cinerea. Mycologia 45: 415-425.
, and D. E. WELLS. 1956. The genus Retinocyclus. Mycologia 48: 865-8719
, and D. E. WILSON9 1967. The nomenclatural status of Coryne. Taxon
16: 35-41.
GRUND, D. W., and K. A. HArriSON. 1967. Nova Scotian fungi. Geoglossaceae.
Canad. J. Botany 45: 1625-1641.
GWYNNE-VAUGHAN,H. C.I. 1922. Fungi. 232 pp. Cambridge.
, and H. S. WILLIAMSON. 1931. Contributions to the study of Pyronema
confluens. Ann. Botany 45: 355-371.
, and . 1932. The cytology and development of AscoboIus mag-
niIicus. Ann. Botany 46: 653-670.
HAHN, G. G., and T. T. AwRs. 1934. Dasyscyphae on conifers in North America.
III. Dasyscypha pini. Mycologia 26: 479-501.
HAKEUER, N. 1964. Bidrag till Sveriges Svampflora II. Geoglossaceer. Svensk Bot.
Tidskr. 58: 337-343.
HANLIN, R. T. 1965. Lamprospora planchonis from Georgia. Mycologia 57: 131-
134.
HARPER, R . A . 1900. Sexual reproduction in Pyronema con[luens and the morphology
of the Ascocarp. Ann. Botany 14: 321-400.
H~ALn, F. D., and F. A. WOLF. 1910, The structure and relationship of Urnula
geaster. Botan. Gaz. 49: 182-188.
HEIM, R. 1966. Quelques ascomycetes remarquables. V. Morilles tropicales (Pacifi-
que sud.). Bull. Trimest. 8oc. Mycol. Fr. 82: 442-449.
HENNINgS, P. 1900. Fungi. Monsunia I: 1-38.
HICC,INS, B. B. 1914. Contribution to the life history and physiology of Cylindro-
sporium on stone fruits. Amer. J. Botany 1: 145-173.
HlascH, H . E . 1950. No brachymeiosis in Pyronema conIluens. Mycologia 42: 301-305.
H6nNEL, F. VON. 1903. Mycologische Fragmente. Ann. Mycol. 1: 396-399.
1907. Fragmente zur Mykologie. Sitzb. k. ak. Wissen. Wein. 116:135 pp.
1909. Fragmente zur Mykologie. Sitzb. k. ak. Wissen. Wein. 118: 275-352.
1917a. Mycologische Fragmente, CXX-CXC. Ann. Mycol. 15: 292-383.
1917b. System der Phacidiales. Ber. Deutsch. Botan. Ges. 35: 416-422.
1918. Fragmente zur Mykologie. Sitzb. k. ak. Wissen. Wein, Math.-nat. K1.
I. 127: 549-634.
1920. Uber Pseudopeziza, Pyrenopeziza, Ephelina, und Spilopodia. Ber.
Deutsch. Botan. Ges. 38: 96-101.
HONEY, E. E. 1928. The monilioid species of Sclerotinia. Mycologia 20: 127-157.
HowAm'IL W. O., and H. G. CHIePINDALE. 1931. Notes on the biology of Coryne
sarcoides Jacq. and C. urnalis Nyl. Mere. Proc. Manchester Lit. Phil. Soc. 75:
47-60.
HUGHES, S . J . 1951. Studies on microfungi. IX. Calcarisporium, Verticicladium and

Hans[ordia. C. M. I. Mycol. Pap. 43: 1-25.
HUTTER, R. 1958. Untersuchungen tiber die Gattung Pyrenopeziza Fuckl. Phytopath.
Zeit. 33: 1-54.
IMAI, S. 1940. T h e Geoglossaceae of Norway. Ann. Mycol. 38: 268-278.
1941. Geoglossaceae Japoniae. J. Fac. Agr. Hokkaido Imp. Univ. 45: 155-
264.
1954. Evellaceae Japoniae. Sci. Rep. Yokohama Nat 9 Univ. 2: 1-35.
1956. Contributions ad studia monographica Geoglossacearum. II. III. Sci.
Rep. Yokohama Univ. 1955: 1-II, 1956: 1-8.
ITO, S., and S. IMAI. 1937. Fungi of the Bonin Islands9 II. Trans. Sapporo Nat, Hist.
Soc. 15: 52-59.
Joy~s, S.G. 1935. T h e structure of Lophodermium pinastri (Schrader ex Fr.) Chev.
Ann. Botany 40: 699-728.
KANOUSE, B . B . 1949. Studies in the genus Otidea. Mycologia 41: 660-677.
KARSTEN, P . A . 1869. Monographia Pezizarum fennicarum. Notis. Sallsk. Fauna. F1.
fenn Forh. 10: 99-206.
9 1871. Mycologia fennica. I. Discomycetes. 263 pp. Helsingfors.
KmLMAN, O. 1883. Zur Entwickelungsgeschichte der Ascomyceten. Acta. Soc. Sci.
Fenn. 13: 5-43.
KILLIAN, K. 1918. Morphologie, Biologia, u n d Entwickelungsgeschichte von Crypto-
myces pteridis (Rebent.) Rehm. Zeit. Bot. 10: 49-126.
KILLIAN, C., and V. LmHITE. 1924. Observations sur le genre Lophodermium. C.R.
Hebdomad. Soc. Biol. 2: 574-576.
KXMBROUGH, J. W. 1966a. T h e structure and development of Trichobolus zukalii.
1966b. Studies in the Pseudoascoboleae. Canad. J. Botany 44: 685-704.
9 1969a. North American species of Thecotheus (Pezizeae, Pezizaceae). Myco-
logia 61: 99-114.
, and R. P. K o ~ . 1967. A synopsis of the genera and species of the tribe
Theleboleae ( = Pseudoascoboleae). Amer. J. Botany 54: 9-23.
, E. R. LUcK-ALLEY, and R. F. CAIN. 1969b. lodophanus, the Pezizeae segre-
gate of "Ascophanus" (Pezizaceae). Amer. J. Botany 56: 1187-1202.
KNAPP, A. 1950. Die europaischen Hypogaeen-Gattungen und ihre Gattungenstypen.
Schweiz. Z. Pilzk. 28: 101-118.
KNIEP, H. 1928. Die Sexualitat der niederen Pflanzen, Differenzierung, Verteilung,
Bestimmung u n d Vererbung des Geschlechts bei den Thallophyten. 544 pp.
Jena.
KOBAYASm, T. 1965. Taxonomic notes on Chloroscyphae causing needle blight on
Japanese conifers. Bull. Gov't For. Exp. Sta. Mergura 176: 55-74.
KOBAYASI, Y., N. HIRATSUKA, R. P. KORF, K. TtmAKI, K. AOSmMA, M. SONEDA, and
J. SUGIYAMA. 1967. Mycological studies on the Alaskan Arctic. Ann. Rep.
Inst. Ferment., Osaka. 3: 1-138.
KORV, R. P. 1951. A monograph of the Arachnopezizeae. Lloydia 14: 129-180.
9 1953. T h e new rules of typification as they affect Sarcoscypha and
Velutaria. Mycologia 45: 296-301.
9 1954. A revision of the classification of operculate Diseomycetes (Pezizales).
VIII Congr. Int. Bot., Rapp. ~: Comm. 18-20: 80.
1956. Daleomyces, Durandiomyces and other sparassoid forms of opercu-
late Discomycetes. Mycologia 48: 711-718.
1957a. Nomenclatural notes. II. On Bulgaria, Phaeobulgaria and Sarco-
soma9 Mycologia 49: 102-106.
1957b. Two bulgarioid genera: Galiella and Plectania. Mycologia 49:
107-111.
9 1958. Japanese Discomycete notes I-VIII. Sci. Rep. Yokohama Nat. Univ.
II, 7: 7-35.
1959. Japanese Discomycete Note IX-XVI. Bull. Nat. Sci. Mus9 Tokoyo.
45: 389-400.
9 1960a. Nomenclatural notes9 IV9 The generic name Plicaria. Mycologia
52: 648-651.
1960b. Jaynea, a new genus of the Pezizaceae. Nagaoa 7: 3-8.
9 1962a. A synopsis of the Hemiphacidiaceae, a family of the Helotiales
(Discomycetes) causing needle-blights of conifers. Mycologia 54: 12-33.
9 1962b. A rare North American discomycete, together with some comments
on the genus Lachnellula, Trans, Mycol. Soc. Japan 3: 47-50.
1963a. Discomycete Flora of Asia, Precursor I: The Scope and Status of
Exploration, Lloydia 26: 16-20.
1969. On four major discomycete works published in 1968. Mycologia
611 442-446.
9 and J. K. RogERs. 1967. A new term, the Schizotype, and the concept of
implicit typification. Taxon 16: 19-23.
KUVVER, E. M. 1902. Studies on Urnula and Geopyxis. Bull. Torrey Botan. Club.
29: 137-144.
LANCV.RON, M. 1952. Pr6cis de mycologie, mycologie g6n6rale, mycologie humaine
et animale, techniques. 2nd Ed. by R. Vanbreuseghem. 703 pp. Paris.
LEGAL, M. 1942. Mode de formation des ornamentations sporales chez quelques
Discomyc~tes opercul~s. C. R. Acad. Sci., Paris. 214: 125-128.
1946a. Mode dfihiscence de asques chez les Cookeina et les Leotia et ses
consequences du point de rue phylogenetique. C. R. Acad. Sci., Paris. 222: 755-
757.
1946b. Les Discomyc6tes subopercul6s. Bull. Soc. Mycol. France. 62: 218-240.
9 1947. Recherches sur les Ornamentations sporales des Discomyc6tes op-
ercul6s. Ann. Sci. Nat., Bot. ser. 11: 73-297.
9 1949. Deux Discomyc~tes mal connu: Ascodesmis nigricans van Tieghem
et ,4scodesmis microscopica (Crouan) Le Gal, non Seaver. Rev. Mycol. I4: 85-99.
1953a. Les Discomyc~tes de Madagascar9 465 pp. Paris.
9 1953b. Les Discomyc~tes de l'herbier Crouan. Rev. Mycol. 18: 73-132.
1957. Le genre Leucoscypha Boud. Bull, Jar. Bot. Bruxelles. 27: 720-728.
1958. Le genre Melastiza Boudier. Bull. Soc. Mycol. France9 74: 149-154.
9 19599 Discomyc~tes du Congo Belge d'apr~s les r6coltes de madame
Goossens-Fontana. Bull. Jard. Botan. de l'Etat. 29: 73-1329
1961. Les Discomycbtes de l'herbier Crouan. Deuxi6me s6rie. Ann. Sci.
Nat. (Bot.) XII, 1: 441-4679
9 1962. Combinaisons nouvelles concernant les genres Galactinia (Cooke)
Boud. emend 9 Le Gal, Scutellinia (Cooke) Lamb. emend I,e Gal et Sarcosorna
Casp. Bull. Soc. Mycol. France 78: 204-216.
9 1963. Valeur taxonomique particuli~re de certains caract~res chez les Disco-
myc6tes sup6rieurs. Bull. Soc. Mycol. France 79: 456-470.
1964. Combinaisons nouvelles concernant le genre Scutellinia (Cooke)
Lamb, emend. Le Gal. Bull. Trimest. Soc. Mycol. France 80: 123-124.
9 1966. Contribution "~ la connaissance du genre Scutellinia (Cooke) Lamb.
emend Le Gal. Bull. Soc. Mycol. France 82: 310-334.
9 and F. MAGENOT. 1956. Contributions h l'6tude des Mollisioid6es. 1. Note
preliminaire; les formes conidiennes. Rev. Mycol. Paris 21: 3-13.
- - , and - - 1958. Contribution ~t l'6tude Mollisioid6es. II. Rev. Mycol.
Paris 23: 28-86.
- - , and ~ 1960. Contribution h l'6tude des Mollisioid6es. III. Rev.
Mycol. Paris 25: 135-314.
- - , and ~ 1961. Contribution "t l'6tude des Mollisioid6es. IV. Rev.
- - , and - - 1966. Contribution h l'~tude des Mollisioid~es. V. Rev.
LINnAU, G. 1897. Pezizineae, Phacidiineae, Hysteriineae. In Engler 8~ Prantl's
Naturl. Pfl-Fam. 1(1): 177-272.
LINr~AEUS, C. 1753. Species Plantarum. 1200 pp. Holmiae.
LOHWAG, H. 1927. Das Oogon als Wesensbestandteil der Geschlechtsorgame im
Pilzreich. Biol. Gen. 3: 699-772.
LUTrRELL, E.S. 1951. Taxonomy of the Pyrenomycetes. Univ. Mo. Stud. 24: 1-120.
9 1955. T h e ascostromatic ascomycetes. Mycologia 47: 511-532.
MAAS GEESTERANUS, R . A . 1955. Notes on Dutch fungi. Fungus 25: 44-52.
9 1965. Geoglossaceae of India and adjacent countries. Persoonia 4: 19-46.
1967. Studies in cup-fungi 9 I. Persoonia 4: 417-425.
MAINS, E. B. 1954. North American species of Geoglossurn and Trichoglossum.
1955. North American hyaline-spored species of Geoglosseae. Mycologia
471 846-877.
9 1956a. T h e relationship of Cudoniella and Helotium. Mycologia 48"
410-419.
9 1956b. North American species of the Geoglossaceae. Tribe Cudonieae.
MALENw G. 1938. Les truffes Europeennes. Rev. Mycol., Mere. Hors-ser. I: 1-92.
MASSEE, G.E. 1895. British fungus-flora. 4: 1-522. London 9
MCCUBBIN, W . A . 1910. Development of the Helvellineae I. Helvella elastica. Botan.
Gaz. 49: 195-206.
MClNTOSH, D. L. 1954. A cytological study of ascus development in Pyronema con-
fluens Tul. Canad. J. Botany 32: 440-446.
MOORE, E . J . 1963. T h e ontogeny of the apothecia in Pyronema dornesticum. Amer.
J. Botany 50: 3744.
9 1965. Ontogeny of gelatinous fungi 9 Mycologia 57: 114-130.
, and R. P. KORF. 1963. T h e genus Pyronema. Bull. Torrey Botan. Club
90: 33-42.
MORAVEC, J. 1968. P~isp~vek k poznfini operkulfitnich discomycetu rodu Cheilymenia
Boud. ~esk~i Mykol. 22: 32-41.
MOREAU, F., and M i r . MORV~U. 1930. Le d6veloppement du p6rith~ce chez quelques
Ascomyc~tes. Rev. G~n. Bot. 42: 65-98.
MOSER, M. 1951. Beitr~ig zur Anatomie der Discomyceten. Das Morchellaproblem.
Sydowia 5: 56-119.
9 1963. Ascomyceten. In H. Gams, Kleine Kryptogamenflora 2a, 147 pp.
Gustav Fischer Verlag. Stuttgart 9
MUNK, A. 1957. Danish Pyrenomycetes. A preliminary flora. Dansk Botan. Ark.
17(1): 1-491.
NANNFELDT, J. A. 1932. Studien tiber die Morphologie und Systematik der nicht-
lichenisierten inoperculaten Discomyceten. Nov. Act. Reg. Soc. Sci. Uppsala. ser.
4, 8: 1-3689
9 1937. Contributions to the mycoflora of Sweden9 4. On some species of
Helvella, together with a discussion of the natural affinities within Helvellaceae
Pezizaceae trib. Acetabuleae. Svensk Bot. Tidskr. 31: 47-66,
1942. T h e Geoglossaceae of Sweden9 Arkiv f6r Botany 30A: 1-67.
1949. Contributions to the Mycoflora of Sweden9 7. A new winter Disco-
mycete, Urnula hiemalis Nannf. n. sp., and a short account of the Swedish spe-
cies of Sarcoscyphaceae. Svensk Bot. Tidskr. 43: 468--484.
NAUNOV, N . A . 1964. Flora gribov Leningradskoi oblasti. Vypusk II. Diskcomitsety.
258 pp. Izdatel'stvo Nauka. Moscow g: Leningrad.
NYLANDEa, W. 1869. Observationes circa Pezizas Fenniae. Not. Fauna F1. Fenn.
10: 1-97.
OBERWlNKLEI~, F., F. DmAGP.anE, and E. MiiLLER. 1967. i)ber Ascocorticium anornalum
(Ell. et. Harkn.) Earle9 Nova Hedwigia I4: 283-289.
OBRIST, W. 1961. T h e genus Ascodesmis. Canad. J. Botany 39: 943-953.
OLIVE, L. S. 1950. A cytological study of ascus development in Patella melaloma.

Amer. J. Botany 37: 757-763.
9 1954. Sexuality of the Ascomycetes. Rapp. Comm. VIII Congr. Int. Bot.,
Paris. 18-20: 67-71.
OTANI, Y. 1967. Note on some cup fungi of the Hyaloscyphaceae collected in
Hokkaido, Japan. Trans. Mycol. Soc. Japan 8: 33-42.
OUELLETTE, G. B., and L. P. MAGASI. 1966. Lophomerum a new genus of Hypo-
dermataceae. Mycologia 58: 275-280.
OVERTON, j. B. 1906. The morphology of the ascocarp and spore formation in the
many-spored asci of Thecotheus pelletieri. Botan. Gaz. 42: 451-492.
PADEN, J . W . 1967. The conidial stage of Peziza brunneoatra. Mycologia 59: 932-934.
, and E. E. TYLUTKI. 1968. Idaho Discomycetes. I. A new genus of the Sar-
coscyphaceae. Mycologia 60: 1160-1168.
, and - - 1969. Idaho Discomycetes. II. Mycologia 61: 683-693.
PALM, B. T. 1932. On Cyttaria Berk. and Cyttariella gen. nov. Ann. Mycol. Berl.
30: 405420.
PEmOON, C . H . 1801. Synopsis methodica Fungorum. 706 pp. Gottingae.
PETER, J. 1962. Aleuria varia (Hedw.) Boudier, Syn. Peziza varia (Hedw.) Ft., Plicaria
varia Hedw. und Plicaria stevensoma Ellis. Zeit. Pilzk. 40: 84-859
PETP,AK, F. 1962. Uber die Gattung Gremmenia Korf. Sydowia 16: 350-352.
PHILLIPS, W. 1887. A manual of the British Discomycetes. 1-462. London.
PLINY (C. Plinius Secundus). A.D. 22. Historia Naturalis. 37 vols.
QUELET, L. 1886. Enchiridion fungoruIn in Europa media et praescrtim in Gallia
vigentium. 352 pp. Lutetiae.
RAHM, E. 1966. Geoglossaceae in Hochtal yon Arosa. Schwiez. Z. Pilck. 44: 165-179.
RA1TVnR, A. 1961. A survey of the distribution of the species of Geoglossaceae in
Estonia. Flor. Mark. 1: 137-145.
RAMAMURTHt, C. S., R. P. KORF, and L. R. BATRA. 1958. A revision of the North
American species of Chlorociboria (Sclerotiniaceae). Mycologia 49: 854-863.
RAMLOW, G. 1906. Zur Entwicklungsgeschichte yon Thelebolus stercoreus Tode.
Botan. Zeit. 64: 85-97.
9 1915. Beitr~ige zur Entwicklungsgeschichte der Ascoboleen. Mycol. Cen-
tralblatt 5: 177-198.
RAMSBOTrOM, J., and F. L. BALFOUR-BROWNE. 1951. List of Discomycetes recorded
from the British Isles. Trans. Brit. Mycol. SOc. 34: 38-137.
RAWLINgS, G . B . 1956. Australasian Cyttariaceae. Trans. Royal Soc. N. Z. 84: 19-28.
R~J~v~s, F. 1967. The fine structure of ascospore formation in Pyronema domesticum.
Mycologia 59: 1018-1033.
REHM, H. 1887-1896. Die Pilze Deutschlands, Oesterreichs und der Schweiz. III.
Abtheilung: Ascomyceten: Hysteriaceen und Discomyceten. In Rab. Krypt-F1.
1(3): 1-1272.
9 1912. Zur Kenntnis der Discomyccten Deutschlands, Deutsch-Ostcrrcichs
und dcr Schwicz. I. Bcr. Bayerinschen Botan. Gcs. 13: 102-206.
REIn, J9 and R. F. CAIN. 1961. The genus Therrya. Canad. J. Botany 39: 1117-1129.
, and . 1962a. Studies on the organisms associated with "snow
blight" of conifers in North America. I. A new genus of the Hclotiales. My-
cologia 54: 194-200.
- - , and - - . 1962b. Studies on thc organisms associated with "snow
blight" of conifers in North America. Some species of the genera Phacidium,
Lophophacidium, Sarcotrichila, and Hemiphacidiura. Mycologia 54: 481-497.
- - - , and ~ 1963. A new genus of the Hemiphacidiaccae. Mycologia 55:
781-785.
- - , and A. FUNK. 1966. The genus Atropellis and a new genus of the Hclotialcs
associated with branch cankers of Western Hemlock. Mycologia 58: 417-439.
~ , and K. A. PIROZYNSKI. 1966. Notes on some interesting North American
fungi. Canad. J. Botany 44: 645-653.
, and . 1968. Critical notes on genera of the Hemiphacidiaceae. I.
Gremmenia. Mycologia 60: 526-531.
RIFAI, M. A. 1968. T h e Australasian Pezizales in the Herbarium of the Royal Bo-
tanic Gardens Kew. Verb. K. ned. Akad. Wet. I1. 57: 1-295.
RIMPAU, R. H, 1962. Untersuchungen uber die Gattung Drepanopeziza (Kleb.) v
H6hn. Phytopath. Zeit. 43: 257-306.
ROSlNSKL M. H. 1953. Two types of spore germination in Sarcoscypha coccinea
(Scop. ex Fr.) Lambotte. Mycologia 45: 302-305.
1956. Development of the ascocarp of Anthracobia melaloma. Mycologia
481 506-533.
SACCAaDO, P. A. 1884. Conspectus generum Discomycetum hucusque cognitorum.
Bot. Cbl. 18: 213-220, 247-256.
9 1889-1931. Sylloge fungorum omnium hucusque cognitorum. 8: 1-1143;
24: 705-1438.
SANCHEZ, A. 1967. T h e sections Aposternium and Microstemium of the genus
Vibrissea (Fungi). J. Agr. Univ. Puerto Rico 51: 79-93.
, and R. P. KORF. 1966. T h e genus Vibrissea and the generic names Lepto-
sporium, Apostemium, Apostemidium, Gorgoniceps and Ophiogloea. Mycologia
48: 722-737.
SANTESSON, R. 1945. Cyttaria, a genus of inoperculate Discomycetes. Svensk Botan.
Tidsk. 39: 319-345.
SATINA, S. 1921. Kistorii razvitiia Phacidium repandum. Zhyrn. Russk. Botan.
Obshchestva. 4: 77-94.
SCHROETER, J. 1893. Die Pilze Schlesiens. In Cohn's Krypt.-F1. Schl. 3(2): 1-256.
9 1897 9 Pezizineae, In Engler, A. und K. PrantL Die naturlicheu Pflanzen-
familien, I, 1: 173-243.
ScniJEvl,, H. 1959 9 Untersuchungen uber Pseudopezizoideae sensu Nannfeldt. Phyto-
path 9 Zeit. 36: 213-269.
SCHULTZ, E. S. 1927. Nuclear division and spore formation in the ascus of Peziza
domiciliana. Amer. J. Botany 14: 307-322.
SEAVER, F. J. 1928 9 T h e North American Cup-fungi (Operculates). 284 pp. New
York.
9 1951. T h e North American Cup-fungi (Inoperculates). 428 pp. New York9
gEBEK, S. 1966. Kli~ k ur~eni smr~ovitych h u b (Morchellaceae) rostoucich v CSSR.
C. C. H. 43: 19-24.
STARB,~CK, K. 1895. Discomyceten-Studien. Bihang Kongl. Svenska Vet.-Akad. Handl.
111, 21(5): 142.
SVR~EK, M. 1954 9 Revise Velenovsk6ho d r u h u rudo Orbilia (Discomycetes). Shorn.
P~'ir. Nfirod. Mus. 10: 1-23.
, and J. KUBI~KA. 1961. Operkulfitnf diskomycety od rybnfka Dvot~igtfi v
jilnich ~echach. ~eskfi Mykol. 15: 61-77.
, and . 1963. Druh~ p~isp~vek k operkulfitnim diskomycetdm z ololi
rybnika Dvof'igt~ v ji~nich ~echach. ~esk~t Mykol. 17: 61-70.
SWlNCLE, D. B. 1934. Fertilization in Ascodesmis nigricans van Tieghem. Amer. J.
Botany 21: 519-545.
TENON, L. R. 1935. A monographic rearrangement of Lophodermium. Ill. Biol.
Monogr. 13: 1-151.
TEm~IE~, C. A9 1942. Essai sur la systfimatique des Phacidiaceae (Fr.) sensu Nann-
feldt (1932)9 Beitr. Krypt.fl. Schweiz 9: 1-999
THIND, K. S., and L. R. BAamA. 1957a. T h e Pezizaceae of the Mussoorie Hills9 I.
J. Indian Bot. Soc. 36: 51-60.
, and 1957b. T h e Pezizaceae of the Mussoorie Hills. IV. J.
Indian Bot. Soc. 361 428-4389
, E. K. CAs~I, and J. s. SETHI. 1958. T h e Pezizaceae of the Mussoorie Hills9
V. Mycologia 49: 831-836.
- - , 51: and833_839.
M'cologiay P" SINCH. 1959. T h e Helotiales of the Mussoorie Hills. II.
9 and S. S. SAINI. 1967. T h e Helotiales of India. VI. Mycologia 59: 467-474.

9 and J. S. SETnI. 1957a. T h e Pezizaceae of the Mussoorie Hills. II. J. Indian
Bot. Soc. 36: 192-206.
, and - - 1957b. T h e Pezizaceae of the Mussoorie Hills. III. Indian
Phytopath. 10: 26~-27.
- - and HARNEK SIN6H. 1964. T h e Helotiales of India. III. J. Indian Bot. Soc.
43: 530-542.
- - and - - 1965. T h e Helotiales of India. IV. J. Indian Bot. Soc. 44:
122-132.
- - and P. SINCH. 1958. T h e Pezizaceae of the Mussoorie Hills. VI. J. Indian
Bot. Soc. 38: 221-232.
- - and - - 1961. T h e Helotiales of Mussoorie Hills. I. J. Indian Bot.
Soc. 40: 295-307.
- - and K. S. WA~ITCIJ. 1964. T h e Pezizales of India. VIII. J. Indian Bot.
Soc. 43: 459475.
- - and . 1966. T h e Pezizales of India. IX. J. Indian Bot. Soc. 4 5 :
384-391.
TnOMeSON, G. E. 1939. A canker disease of poplars caused by a new species of
Neofabrae. Mycologia 31: 455-465.
TULASNE, L. R., and C. TULASNE. 1865. Selecta fungorum carpologia. T h e Imperial
Press, Paris. 3: 1-206.
, and . 1866. Note sur les p h f n o m ~ n e s de copulation que pr6sentent
quelques champignons. Ann. Sci. Nat. Bot., V. 6: 211-220.
VAN T1EGXJEM, P. 1876. Sur de d6veloppement du fruit des Ascodesmis, genre nouve
de l'ordre des Ascomyc~tes. Bull. Soc. Botan. France 23: 271-279.
VOORHEES, R. K. 1939. T h e validity and morphology of Tryblidiella species. My-
cologia 31: 113-123.
WEBS~R, J., M. A. RIVAI, and SAMY SL-ABYAD. 1964. Culture observations on some
Discomycetes from b u r n t ground. Trans. Brit. Mycol. Soc. 47: 445-454.
WEHMEYER, L. E. 1966. Development of the ascocarp in Cryptomycina pteridis.
WEIss, F. 1940. Ovulinia, a new generic segregate from Sclerotinia. Phytopathology
30: 236-244.
WHETZV~, H. H. 1926. N o r t h American species of Sclerotinia. I. Mycologia 1 8 :
224-235.
1928. North American species of Sclerotinia. II. Two species on Carex,
S. duriaeana (Tul.) Rehm, and S. longisclerotialis n. sp. Mycologia 21: 5-32.
1937. Septotinia, a new genus of the Ciborioideae. Mycologia 29: 128-146.
1942. A new genus and new species of brown-spored inoperculate Disco-
mycetes from Panama. Mycologia 34: 584-591.
1943. A m o n o g r a p h of Lambertella, a genus of brown-spored inoperculate
discomycetes. Lloydia 6: 18-529
1944. A new genus of the Sclerotiniaceae. Farlowia 1: 483-488.
9 1945. A synopsis of the genera and species of the Sclerotiniaceae, a family
of stromatic inoperculate Discomycetes. Mycologia 37: 648-714.
1946. T h e cypericolous and juncicolous species of Sclerotinia. Farlowia 3:
385-437.
WHITE, N . H . 1954. T h e development of the ascocarp of Cyttaria gunnii Berk. Trans.
Brit. Mycol. Soc. 37: 431-436.
WroTE, W . L . 1941. A m o n o g r a p h of the genus Rutstroemia (Discomycetes). Lloydia
4: 153-240.
1942a. Studies in the genus Helotium. I. A review of the species de-
scribed by Peck. Mycologia 34: 154-179.
1942b. Studies in the genus Helotium. II. Lachnum pygmaeum and the
status of the genus Helolachnum. Amer. Mid. Nat. 28: 512-523.
9 1943. Studies in the genus Helotium. III. History and diagnosis of cer-
tain European and North American foliicolous species. Farlowia 1: 135-170.
KIMBROUGH" CLASSIFICATION OF DISCOMYCETES 161
9 1944. Studies in the genus Helotium. IV. Some miscellaneous species9

Farlowia 1: 599-617.
, and H. H. WnETZEL. 1938. Pleomorphic life cycle in a new genus of the
Helotiaceae. Mycologia 30: 187-203.
WmTrHOUSE, H. L. K. 1949. Heterothallism and sex in the fungi. Biol. Rev. 24:
411--447.
WHITN~V, H. S., and J. R. PARMETER. 1964. The perfect stage of Rhizoctonia hiemalis.
WojEwooA, W. 1966. Morchellaceae zebrane w poludniowej Polsee w latach 1962-
1965. Fragm. flor. geobot. 12: 205-208.
WOLf, F. A. 1958. The conidial stage of Lamprospora trachycarpa (Currey) Seaver.
J. Elisha Mitchell Sci. Soc. 74: 163-1669
9 1959. Mechanism of apothecial opening and ascospore expulsion by the
cup-fungus Urnula craterium. Mycologia 50: 837-843.
9 1963. The disc fungus Sphaerospora hinnulea (Berk. & Br.) Massee and
its conidial stage9 J. Elisha Mitchell Sci. Soc. 79: 156-159.
- - , and F. T. WOLF. 1947. The Fungi. Vol. I, 437 pp., Vol. IL 538 pp. John
Wiley and Sons. New York.
ZUKAL, H. 1886. Mycologische Untersuchungen. Denkschr. k. ak. Wissen. Wien 51:
21-36.

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THE BOTANICAL REVIEW

VOL. 36 APRIL-JUNE, 1970 No. 2

A HISTORICAL REVIEW ................................................................................ 92

on the configuration of the hymenium. The Cupulati was divided into

~, $3 NIOI=IVlAI S3"'IAO ~13dONI

Dennis relied heavily on Nannfeldt for an over-all system for major

mycetes, compliments beautifully the cytological studies by Berthet. This

Here, ascogonia developed early and the ascogenous hyphae proliferated

to many-celled coil. An antheridial element may be present or absent.

There were four pairs of chromosomes in the primary nucleus of the

those of Thelebolus (Fig. 1F), in the process of maturation an obvious

FIc. 2. Scheme of the developmental types in Ascobolaceae. The gametophytic system

linia, and both binucleate and plurinucleate spores in members of the

T H E ORDERS AND FAMILIES OF DISCOMYCETES

Boudier (1885) Seaver(1928) Clements & Shear (1931) Le Gal (1947)

ever, with the appearance of three m a j o r works dealing primarily with

writers and the relative importance given to various structures. Fitzpatrick

monographic treatment of the family. Dennis (1968) treats more than

follows: Asci operculate, or very rarely suboperculate, thin walled,

the limits of the Helvellaceae. Superficial similarities between members

ornamented, and the paraphyses usually become encrusted with dark

drical, dehiscing by an operculum, bilabiate split, irregular tear, or apical

condition of vegetative and reproductive cells and the basic chemical

b. Pluriapothecial stromata superficial to slightly immersed; stromata

CI~USSEN, P9 1905. Zur Entwicklungsgeschichte der Ascomyceten. Boudiera. Botan.

1908. T h e Geoglossaceae of North America. Ann. Mycol. 6: 387477.

1968. T h e genera of the Operculate Discomycetes. A reevaluation of their

GAUMANN, E . A . 1926 9 Vergleichende Morphologie der Pilze. 626 pp. Jena.

HUGHES, S . J . 1951. Studies on microfungi. IX. Calcarisporium, Verticicladium and

OLIVE, L. S. 1950. A cytological study of ascus development in Patella melaloma.

9 and S. S. SAINI. 1967. T h e Helotiales of India. VI. Mycologia 59: 467-474.

9 1944. Studies in the genus Helotium. IV. Some miscellaneous species9

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