Laboulbeniales
(Fungi, Ascomycetes)
ISABELLE I. TAVARES
MYCOLOGIA MEMOIR NO. 9
Published for
The New York Botanical Garden
in Collaboration with
The Mycological Society of America
J. CRAMER PUBLISHER
IN DEN SPRINGAECKERN 2
3300 BRAUNSCHWEIG, GERMANY
1985

Author's Address
University Herbarium
Department of Botany
University of California
Berkeley, California 94720
U.S.A.
© 1985 The New York Botanical Garden
LIBRARY OF CONGRESS CATALOG CARD NUMBER 84-14900
Printed in Germany
28th FEB 1985
by Strauss & Cramer GmbH, 6945 Hirschberg 2
ISBN 3-7682-1389-7 (US: 0-89327-262-0)
CONTENTS
Dedication 7
Introduction 9
Historical Background and Recent Trends in Research 15
Development, Morphology, and Sexuality 27
Introduction 27
Ontogeny of Laboulbenia ;... 34
Ontogeny of Herpomyces 44
Ascus Development 54
Laboulbenia 54
Herpomyces 57
Parts of the Thallus 60
Receptacle 60
Perithecium 66
Appendages 74
Antheridia 78
Fertilization and Sexuality 83
Classification of the Laboulbeniales 89
Introduction 89
Comparison of Classification Systems 90
Classification, Diagnoses, and Comparisons 93
Generic Key to the Laboulbeniales 102
Generic Taxa (in alphabetical sequence) 130
Origin and Distribution 351
Brief Outline of Paleontological History 351
Position of the Laboulbeniales among Fungi 354
Proposed Phylogeny and Host Relationships of Ancestral Laboulbeniales 357
Distribution of the Laboulbeniales 360
Introduction 360
Distribution on Host Groups 361
Geographic Patterns of Distribution 365
The Laboulbeniales Parasitizing Trechinae 370

Acknowledgments
377
Bibliography 379
Glossary 425
Interpretation of Figures 429
Alphabetical Guide to Abbreviations 431
Collecting, Mounting, and Staining 435
Figures 443
Plates 487
Index to Hosts 599
General index 617
i
i
7
DEDICATION
This publication is dedicated to Dr. Ralph Emerson for his exciting
introduction to the Laboulbeniales, to Dr. Lee Bonar for offering me
the freedom to pursue a subject of my own choosing for the thesis
written in partial fulfillment of the requirement for a Ph.D. degree, to Dr.
William C. Snyder for encouraging me to produce a complete study of
the order, to Dr. Hans N. Hansen for his encouragement and interest in
the early part of the investigation, and to Dr. Herbert L. Mason for the
inspiration of his discussions of speciation, phylogeny, and
geographical distribution.
Above all, this publication is dedicated to Dr. Roland Thaxter. His
monumental work on the Laboulbeniales may be impressive to the
general reader, but it is the investigator who delves deeply into this
group who can really appreciate the thoroughness, vast extent, and true
significance of his work, even though he can also understand its
limitations and errors. For me, its chief significance lies in its value as a
point of reference from which future investigations must proceed. It is
the major accumulation of knowledge that should be sifted through and
reexamined; it contains the ideas that should be followed up with
investigations using the improved techniques available to us today.
Finally, the contributions of entomologists and others who are not
mycologists should not be slighted. They have done much to advance
our knowledge of the Laboulbeniales.

9
INTRODUCTION
The Laboulbeniales are unique among the Ascomycetes because of
their habit of growth on the integument of living arthropods, especially
insects, and their compact thalli, consisting of a reduced hyphal system
organized into a receptacle bearing one or more perithecia and
appendages. Various taxa in this large order, which includes many hundreds
of species, are characterized by striking morphological features, such as
the compact cluster of perithecia and appendages borne at the broad,
flattened apex of the massive thallus of Zodiomyces and the long, coiled
perithecial neck and peculiar secondary feet of Thaumasiomyces scabel-
lularius Thaxter. Within a single genus such as Laboulbenia there may
be considerable diversity of form, resulting from variation in size,
shape, and color of cells, as well as in the branching of the appendages.
Typically, there is a distinctive black foot at the base of the thallus in
the Laboulbeniales, although this is often reduced in size or lacking in
taxa having conspicuous haustoria. The development of the asci is
similar to that of the majority of Ascomycetes, although ascogenous
hyphae do not occur and there is no modification of the apices of the
asci. There are no anamorphs. Thalli develop only from ascospores.
The outer wall of the spore usually swells strongly when it emerges
from the ascus, especially around the foot (the foot may also enlarge
somewhat within the ascus). The viscidity of the spore surface makes
detachment difficult (Richards and Smith, 1955a). It is probable that
spores in soil, such as those believed to infect beetles in Lindroth's
experiments (Lindroth, 1948), could become attached to hosts only under
humid conditions. Only the foot is adherent in genera like Laboulbenia
(cf. the two attachment points in Herpomyces periplanetae Thaxter),
indicating a greater viscidity there. The outer spore wall (or gelatinous
envelope), which protects the spore from desiccation (Thaxter, 1896),
persists as a thin, tough membrane surrounding the growing thallus
(Benjamin and Shanor, 1950b).
The spore in the Laboulbeniales is bicellular throughout the order
(Tavares, 1961). The cells of the spore are distinctly unequal in length
in most genera, with the larger of the two cells being uppermost within
the perithecium and assuming the basal position at germination.
Germination normally takes place on the integument of a host, or on
a seta protruding from it, although it sometimes occurs on the outside
of a thallus or within a perithecium (see Thaxter, 1896, PI. 5, fig. 1;
Benjamin and Shanor, 1950b). In a germinating spore remaining inside
an old perithecium of Laboulbenia gyrinidarum Thaxter, I observed a

10
MYCOLOGIA MEMOIR NO. 9
black basal spot immediately adjacent to the inner surface of the peri-
thecial wall; possibly some absorptive activity had been initiated.
The integument of insects consists of a thin outer layer, the epi-
cuticle; a thick, laminated procuticle (composed of protein and chitin),
through which pore canals pass outward; an epidermal cell layer; and a
thin basement membrane, which separates the cuticle from the body
cavity or hemocoel. The epicuticle consists of four layers: an outer
varnish-like cement layer, which is secreted by the dermal glands; a wax
layer, produced by the epidermal cells; and a thin cuticulin and thick
underlying homogeneous layer, which together make up the protein
epicuticle (it contains no chitin) (see Patton, 1963, Locke, 1974; also see
Richards, 1954). (See Hughes [1959] and Vitzthum [1940-1943] for
equivalent terms used for cuticle in Acarina; see Thor [1931] for structure of
the integument in Acarina and Langner [1937] for the integument of
Diplopoda.) There is some variation in the relative thicknesses of the
layers in different insects and in different parts of the body, such as the
hard sclerites and soft interconnecting membranes. The exocuticle,
which becomes darkened and hardened, may be very thin in larvae,
whereas it may occupy almost the entire thickness of the procuticle in
adult beetles (Locke, 1974). After the last molt, the epidermal cells may
degenerate, forming a thin protoplasmic layer in which the cell areas are
indicated primarily by nuclei (Steinhaus and Tanada, 1971). (See Ta-
vares, 1979.)
Pressure on the perithecium causes spores to emerge through the
ostiole. When the spores are deposited horizontally on the surface of the
host, which is usual in Laboulbenia, the germinating thalli become
attached by the side of the foot. Generally, the lower side of the foot
enlarges considerably, so that the apex of the spore is raised from the
surface of the integument (Thaxter, 1896). The small amount of foot
inflation occurring in Herpomyces ectobiae Thaxter, which germinates on
large setae of its host, is accompanied by no appreciable alteration of
the orientation of the primary axis of the thallus (i.e., the longitudinal
axis of the spore) with relation to the longitudinal axis of the seta. Other
taxa may take an erect position with relation to the setae on which they
germinate — for example, Troglomyces and Laboulbenia marina Pi-
card. In the latter, Picard's illustrations (1908a) show that the upward
curvature from the horizontal position typical of the genus takes place.
In the genera in which the cells of the spore are unequal in size, the
larger cell of the spore (that uppermost in the perithecium) apparently
always forms the foot, even when germination begins inside the
perithecium. However, in Herpomyces, where the cells of the spore are
essentially equal in size, there is some question as to the factors
determining which of the two cells will give rise to the foot. In this
TAVARES: INTRODUCTION
11
genus, the thalli that occur on setae grow with the foot directed toward
the base.
Because they observed that the apices of the thalli of Herpomyces
stylopygae Spegazzini projected toward the distal end of the host
antenna, Richards and Smith (1955a) assumed that either the orientation
of the thalli is determined by their positions or that only those thalli
extending in the proper direction will develop. It is possible that the an-
tennal surface itself may exert an influence on polarity (see young male
illustrated in Tavares, 1966, Fig. 11). There is no definite evidence that
equal-celled spores exhibit a predetermined polarity. It is possible that
they develop in the normal way, regardless of which end points
downward if they germinate on a suitable substratum. However, a four-celled
thallus of H. ectobiae, attached by the foot to the tip of a seta, bore
from its free, abnormally developed upper end a peculiar extension
resembling a haustorial protrusion.
It is probable that almost all spores become attached to the host with
the end that is uppermost in the perithecium either pointed downward
on a seta or toward the front on the surface of the insect. If an insect
moved forward past another insect (exerting sufficient pressure on a
perithecium on the other insect to squeeze out a spore), it could pick up
the spore by the tip as it continued forward, drawing the spore out of
the ostiole so that it lay flat against the surface with the apex (the part
that leaves the perithecium last) in the rear. The same thing probably
occurs when the spore is detached from the ostiole by a seta on a passing
insect — the part that emerges first would become attached and as the
insect moved away, it would pull away the rest of the spore, so that it
lay with the apex pointed toward the distal end of the seta.
After the first two divisions in each of the two unequal cells of the
spore in Laboulbenia, the shape of the thallus becomes radically altered
by considerable expansion, accompanied by the extension upward of the
primary appendage and the formation of the lateral protuberance that
becomes the perithecium. In Herpomyces, the shape of the spore is not
altered (although the female spore may enlarge somewhat), the thallus
growing from extensions of the primary axis — antheridial branches or
receptacles in the males and secondary receptacles bearing perithecia in
the females. In monoecious genera having well-developed secondary
receptacles, such as Rhachomyces, there may be little expansion of the
limited number of cells in the primary axis, which is coincident with that
of the spore.
The relation of the fungus to the host has been a subject of
speculation since the discovery of the Laboulbeniales (see Robin, 1853; Ist-
vanffi, 1895a; Cavara, 1899; Thaxter, 1908, p. 227; Scheloske, 1969, p.
42; Tavares, 1979). Picard (1908a) observed that Laboulbenia marina

12
MYCOLOGIA MEMOIR NO. 9
had no visible effect on setae and elytra, both of which he believed to be
incapable of regenerating cuticle. He suggested, therefore, that
nutrients might be obtained by the fungus from the fluid, waxy surface
layer of the cuticle, which he thought was nutritionally comparable to
the fat body utilized by Trenomyces. Cepede (1914), seeing no effect on
the integument as a result of infection by L. blanchardii Cepede,
favored this view, as did Lepesme (1946). In support of it, Cepede cited
Robin's observation (1853) that a mucous material favoring fungus
growth may appear on the integument of insects attacked by destructive
parasites. This was presumed to have been produced by the integument.
Robin, himself, did not associate this phenomenon with the Laboul-
beniales, however.
Wigglesworth (1961) reported that if the upper two layers of the epi-
cuticle are removed by gentle abrasion, the underlying epidermal cells
may be induced to secrete more wax to repair the injury. Such a
phenomenon would help to support the theory of the use of the surface wax
for nutrition.
In Periplaneta americana (L.), wax presumably is secreted throughout
life (Kramer and Wigglesworth, 1950), probably by the oenocytes of the
body cavity, which may lie just beneath the epidermal cells in the adult.
Oenocytes also may be incorporated into the fat body. Because the
haustorium produced by the fungus parasitizing P. americana
penetrates into the epidermis (see Tavares, 1979), the wax layer (a soft, thick
grease layer [Ebeling, 1974]) is clearly not the primary source of
nutrients.
The ability of spores of Herpomyces to divide on waxy surfaces
(Richards and Smith, 1954) and of thalli of Fanniomyces to reach the
spermatial stage on detached fly wings placed on brain-heart infusion
agar (Whisler, 1968) indicates that some growth can occur in the
absence of living insect cells or hemolymph. It is possible that there is some
utilization of surface secretions and cuticular substances during early
growth and penetration of the cuticle (Benjamin, 1971; Scheloske,
1969).
Except in Herpomyces, in which a small haustorium emerges from the
foot, Thaxter (1896,1908) believed that taxa having a black foot
directly absorb nutritive materials through it from the host cuticle (cf. the
central, clear area of the foot of L. hagenii Thaxter [Thaxter, 1896] with
the pale area in the cuticle below the foot of L. borealis Spegazzini
[Tavares, 1979]). Hydrolysis of chitin by the fungi was proposed by
Picard (1908b), although chitin actually is present in a relatively small
proportion in the procuticle of some hosts (Wigglesworth, 1961; Hack-
man, 1974). Picard (1908b) suggested that epidermal cells secreted more
abundantly at the point of attack, repairing the damage so that none
TAVARES: INTRODUCTION
13
was evident (see discussion of wound healing, Wigglesworth, 1961; also
see Cepede and Picard, 1908, Chatton and Picard, 1909).
Conspicuous haustoria occur in several genera of the Laboulbeniales,
including Arthrorhynchus (Blackwell, 1980b), Herpomyces (PI. 10, b-
c), Hydrophilomyces (PI. 31,e), Rhizomyces, and Trenomyces (Chatton
and Picard, 1909; Meola and DeVaney, 1976; Tavares, 1979; Meola and
Tavares, 1982). The haustorium of Stigmatomyces (Peyritsch, 1871,
1873; Dainat and Manier, 1974) and of Hesperomyces (Tavares, 1979)
penetrates the host integument and expands into a small bulb below the
cuticular layers. Whereas the haustorium of Herpomyces terminates in a
large bulb within the well-developed epidermal cell layer of the host
(Richards and Smith, 1956), an extensive, slender haustorial system
(rhizomycelium) radiates from the small haustorial bulb of
Hesperomyces into the hemocoel of the host (Kamburov et al., 1967b). The
haustoria of the Laboulbeniales apparently are enucleate and nonsep-
tate (Blackwell, 1980b, Meola and Tavares, 1982). Although Colla
(1926a) reported short, sinuous haustoria with dense, granular
cytoplasm in the integument subtending the black foot of Laboulbenia
vulgaris Peyritsch, her photographs did not show these structures
satisfactorily (cf. Carbonelli Tonghini, 1913).
Although he had concluded earlier (Thaxter, 1914b) that all fungus
parasites of insects obtained their nutrients from the circulatory systems
of their hosts, Thaxter (1931, p. 115) later repeated his earlier
supposition that absorption takes place through a thin osmotic foot
membrane applied to the pore canals of the host cuticle. That species of
Laboulbenia parasitizing Coleoptera actually take up material from the
hemolymph was demonstrated by Scheloske (1969). When Nile blue
sulfate became sufficiently concentrated within an elytron, the dye
could be seen to rise into an attached thallus.
A relationship with pore canals was proposed as early as 1873 by
Peyritsch, who reported seeing one joined to the foot of L. vulgaris.
Absorption could probably take place more readily in the region of the
pore canals, whether or not a conspicuous haustorium is present. In
most integuments, the cuticle is perforated by pore canals that run up
to, but not through, the epicuticle (Locke, 1961). In some insects, an
extension of the epithelial cell may remain within the canal. In others,
the canal may become clogged, at least at certain times of life (Richards,
1951; Beament, 1964). In the abdominal cuticle of Periplaneta
americana, Kramer and Wigglesworth (1950) reported that the pore canals
went up as far as the cement layer (however, it is probable that they did
not actually penetrate the thin epicuticle). In this genus, the cytoplasm
apparently remains within the pore canals (Wigglesworth, 1961). The
extremely thin layer of epicuticle that intervenes between the foot of the

14
MYCOLOGIA MEMOIR NO. 9
thallus and the pore canal could easily be penetrated by an absorptive
process so small as to be virtually undetectable (cf. Cooke's suggestion
[1977] of absorption through submicroscopic penetrating structures).
Early workers had conflicting opinions about the effect on the host.
For example, Peyritsch (1871) never saw any evidence of damage by the
fungifs, whereas Karsten (1869) reported resorption of muscle sheaths in
the thorax of the host of Stigmatomyces baeri (Knoch) Peyritsch. Black-
well (1980b) reported that the haustorium of Arthrorhynchus extends
into the skeletal muscles of the dipteran host. The tergosternal muscles
of the abdomen are among the most frequently invaded tissues of the
louse hosts of Trenomyces histophthorus Chatton & Picard (Meola and
DeVaney, 1976). These authors showed that the muscle striations
become diffuse (see also Meola and Tavares, 1982). Extensive damage
to fat body cells was reported by Chatton and Picard (1909) (see also
Miiller, 1927). The nuclei of the epidermal cell layer near the haustorial
bulbs of Herpomyces periplanetae lose their normal ability to stain with
iron haematoxylin (Tavares, 1979, Fig. 2). The pathogenic effects
on the hosts of the Laboulbeniales have been discussed in detail by
Benjamin (1971) and Scheloske (1969) (see also Strandberg and Tucker,
1974). The effect of the Laboulbeniales on their hosts should be
compared with that of animals, rather than with that of other fungi,
because the Laboulbeniales are dependent on the life of the host for
survival (Scheloske, 1969; see also discussions by Rogers, 1962,
pertaining to a balance between harmful actions of a parasite and the capacity
of the host to resist; see Frank, 1982).
TAVARES: HISTORICAL BACKGROUND
15
Historical Background and Recent Trends in Research
In the seventy-five years that have elapsed since the second volume of
Thaxter's monograph on the Laboulbeniales was published in 1908,
little has been added to his basic outline of their characteristics, and to
his enumeration and evaluation of the contributions of earlier observers
of the group. However, correction is necessary of certain
misconceptions that have arisen in the literature because of errors and
omissions in his historical discussion of 1896 (Thaxter, 1896). In addition,
the early publications show the evolution of our knowledge of these
unusual fungi. Those who are introduced to their study for the first time
will encounter some of the same problems in interpreting anatomy that
others have faced.
Despite the large body of descriptive literature published in the field
of entomology during the latter part of the eighteenth century and the
first half of the nineteenth century, it was not until 1849 that the
existence of the Laboulbeniales was made known to the scientific
world — first, through a paper by Auguste Rouget (1850), which he
submitted for reading before the Societe Entomologique of France on
May 23, 1849, in Paris, and second, by Ferdinand Schmidt at a meeting
of the Wissenschaftsfreunde in Laibach (Ljubljana), Yugoslavia, on
July 20, 1849. The first published record of the Laboulbeniales was
incorporated in the proceedings of the meetings of the latter society in the
Illyrischen Blatt no. 60, 1849, according to Mayr (1853).
An account of the 1849 meetings of the Freunden der Naturwissen-
schaften in Laibach, presented before the society in Vienna in
December, 1849, by von Hauer (1850), included Schmidt's comments on
the structures that he had found on "Nebria stentzii" (ICarabus stentzii
Villa & Villa, a synonym of Carabus scheidleri Panzer [Coleoptera,
Caraboidea, Carabidae], a genus not subsequently reported as a host;
the specimen may have been misidentified). Together with Heinrich
Freyer (mentioned also by Gratzy, 1898), Schmidt had discovered
parasitic plants that resembled bunches of setae. Each consisted of a dark,
broad, basal part having a hairlike prolongation on one side and a fruit
capsule on the other.
The same hairlike structures, observed both on "Nebria stentzii" and
Nebria brunnea Duftschmid, were thought by Mayr (1853) to be chi-
tinous. Apparently he believed that they were outgrowths of the insect
integument. Nevertheless, he observed that forms on younger insects
were always hairlike, whereas on older individuals, additional stages
were present in which a swelling, of varying shape, developed laterally
on the lower part of the hair.

16
MYCOLOGIA MEMOIR NO. 9
Laboulbene and Follin (1848) believed that a powdery substance on
the integument of certain Coleoptera (Lixus, Buprestis) was composed
of fungus spores and that these were the first fungi to be found on
normal, healthy insects. Because they had not published more on this
subject, Rouget submitted the observations that he had first made in 1840
of parasites on the beetle, Brachinus crepitans L. Robin's statement
(1853) that Laboulbene, a long time before, had observed Laboulbenia
has apparently been interpreted as indicating that Laboulbene and
Rouget discovered the fungi at about the same time (see Cepede and
Picard, 1908). The omission of these fungi from their discussion of
various organisms living on insects by Laboulbene and Follin (1848)
indicates that if Laboulbene had observed them, he had no conception
of their nature before Rouget's paper was presented.
At first, Rouget believed that the protuberance he saw on Brachinus
was a supernumerary antennal segment. He (Rouget, 1850) later
published remarkably accurate drawings of what he recognized as
organisms living at the expense of their hosts, together with a description
of their structure.
Recognition of these parasites as fungi by Robin (1852) led to the
establishment of the genus Laboulbenia (Robin, 1853), which was
placed in the family Pyrenomycetes. However, Robin referred to the
spore-producing structure both as a sporangium and as a perithecium.
Montagne and Robin (Robin, 1853) made no clear distinction between
asci and mature spores, making it difficult to evaluate their
observations. They misunderstood the perithecial wall structure in L. rougetii
Montagne & Robin. In L. guerinii Robin (Robin, 1853), it was believed
that filaments intermingled with the spores within the sporangium. The
tenacity of the foot in Laboulbenia was mentioned, as well as the
tendency of the spores to germinate in pairs, presumably developing a
common foot. Because attachment of the foot was superficial, it was
assumed that the thallus must derive nutritive substances from the air.
Fungi resembling L. guerinii were found by Hagen (1855) on nymphs
of Macrotermes bellicosus (Smeathman) var. mossambicus (Hagen) (as
Termes [Isoptera — termites]) from Mozambique. Then specimens on
wingless dipteran parasites of bats were described as Arthrorhynchus,
an acanthocephalan worm, by Kolenati (1857). The short antheridial
branch of the mature thallus was interpreted as a doubled-over spiny
snout, with a basal adhesion disk; the inner surfaces of the outer wall
cells of the perithecium were apparently mistaken for intestinal walls.
Diesing (1859), in his revision of Rhyngodeen, listed Arthrorhynchus as
a doubtful member of the group; Kolenati's illustrations, which were
included, showed the minor "caudal" variation (at the apex of the
perithecium) distinguishing "male" from "female," as well as reproductive
TAVARES: HISTORICAL BACKGROUND
17
structures in the "male" in the position of the young asci, which are
lateral in Arthrorhynchus. No further publications on these organisms
appeared until 1868, when Knoch (1868) described Laboulbenia baeri
Knoch, which occurred in large numbers on houseflies at the time of a
cholera epidemic and was thought by some to be a possible cause of this
disease.
Karsten (1869) noticed that the distribution of Stigmatomyces muscae
Karsten (the same fungus) was related to mating activities of the hosts.
He concluded that the trichogyne and the spermatia produced by the
antheridial branch have a sexual function in this species and recognized
their similarity to the reproductive organs of the Florideae. His account
of the early development of the spore is incorrect. He misinterpreted the
sequence of development; what he thought was a thallus having a
rounded germ cell in the venter of the archegonium and spherical
bodies around the antheridia actually was undergoing perithecial
elongation and initiation of asci, whereas what he believed to be a later
stage, in which the perithecium was erect and the trichogyne was lying
over the antheridial branch, actually represented a much younger
specimen. Because he apparently thought that the spores were formed in a
germ cell, Karsten (1869) placed Stigmatomyces in the Mucorinee.
In his review of Sorokin's description (1871) of Laboulbenia pitra-
eana Sorokin, de Janczewsky (1872) doubted that it was distinct from
Stigmatomyces muscae (see also Hoffmann, 1871). Sorokin (1871)
thought it probable that an internal mycelium was formed within the
body of the host. He mistakenly thought that mycelia on the surface of
the host had developed from spores of L. pitraeana. He (Sorokin, 1883)
subsequently observed that the Laboulbenia spores became attached
externally to the host and the thalli developed on the surface.
The most thorough study of this group of fungi prior to the work of
Thaxter was undertaken by Peyritsch when he began his extensive
observations in 1871 on the structure and development of the housefly
Laboulbenia, which he named L. muscae Peyritsch. He discussed the
appearance of the asci within the perithecium, spore germination and
discharge, haustorium formation, and the early cell divisions taking
place during growth of the thallus (Peyritsch, 1871). Because he never
saw a trichogyne covered with rounded cells like those described by
Karsten, he doubted that spermatial fertilization took place. He
believed that the spherical protrusions at the tip of each branchlet of the
appendage did not absciss and that fertilization might be caused by mere
contact of a branchlet with the trichogyne. Because he saw no abscission
of cells on what he thought were male structures in species of
Laboulbenia on beetles, he assumed (Peyritsch, 1873) that fertilization must be
effected by the tips of certain appendage branches called pollinodia.

18
MYCOLOGIA MEMOIR NO. 9
Peyritsch (1871) brought together all but one of the organisms that
were believed to belong to the genus Laboulbenia; L. pilosella Robin
(Robin, 1871) was omitted. Arthrorhynchus was redescribed by
Peyritsch (1871) as L. nycteribiae Peyritsch (see also Brauer, 1870), whereas
the species probably first observed by Schmidt in 1849 (see von Hauer,
1850) was described as L. nebriae Peyritsch. Later, Peyritsch (1873)
established the family Laboulbeniaceae in the Ascomycetes. In the
descriptions and figures of the new species, many important details,
particularly of internal structure, were not clearly presented or were
overlooked or misinterpreted.
Observations on laboratory colonies of houseflies permitted Peyritsch
(1875) to determine that Stigmatomyces matured in 10-14 days. He
studied the relation of sexual activity to transmission of infection. It was
noticed that the fungi would not grow on larvae or pupae nor would
they grow on adults of the other species of Diptera that Empusa muscae
(Fr.) Cohn parasitized. As the first to study the Laboulbeniales in the
field, Peyritsch (1875) concluded that they overwinter on the host and
form no resistant spores. Although he failed to make accurate drawings
of their structure, he laid a sound foundation for our understanding of
the biology of these fungi.
The difficulties involved in observing structural details were not
solved until the work of Faull (1911, 1912) showed the value of cyto-
logical staining. It is probably safe to say that Peyritsch's failure to
represent anatomical characters accurately has been repeated to some
degree by almost every investigator of the Laboulbeniales.
The Laboulbeniaceae were called doubtful Ascomycetes in de Bary's
survey (1884,1887) of fungus morphology. He apparently utilized
Peyritsch's interpretation of the fertilization process to support his own
doubt about sexuality in these fungi. In reply to de Bary's disposition of
the Laboulbeniaceae, Karsten (1888) reiterated his former opinion that
the venter of the archegonium (perithecium) contained only a single
egg-cell in which spores were formed.
Peck's unfamiliarity (1885) with European studies on Stigmatomyces
resulted in the description of Appendicularia entomophila Peck from a
species of Drosophila (Balazuc [1974] suggested that the species was
misidentified). The fungus was believed to be a Deuteromycete, possibly
related to Sphaeronaema. Gercke (4886) observed similar specimens on
Drosophila funebris Fabricius that were also Stigmatomyces.
The first report of Laboulbenia on a group other than the Insecta —
the Acarina or mites in the Arachnida — was published by Berlese
(1889). He believed it unlikely that the paraphyses (appendages) were
male fertilizing branches because they do not resemble those of other
cryptogams.
TAVARES: HISTORICAL BACKGROUND
19
A careful, systematic study of the Laboulbeniaceae was begun by
Roland Thaxter in 1890, when he published the first in a series of papers
describing hundreds of new species and discussing various aspects of
their biology (Thaxter, 1890, 1891, 1892, 1893, 1894, 1895). Having
observed spherical bodies in and around the apices of the antheridia, he
(Thaxter, 1890) cautiously accepted Karsten's view of fertilization,
recognizing that a form of sexual reproduction definitely was present.
Corroboration of emission of spermatia from the antheridia and
formation of ascospores within the asci followed (Thaxter, 1891). The
proposal was made that the nonmotile male cells fertilized the carpogonium
through a trichogyne and an intervening axial cell (Thaxter, 1894), the
placement of the sexual structures being conducive to self-fertilization.
The carpogenic cell was seen to divide into three cells, the middle one
becoming an ascogenic cell, either directly or after further division. The
major subdivisions within the Laboulbeniaceae (now including 23
genera) were based on the type of antheridium. Emphasis was placed on
the persistence of the gelatinous envelope surrounding the thallus and
on its independence from internal cell divisions (Thaxter, 1894).
Thaxter's first monographic volume on the Laboulbeniaceae, which
appeared in 1896, summarized and enlarged upon all previous work; it
has served as the basis for all subsequent studies of the group.
Succeeding volumes appeared in 1908, 1924, 1926, and 1931 (see Benjamin's
descriptions'[1971]), preceded by preliminary diagnoses (Thaxter, 1899,
1900,1901a,b, 1902,1905).
After Thaxter's work became known, a number of short papers were
published by various authors, either describing one or two new species
(Giard, 1892; Istvanffi, 1895a,b; Spegazzini, 1902) or reporting new
hosts or localities (Speiser, 1901b; Rick, 1903; von Beck, 1903). Thaxter
(1896) and other authors have stated that Istvanffi expressed the
opinion that the external thalli in the Laboulbeniales are derived from a
vegetative portion growing within the body cavity of the host. This
remark did not appear in Istvanffi's publication (1895a) (see de Janc-
zewsky, 1872, on Sorokin's work); neither did he indicate anything,
either in his description or in his drawings, other than a superficial
attachment of the thallus by means of a blackened foot. Although he
did state that he believed that the fungi lived both on and from the body
of the host, the statement appears to have been made in the sense of
designating the source of nutrients. In fact, he hesitated to accept the
idea of parasitism, because Stigmatomyces had been shown by Peyritsch
to have no deleterious effect on houseflies (similar views were held by
Cavara [1899]; cf. also Rick [1903]; see Tavares [1979]).
For several years French publications appeared that emphasized the
biological and phylogenetic aspects of these fungi. Although their
factual contribution, other than the addition of new species and genera

20
MYCOLOGIA MEMOIR NO. 9
to the flora, was not great, the authors proposed several new theories and
approaches to the biological problems; they also revived
experimentation. The damage caused to the host's fat body by the large
haustorium of Trenomyces histophthorus Chatton & Picard prompted
Chatton and Picard (1909) to consider possible physiological differences
between species with visible haustoria and superficially inserted species.
Cepede (1914), following Mirande's example (1905), tested
unsuccessfully for glucose in integument, although he found it in the
foot of Laboulbenia blanchardii (see Richards [1951], Hackman [1974]
regarding cuticle components; cf. Errera [1905]).
Picard (1909) added to the observations that had been made by
Thaxter (1896) on extreme location specificity of species parasitizing
aquatic insects. In a series of infection experiments, Cepede and Picard
(1907,1908) demonstrated host specificity both at the genus and species
level. They suggested that the polymorphic species Laboulbenia elon-
gata Thaxter (a synonym of L. flagellata Peyritsch) might include
several distinct species and believed that this could only be determined
by cross-infection experiments. They concluded that there is almost
always a narrow specificity with respect to the host. The suggestion was
made that two factors probably influence specificity: the effect of the
physical and chemical properties of the integument on the attachment of
the fungus, and the nutritional requirements of the parasite (Cepede
and Picard, 1908). It was Picard's opinion (1913b) that fungi living on
distantly related hosts probably should be regarded as different species.
(All of these views should be compared with the data published by Sche-
loske, 1969).
Whether morphological differences are due to environmental or to
hereditary factors was a question that arose both in connection with a
new species of Misgomyces on Bledius (a similar species parasitized
Dyschirius, a beetle preying on Bledius) and with the proliferation of
additional perithecia and appendages in Laboulbenia proliferans
Thaxter. The tendency to produce supernumerary perithecia was assumed by
Picard (1913b) to be hereditary.
On the species level, Cepede (1914), like Picard, favored treating as
separate species fungi having different hosts and slight morphological
differences; place of germination on the host was also thought to be of
importance in delimiting species. In considering the Laboulbeniales as a
whole, he believed that they merited a special place among the fungi;
therefore, he proposed the name Phycascomycetes, because of the
similarity of the group both to the Ascomycetes and to the red algae. Cepede
and Picard (1908) had already accepted this similarity as clear evidence
of the close relationship of the Laboulbeniales to the Florideae, and had
dismissed the lack of chlorophyll as an adaptation to parasitism.
TAVARES: HISTORICAL BACKGROUND
21
Chatton and Picard (1909) pointed out the structural difference
between those dioecious genera with compound antheridia in which either
antheridia or perithecia arise in a linear sequence from the lower
receptacle and those genera, such as Laboulbenia, in which antheridia
arise only from the apical cells of the thallus and the perithecia are
produced by the lower receptacle. They believed that a classification system
based on manner of development, which becomes evident early in
ontogeny and therefore represents a basic evolutionary trend, would be more
indicative of phylogenetic relationships than Thaxter's classification
based on antheridial types. Chatton and Picard had virtually no
knowledge of the early development of the different genera, but they were
able to recognize the importance of characteristics of the receptacle in
taxonomy. Considerable insight into the essential taxonomic
characteristics was indicated by Picard's (1913b) suggestion that Herpomyces
was ancestral to Dimorphomyces and that the distinction between the
Peyritschiellaceae and Laboulbeniaceae was overemphasized.
The preliminary note published by Faull (1906b), in which he
reported syngamy and three subsequent nuclear divisions in the asci of
some Laboulbeniales, established without question the ascomycetous
nature of the group. Chatton and Picard (1909) believed that studies
such as Faull's (1906a,b), demonstrating delayed nuclear fusion in the
ascus, merely shortened the gap between the Ascomycetes and the
Laboulbeniales, but did not eliminate it. Although brief notes on other
species appeared earlier (Faull, 1911), only Faull's work on Laboulbenia
gyrinidarum and L. borealis (as L. chaetophora) was published in its
entirety (Faull, 1912). Faull (1912) reported the sequence of divisions of
the trichophoric cell and carpogenic cell which are outlined for
Laboulbenia in the present paper and reported also by Benjamin (1968a)
for Symplectromyces; it probably is characteristic of the suborder.
A long series of regional investigations began to appear after 1900
(see annotated bibliography; see discussion by Benjamin, 1971). Taken
individually, most of them were rather unimportant; however, when
considered together, they added an impressive amount of information
on the distribution of these fungi, particularly in Europe and North
Africa. Reports included those on Laboulbeniales occurring in Hungary
(Bokor, 1926), Latvia (Briedis, 1934), West Africa (Spegazzini, 1914a),
East Africa (Poisson, 1929), and Europe, North Africa, and the Near
East (Maire, 1912, 1916a,b, 1920; Picard, 1912, 1913a,b, 1917; de
Peyerimhoff, 1910; Lepesme, 1939; see also Carbonelli Tonghini,
1913). Freshly collected hosts (usually from a small number of
localities), as well as museum specimens, were examined for these fungi.
During his surveys of the Laboulbeniales parasitizing Argentinian
and Italian insects, Spegazzini (1912, 1914b, 1915b, 1917) briefly

22
MYCOLOGIA MEMOIR NO. 9
studied ontogeny, longevity, effects of temperature and humidity on
growth, and host-parasite relationships. Although he (Spegazzini, 1912)
remarked that intergradation of morphological types made it difficult to
determine the limits of species and that these variations might either
represent hybrids or be the result of the position of the fungi on the
insects, he nevertheless described several newtaxa on the basis of slight
morphological differences. Spegazzini (1917) later became convinced
that the amount of host specificity and the influence on the parasite that
is exercised by its position on the host differ according to the species of
fungus. Speciation and specificity were problems that Spegazzini
believed could be solved best by artificial cultivation; however he was
successful only in growing fungi experimentally on the cockroach,
Blatta orientalis L. He (Spegazzini, 1914b) introduced an artificial
system of characters and published a key to the structural groups in
Laboulbenia, breaking down the genus into subgenera, sections, and
subsections. However, the characters he used had little taxonomic
significance.
The difficulty of reconciling the lack of any visible effect of the
parasite on the host cuticle with the apparent absence of haustoria in
many species led Spegazzini (1917) to suggest that the fungi obtain their
nutrients from the external environment, possibly through pigments.
The problem of the absorptive mechanism of the fungus interested a
number of other investigators, including Carbonelli Tonghini (1913),
Boedijn (1923), Colla (1926a), Mercier and Poisson (1927), and Poisson
(1930).
During her survey of the Laboulbeniales of Italy, Colla (1925)
recognized that they were reminiscent of Puccinia in the manner of formation
of species through the modification of biological forms into stable
morphological types. A wide range of variations of Laboulbenia cone-
glanensis Spegazzini living close together on their hosts could have been
segregated into a series of subspecies had they been situated on separate
hosts or on different parts of the same host (Colla, 1926b). Pursuing
further the problems of variation in form, Colla (1933) distinguished
three forms of L. rougetii, as well as intermediate specimens. Although
the three forms were distinct, regardless of their position on the host,
position could cause some variability. Colla (1933) suggested
experimental study to determine whether these forms should be designated as
subspecies.
Because of slight differences that he observed in thalli of Laboulbenia
fasciculata Peyritsch from various hosts, Poisson (1930) suggested that
such ubiquitous species might permit determination of the amount of
influence exercised by the host on the structure of the fungi. Like
Spegazzini, Colla (1934), as a result of several years of observation,
TAVARES: HISTORICAL BACKGROUND 23
finally reached the conclusion that a great number of the described
species and genera would be placed in synonymy if an adequate number
of specimens were studied and if culture experiments were carried out.
The records of Laboulbeniales parasitizing European arthropods
were augmented by a series of regional studies and collection reports in
many countries; among them were Rumania (Lepesme, 1941b; Bechet
and Bechet, 1960a,b), U.S.S.R. (Blagoveshtchensky, 1950, 1951, 1959;
Bechet and Bechet, 1960b; Lunkashu, 1970), Yugoslavia (Bechet and
Bechet, 1970), West Germany (Poelt, 1952a,b; Scheloske, 1969);
Czechoslovakia (Fassatiova and Fassati, 1956), Poland (Banhegyi,
1964; Samsinakova, 1960b; Nowasad, 1973 [for Majewski, see
annotated bibliography]); Great Britain (Donisthorpe, 1942a,b; Blair, 1947,
1949; Shaw, 1952a, b; Green 1954), and Italy (Rossi and Cesari Rossi,
1981), as well as Finland (Hulden, 1983).
New host and locality records have been published for other areas
also. These have been based on museum specimens (partly obtained
by various expeditions), on the collections of perceptive entomologists,
and on the authors' own collections. Examples are those from tropical
Africa (Lepesme, 1945; Balazuc, 1971a, 1972, 1973d, 1975d-f), Japan
and Taiwan (Kishida, 1929 [1930?]; Ishikawa, 1966), Indonesia and
India (Jeekel, 1959; Samsinakova, 1960a; Batra, 1963; Balazuc, 1972,
1973d), Central and South America (Lepesme, 1942c; Poisson, 1954;
Sugiyama, 1972; Balazuc and Demaux, 1973; Balazuc, 1975d, f,
1977a), the subantarctic islands (Lepesme, 1947), Cuba (Krejzova and
Weiser, 1968; Balazuc, 1975d, f); Borneo (Middelhoek, 1957; Balazuc,
1977a), Philippines (Balazuc, 1973d), Malaya (Balazuc, 1977a), North
Africa (Maire and Werner, 1937; Lepesme, 1939; Balazuc, 1975e), Juan
Fernandez Islands (Balazuc, 1975e), and Mexico (Rossi and Cesari
Rossi, 1977c).
Laboulbeniales were observed quite early on Diplopoda (millepedes).
Verhoeff (1897) published an illustration showing what may have been
immature specimens of Laboulbeniales on a millepede. Mature
specimens were observed by him (1916) on males of another species, but
as before, he did not recognize them as Laboulbeniales. He suggested
that a ramification from the foot, which he interpreted as the spore
from which the fungus grew (Schellenberg [1923] pointed out this
error), penetrated through^ the pore canal into the exoskeleton of the
host. Verhoeff recognized that the fungi must be thrown off during the
molting period, although the great length of this period in millepedes
would probably permit reinfection. Schellenberg (1923) suggested that
the mature specimens found by Verhoeff in 1916 were a new species of
Laboulbenia (Jeekel [personal communication] reported thalli on
anterior legs of Pachyiulus that may have been Laboulbenia). (See also

24
MYCOLOG1A MEMOIR NO. 9
Brolemann [1920], Verhoeff [1926], Manfredi [1931], Colla [1932a],
Schubart [1934,1945], Scheloske [1969], and Rossi and Balazuc [1977].)
Several reports were published on the occurrence of the Laboul-
beniales on ants, beginning with that of Wheeler (1910) (see Blum, 1924;
Baumgartner, 1934; Batra, 1963; Benjamin and Shanor, 1950b; also Be-
quaert, 1920,1922, Cole, 1935,1949, Donisthorpe, 1912,1942c, Smith,
1917, 1928, 1946, 1961, and Judd and Benjamin, 1958). To Hagen's
report (1855) of a Laboulbenia on nymphs of termites were added those
of Buchli (1966) and Rossi (1974); the stage of development was
indicated only by Hagen (see also Kimbrough et al., 1972; Rossi and
Cesari Rossi, 1977a). New mite parasites were added by Baccarini
(1904), Paoli (1911, 1912), and Sibilia (1927). Vitzthum (1925) reported
a parasite on a mite that his figure showed to be Rickia. (See also Elsen
and Fain [1973], Lindquist [1961], Blum [1924], Balazuc [1971f], Hoyt
[1963], Balazuc [1978], Blackwell and Kimbrough [1976], Strand-
berg and Tucker [1974], and Theodorides [1955].)
A higher rate of infection of beetles took place when Lindroth (194S)
placed them in soil containing spores of Laboulbenia than when he
exposed them to infected beetles (see caution about experimentation
[Baumgartner, 1927]). Nevertheless, Lindroth concluded that infection
in nature is chiefly through contact between insects (see also Arwidsson,
1946). Later, Boyer-Lefevre (1966) suggested that the clay of the caves
might play a role in the transmission of Rhachomyces parasitizing
cavernicolous Trechidae.
A deleterious effect on salt-marsh beetles was attributed by Bro
Larsen (1952) to infection by Laboulbeniales (see discussion by Benjamin,
1971). A reduced survival period was observed among infected coccinel-
lid beetles in citrus groves by Kamburov et al. (1967b), although Apple-
baum et al. (1971) detected no appreciable change in metabolism in
parasitized beetles (cf. also Kehat et al., 1970). A reduced infection rate
among salt-marsh beetle hosts exposed to higher salinity was observed
by Bro Larsen (1952), as well as by Dainat (1973) during her studies of
Diptera parasitized by Stigmatomyces.
A revival in the United States of interest in the Laboulbeniales began
with a brief note on the dioecious nature of Laboulbenia formicarum
Thaxter on North American ants (Benjamin and Shanor, 1950a, b).
Studies by Benjamin and Shanor (1952) on Laboulbenia parasitizing
Bembidion demonstrated that position specificity was the result of
transmissional rather than physiological limitations. The histerid beetle-
ant-mite relationship reported by Blum (1924) suggested to Shanor
(1955) that an entirely different related fungus possibly occurs as a
saprophyte in ant nests.
TAVARES: HISTORICAL BACKGROUND 25
Histochemical comparisons were made by Richards (1954) between
the cell walls of Herpomyces and insect cuticle. Photographs of the
large haustoria produced by Herpomyces were published by Richards
and Smith (1956). Hesperomyces virescens Thaxter was later found by
Kamburov et al. (1967b) to have an extensive haustorial system that was
different in structure (cf. PI. 16,c).
The similarities between species of Laboulbenia occurring on a tribe
of carabid beetles and some species parasitizing other tribes suggested to
Jeekel (1959) that the tribes were related to some extent (cf.
observations of Rossi and Vigna Taglianti [1979] on Rhachomyces). In
addition, similarity between species on a group of hosts indicated to him
that the species were related and were host-specific. On the other hand,
a combination of differences and similarities could be evidence of
parallel evolution. Morphological differences between the groups of fungi on
different host tribes showed that a character such as a stalked peri-
thecium appeared independently in different phylogenetic lines; other
characters, such as similar appendage structure, indicated close ties
between species of Laboulbenia (Jeekel, 1959).
When Benjamin (1955, 1968a, b, 1970) described Neohaplomyces,
Balazucia, Prolixandromyces, and Sandersoniomyces, he compared
them carefully with related genera, including ontogenetic details
whenever possible. He (Benjamin, 1967) included an extensive
discussion of the groups of Hemiptera parasitized by Laboulbeniales in his
publication on semiaquatic Laboulbenia. When comparing related
genera to Sandersoniomyces, Benjamin (1968a) included photographs
of the development of the asci of Symplectromyces vulgaris (Thaxt.)
Thaxter.
Although drawings may show details that cannot be clearly seen in
photographs (they may also include errors), photographs frequently
reveal structural characteristics of which the investigator is not yet
aware. The photograph shown by Strandberg and Tucker (1974) of a
thallus of Filariomyces forficulae Shanor on a mite reveals it to be
abnormal when compared to the thalli from the earwig host.
Photographs were published by Sugiyama (1973), in addition to drawings;
they were trimmed in such a manner that a large number could be placed
on a single plate (for traditionally arranged photographs, see Rossi
[1975], Rossi and Cesari [1976], and Rossi and Cesari Rossi [1977b, c,
1978]).
It is essential that electron microscopy be utilized more fully in the
study of the Laboulbeniales. Hill (1977) not only provided important
information about ascospore production in Herpomyces, but his
illustrations revealed ultrastructural details for the first time in this order.

26
MYCOLOGIA MEMOIR NO. 9
Keys to genera of the Laboulbeniales were published by Benjamin
(1971, 1973b); keys to Japanese species were included by Sugiyama
(1973, 1977). Such keys should be useful for other areas as well,
particularly if precise details of structure and several different
characters are used. A key that includes only a small proportion of the
taxa in a given area may be confusing if only enough characters are used
to separate these taxa, but not enough to distinguish them from other
taxa that may occur in the area. Host indices are also helpful, such as
that included by Sugiyama (1973); particularly useful is the inclusion of
tribes to enable comparison of species on different groups of hosts.
References are helpful for insect names that have undergone recent
changes.
During an intensive study of flies parasitized by Stigmatomyces in
southern France, Dainat (1973) recorded details about habitats and
seasonal fluctuations. A more extensive study of surface beetles in a
variety of biotopes in West Germany (Scheloske, 1969) provided
information about the percentage of infection at different seasons, the
proportion of young beetles present to the number of infected
individuals at different times of the year, the sexes of infected insects of
different ages at different times of the year, and the age of the fungi
from each sex during the various seasons, as well as other data. Similar
studies of different groups of hosts should provide a better
understanding of host-parasite relationships.
DEVELOPMENT, MORPHOLOGY, AND SEXUALITY
Introduction
Despite the many descriptions and illustrations that have been
published, only a few species of the Laboulbeniales have been carefully
studied. Investigators have always had difficulty in understanding
structure and development in this'order. In addition, a number of
misconceptions about perithecial structure arose among the earlier observers
because of the difficulty of making accurate observations of unstained
specimens. Of the two species studied by Robin (1853), only Laboul-
benia rougetii was sufficiently hyaline to enable him to see the
perithecial contents. After the spores were expelled, walls of the outer cells
of the perithecium were more easily observed, but were misinterpreted
as a branched, flexible tract composed of a viscous substance and
occupying the center of the perithecium. In a figure of a younger
perithecium, in which the first transverse divisions of the basal wall cells had
occurred, the longitudinal walls separating the three wall cells that were
visible were* referred to as parts of this viscous tract. Robin was
convinced that these structures were not walls because of their flexuosity
and vague outlines. He believed that the tract functioned as a support
for spores.
This reticulum of walls remaining after spore discharge was also
observed by Peyritsch (1873) in Heimatomyces paradoxus Peyritsch on
Laccophilus sp., as well as in Stigmatomyces baeri (Knoch) Peyritsch. It
was his opinion that the entire contents of the perithecium, with the
exception of the parts used up in spore formation, were transformed
into slime; the remaining framework thus represented the outer
membrane of one large cell. However, intact cells were observed in the neck
region.
The cellular arrangement of young specimens of Laboulbenia guerinii
led Robin (1853) to conclude that the perithecium arises as a lateral
outgrowth of the third cell of the thallus rather than the suprabasal
cell. In Stigmatomyces baeri, Peyritsch (1873) described the perithecial
initial as one of the cells resulting from the diagonal division of the third
cell, the other daughter cell becoming the basal cell of the appendage.
On the other hand, Istvanffi's figure (1895a) of this stage in
Laboulbenia gigantea Istvanffi indicated that the perithecium arises from the
suprabasal cell of the thallus.
27

28
MYCOLOGIA MEMOIR NO. 9
The formation of the perithecial walls puzzled the early investigators.
In figures of Istvanffi (1895a) and Baccarini (1904), the wall cells and
the procarp were labelled as asci at the trichogyne stage. Peyritsch
(1873) believed that the perithecial initial of Laboulbenia became at first
a simple cell row (apparently he included the stalk cells); the lower part
then was divided into a many-celled tissue and the central cell row ended
in a one-celled outgrowth (the trichogyne). A series of wall cell tiers was
later formed.
Because of Peyritsch's confusing descriptions and figures of the
young perithecia of Laboulbenia, de Bary's account (1887) of the
development of the perithecium in the Laboulbeniaceae indicated that
the perithecium divided by successive transverse divisions into three
tiers of one cell each, each tier in acropetal succession dividing
longitudinally into an axile and several parietal cells. Despite Thaxter's
descriptions (1896) of perithecial development in Laboulbenia and Stig-
matomyces, Spegazzini (1917) apparently retained the same erroneous
conception of the origin of the wall cells. On the whole, however, most
authors have accepted Thaxter's interpretation of hyphal growth from
the stalk cell and the secondary stalk cell of the perithecium.
In 1896, Thaxter published a series of drawings showing the ontogeny
of Stigmatomyces baeri, as well as the origin of the secondary stalk and
basal cells of the perithecium and carpogenial cell development up to the
time of ascus formation in Laboulbenia flagellata. No observations
were made on the nuclei of either taxon or on spore development in
Laboulbenia, however. He also illustrated stages in perithecial
development of Enarthromyces. He also showed how the perithecium of Ce-
ratomyces mirabilis Thaxter is formed from two intercalary cells of the
primary axis of the thallus. Later, in 1908, Thaxter illustrated the
arrangement of cells in the pseudop'erithecium of Coreomyces and
showed that perithecial ontogeny in Peyritschiella geminata Thaxter is
similar to that in Enarthromyces and Laboulbenia. The phylogenetic
significance in Euzodiomyces (see Benjamin and Shanor, 1951) and
Colonomyces (Benjamin, 1955) of the production of a perithecium by
intercalary cells of an appendage and the significance of a central
upgrowth from a lateral perithecial cell in Herpomyces (see Richards and
Smith, 1955b) were not recognized by these authors. Thaxter's use of
antheridia as the basis of his classification system apparently has caused
others to overlook the potential importance of other characters.
A comparison of the ontogeny of Laboulbenia flagellata Peyritsch,
on Caraboidea, with that of two species on Gyrinidae, L. gyrinidarum
Thaxter and L. borealis Spegazzini, shows the contrast between the
generalized form of the genus and one of the modifications — that of
theJL. variabilis Thaxter group, found on numerous Caraboidea, as well
TAVARES: DEVELOPMENT, INTRODUCTION
29
as on Gyrinidae. Thaxter (1908) referred to the species of the L.
flagellata group as the normal form of the genus, around which other groups
of species were clustered. Similarly, Herpomyces ectobiae Thaxter and
H. periplanetae Thaxter on Blattaria are members of two different
structural groups within the genus, in which the basic structure of the
perithecium is totally different from that of Laboulbenia.
LABOULBENIA FLAGELLATA. This species has been reported
from at least 40 host genera (Peyritsch's report [1873] on Bembidion is
doubtful and his thallus from Patrobus was not this species). Picard
(1913b) suggested that some of the thalli referred to this species should
be placed in closely related species that have already been found on the
host genera (see discussion by Balazuc, 1974). Colla (1934) concluded
that a large number of species should be placed in synonymy with L.
flagellata. She segregated them into two groups, one with the insertion
cell detached from the perithecium and one with the two joined
together. Thaxter (1896) wrote of L. elongata Thaxter (later put into
synonymy with L. flagellata) that it was a good example of the variation
caused by the nature of the host and the position of the thallus on the
host. Infection experiments should demonstrate the capacity of spores
to germinate on different host genera or different parts of the body.
Morphologically, L. flagellata is characterized by a wedge-
shaped celr V, separated by a short diagonal septum from cell IV, and
by inner and outer basal cells of approximately equal size (PI. l,a,b).
The color is pale to dull brown, never red-brown as in L. rougetii.
In a population of L. flagellata from Alpine Road near Portola Road,
San Mateo County, California (I. Tavares and Francia Chisaki,
November, 1956), the greatest incidence of thalli was on the anterior and
middle dorsal surfaces of the elytra of the host, Agonum ovipennis
(Mannerheim) (syn. Platynus ovipennis [Caraboidea, Pterostichidae]).
Thalli also grew on the posterior part of the dorsal surface of the elytra,
on the upper side of the pronotum, and on the legs. A few thalli were
found on the head, the bases of the antennae, and on the ventral side
of the body. The distribution pattern indicated that the spores were
dislodged from the perithecium in groups. Large clusters of young thalli
were seen which apparently had grown from adherent groups of spores.
Spores also germinated singly or in pairs; the feet of members of a pair
were either arranged side by side or contacted one another at various
angles. Germination was not dependent on the presence of punctae, but
occurred at random on the integument.
Mature thalli exhibited considerable difference in shape, although
there was little variation in overall length (ca. 320 /tm) and the
relationship between perithecium and receptacle (PI. la, b). Young
specimens were very similar in appearance, although at some stages of

30
MYCOLOGIA MEMOIR NO. 9
development there were slight differences in size (Figs. 3,b,c; 4,a). (See
glossary, abbreviations, figure interpretation.)
LABOULBENIA GYRINIDARUM and L. BOREALIS. Several
species of Laboulbenia bearing black-septate appendages have been
described from aquatic beetles of the family Gyrinidae. The dark color of
the outer walls of these fungi (as well as the multiplication of cells and
appendages of the upper receptacle) obscures their cellular structure, so
that it may be only when young thalli are available for study that an
understanding of their anatomy can be obtained.
Laboulbenia gyrinidarum was described from Gyrinus species by
Thaxter (1892) and L. chaetophora Thaxter was found on Dineutus soli-
tarius Aube" (Thaxter, 1905). The latter species is extremely large
(800 fim in height) and has a broad-based, conical perithecium with
spiralled outer wall cells and a straight receptacle base (PI. 6,c). The
perithecial apex bears a dark spine 40 /tin long, as well as a slightly
shorter protrusion. Thaxter (1908) pointed out that two distinct forms
occur on North American gyrinids — in one of them, the perithecium
is narrow, with straight wall cells and short, rounded, apical
projections, the receptacle is usually curved at the base, and the insertion
area of the appendages is small. In the other form, the general
proportions are somewhat similar to those of L. chaetophora, although the
thalli are smaller. Thaxter (1908) referred the latter thalli to L.
chaetophora.
In general appearance, the thalli I studied were usually intermediate
in size and somewhere between these two forms in shape (PI. 5,b). On
the basis of most of the characters used by Thaxter to distinguish the
two species, therefore, one might conclude that the two forms simply
represent extremes within a species in which there is a great amount of
variation.
However, a long, dark, spathulate outgrowth (the umbo) occurs
opposite the elongate spine of L. chaetophora on Dineutus (Thaxter,
1908, PI. LXVII, Fig. 19) and a similar outgrowth (ca. 5-6.25 /tm long)
is also found on specimens on Gyrinus that are referable to L.
chaetophora on the basis of other characters, such as those mentioned
previously. On the other hand, the protrusion in this position in thalli that
are referable to L. gyrinidarum is less than half the length of that in the
other taxon (they are ca. 1.75-2.75 /tin long). In most of the thalli of L.
gyrinidarum observed in my study, this projection was opposite two
large hooklike appendages (unci) (PI. 4,b,c). In the type of L.
gyrinidarum (Thaxter, 1896, PI. XXII, Fig. 31) (PI. 5,a), these unci are
reduced to short, rounded extensions about the same size as the umbo. In
some thalli from Gyrinus resembling L. chaetophora, the long spine
breaks off, or perhaps fails to develop. The adjacent short uncus varies
TAVARES: DEVELOPMENT, INTRODUCTION
31
in size and shape, but it is usually quite inconspicuous. However, the
long, dark umbo is always present (PI. 5,d) and serves to distinguish
these thalli from those with the short umbo, regardless of the character
of the other outgrowths.
The umbo is located over the wall between the terminal perithecial
cells on one side (PI. 4,b). However, it originates above one wall cell
row, not over both. It gives the appearance of a structure composed
entirely of wall material. There is no strand of cytoplasm entering it.
The unci and the spine — one anterior, one posterior — are extensions
of the cells opposite (PI. 4,c), into which cytoplasm passes, at least at
early stages of development. In addition, there is sometimes a low hump
visible between the level of the trichogyne stump and the perithecium
apex (cf. Pis. 4,b-e, 5,a-d).
In my study, it was observed that thalli growing on the elytra margins
of Gyrinus were L. gyrinidarum, whereas those on the abdomen tip
almost always resembled L. chaetophora. These observations were more
or less comparable to those of Faull, who found the L. chaetophora
form most frequently on the abdomen and L. gyrinidarum most often
on the elytra (Faull, 1911, 1912). Infection experiments should be done
to determine whether spores of the former taxon will germinate readily
on elytra, and if so, whether the perithecial apex is affected by the
difference in position.
Balazuc (1971c) suggested that specimens on Gyrinus which had been
referred to in the literature asX. chaetophora were probably L. borealis.
Spegazzini (1915a) indicated that the suprabasal cell (II) of L. borealis
is short and broad and the perithecial apex terminates in a prominent
black outgrowth; opposite this structure are two colorless subspinose
processes. The thallus that he illustrated (Fig. 13, p. 468) had these
characteristics. No thallus identical to his drawing is present on the
type slide, which bears a specimen referable to L. gyrinidarum (having
two short unci and a broad cell II), as well as a thallus with a narrow cell
// and a solitary dark apical outgrowth opposite a short umbo. The
third thallus, which is hereby designated the lectotype (assuming that
Spegazzini's collection does not include any thalli in which the spine is
colorless and perhaps has broken off), has a long umbo, a dark spine,
and a small, pale outgrowth, as well as a short, broad cell II; this thallus
is broken, like that of Spegazzini's drawing (1915a), judging from the
space he showed between the apical outgrowths. It is only on a thallus
having a long, dark umbo and two other outgrowths that one may find
three prominent structures of the size shown by Spegazzini. Well-
developed thalli of L. gyrinidarum only have two prominent
outgrowths. Although Spegazzini reported finding the taxon on elytra, the
nature of the mixed collection suggests that one of the thalli may have

32
MYCOLOGIA MEMOIR NO. 9
come from the abdomen of the host. The type locality, given as Bezu-
lian, N. America, may actually be Brigantine, a suburb of Atlantic City,
New Jersey, which is within the range of the host, Gyrinus borealis
Aube.
Spegazzini did not compare L. borealis with L. gyrinidarum. He
contrasted the shape of the broad // cell with the narrow // cell of L. gy-
rinicola Spegazzini, a European species. Some of the thalli in my
collection have either short /// and VI cells or a short cell //. However,
the specimen shown in PI. 5,c has a tall, narrow //. Apparently there is a
certain amount of variation in cell shape. Spegazzini's illustration shows
rather small IVand Vcells.
The most constant characteristic of L. borealis, the commonest taxon
on the abdomen of Gyrinus in the United States and Canada, is the long
umbo. It is possible that L. borealis is more closely related to L. chaeto-
phora than it is to L. gyrinidarum. Each of the populations that I have
studied, which parasitized Gyrinus plicifer LeConte (stream above Lake
Anza, Tilden Regional Park, Contra Costa County, California, I.
Tavares, 1948-1955; below Alpine Dam, Marin County, California, I.
Tavares, September, 1957; and near Lagunitas, Marin County, I.
Tavares, October, 1956), have included thalli with slightly different
kinds of apical outgrowths (cf. Balazuc's comments [1971c] on L. gy-
rinicola).
Although it is difficult to determine how environmental factors might
influence the development of the perithecial outgrowths — unless
especially favorable growth conditions contribute to more extensively
developed structures (Scheloske, 1976a) — other characteristics of thalli
are definitely affected by the habitat. The cell length and amount of
curvature in cells / and // are clearly related to the position of growth
(PI. 6,d). Thalli on the tip of the abdomen tend to extend straight
outward, whereas those on the elytra margins and the sides of the abdomen
just beyond the rear edge of the elytra tend to curve forward as they
grow, as a result of the curvature of the base of the receptacle. As the
thallus grows erect from the horizontal position in which it germinates ^
the configuration of the substrate, the presence of adjacent
obstructions, and probably also the pressure of water currents influence its
form. It is clear that size of the thallus also can be affected by the
environment — thalli growing in locations where a better supply of
nutrients is available tend to be larger in size and perhaps have a more
luxurious growth of appendages. In addition, one member of a pair
growing together is often stunted (PI. 6,d). Although the amount of
curvature of the perithecial wall cell rows depends to some extent on the
growth conditions, it probably has a definite relationship to the width of
the base of the perithecium.
TAVARES: DEVELOPMENT, INTRODUCTION
33
The majority of thalli on Gyrinus plicifer were found on the dorsal
surface of the posterior lateral margins of the elytra. Many were also
found on the outer dorsal margins of the tip of the abdomen. Rarely,
beetles were collected on which the fungi grew on the legs or on the
dorsal surface of the pronotum. As Faull (1912) pointed out, L.
gyrinidarum and L. borealis are not clearly distinguishable when immature.
However, the assumption may be made that those occurring on the
elytra are L. gyrinidarum and that those on the abdomen are mostly L.
borealis. Occasional mature thalli of L. gyrinidarum have been seen on
abdomens.
HERPOMYCES ECTOBIAE and H. PERIPLANETAE. Unique
among the species of Herpomyces is H. ectobiae, in which the male
thallus, instead of being of limited size, grows in the same manner as the
female. Herpomyces ectobiae occurs on the common German
cockroach, Blattella germanica (L.), primarily on the antennae. Thalli are
frequently found on the legs also and sometimes they occur on the
caudal cerci and the maxillary and mandibular palps. Less
frequently they grow on the abdomen and upper wing surface. On the an-
tennal annulations, thalli are usually found on the long, distal setae and
the larger of the setae proximal to them. Very often, these setae are
covered with long rows and clusters of germinated spores. Of the few
spores seen apparently lying on the surface of the antennal integument,
none appeared to have progressed beyond the four-celled stage.
However, because the annulation is covered by closely spaced bands of
minute setae, it was impossible to see whether the germinated spores
were attached to one of the setae or to the surface of the antenna itself.
The ostiolar apertures of the perithecia extend far out toward the
apices of the larger setae. The spores are probably detached from the
perithecia by the setae during contact between antennae, or between
antennae and other parts of the body, such as the mouthparts. The
spores, which like those of Laboulbenia, are viscid, rarely fall to the
bases of the setae, adhering instead to their sides.
The most distinctive characteristic of Herpomyces periplanetae Thax-
ter, parasitic on Periplaneta americana (L.), is the presence of a broad-
based, more or less triangular, multicellular shield growing at one side
of the female thallus. In Thaxter's original description (1902), he
mentioned thalli without shields growing on conspicuous setae and
thalli with shields growing directly on the integument. Shields of
individuals growing on Blatta orientalis L. (syn. Stylopyga orientalis)
were usually much smaller than those on Periplaneta americana,
according to Thaxter, who noticed that they were frequently absent in
thalli on the former host. Spegazzini (1917) concluded that there were
sufficient differences between the fungi occurring on these two host

34
MYCOLOGIA MEMOIR NO. 9
species to warrant separation of the form on Blatta as Herpomyces
stylopygae (see Tavares, 1966). In addition to a more slender shield,
Herpomyces stylopygae has a perithecium with parallel sides and narrow
neck cells, whereas H. periplanetae has a wider, rounded venter and
some neck cells that are wide in section.
All intact thalli observed, except some small four-celled individuals,
had germinated on minute setae of the antennae (PI. 20,b; Fig. 16,a,b)
(those illustrated by Thaxter [1908] on large setae are clearly H.
stylopygae). It is possible that further growth does not occur in spores
attached to large setae because the foot haustorium is unable to
penetrate the setal wall. Although Richards and Smith (1954) did not
indicate the exact mode of growth of the thalli of H. periplanetae
that they were able to grow on Blatta, one may assume that they were
shield-forming thalli on small setae; in such thalli, the distinctions
between the two species should be quite noticeable. Because H.
stylopygae is able to germinate on large setae of its host, it may be assumed
that the haustorium of the foot is capable of penetrating a thicker wall
than that of H. periplanetae. It is difficult to comprehend their inability
to induce H. stylopygae to germinate on Periplaneta; possibly there is a
physiological barrier.
Ontogeny of Laboulbenia
On one side of the prominent swelling near the base of the spore of
Laboulbenia gyrinidarum and L. borealis, a small circular area may
appear within the thick cell wall close to the cytoplasm (Fig. 8,a). At
first, this area stains lightly with acetocarmine. In slightly older spore-
lings, a dark-staining, blackening disk is present on the outer surface
of the narrow inner wall of the cell, whereas a yellowish brown granular
ring lies outside it near the outer surface of the outer spore wall.
Although these two circular areas appear to be almost the same diameter
in some thalli, in others the inner disk is much smaller and appears to be
surrounded by the ring in face view. The relationship between the two
dark areas may vary, although they always seem to be close together.
The cytoplasm above the darkening base of the foot begins to protrude
into the spore wall beyond the disk (although the disk may appear when
the spore is inside the perithecium, the swelling of the foot may be, in
part, a response to the substrate on which the spore germinates). The
ring resembles those that encircle haustorial apertures in other taxa.
However, the cytoplasm seems to approach the surface of the thallus
only above the ring, where the outer wall flattens out and becomes
appressed to the integument of the host. The cytoplasm presses against
this surface over a wide area, rather than entering the swelling as a
TAVARES: LABOULBENIA ONTOGENY
35
narrow haustorial strand. The blackening disk appears to become lens-
shaped; in some thalli, the inner part of the disk is lighter in color and
takes up acetocarmine, whereas the outer part is black.
Meanwhile, sparse, yellowish brown granules may appear over the
entire wall of the foot, or the darkening may be denser and is limited to the
base of the foot at first (Fig. 8,b). In some thalli, dark extensions
emerge from the disk-ring region, which becomes more indistinct. The
base of the foot usually becomes blackened first, with the black area
extending upward around the swelling of the side of the foot (Fig. 8,c).
The thinner wall of this enlargement remains clear until a later stage of
development. The foot increases in size at first, but as the rest of the
thallus grows, its relative size decreases markedly (PI. 6,d; Fig. 8,d; cf.
PI. l,a,d).
Early stages of foot development were not observed in Laboulbenia
flagellata. On the side of the foot that is attached to the integument,
a swelling develops that elevates the upper part of the germinating
spore. The upper curved surface of the swelling blackens, leaving a
conspicuous, clear area inside this dark margin (PI. l,d). The wall of
this area becomes filled with dark granules. The sequence of the two
divisions in the basal cell of the spore was not determined in the
numerous thalli having three lower cells and a primary appendage of a
few cells (PF. l,d). Nuclear division in cell c results in the formation of
cells d and // (Fig. l,a). The insertion cell e and the basal cell of the
outer appendage (/) are formed. Cell e becomes triangular in section and
cuts off cell g anteriorly (Fig. l,b). The cell wall of e turns yellowish,
then gradually blackens, obscuring the cell contents (PI. 2,b; Fig. 2,b).
Lateral branches may arise from other appendage cells (Pis. l,e, 2,b).
Cell g divides to form the inner appendage (PI. 2,b; Fig. l,c); sometimes
it produces a lateral branch directly (Fig. 2,a). The upper wall of cell/
is darkened in some thalli (PI. 2,c, Fig. 1 ,a).
in L. gyrinidarum and L. borealis, the first division within the
germinating spore usually occurs in the small upper cell, which develops
into the primary appendage. The first two septa formed in the
appendage are oblique and undulating (Fig. 8,c). These septa soon turn
brown; later they appear to be black at the outer margin. A prominent
small central pore is present in each septum. The next division usually
occurs in the lower spore segment; two septa are formed, dividing it into
three cells, as in L. flagellata. The sequence of division varies.
When the first two septa in the appendage have become quite dark, a
third is formed just above the original spore septum, cutting off cell e
below and cell / above. The middle appendage septum becomes very
elongate and oblique (Figs. 8,d; 9,a); its pore remains near its base (Fig.
9,b). The primary appendage elongates and becomes multicellular

36
MYCOLOGIA MEMOIR NO. 9
above the upper dark septum, the upper cross-walls remaining hyaline;
a prominent lateral branch grows out anteriorly between the two black
septa (Fig. 8,d).
Meanwhile, the septum between cells e and / may blacken, but it
usually remains unmodified (Figs. 8,d; 9,c). Division of cell e occurs,
resulting in the formation of cell g (Fig. 8,d; the septa between e and
cells g and g', which is formed later [Fig. 10,a], do not darken). Cell
/cuts off an anterior cell; the septum, which soon darkens, is located at
right angles to the elongated black septum above cell /. The newly
formed cell then divides; the resulting septum, which is sometimes
dome-shaped, may intercept the previously formed septum at or near its
junction with the elongated septum, or it may extend to the latter
septum instead (Figs. 8,d; 9,a). The second anterior septum (from
division of cell/) blackens and thickens (Figs. 8,d; 9,a); another black
septum may be formed above the second one, adjoining the upper end of
the elongated septum. In the formation of the dark septa of the
appendages, the narrow initial wall darkens, so that in lateral view, the dark
region appears to be bordered by a wider hyaline wall layer; the outer
edge appears darker also (PI. 6,b). Later, the entire septum becomes
blackened (see also Faull, 1912). Appendage branches may grow out on
the anterior side of each of the cells between/and the uppermost black
septum. The number of branches varies, as well as the sequence of
appearance. Branches are often broken off; proliferation of a new
hypha may occur through the broken end of the main axis of the
appendage.
After cell d has been formed in L. flagellata, cell b undergoes
division, first cutting off cell V and later cell IV (PI. l,e; Figs. l,b,c [cf.
PI. 6,a; Figs. 8,d, 9,a]). In some older thalli (PI. 2,b, left), there may be
two nuclei (or possibly two cells) in the position of cell V; however,
in almost all thalli, only one nucleus is present, even at much later stages
of development.
In L. gyrinidarum and L. borealis, cell V cuts off an elongated cell
{Va) at its apex (Fig. 9,a); after a second division in V, a smaller cell,
triangular in section (Vb), is formed. Sometimes both cells Va and Vb
are triangular in section and each arises at one side of the top of cell V.
Cell Va soon produces the first secondary appendage, the second arising
from cell Vb (Fig. 9,b). During initiation of each secondary appendage,
the septum formed during cell division soon darkens so that the small
central pore is readily visible. Meanwhile, the first g cell produces an
appendage (Fig. 9,c). By the time a large number of appendages have
been produced by cells Va, Vb, g, and g', only a black stump remains
of the main axis of the primary appendage (Figs. 9,c; 10,a). The
diameter of this portion of the appendage increases to almost 12 /tin,
TAVARES: LABOULBENIA ONTOGENY
37
whereas it measures only about 6.5 /tin when the dark septa are first
formed. Cell IVproduces a small upper cell (IVa), which is triangular in
vertical section and resembles cell F(Fig. 9,c). Another, much smaller
cell (IVb) is then formed next to it (Fig. 10,a). In some thalli, the order
of appearance of these two cells appears to be reversed. Secondary
appendages later arise from cells IVa and IVb.
Only the appendages arising from the g cells are comparable to the
anterior branches of the primary appendage of L. flagellata, in which
there are no secondary appendages (see Fig. 2,a).
While cell division takes place in the upper receptacle, cell d divides
to form cells h and / (Figs. l,b; 9,a). Division of cell h results in the
formation of cells j above and k below (Figs. l,c; 9,b). Cell k extends
upward posterior toy and divides into Wand m (Fig. 9,d [this division
was not observed in L. flagellata, but cf. Fig. l,c]). Celly then cuts off
cell n from its broad, upper part (Fig. 10,a) on the side of the peri-
thecium opposite cell m. A second division of cell j results in the
formation of cells n' and VII (Fig. 10,a). This division is followed by the
formation of cells o and o' from cells n and m (Figs. 2,b,c) and the
division of cell / into the trichogyne initial and the carpogenic cell.
The orientation of the cells of the perithecium during growth is not
constant wijh respect to the side of the perithecium on which they are
formed. In the thalli shown in PI. 2,b (center) and PI. 2,a, cell m is on
the upper side, whereas cell n is at the lower surface. In the thallus
illustrated in Fig. 2,a, which is oriented in the same way, these two cells
have been cut off on sides opposite to those in the above-mentioned
individuals. Apparently, cell k may grow upward along either side of the
broad part of celly, which becomes separated as cell n. The n' cell then
develops beside cell m, on the side of cell VII opposite cell n. On the side
next to m, cell VII is vertically oblong in section, but it broadens out
toward the other surface into a cell that is roughly triangular in
longitudinal section (Fig. 2,c). These variations in shape are apparently
caused by the pressures exerted by the adjacent cells within the outer
wall of the perithecium. Cell VII directly underlies the procarp (Fig.
2,a).
The trichogyne cell of L. flagellata divides into a multicellular,
branched structure. Destruction of its terminal cells results in the
formation of lateral branches below the deteriorating cells. The
cytoplasm in the actively growing apices is dark-staining. The tips of the
branches sometimes appear to grow toward the orifices of the antheridia
with undulating movement, so that the branches are spiralled.
The first divisions of the trichogynes of L. gyrinidarum and L.
borealis were not observed. The basal cell of the trichogyne is typically very

38
MYCOLOGIA MEMOIR NO. 9
short; its upper septum is blackened (Figs. 10,b-d). The cells of the
trichogyne are very similar to those of the appendages. The short lower
cells are separated by black septa; the terminal cells may become very
long and tenuous. By the time the first inner wall cell appears, the
trichogyne may consist of three or four cells. In some thalli, a dark disk
and a pigmented area appear near the base of the lower trichogyne
cell (Fig. 10,d); in others, the disk appears just below the dark septum.
The entire wall of the lowest cell eventually darkens.
An intercalary division of the carpogenic cell (cp) results in the
formation of the trichophoric cell (tc), which may appear while the o cells
are still being produced. Prior to division in L. flagellata, cp extends
upward so that the trichophoric cell occupies the same position as the
initial trichogyne cell with respect to the insertion cell (Fig. 2,c). A
different sequence of cell formation was reported in Stigmatomyces baert
(Thaxter, 1896), in which the trichophoric cell is produced before the
trichogyne; however, in Stigmatomyces the trichogyne cell is a small cell
cut off the apex of the trichophoric cell.
The basal wall cells o and o' elongate until they extend slightly above
the apex of the carpogenic cell (Fig. 3,a; cf. Figs. 7, 10,b). At the same
time, cell n' enlarges and divides, producing cell o", while cell n cuts off
the small cell o'" (Fig. 3,a; o cells designated in Fig. 7). Thus, one of the
two adjacent wall cell rows arising from the wedge-shaped n cell is on
the free (anterior) side of the perithecium, whereas the other is on the
posterior side. The arrangement of the 4 rows of outer wall cells is 2
posterior and 2 anterior; these cells enclose the carpogenic cell on all
sides (Fig. 3,a).
The inner wall (parietal) cells, which are much narrower than the
outer wall cells, are also cut off from cells, m, n, and n' (Figs. 3,b,c;
4,a,b; PI. 2,c); they become arranged alternately with the outer wall
cells, one being anterior (one of those from cell n), one, posterior (that
arising from m) (Fig. 3,b), and two, lateral (one each from n and h5) (PI.
2,c). Thus, cell n gives rise to 2 inner and 2 outer wall cell rows. The first
of the inner wall cells of L. flagellata arises as an upward extension of
the inner edge of the upper surface of cell m (PI. 2,a), whereas the
second is produced by n' (Fig. 3,c). In L. gyrinidarum and L. borealis,
the initial inner wall cell may arise either from cell n'orm (Figs. 10,b,c).
Meanwhile, cells o and o' have each divided to form an apical wall
cell (w). In L. flagellata, cells o" and o'" divide at approximately the
time that the second inner wall cell is formed (Fig. 3,c, specific o cells
not indicated; see Fig. 7). The third and fourth inner wall cells then arise
from the upper part of the wedge-shaped cell n. The inner wall cell
nuclei are flattened, so that they appear to be oval in lateral view (Figs.
4,b, 5,a).
TAVARES: LABOULBEN1A ONTOGENY
39
The septum between the trichophoric and carpogenic cells in L.
flagellata remains narrow. It has the appearance of a recently formed
wall until its disappearance. The carpogenic nucleus may appear to be
in prophase before the septum breaks down, at a time when there is no
evidence of division in the trichophoric cell. On the other hand, there
may be two nuclei in the trichophoric cell before there is nuclear division
in the carpogenic cell. In the few examples of this stage that were seen,
one of the nuclei in the trichophoric cell resembled the deteriorating,
vacuolate nuclei of the lower trichogyne cells, whereas the other more
closely resembled the carpogenic cell nucleus (i.e., appeared to be in
early prophase). Divisions were not seen.
The disappearance of the septum between the trichophoric cell and
the carpogenic cell in L. flagellata takes place just before cell division in
the inner wall cells. In L. gyrinidarum and L. borealis, the septum
breaks down during elongation of the inner wall cells (Fig. 10,d). A
number of three-nucleate trichophoric-carpogenic cells were observed in
L. flagellata; the three nuclei were similar in size or, as in PI. 3,a, one of
them was larger than the others. In none of these three-nucleolate cells
was a deteriorating nucleus seen, such as that mentioned above in the
trichophoric cell. In the majority of thalli of L. gyrinidarum and L.
borealis that had a fused trichophoric-carpogenic cell, there were only
three nuclei present. This indicates that either the nucleus of the
trichophoric cell or that of the carpogenic cell had divided.
In L. flagellata, the four inner wall cells divide when their apices have
extended almost to the tips of the outer wall cell rows. Their narrow
nuclei are located approximately in the center of each cell; after
division, the upper daughter nucleus moves into the upper third of the
cell; the lower may move downward somewhat. A septum cuts off a
small terminal cell (x) (Figs. 4,c; 5,a). The inner wall cells do not always
divide concurrently (Fig. 4,c; PI. 3,a). At this time, another cell is
formed at the apex of each of the outer wall cell rows; these divisions do
not occur simultaneously (PI. 3,a).
A septum isolates the uppermost nucleus of the row of four nuclei
resulting from divisions taking place in the fused trichophoric-
carpogenic cell of L. flagellata (fc*) (Fig. 4,c); the lowermost nucleus is
segregated in an inferior supporting cell (Fig. 5,a). Formation of either
of these two cells may precede the other. Apparently, two divisions of
the binucleate central cell usually follow (these two nuclei presumably
include one from the carpogenic cell and a trichophoric or male
nucleus); as a result, a binucleate superior supporting cell is formed above
the ascogonium {am) and a binucleate secondary inferior supporting cell
is formed below the ascogonium. This sequence of development is
essentially the same as that reported by Faull (1911, 1912) in L.
gyrinidarum and L. borealis (as L. chaetophora).

40
MYCOLOGIA MEMOIR NO. 9
The secondary inferior supporting cell of L. flagellata may extend
upward; the upper nucleus migrates into the upgrowth (PI. 4,a). This
outgrowth later is cut off by a septum (Fig. 6). It is not known how
frequently this extension is formed, because few thalli permit
observations of internal details at this stage of development. Such an
outgrowth was not reported by Faull (1911, 1912) in either of the two
species he studied.
The ascogonium retains a deeper stain than the other cells of the
central cell row. The non-functional cells exhibit varying degrees of
deterioration of cytoplasm. Their chromatin tends to become more
heterogeneous in appearance.
Another division of inner and outer wall cells takes place, resulting in
the formation of the w' and x' cells. The terminal cells begin to grow to
one side of the deteriorating trichogyne (Fig. 5,b). At the stage of
development represented by Fig. 6, the trichogyne is located
approximately at the level of the center of the w' cells. It marks the position of
the last outer wall septum, which is formed at maturity. In L. gyrini-
darum and L. borealis, the short basal cell of the trichogyne remains
embedded in the perithecial wall. Although the distal cells usually break
off, sometimes intact cells remain (PI. 4,e).
Supernumerary nuclei appear in the subterminal cells of some of the
vertical rows of inner and outer wall cells in L. flagellata (Fig. 5,b).
Multiplication of nuclei may also take place in the lowermost outer wall
cells; presumably these divisions occur later.
The w cells of L. flagellata elongate until they are more than twice as
long as the o cells. Because of the blackness of the perithecial apex, it is
not always possible to see whether an additional division of the wall cells
has taken place. However, in one thallus, small cells had been
produced at the tips of the w' cells even though division of the x' cells had
not occurred.
In L. flagellata, both of the pairs of outer wall cell rows are about the
same size; the n cell is both larger than and different in shape from the
n' cell. In L. gyrinidarum and L. borealis, there is little difference in size
and shape between these cells. The smaller size of cell n apparently may
influence the size of the wall cells produced by it. In some thalli
observed, o and w cells formed by cell n were as little as 1/2 as wide as
those from the m and n' cells. These size differences might have some
effect on the amount of wall spiralling.
When ascus production begins in L. flagellata, terminal swellings are
formed on the w' cells (Fig. 6). The two posterior cells broaden apically,
while the anterior cells extend upward very slightly and remain
acuminate at the tip. As the perithecium matures, its outer wall turns a
TAVARES: LABOULBENIA ONTOGENY
41
yellowish brown color and its apex blackens. The persistent base of the
trichogyne may emerge at the base of the black apex (PI. l,b).
The older parts of the outer wall of the perithecium and receptacle are
quite dark by the time the ascogenic cells are formed in L. gyrinidarum
and L. borealis. By contrast, the newly formed wall of the perithecial
apex is hyaline. As the w' cells elongate, a cleft forms at the perithecial
apex along the line of the cell walls between the outer wall cell apices. As
the outer wall cell rows grow in length, the inner wall cell rows also
elongate and divide to form the same number of cells. The lowermost inner
wall cells become extremely attenuated because of the lateral pressure of
the expanding mass of asci. The terminal wall cells contain
homogeneous cytoplasm, whereas that of the subterminal cells becomes quite
alveolar in appearance and the cells may become multinucleate.
During formation of the terminal perithecial outgrowths in L.
gyrinidarum and L. borealis, two apical outer wall cells extend upward while
the other two broaden slightly. The terminal wall above the latter
thickens across the intervening septum. It protrudes upward and
darkens. Meanwhile, the paired extensions characteristic of each species
develop. Sometimes there is a small lateral outgrowth; blackened areas
develop in an irregular pattern.
As the outgrowths develop, pigmentation appears at the junctions of
the walls and on ridges on the terminal outer wall cells; later, dark spots
are formed on the outgrowths. Eventually the w' cells probably divide
to form an additional row of outer wall cells, although the darkness of
the perithecial wall makes it difficult to determine whether this division
is a constant occurrence.
Even before procarp division (Fig. l,a), the primary appendage of L.
flagellata may have broken, resulting in a forked appendage if a large
lateral branch is present. Lateral branches may also break off (PI. 2,a);
proliferation of a new hypha within the old walls of the broken cell
often follows. This breaking and proliferating process may be repeated
several times. The cytoplasm in the proliferating cells is more
homogeneous and darker staining than that in the cells from which they arise.
The outer walls of these cells are considerably thinner than those of the
broken cells through which they grow.
In L. flagellata, the first phialides are borne on the inner appendage,
which eventually becomes highly branched, with clusters of short
terminal branchlets bearing antheridia (PI. 3,a). One or a few of the
branchlets in a group may remain undifferentiated, capable of
continued division into additional vegetative cells or phialides. Sessile
phialides sometimes develop on the large, lower cells of the outer branch of
the primary appendage (PI. 2,a), where they may occur in pairs. Quite

42
MYCOLOGIA MEMOIR NO. 9
late in development, large antheridial branches may arise on the
lowermost outer primary appendage cells, when only a short stump may
remain of the original primary appendage. These branches always
appear on the anterior side, toward the perithecium. This tendency may
be seen also in the formation of new branches of the inner (anterior)
appendage, which may occur long after the trichogyne has disappeared
and the perithecium has extended upward past the trichogyne stump.
The antheridia seem to undergo a rather short period of spermatium
production; branches characteristically bear empty antheridial walls, as
well as old antheridia through which new cells are proliferating (PI.
3,b). These proliferating cells may extend into empty walls, or they may
grow up through cells that still retain either cytoplasm and nucleus or
cytoplasm alone.
The first spermatium is apparently cut off by a broad septum formed
in the short broad neck of the phialide. During this process, the narrow,
refractive inner wall of the phialide pulls away from the refractive outer
wall layers in the portion of the antheridium that becomes the
spermatium. The tip of the cell breaks down, forming an exit pore.
Subsequent spermatia are formed at a slightly lower level by the upgrowth of
the antheridial protoplast into the neck region. Simultaneously, a collar
is laid down on the inside of the outer layers of the antheridial wall. This
collar fills the space between the broad upper part of the antheridial
protoplast and the broadest part of the spermatial upgrowth when the
latter is fully extended. It may stain faintly with acetocarmine (PI. 3,f).
The spermatial extension may become quite long before nuclear
division occurs in the phialide, or it may develop concurrently with the
migration of the upper daughter nucleus into its tip. At telophase, the
daughter nuclei become widely separated (PI. 3,b). The lower nucleus
remains in the base of the antheridium. The chromosome number
appears to be n = 4, as in L. gyrinidarum and L. borealis (Faull, 1912 — L.
borealis referred to as L. chaetophora); two of the chromosomes are
large, one is medium-sized, and one is small. When the upper nucleus is
in the fully extended tip of the upgrowth, a septum grows across the
isthmus, leaving a central pore. As the spermatial wall separates from
the apex of the antheridial protoplast, it may be pulled downward at the
sides, probably because of a delay in separation of the two layers. At the
same time, the antheridial protoplast rounds up at the apex and retracts
preparatory to forming a subsequent upgrowth.
Occasionally, the normal pattern of spermatium formation is
disrupted and teardrop-shaped spermatia are formed that lack some of the
chromatin of the nucleus or are completely enucleate (PI. 3,c). The rest
of the nuclear material rounds up into a mass in the antheridium. At
times, there are two or three masses of chromatin in an antheridium.
TAVARES: LABOULBENIA ONTOGENY
43
That there may be at least six spermatia formed by a single phialide was
indicated by an antheridium that was observed in which an apical pore
did not form to release the spermatia.
Only a thin layer of cytoplasm surrounds the ovoid to ellipsoid
nucleus of the spermatium. At first, most of the cytoplasm is at the base,
but it appears to become more evenly distributed later, as the
spermatium rounds up at the base. The nuclei are very homogeneous in
structure; the entire spermatium stains more heavily with acetocarmine
than does the antheridium. The spermatium possesses a thin wall of the
same thickness as that surrounding the antheridial protoplast.
By maturity, some of the upper receptacle cells of L. gyrinidarum and
L. borealis are covered with large numbers (10-20) of small, black, basal
septa; most of the branches distal to these cross-walls have entirely
disappeared. With age, a brown suffusion develops in the walls of these
cells around the dark septa. The basal secondary appendage cells may
produce small branches; as more branches are formed, these cells
enlarge and darken. The second and third cells of each secondary
appendage also enlarge. A fascicle of appendages may be formed from
the upper side of each enlarged cell. A multiplication of branches may
also occur on the lower cells of the primary appendage.
The appe.arance of the appendage system varies; although there are
undoubtedly genetic factors involved, mechanical influences and the age
of the thallus are probably responsible for much of this variation. Some
of the terminal cells of the secondary appendages (and primary
appendage branches from the g cells) are very broad and short, with small
papillate apices. Others are long and narrow, with walls that are much
thinner than those of the basal parts of the appendage. In the Alpine
Dam population, secondary appendages in immature thalli either
terminated in short cells or exceedingly long, attenuated extensions which
were sometimes over 100 /tin in length, instead of the normal terminal
cell length of 4-5 /an and intercalary cell length of 5-8 /an; the width of
these long cells was about 3 /on for most of their length, instead of the
normal appendage width of 5-6 /an. Some of the apical extensions were
multicellular with hyaline septa. Occasionally, the terminal cell of an
appendage may be antheridia-like in shape. In thalli from the Alpine
Dam population, division of some of the terminal cells resembled
spermatium formation very closely. Small cells were cut off squarely, as
are many spermatia in L. flagellata. The cell nucleus migrated up into
the daughter cell, leaving the mother cell enucleate. The daughter cells
in these appendages differed from spermatia in having more cytoplasm
below or surrounding the nucleus.
When breakage occurs, some appendages produce long, lateral
upgrowths which originate just below the black septa of the intercalary

44
MYCOLOGIA MEMOIR NO. 9
cells. The apical cell wall of an appendage may bear a discoid, brown
area, either terminal or somewhat lateral in position; this occurrence is
rare. In one thallus observed, a lateral extension arose from just below
such a terminal disk. The upgrowths appear to arise as narrow papillae.
As each one elongates, it may enlarge in the lower part, while still
retaining the narrow papillate apex, or it may maintain approximately the
same diameter throughout its length. Such extensions are narrower in
diameter than normal appendage cells and may be three or four times
their length. In thalli in which most of the appendages have broken off,
many lateral branches may be present; in those in which breakage is
rare, there may be only one or two of these branches. The nucleus may
remain either within the appendage cell itself or move into the base of
the lateral extension. Sometimes the nucleus migrates later up into the
branches and septation occurs; the basal cells eventually appear to lose
their protoplasts. The growing tips are the most heavily staining
portions of the appendages.
Ontogeny of Herpomyces
Spores almost always germinate in pairs. Usually only one or two
well-developed thalli occur on each of the large setae on which the
spores of Herpomyces ectobiae germinate, the remaining spores being
arrested in development. Germination of this species may take place
anywhere on the surface of the seta; the position, however, determines
to a considerable extent the length of the secondary receptacle, which is
produced as a lateral extension from the suprabasal cell of the primary
receptacle (PI. 11,a). Reproductive structures are usually formed near
the base of the seta; if germination occurs at a considerable height from
the base, the secondary extension may grow to a length of about 50 /xm
before it produces a perithecium or antheridial tuft.
Normally, the secondary extension is initiated from the side of the
thallus opposite the haustorium of the foot, but sometimes it arises at an
angle of 90° to one side of the haustorium. The outgrowth is composed
of a row of cells which later become arranged in an irregularly double
series as a result of intercalary divisions. From the bases of the
secondary receptacle cells, haustoria penetrate the cuticle of the setal wall or
that of the integument (see PI. 11 ,c).
The secondary receptacle, which may become bifurcate, grows down
toward the base of the seta in large thalli and out over the surface of the
integument. A single thallus may be very large and extend almost the
entire length of an antennal annulation. Males seem to develop less
often than females into extensively spreading thalli, especially when
both male and female individuals develop on the same seta.
TAVARES: HERPOMYCES ONTOGENY
45
Although both ends of the germinating spore of if. ectobiae blacken,
development of the foot precedes that of the apex. The foot undergoes
little expansion, although it becomes slightly larger than the spore apex.
At first, the tip of the blackened foot is hyaline, but it later darkens. At
the upper edge of the foot, on the side toward the seta, a brownish black
ring surrounds a clear area (Fig. 1 l,a); the haustorial opening is located
in the lower part of this clear area. The convex ring later darkens,
sometimes before further cell division occurs. The blackening of the spore
apex begins with the appearance of small dark spots; in one specimen
observed, a short black streak occurred on one side of the apex; the
color diminished toward the other side. Later, this black area extends
around the apical region, leaving the tip hyaline for a short time. The
time of occurrence of the blackening of the apex varies. In one thallus
seen, in which the secondary receptacle had already been formed, there
was no evidence of apical darkening.
After it divides into a four-celled primary axis, the male of H.
ectobiae produces antheridial branches from the two upper cells (PI. 11,a),
as well as a lateral extension from the suprabasal cell. These branches
arise from the side of the thallus opposite the seta. The suprabasal
extension usually grows downward only a short distance before cell
division occurs. If germination has taken place near the base of a seta, the
extension usually grows to the base, then forms a vertically elongate
secondary receptacle cell on its free side (PI. ll,a,b). The secondary
receptacle cell divides, cutting off an antheridial tuft primordium
apically.
If germination occurs high on the seta, the suprabasal cell extension
may grow a considerable distance before producing antheridial tufts.
The outgrowth usually becomes separated from the suprabasal cell by a
septum not far from its point of origin. The secondary receptacle, which
extends down the seta, is composed of elongate cells; the long axes of
these cells are usually more or less perpendicular to the setal axis.
At the same time that the extension is growing downward, the
antheridia or antheridia-bearing branches of the primary appendage are
initiated (PI. lla.b). These branches may arise before there is any
evidence of the secondary receptacle. On the other hand, thalli with two or
three tufts have been observed in which spermatium production has
proceeded for some time, even though no spermatia have been formed
on the primary appendage. The factors influencing the manner of
development of the male are not known.
The first branch of the primary appendage usually arises beside the
black apex as an upward extension of the terminal cell (PI. 11,a); the
black apex later becomes displaced laterally. The branch, which is
walled off (PI. ll,b), may divide again to form a terminal antheridium

46
MYCOLOG1A MEMOIR NO. 9
and a stalk cell. Lateral extensions from below the upper septum of the
stalk cell may produce additional antheridia; the branchlets are usually
two-celled. Other branches may arise from the terminal cell of the
primary appendage as well as from the subterminal cell (PI. 11 ,b).
Each secondary antheridial tuft develops as a result of extensive
division of a primordial cell (pa) cut off apically by one of the secondary
receptacle cells (Figs. ll,b,e). Following nuclear division, an oblique
septum divides the primordial cell into a stalk cell and an apical cell
(Fig. ll,c). An oblique wall is also formed following the first nuclear
division of the apical cell, so that the two lower cells are approximately
at the same level (Fig. ll,d). Because subsequent septa are also
diagonal, a double series of cells is formed below the apical cell (Fig.
ll,e). The main axis grows in a curve toward the primary receptacle.
Lateral outgrowths are produced by the axis cells on the concave side of
the cell row (Pis. ll,c; 12,a; Fig. ll,e). Usually no branches are formed
by the stalk cells (Jbb, be) until late in development (Fig. 11,0- Tne
number of axis cell tiers varies at the beginning of branch initiation; it is
usually three. When branching is delayed until the axis is longer, the
lowest branch may be formed by the third cell from the base and the
young antheridial tuft is more slender in appearance.
The two or three lowest cells of the main axis of the tuft are larger
than those above, the upper cells being narrower and more elongate.
The basal cells of each lateral branch are larger than the cells arising
from them. Thus, the mature tuft (PI. ll,c) consists of a curved main
axis of seven to nine pairs of cells, with a single apical cell and with
crowded branches arising from most of the double row of cells. Usually
each branch consists of one or two large basal cells surmounted by an
attenuated phialide. As the branch elongates, its apex narrows; the
young branches produced at the upper, lateral margins of all of the
intercalary cells of the antheridial tuft are much shorter and broader
than fully developed branches (PI. 11 ,c; Fig. 11,0-
At maturity, the phialides of H. ectobiae are very elongate, gradually
tapering toward the apex. Following the production of the first sper-
matia, the apical wall is ruptured, forming an exit passage. The sper-
matia are oblong, with a very thin cell wall and a narrow layer of
cytoplasm surrounding the nucleus. The spermatium nucleus, which may be
either spherical or elongated, is usually apical at first, later becoming
central in position. A gradual constriction just below the apex of the
protoplast of the phialide results in the separation of the spermatium.
The spermatia may be either truncate or rounded at the base. As the
spermatia are produced, the antheridial protoplast becomes
progressively shorter. Although the total length of antheridial walls in the
secondary tufts ranges from 13-24 jtm, the length of the protoplast may
TAVARES: HERPOMYCES ONTOGENY
47
be reduced to about 4 pxn after many spermatia have been produced.
The open apex of the antheridium may either be constricted or slightly
expanded. Two or three spermatia may be present in the lumen of the
antheridial neck.
The thalli of H. periplanetae germinate on minute setae; females and
males are commonly paired (Fig. 16,a). Occasional solitary females
occur; either they have germinated alone or associated males may have
become detached. Female thalli bearing five perithecia are frequent.
Usually several developmental stages are present, although in some
thalli, all perithecia are the same age (PI. 13,c). Sometimes broken peri-
thecial bases are present. Very young stages are extremely rare in thalli
bearing mature perithecia.
Like those of Herpomyces ectobiae, the spores of H. periplanetae are
divided into two cells almost equal in size. During growth of the female
thallus, the spore tips remain narrowly acuminate even though the
primary axis doubles in width during the four-celled stage (to ca. 3 /an).
In the male, no appreciable growth takes place at this stage (Figs. 16,a,
b). The upper (fo') and lower (fo) attachment disks may appear at the
two-celled stage; the lower disk may be fully developed while the upper
one is totally lacking or consists of an undifferentiated dark spot. Both
disks are adherent, although the upper one rarely remains attached;
usually growth of the thallus carries it beyond the tip of the seta on
which germination has taken place.
The young male consists of a primary axis of two primary receptacle
cells and two primary appendage cells; the basal and subapical cells are
very long, whereas the suprabasal cell is conspicuously short. Each of
the three upper cells may produce lateral antheridial branches, which
arise as outgrowths just below the upper end of the cell (Fig. 16,b).
Although branches from the suprabasal cell are uncommon, they
apparently arise early in development; those observed were broken off
or their antheridia had already deteriorated. Probably they arise later
than the branches from the upper two cells, but undergo no repeated
branching, so that they disappear while those above are still developing.
(Compare H. paranensis Thaxter and H. stylopygae, Tavares, 1965,
1966.)
In the formation of the first branch from the terminal cell, an upward
extension grows from one side of the spore apex (PI. 14,c), which is
thereby displaced laterally until it finally becomes located on the wall of
the lower half of the apical cell (Fig. 16,c). Two or three branches may
arise from this cell, none of them from the side toward the seta. One or
two branches may develop from the subapical cell (Figs. 16,b,c). In one
thallus observed, what appeared to be the initial outgrowth of a branch
arose laterally just above the basal septum of the apical cell, rather than
in the upper lateral position of all other branches.

48
MYCOLOGIA MEMOIR NO. 9
The first branchlet from each branch usually arises from its basal cell.
The terminal segment of each branch consists of a stalk cell and a long,
slender antheridium (Fig. 16,c). After the spermatia are formed and the
protoplast of the antheridium disappears, the empty cell wall remains.
There seems to be a tendency for the antheridial stalk cell to grow out
laterally below the empty antheridium to form a new branch.
In the female of if. ectobiae, the perithecium arises as a broad, dome-
shaped cell from the upper side of the secondary receptacle (Fig. 12,a).
This cell divides into two superposed cells separated by an oblique wall
(Fig. 12,b). The lower cell divides and a diagonal septum is formed
(Fig. 12,c). A third division, in the upper cell, results in a four-celled
structure two cells in height (Fig. 12,d). The upper two cells elongate
and divide again by vertical septa, resulting in the formation of the
initial four outer wall cells of the perithecium; the divisions are not
simultaneous (Figs. 12,e,f). Usually, there is no division of the lower
pair of cells until after trichogyne formation, although some thalli were
seen in which three flattened lower cells were present.
The next division is by horizontal septa in the four wall cells (Fig. 12,
g). The lower tier of wall cells remains about the same shape, while the
upper tier elongates and its cells divide (Fig. 13,a). A number of thalli
were seen in which the cell divisions during the early stages of perithecial
development did not follow the normal pattern. In one perithecium^
there were three outer wall cells in each of the two vertical rows on one
side. On the other side, three cells were present; there was a small basal
cell and an elongated upper cell in one row, but what would normally
have been the fourth wall cell row consisted of a single large cell.
In the four-celled female thallus of H. periplanetae, the two middle
cells are short and broad; the apical cell is sharply pointed, and the basal
cell rather broad and elongated. The suprabasal cell grows downward
(Fig. 16,a) to form a perithecial initial (PI. 14,a). Meanwhile the
primary axis widens above to 6-7 /mi, so that the apical cell becomes
broadly dome-shaped. The female axis at this stage is considerably wider
than the male. The primary axis of the female differs from that of H.
stylopygae in having a broader, shorter, more rounded apical cell.
Transverse division of the suprabasal cell (at times, of the subapical
cell of the primary appendage also — see Fig. 16,b) takes place,
resulting in the formation of a vertical row of several cells, each of which
turns downward at a sharp angle in a slightly different direction, but
never on the side of the primary receptacle on which it is attached to the
seta (Figs. 16,b,e). Rarely the subapical cell may also grow downward
(PI. 14,c; Fig. 16,b). Typically, each downgrowth produces a one-celled
upgrowth (PI. 14,a), which soon divides transversely to form a terminal
perithecial primordium (pa) and a secondary receptacle primordium
TAVARES: HERPOMYCES ONTOGENY
49
(Fig. 16,e, left). These cells divide (Fig. 16,b,d), so that there are soon
three secondary receptacle cells (Fig. 16,e) and two basal cells (bb, be)
below the terminal cell, which will produce the perithecium itself (Figs.
16,d,e; PI. 14,b). Cell be is cut off first by a diagonal septum (Fig. 16,b),
then cell bb is cut off by another (PI. 14,b) that intersects the first. The
terminal cell then undergoes three vertical divisions to produce the four
basal wall cells (wl) of the perithecium (Fig. 16,e). About the same time
that the wl cells divide transversely to form tier w2, cell bb forms cell bd
below (Fig. 17,b). In the perithecium shown in Fig. 17,a, cell bb is at the
upper level, with cell be at left below and cell bd at right below.
When there are two tiers of outer wall cells in Herpomyces, one of the
cells of the lower tier (ci) typically produces an extension that grows
upward centrally. By the time the extension has reached two-thirds of the
way to the apex of the perithecium in H ectobiae (Fig. 13,a), nuclear
division may have occurred; the upper nucleus may either move toward
the end of the extension or remain near its base. Meanwhile, cell
division in the terminal wall cells results in the formation of a third
tier of cells (Fig. 13,b). These divisions do not always take place
concurrently (Fig. 13,a).
There seems to be considerable variation in the time of division of the
central cell in H. ectobiae. In some thalli there is only a slight upward
extension of the inward-growing cell in three-tiered perithecia, whereas
in others, a constriction between the central upgrowth and the basal cell
is apparent at this time (see Fig. 13,b). In many older perithecia, there is
no separation between the procarp area and the ci cell (Figs. 13,c, 14,a,
b; cf. Fig. 13,a). When there are four tiers of outer wall cells in the
perithecium in Herpomyces, the central upgrowth may extend from the
apex of the perithecium as a trichogyne (Fig. 17,c; cf. H. paranensis, H.
stylopygae [Tavares, 1965, 1966]). The size of the nucleus that appears
in the trichogyne of H. ectobiae (Fig. 13,c) indicates that it probably
arises by division of the upper central cell nucleus. However, it is
possible that it is a male nucleus.
A septum forms at the base of the nucleated trichogyne of if. ectobiae
(PI. 12,c; Fig. 14,a). After the cytoplasm of the trichogyne becomes
separated from that of the carpogonium, its deterioration begins. The
base of the trichogyne becomes very dark when stained with acetocar-
mine. Its cytoplasm shrinks upward within the outer wall; pronounced
elongation and some branching may occur (PI. 12,d). By the time
another wall cell division takes place, the trichogyne has usually broken
off, leaving a flat, dark-staining cap or a short section of empty wall at
the apex of the perithecium (Fig. 14,b).
No nucleus was seen in the trichogyne of H. periplanetae (cf. also H.
stylopygae). Instead, a septum divides the central cytoplasmic mass at

50
MYCOLOGIA MEMOIR NO. 9
the level of the upper septum of tier w3, cutting off the upper nucleated
portion as the ap cell (Fig. 17,c; cf. Figs. 14,a,c). Very often, no nucleus
was seen in the first central upgrowth (cp) between the upper septum
and what appeared to be a septum below that separated this upgrowth
from the ci cell; presumably, the nucleus deteriorated or else the
protoplast was actually continuous with the ci cell.
A second cell (w3cT) appears in the third tier of the ci cell row in Her-
pomyces ectobiae (Fig. 13,b). Cell w3c' is the inner daughter cell
resulting from a longitudinal, slightly oblique division in the subterminal cell
of the ci wall cell row. It may be formed before the divisions that result
in the four-tiered perithecium (Fig. 13,b). The perithecium apparently
remains in the four-tiered stage for a long time, because no further
extension occurs until after the four inner wall cells are formed and
nuclear division takes place in the centrum.
Meanwhile, a division may take place in the basal cell underlying the
ci cell row; a vertical septum is formed between the two daughter cells
bb and bb' (Fig. 14,a). However, the basal cell usually does not divide
until after the inner wall cells have started to grow (Fig. 14,b).
As the first inner wall cell grows upward from the w3c' cell, a second
one arises either from the subapical cell of the opposite outer wall cell
row or from one of those adjacent to the ci cell row (Fig. 14,a). At the
same time, an extension of the second subapical cell (w3c) in the ci row
may appear. Such an extension may reach more than half the height of
the adjacent terminal outer cell before nuclear division occurs (Fig. 14,
b, lower right inner wall). Following division, one daughter nucleus
moves into the extension; a septum is later formed (Fig. 14,b, upper
right inner wall). Typically, the first and the third or fourth inner wall
cells originate in the ci cell row, whereas the other two arise from the
opposite row and from one of the other rows.
In the perithecium shown in Fig. 15,b, no inner wall arises from the
w3 cell at upper left; the inner walls from the neighboring cells grow into
the area that such a cell would occupy. It seems possible that the place
of origin of the inner wall cells of varying position may be influenced by
the position of the extra cell of the ci cell row. A fifth inner wall
extension at right, shown in Fig. 15,b, may represent, together with the
adjacent extension, a single upgrowth with an elongated dividing
septum or it may be a free fifth extension. There is sometimes an
upward extension, the base of which reaches the outside of the perithecium
in the same place where w4c' occurs in H. periplanetae.
Cell w4c' is a narrow cell cut off at the base of the w4 cell in the ci
cell row of H. periplanetae (cf. also H. stylopygae [Tavares, 1966]).
It appears when three of the third-tier cells of the broadening four-tiered
TAVARES: HERPOMYCES ONTOGENY
51
perithecium grow inward and upward to form inner wall cell initials.
The w4c' cell grows inward and upward to form the fourth inner wall
cell row. However, its nucleus does not divide until the other inner wall
cell rows divide to form the second tier of cells.
While the four inner wall cells of H. ectobiae are extending toward
the apex of the perithecium, a second central upgrowth (cp') arises from
ci (Fig. 14,a,b); although the first upgrowth (cp) appears to become
separated by a basal septum (Fig. 14,b) from the initiating ci cell, it is
probable that the protoplasts remain joined together at the base for
some time (cf. the basal connection between ascogenous cells shown by
Hill, 1977, in Fig. 6).
The secondary upgrowth may appear in H. periplanetae at an earlier
stage of development (Fig. 17,c; cf. PI. 14,d). Sometimes the ci nucleus
migrates up into cp' in this species; presumably, after nuclear division
one of the daughter nuclei moves downward into the ci cell. A small
apical cell (ap') is formed at the top of cp' (Fig. 18,a).
Presumably, ci is dividing to form a third upgrowth (cp") in the
perithecium of if. ectobiae shown in Fig. 14,c, and what appears to be a
binucleate cell above the dividing nucleus is cp'. However, it is possible
that this binucleate cell has resulted from nuclear division in cell cp,
because no other nucleus is visible that can be identified as that of cp. If
the protoplast of cp is not walled off from that of ci, its nucleus could
migrate downward into the active protoplast of ci and participate in
further divisions. No nucleus was detected in the cytoplasmic mass in
the upper left part of the centrum in the perithecium shown in Fig. 14,c
(cf. Hill, 1977, Figs. 3 and 5, which show the residual portion of the
centrum surrounding ascogenic cells at a later stage of development,
and Figs. 8 and 18, which show the fibrous appearance of this matrix).
The elongate protoplast in the upper right of the centrum of H.
ectobiae shown in Fig. 14,c has a slightly smaller nucleus than those of the
lower part of the centrum; this cell has been designated an ap cell despite
its unusual form. However, its nucleus is larger than those of cells ap,
ap', and ap" in other perithecia (see Fig. 15, a,b,c); the latter are
comparable in size to spermatial nuclei. It is probable that the small cell ap'
terminates the upgrowth containing the large upper nucleus labelled ce
in Fig. 15,a (as in Fig. 15,b, it was not possible to detect the outlines of
the protoplasmic masses in the centrum); the lower ce nucleus probably
was recently formed by division of the ci nucleus.
It was shown in H. paranensis (Tavares, 1965, Fig. 8) and H.
stylopygae (Tavares, 1966, Fig. 7) that ap' and ap" terminate secondary
upgrowths from the ci cell. Possibly the ap' and ap" nuclei are formed by
division of male nuclei, the sister nuclei moving downward and entering

52
MYCOLOGIA MEMOIR NO. 9
later upgrowths from ci (cf. the upgrowths below and at left in Fig. 7,
Tavares, 1966; in the perithecium shown, division of the ci nucleus in
the protoplast subtending ap" has not occurred — there is no nucleus
comparable to that of cp'). Conjugate division conceivably takes place
during the formation of these large upgrowths; it does not appear to be
occurring in earlier stages (it seems unlikely than another dividing
nucleus is obscured by the nuclei in the wl tier in Fig. 14,c). On the other
hand, it is possible that conjugate divisions do not take place until asco-
genic cells are formed (Fig. 15,c).
As further nuclear divisions occur in H. ectobiae, binucleate cells
appear at the base of the centrum (Fig. 15,c; cf. Tavares, 1965, Fig. 8,
and Hill, 1977, Fig. 3). These cells function as ascogenic cells; they
extend approximately the height of the second tier of outer wall cells.
The more elongate binucleate cells above them probably function as asci
if they do not deteriorate (it is possible that their nuclei are not dicaryo-
tic, but result from division of a single nucleus) (Fig. 15,c; cf. Fig. 18,b).
A septum later separates the ci cell from the centrum; the ci cell may
become binucleate (Fig. 15,c). Small cells appearing nearby conceivably
might be formed during divisions of dicaryotic nuclei and be
comparable to the inferior supporting cell of Laboulbenia, just as the ap' and
ap" cells may be the equivalent of the reformed trichophoric cell in that
genus. Further division of the basal cells of the perithecium may take
place. In the perithecium shown in Fig. 15,b, cell bb has produced cell
bb". In most perithecia, however, additional divisions, if they occur at
all, are delayed until ascospore formation.
Meanwhile, elongation and narrowing of the perithecial neck takes
place, accompanied by an increase in the number of inner and outer wall
cells in this region. At the same time, the asymmetry of the perithecium
becomes more pronounced (Fig. 15,c). Growth of the long, curved neck
during ascus production results in the formation of six tiers of inner wall
cells and nine tiers of outer wall cells. The ostiole of the perithecium is
essentially terminal. The outer wall cells in the venter of the perithecium
become so thin that they can hardly be detected (however, Hill's
electron micrograph [1977, Fig. 15] shows that they persist).
In H. periplanetae, the same number of outer and inner wall cells are
formed; however, a subterminal spine is produced from one outer wall
cell row (PI. 13,a; cf. H. stylopygae [Tavares, 1966]). Cell ci cuts off
another cell (cir) in the same tier (cf. small cells at base of centrum, Fig.
15,c). Later, the lower part of the centrum is filled with a number of
fusiform cells. The binucleate ascogenic cells (two are shown in Fig. 18,b)
are flanked by binucleate and uninucleate asci (in the latter, the nuclei
have presumably fused). In older perithecia, long rows of asci rise above
the ascogenic cells (PI. 14,f), which are generally 3-4 in number.
TAVARES: HERPOMYCES ONTOGENY
53
The basal cells may become multinucleate in H. ectobiae (PI. 15,d).
Some solitary nuclei had two nucleoli. In one cell there was a ring of
nucleoli, each surrounded by a small amount of nucleoplasm and part
of a highly lobed nuclear membrane. Often there appeared to be two
cells lying one over the other; these may either have been recently
divided cells or fused cells. The number of cells increases considerably
in the basal cell — wl tier zone (see PI. 15,c,d) and it is difficult to
determine the origin of cells in flattened perithecia. The basal cells are
generally more darkly stained and irregular in shape.
In H. periplanetae, the number of secondary receptacle cells
subtending perithecia of a particular stage of development varies; the number is
probably greater in the first-formed perithecium of the thallus, which
presumably forms the large enveloping shield. Each secondary
receptacle cell narrows at the base into one or more haustoria that
penetrate the host cuticle. Only one of the three sr cells undergoes
division; this cell forms the entire shield — the dividing cell being carried
outward on the end of the shield as the number of narrow shield cells
increases (Fig. 17,c). Cell bd fuses with the underlying secondary
receptacle cell. In one thallus, a pore was observed between cell bb and the
wl cell opposite ci, but these pores are not as conspicuous as they are in
H. paranensis (see Tavares, 1965).
The penetration of a haustorium into the setal cavity is evident in PI.
1 l,c below the attenuated base of the upper antheridial mass in H.
ectobiae. The bases of the elongate secondary receptacle cells gradually
narrow downward into haustoria; more than one haustorium may
extend from the base of each of the cells growing along the seta (Pis.
ll,d; 12,a; Fig. ll,f). The small haustoria that penetrate the setal walls
appear to expand inside; because haustoria stain very lightly, it was not
possible to determine the extent of haustorial growth within the setae.
Whether the cytoplasm of each haustorium extends downward
separately or whether the cytoplasm of adjacent haustoria intermingles is
not known; nor is it known how far down the seta haustoria extend.
It can be seen, however, that setae bearing fungi show the downgrowth
from their bases of a strand of homogeneous cytoplasm that is absent
below setae not bearing fungi.
Haustorial bulbs that resemble those reported by Richards and Smith
(1956) in H. stylopygae occur in the epidermal cell layer of the host. A
strand of fungus protoplasm passes down the center of the seta in PI.
10,c from a thallus attached to the seta (not in view); it spreads out into
a haustorial bulb below the setal socket. It cannot be determined
whether the strand entering the setal socket from the left fuses with the
central strand. Apparently at least one haustorium is formed at the base
of each secondary receptacle cell; each forms a clear vertical channel

54
MYCOLOGIA MEMOIR NO. 9
that is visible in the dark outer layer of the host cuticle. The base of a
cell of a large male thallus, shown in PI. 10,b (center), becomes
attenuated and probably extends as a haustorium into the adjacent setal
socket; at the base of the seta, the haustorium enters the epidermal cell
layer of the host. In this thallus, the fungus cytoplasm, which is more
homogeneous in appearance than that of the insect cells, spreads out
into a large network in the epidermis. There also appears to be a thread
of cytoplasm approaching this haustorium down the right side of the
setal socket; it is impossible to determine whether this cytoplasm forms
part of the haustorium.
Ascus Development
Laboulbenia
In Laboulbenia flagellata, L. borealis, and L. gyrinidarum, asci are
formed in a vertical series from each of the two binucleate ascogenic
cells (PI. 7, a-c, e). As the asci enlarge, they extend upward within the
perithecium; their bases become attenuated until finally they break
away from the subtending ascogenic cells. During ascus formation, the
slightly elongated, often broad-based or dolabriform ascogenic cell
extends vertically or diagonally upward (Fig. 6, PI. 7,b). Presumably, the
two nuclei divide shortly before the ascus is formed; I have seen no four-
nucleate ascogenic cells. In an ascogenic cell, probably of L. borealis,
which was undergoing conjugate division, Faull (1912, PI. XXXVIII,
Fig. 37) showed the spindles arranged so that the daughter nuclei would
be at opposite ends of the cell. No septum was indicated between the
recently formed prefusion binucleate ascus portion and the remainder
of the undivided ascogenic cell.
The ascus is separated from the upper side of the broad base of the
ascogenic cell by a short horizontal septum, according to Faull (1912;
PI. XXXVIII, Fig. 38). It seems probable, however, that when one of
the ascogenic cell nuclei is located in the upper part of the cell (see PI.
7,b), the young ascus is separated laterally by a vertical furrow from the
upper part of the cell and by a horizontal septum from the broadened
basal part. What appear to be three-nucleate ascogenic cells in situ
(Fig. 6) in L. flagellata are probably binucleate ascogenic cells, each
with an ascus in which nuclear fusion has occurred or is about to take
place (the narrow septum is hard to detect through the wall cell layers of
the perithecium). A comparison of the ascogenic cell nucleus in PI. 7,b
TAVARES: ASCUS DEVELOPMENT, LABOULBENIA
55
with the small nuclei in the binucleate ascus in PI. 7,a shows a marked
difference in size; the fusion nucleus in the previously formed ascus
at its right is closer in size.
Obviously, the nuclei of the ascogenic cell rapidly increase in size
after division, forming large nucleoli. By contrast, the nucleoli in the
binucleate prefusion asci are so minute that they can rarely be detected.
After nuclear fusion, the nucleolus is clearly visible and may be seen (PI.
7, a-c) attached to the chromatin mass by a double thread.
The asci almost encircle the ascogenic cell; in PI. 7,e, the youngest
ascus lies over the ascogenic cell, the next are at the right, and the oldest,
at the left. Similarly, in PI. 7,b, the youngest ascus lies over the
ascogenic cell; three young asci are attached close together under the
ascogenic cell where it becomes narrow; the large ascus lies below. Isolated
ascogenic cells bear a scar where recently formed asci were attached.
Nuclear fusion was not observed.
Before meiosis, the nucleus grows to about 13 /an in diameter. Diplo-
tene and diakinesis were not recognized in meiotic prophase in
Laboulbenia, although they were seen in Herpomyces. The first meiotic
division appears to be intranuclear (PI. 7,d; cf. Faull [1912]); a clear area
surrounds the metaphase plate and the nucleolus is in a lateral position
within this area (cf. nucleolar position in Pyronema, Figs. 8-9, Hung
and Wells [1977]). Faull (1912) observed that in anaphase I the spindle
elongated greatly, parallel to the long axis of the ascus (see his PI. XL,
Fig. 54). In his preparations, the spindle elongated very little during
anaphase II, so that the daughter nuclei remained close together. Faull
(1912) found that smaller central bodies and more sharply pointed
spindles were present during the second division, which was also
intranuclear (see his PI. XL, Fig. 55, which shows the nucleolus within a
lateral lobe surrounded by the nuclear membrane; cf. Fig. 10, Hung and
Wells [1977] and observations on somatic mitosis in Cochliobolus,
Huang et al. [1975]). Rapid succession of the first and second divisions
(Faull, 1912) would account for my failure to observe binucleate asci —
perhaps because no spindle elongation took place (cf. Zickler, 1970).
A nucleolus was reported by Faull (1912) in each nucleus at the
binucleate stage. The nucleoli of the four nuclei resulting from the second
division have a central, clear area (cf. PI. 8,a with Faull, 1912, PI. XL,
Fig. 57). The arrangement of the four nuclei in PI. 8,a suggests that the
spindle formed during the first division elongated transversely, although
the lateral disposition of the nuclear pairs could have resulted from
elongation of the second division spindles. By contrast, the division I
spindle in the ascus shown in PI. 8,b would have elongated in a
longitudinal direction.

56
MYCOLOGIA MEMOIR NO. 9
My observations confirmed Faull's report (1912) of a chromosome
number of n = 4 in L. borealis (as L. chaetophora), although there is
some uncertainty because of the absence of suitable preparations. There
appear to be three large and one small chromosomes in L. gyrinidarum
and L. borealis (Pi. 7,d). The chromosomes are somewhat similar in L.
flagellata (PI. 3,d).
During the third nuclear division, the chromosomes appear to be
much longer and narrower; the nucleoli are persistent as in the first
division. In the ascus shown in PI. 8,b,c, three of the four nuclei are in
anaphase; there appears to be a telophase in the lower, central part of the
ascus (PI. 8,c). Apparently the nuclear membrane breaks down before
metaphase is completed.
The spindle of the third division becomes very elongate, resulting in
wide separation of the daughter nuclei at telophase (PI. 7,e, right). Four
of the nuclei remain in the middle of the ascus, together with the
persistent nucleoli, which apparently disappear shortly after telophase. In the
eight-nucleate ascus shown in PI. 7,e, the old nucleoli are not present.
The four nuclei that move to the apex of the ascus during anaphase are
not included in ascospore formation (PI. 8,d); nucleoli are formed in
these supernumerary nuclei, but the chromatin remains very
heterogeneous in appearance. These nuclei persist at the apex or side of the
ascus after spores are formed, later breaking down into loose
accumulations of chromatin material. The four deteriorating nuclei remain in a
close group.
During spore formation a nucleolus appears near one end of the
elongate nucleus (PI. 8,d). Four large spindle-shaped spores are delimited
within the ascus; the fibrous appearance of the spores is conspicuous in
PI. 8,d (cf. FauU [1912, PI. XL, Fig. 64] and the observations of
Schrantz [1967] on ascospores of Pustularia). After the spores are
formed, they may shorten and become almost spherical in shape (cf.
Faull, 1912). A distinct wall separates them from the remaining
cytoplasm of the ascus (Pis. 8,e; 9,a; cf. paired prospore membranes of
Herpomyces [Hill, 1977, Figs. 26-27, 31-34]).
By the time their nuclei divide, the spores have extended almost the
entire length of the ascus, which has also enlarged (from ca. 70 /tin at
first metaphase to approximately 90 /on). The elongation of the spindle
results in the relocation of the lower daughter nucleus in the base of the
spore, where it is separated in a small basal cell by a septum.
Apparently, the nuclei of the prospores move downward away from the
deteriorating nuclei (see PI. 9,a-d). Meanwhile, in some asci the chromatin
of the supernumerary nuclei becomes completely dispersed throughout
the apical cytoplasm of the ascus.
TAVARES: ASCUS DEVELOPMENT, HERPOMYCES
57
At maturity there remains only a narrow margin of epiplasm in the
ascus beside the spores; the ends of the ascus extend far beyond the tips
of the spores. The mature spores become displaced to the sides of the
perithecium at a low level (PI. 5,b), as well as toward the apex. Younger
asci are continually formed, forcing the maturing asci upward and
outward. The mature spores appear to be arranged in fours; however, the
ascus walls could not be detected in intact perithecia. Faull (1912)
indicated the presence of empty ascus walls. The walls had previously
been thought by Thaxter (1896) to be absorbed, leaving the spores free
within the cavity of the perithecium. It is probable that they eventually
are either ejected as spores are discharged or deteriorate within the
perithecium.
Herpomyces
Ascus development in Herpomyces ectobiae and H. periplanetae
differs in a number of ways from the process as it occurs in Laboul-
benia. The primary difference between the asci of the two genera is the
number of spores produced. The position of the spore septum also
differs — those of the eight spores in the ascus of Herpomyces are median,
rather than located near the lower end as they are in the four spores of
the Laboulbenia ascus (the unusual position of septa in the micrographs
of Hill [1977] is the result of oblique sections — personal
communication) (see PI. 15,a). In addition, the asci are produced alternately on
one side, then the other, in Herpomyces, rather than in a single series as
in Laboulbenia. The common longitudinal walls that join consecutive
asci together on the side of the ascogenic cell persist; they can be
detected between mature, spore-filled asci (Hill, 1977, Fig. 15) (however,
asci can easily be separated by flattening the perithecium).
Hill (1977) showed the formation of vertical septa separating the asci
produced by a single ascogenic cell (Figs. 8,14); his Fig. 12 illustrated
the process of septum formation. One can interpret this figure as
follows: the cell in the lower half is a recently formed ascus; the wall that is
growing out from the junction of the walls of this ascus will cut off the
next ascus. The appearance of the vertical rows of asci in PI. 14,f
indicates that the asci are produced more or less in two rows, one from
each side of the ascogenic cell. As Hill's micrographs (Figs. 14, 15)
showed, there may be three or four asci at each level because of
overlapping (see wall enclosing four spores in upper part of section, Fig.
15 — because this section is near the end of the ascus, all eight spores are
not included).

58
MYCOLOGIA MEMOIR NO. 9
Ascogenic cells producing asci by longitudinal septation tend to be
rhomboidal or trapezoidal in shape (PI. 15,c); sometimes they are more
reniform (PI. 15,d, left). They may be quadrinucleate or binucleate.
Elongate ascogenic cells may be almost dolabriform as in Laboul-
benia — such cells are separated from the asci they produce by a short,
horizontal septum (cf. PI. 15,d, right). The asci formed tend to be
broader and shorter than those separated only by a longitudinal septum
from the ascogenic cells; in both types, the final separation seems to be
at the base (see PI. 15,c, center left) (cf. H. paranensis, Tavares, 1965,
Fig. 8). The orientation of the spindles and the position of the four
nuclei in the ascogenic cell probably determine the shape of the cell and
its asci. The differences in ascus formation may also be related to the
numbers of ascogenic cells and asci produced, as well as to the source of
ascogenic cells. In Laboulbenia they arise from one of the median di-
caryotic cells in the carpogenic cell row. In Herpomyces at least one of
the ascogenic cells is among the last cells formed by the carpogonial
initial (Hill, 1977, Fig. 3, showed Woronin bodies on either side of a
connection between an ascogenic cell with two visible nuclei and the
carpogonial initial cell [labelled as a basal cell, but actually one of the
first tier of outer wall cells]). The high position of the nuclei in this
ascogenic cell should be compared with that of pairs of nuclei in the centrum
in Figs. 15,b, c.
Although Hill (1977) labelled the centrum cells in his Fig. 4 as asco-
genous cells, the thickness of the wall cells and small diameter of the
section indicate that the centrum is probably filled with upgrowths from
the ci cell (cf. the nuclei in Fig. 14,c). The older perithecium shown in
Hill's Fig. 3 has probably just produced the last ascogenic cell because
of the lateral position of the intercellular connection. It seems probable
that each of the ascogenic cells has been joined to the adjacent one, as
he has indicated in Fig. 6; this may also have been true of the earlier
carpogonial upgrowths. At some point dicaryotic cells are formed that
are ascogenous; possibly the perithecium must reach a certain size
before the ci cell begins to produce the basal layer of ascogenic cells rather
than more elongate upgrowths. A five-tiered perithecium of H. stylo-
pygae (Tavares, 1966, Fig. 7) had not yet begun to produce ascogenic
cells, whereas by the six-tiered stage, H. ectobiae (Fig. 15,c) had
produced one and by the seven-tiered stage, H. paranensis (Tavares, 1965,
Fig. 8) was producing asci from a solitary ascogenic cell (it should be
noted that the division figures in the latter perithecium may be meiotic
metaphases; however, it is not certain that a wall actually separates
these two protoplasts — the metaphases may represent a conjugate
division within an ascogenic cell). By the time the perithecium of H. peri-
planetae is nine tiers in height, two ascogenic cells may be present (Fig.
18,b).
TAVARES: ASCUS DEVELOPMENT, HERPOMYCES
59
In Herpomyces, many prefusion binucleate asci 8.5-10 jtm in length
were observed, each nucleus containing one nucleolus (PI. 15,c,d).
Apparently the nuclei approach one another and fuse, the nucleoli fusing
later (cf. Hung and Wells, 1977). A number of asci were seen in which
the two nuclei were together rather than at opposite ends of the cell.
Young uninucleate asci were observed in which two nucleoli were
present; in some of these asci the two nucleoli were located at either end of
the elongate nucleus. In later stages, the two nucleoli are closer together
in position. The fusion nucleus in H. periplanetae is frequently long and
elliptical in shape; often a rather small nucleolus is located at one end.
The meiotic prophase appears to be of long duration, because each
perithecium contains many asci in this stage. Diakinesis was observed in
H. periplanetae (PI. 15,a), which has a chromosome number of « = 4;
one chromosome is considerably smaller than the other three. The
chromosome number in H. ectobiae was not determined. During
meiotic metaphase, the nuclear area in acetocarmine preparations of H.
periplanetae is clear and there is a short marginal zone resembling spindle
fibers near each end. Probably the nuclear membrane is present during
the first meiotic division. The nucleolus persists throughout the
division.
Nuclei of H. periplanetae may be situated far apart in the ascus
following the first meiotic division (cf. Faull's observations [1912] on
Laboulbenia). The nucleoli usually seem to be reformed at this time. The
second division may take place in a plane perpendicular to the first,
resulting in the formation of two daughter nuclei near each end of the
ascus. However, in some asci, the nuclei in the four-nucleate ascus are
disposed in an irregular vertical series. Following the third division (PI.
15,b, left), there may be a tetrad of nuclei in each end of the enlarging
ascus. When the nuclei are closer together in the four-nucleate stage,
they appear to be more closely grouped in the eight-nucleate stage. Prior
to spore formation, the eight ascus nuclei elongate (PI. 15,b, right).
Three elongate masses of chromatin (probably the larger chromosomes)
were visible in some nuclei in the asci at this stage; no nucleoli were
observed in these nuclei. About the time of spore delimitation,
metaphase takes place. By telophase, the elongate - daughter nuclei have
moved to opposite ends of the spores. A septum is later formed
separating the two halves of each spore. Subsequent to nuclear division,
contraction of asci seems to take place. Early in spore development, the
same ascus may contain short ascospores approximately 5.5 pan long
and elongated spindle-shaped ascospores 9-10 /tin long (PI. 15,a). Later
the spores elongate until at maturity they reach 11-13 /xm in length.

60
MYCOLOGIA MEMOIR NO. 9
The Parts of the Thallus*
The Receptacle
The concept of the receptacle introduced by Thaxter (1896) (all cells
from the lower spore segment except those comprising the perithecium)
is preferable to his (Thaxter, 1908) limitation of the primary receptacle
to the 2 lower cells of the primary axis of the thallus (cells / and II). The
5 receptacle cells of the typical mature thallus of Laboulbenia were
called a pseudoreceptacle by Thaxter (1908) and he considered cells ///,
IV, and V to be the base of the appendage, which is adnate to the
developing perithecium in most species of Laboulbenia. In a species in
which these 3 cells are free from the perithecium, such as L. brachi-
onychi, the later concept of what actually constitutes the appendage
would be more applicable.
The primary septum, which separates the upper and lower spore
segments, can often be detected by its thickness, the amount of
constriction, slight darkening (although strongly blackened septa are rarely
primary septa), or the amount of displacement of the outermost edge of
the septum (which forms a ring in the outer part of the thallus wall).
At the time that the spore septum is formed, a wall layer is laid down
throughout the two cells (see Hill, 1977, Fig. 37) that turns inward at the
septum. A triangular space is formed that is bordered on the outside by
the outer wall of the spore. The position of the spore septum may be
determined by looking along the side of the thallus to see which septum
extends to the thin, outermost wall layer, where no other layer
intervenes. A comparison of a series of sporelings with older thalli, using
measurements of cell lengths (allowing for some elongation) will often
indicate the position of the primary septum. The primary appendage
is often considerably narrower than the upper receptacle, although its
lower cell may be as wide as the ///cell.
Although all genera have not been carefully studied, it is believed that
the lower spore segment generally divides into 3 cells (/, //, and III),
even though further division may occur later. Exceptions include genera
in which there is no division into separate // and /// cells (see Amorpho-
myces, Rhizopodomyces). In Chitonomyces, the derivatives of the
undivided //-/// cell of the sporeling are separated by cell walls that are less
distinct than those that separate them from the primary appendage and
from cell /(PI. 34,g, 35,b,c).
♦For authors of taxa, see generic section.
TAVARES: RECEPTACLE
61
Although there is a separate /// cell in Dimeromyces isopterus (the
spore septum lies just above a tall /// that is superposed upon a tall //
and perithecia-forming cells produced by / — see Fig. 9, 20, Tavares,
1979) and probably also in Polyandromyces, there apparently is an
undivided //-/// in Dimorphomyces muticus, in which the two terminal
cells of the primary axis are subtended by the spore septum (Tavares,
unpublished). It has not been determined whether the failure of cell II-
III to divide is common in the Dimorphomyceteae or whether it is
confined to species of Dimorphomyces.
In the more primitive genera (see Fig. 21 and arrangement of genera
in the key), cells // and /// (or at least II) are subdivided (indeterminate
receptacle — Thaxter, 1908), usually by horizontal septa. In many of
the more highly evolved genera, such as those in the Stigmatomycetinae,
there are only 3 cells in the determinate receptacle, but their positions
with respect to one another may vary. The following patterns may
occur:
// and /// parallel or with diagonal septum — Acompsomyces, Auto-
phagomyces on Anthicidae, Corylophidae, Cryptophagidae, and Phala-
cridae, A. poissonii, Cryptandromyces, Diclonomyces, Distolomyces,
Hesperomyces, Ilytheomyces, Nycteromyces, and Sphaleromyces. The
manner in which the parallel septa are formed in the sporeling is shown
in Diclonomyces (PI. 45,d-f).
//and VIparallel — Acallomyces, Diphymyces, andPolyascomyces.
I and // parallel or with oblique septum — Acompsomyces lasiochili
(with VI enclosed), Acrogynomyces (with VIin most species enclosed by
VII, II, I, and III), most species of Corethromyces on Rugilus, Eucan-
tharomyces, Ilyomyces (with /// and // on opposite sides and /// and /
separated by a slightly diagonal septum), and Prolixandromyces.
HI and VI parallel — Apatomyces, Laboulbenia, and Stigmatomyces
(in all except a few species of Laboulbenia, these two cells are adnate, at
least in part).
// and /// extending down on either side of I nearly to base — Stem-
matomyces and some species of Synandromyces.
I, II, and /// superposed — Autophagomyces microveliae, Autopha-
gomyces on Pselaphidae, Corethromyces species, Rhizomyces, Zeugan-
dromyces, and Gloeandromyces.
The // cell undergoes multiplication in many genera. There are several
genera with long, uniseriate receptacles — examples are Cochliomyces,
Enarthromyces, Ecteinomyces, Filariomyces, Hydrophilomyces,
Chaetomyces, Ormomyces, Plectomyces, and Drepanomyces. In
addition, there are some species with similar receptacles, such as Botry-

62
MYCOLOGIA MEMOIR NO. 9
andromyces heteroceri and Chaetarthriomyces coelostomalis, that
belong to genera having species with short receptacles.
In many genera, the lower receptacle (below the perithecium) is
biseriate or multiseriate. In Asaphomyces and Dermapteromyces, the
appearance of a biseriate receptacle results from the receptacular stalk
cells (Pis. 42,c; 44,b-d). Although there appear to be vertical septa in the
receptacle of some species of Dixomyces, the receptacle is actually
composed of a branching row of // and III cells. In D. perpendicularis, II
divides to form one or two receptacle cells that subtend the stalk cell of
the perithecium (PI. 41,a). A row of cells extends upward beside the
base of the perithecium, where it divides again in the upper receptacle
(the upper cells, at least, are derived from cell III).
Rhipidiomyces is unusual in having a receptacle
consisting of elongate, vertically oriented cells (PL 42,e); the predominant cells
in Diaphoromyces are also long and vertically oriented. In Rickia,
Homaromyces, Benjaminella, and Euzodiomyces, on the other hand,
the // cell area is divided into shorter cells. Abundant horizontal
divisions, with regular vertical septations, are characteristic of Kainomyces,
Blasticomyces fastigiatus, and Columnomyces. In Peyritschiella, there
are vertical septa in each tier of receptacle cells. Multiseriate thalli in
most genera are flattened (cf. Columnomyces). In Zodiomyces, there is
a single dorsal and ventral cell layer covering the central core in young
thalli. There are 2 cell layers in the receptacles of Kainomyces,
Peyritschiella, and Smeringomyces species on Sepedophilus; there are at
least 2 layers in Euzodiomyces. There is only 1 layer in Blasticomyces
fastigiatus, Benjaminella, Tettigomyces, Rickia, Diaphoromyces,
Amphimyces, and Rhipidiomyces. In Amphimyces, there may be 2
partially overlapping axial cells in vertical rows; these cells produce rows
of cells extending toward opposite sides of the thallus. Horizontal septa
are formed in the lower part of the receptacle (PL 42,k-m). Although
there may be some overlapping of receptacle cells in Tettigomyces as
new cells are formed, there are not two distinct layers of cells.
In Rickia wasmannii the receptacle may consist of a central axis of
elongate cells, somewhat irregularly arranged, with short side branches,
each having an elongate basal cell that is more or less parallel to the
central axial cell. Sometimes there is forking of the vertical cell row
(cf. Chadefaud's description [1960, pp. 490-491] of the structure of
Rickia, in which he asserts that there are three separate axes arising
from the base of the receptacle).
A series of divisions of cell /// in Hydraeomyces results in the
formation of secondary appendages and antheridia. Cell III also divides
in Misgomyces, many species of Mimeomyces, and in most species of
Laboulbenia. There are apparently a number of divisions of /// in
TAVARES: RECEPTACLE
63
Chaetomyces, Amphimyces, and Hydrophilomyces, but in most genera
they are limited in number if they occur at all. A single division of ///
takes place in Ceratomyces mirabilis (PL 22,c) and probably in other
genera of the Ceratomycetaceae. It has not yet been determined whether
/// divides horizontally in Ecteinomyces. The cells formed from cell ///
are most clearly set off in Kainomyces (PL 26, c-e). In some species of
Dixomyces, the upper receptacle is quite similar to that of Misgomyces
(PL 40, c-e). There is a Ilia cell just below the spore septum in Chi-
tonomycesaethiopicus (PL 35,e).
The cells of the secondary receptacle lie outside the primary axis of
the spore. There appear to be four basic types of secondary receptacles:
1. At least one secondary axis is formed by the separation of terminal
cells from an elongation of cell I in Dimorphomyces.
2. The distinctive primary axis, with its appendage, remains clearly
visible, the secondary receptacle growing out laterally (examples are
Herpomyces, Dipodomyces monstruosus).
3. The primary appendage remains as a small or large branch near the
base of the thallus, but it may be similar to secondary appendages and is
inconspicuous (examples are Histeridomyces, Scaphidiomyces, Clono-
phoromyces, Rhachomyces, Monoicomyces, Eumonoicomyces, Comp-
somyces, Balazucia, Cucujomyces). In Rhachomyces particularly, the
relative sizes of the primary and secondary axes are so different that the
primary axis is visible only as a small basal branchlet similar to many
borne along the secondary axis.
4. A suprabasal cell complex develops from cell //and the position of
the initial primary appendage is obscured. There is a variable amount of
development of secondary axes and thus the receptacle cannot be
considered as merely a primary axis in which divisions in two or three planes
have taken place (see Clematomyces calcaratus [PL 27,e]; cf. Pis. 27,d;
28,a). Although an irregular mass of cells forms the receptacle in most
genera in the Teratomycetinae, in Sandersoniomyces secondary axes are
clearly visible. In Carpophoromyces the secondary receptacular axes are
elongated and come from the suprabasal cell complex irregularly at
different levels.
Although the male thallus of Amorphomyces falagriae was
recognized by Thaxter (1896) as possessing two receptacle cells, the receptacle
of the female was reported to consist of only 1 cell. His
misunderstanding of the origin of the perithecium apparently led him to mistake the
suprabasal cell of the receptacle for one of the basal cells of the
perithecium, whereas recognition of the presence of the rudimentary
appendage (Tavares, 1961, 1970) makes the position of the suprabasal cell
quite clear. This cell appears to be an undivided //-/// cell (Tavares,

64
MYCOLOGIA MEMOIR NO. 9
1970) (cf. Rhizopodomyces). In Amorphomyces and Rhizopodomyces,
the perithecium arises as a lateral outgrowth of the suprabasal cell as in
Laboulbenia (cf. Thaxter, 1896, 1931). In Amorphomyces, the thallus
consists only of the two receptacle cells, the perithecium, and the
rudimentary appendage; its primary axis is essentially the equivalent of the
4-celled axis of the female thallus of Herpomyces ectobiae.
A 2-celled primary receptacle is clearly seen in the female and male
thalli of Herpomyces paranensis (Tavares, 1965) and H. ectobiae, as
well as in the male of H. periplanetae. However in the female of H. sty-
lopygae (Tavares, 1966) on small setae and the female of H
periplanetae, each of a series of primary axis cells functions in the same way as
the suprabasal cell of the primary receptacle of H ectobiae. However,
only one perithecium is produced by each of these cells. Many may be
formed from the suprabasal cell in H. ectobiae (this cell produces the
entire secondary receptacle in both sexes).
When H. stylopygae grows on large setae, the secondary receptacle of
the female consists of a vertical series of cells as in H. ectobiae; no shield
is formed (Richards and Smith, 1955b). The thallus usually bears only
one perithecium.
Species of Laboulbenia in which cell /// remains undivided may be
considered to be reduced forms. In addition to regularly occurring extra
cell divisions, such as those in L. gyrinidarum, L. partita, and L. hing-
stonii, there are occasional anomalous divisions in Laboulbenia that
result in abnormal thalli (cf. PI. 2,c). Picard (1913b) observed that
proliferation of cell // in L. proliferans resulted in the formation of a
secondary perithecium as in genera normally producing more than one
perithecium.
Other unusual conditions of the receptacle that sometimes occur in
the Laboulbeniales include the development of sterile, antheridial, or
perithecial branches from any cell of the receptacle except cell I, the
absence of certain cells, and altered relationships between cells (Colla,
1934) (see Balazuc, 1976, on teratological forms of Laboulbenia and
Ormomyces). The abnormal specimens of L. proliferans that Middel-
hoek (1951) observed suggested to him that genera with a limited and
constant number of receptacle cells are the most advanced phylogene-
tically. He suggested that abnormalities should help to clarify
relationships between various groups. Abnormal multiplication of receptacle
cells possibly represents a reversion to an ancestral receptacular type
(Middelhoek, 1957).
Although cell / does not usually divide, in the upgrowth from the
foot in Kyphomyces ansatus there appear to be septa, although
specimens have not been stained to determine whether several cells are
actually present. There is no indication that there is cell division in the up-
TAVARES: RECEPTACLE
65
growth from / in other species of Kyphomyces, although in K. thino-
charinus there seems to be the beginning of a row of dark rings
developing around the upgrowth. I have seen no evidence of cell division in the
black foot upgrowths of species of Meionomyces or Corethromyces
(cf. Pis. 32, b-d; 44, f-j). The secondary foot in Dipodomyces mon-
struosus does not undergo division; no septa were detected in the
secondary feet of Thaumasiomyces. Nuclear staining should reveal whether
division occurs in the upgrowths from cell / in Kleidiomyces.
Cell / contributes to the formation of the perithecia in Dimorphomy-
ces (cf. Thaxter, 1896, 1924). The presence of the youngest perithecium
on the lowest perithecium-producing cell in the series in Dimeromyces
shows that Thaxter (1924) was correct in his belief that these cells are
separated from the top of cell /. In addition, it is sometimes possible to
see a flat cell separated from the upper end of/by a very narrow septum
(cf. also Polyandromyces). In Nanomycesperpendicularis, a small cell,
presumably arising from cell /, becomes laterally adnate to the
suprabasal cell of the primary axis; it produces the perithecium.
The perithecia of Trenomyces are produced by cells derived from //.
Possibly, the ability of cell / to produce cells from its upper end was lost
in this genus of the Dimorphomyceteae when / began to enlarge and
produce a massive haustorium.
The absence of daughter cell production by the basal cell in most
genera and the lack of branch formation by cell / in males of
Herpomyces (Tavares, 1965, 1966) suggests that the modification into an
attachment structure in some way renders this cell incapable of division.
It is possible that in some genera cells may be formed from / that are
indistinguishable from those produced by subdivision of cell //. Such
divisions might be detected by study of sporelings, comparison of cell length
and cell wall thickness, and observation of the general configuration of
the receptacle.
In Chitonomyces and Hydraeomyces, in which division of cell II-III
is delayed, cell / cuts off a small flat cell la in the young sporeling (C.
aethiopicus [PI. 35] is an exception). In Hydraeomyces two flat cells are
formed at the upper end of cell / shortly after germination (PI. 34,c).
In some thalli of Diphymyces bidentatus, tnere appears to be an
intercellular pore between cells / and VI, which suggests that VI is produced
by / rather than by // (the presence of a pore is indicated by a small
outward bulge of the cytoplasm from each of two adjacent cells; it is
usually in the middle of the septum). However, in one thallus observed
there was also evidence of a pore between // and VI, as well as one
between / and //. Probably a secondary pore is formed between / and
VI. In closely related D. silphidarum, VI is clearly formed by division of
//.

66
MYCOLOGIA MEMOIR NO. 9
Often the protoplast in cell / seems to deteriorate in mature thalli of
various genera, but there is sometimes upward proliferation of the cell.
Under unusual circumstances, such as the necessity of reaching a more
advantageous place for penetration, the cytoplasm from the lower part
of the thallus may emerge to form a secondary foot; this apparently
occurs in Thaumasiomyces and Dipodomyces monstruosus. In D.
monstruosus cytoplasm from the lower part of the basal spore segment
forms a secondary foot; the upper part of the same cell (PI. 29,b) grows
into a secondary receptacle.
Unicellular buffer-organs (so called by Thaxter, 1931, presumably
because of their function of maintaining the position of tall thalli during
the movements of the host) are formed near the base of the thallus in
species of Hydrophilomyces (see Majewski, 1974), presumably from cell
II. Uniseriate, multicellular buffer outgrowths occur in Kyphomyces
rhizophorus, whereas those of Zodiomyces are multiseriate. In Scelo-
phoromyces and Osoriomyces the uniseriate, multicellular
downgrowths (PI. 33) are similar in position, and presumably also in
function. Nonseptate outgrowths of cell / extend laterally from the base
of the female in Rhizopodomyces (Benjamin, 1979). Thaxter's figures
(1924, 1926) show numerous examples of appendages in a normal
position on the receptacle which probably have the same function as the
outgrowths near the receptacle base.
In the massive thallus of Zodiomyces there is a central core of
modified cells, the walls of which stain darkly in acetocarmine. Their thick
walls indicate that they serve as support, but it should be determined
whether the core may also facilitate transport of nutrients.
Ovals, which Thaxter (1908) compared with the stigmata of insects,
occur in the walls of secondary receptacle cells of some species of
Rhachomyces. He suggested that they might act as valves to relieve
tension caused by sudden flexion. Their structure is unknown and
should be studied with electron microscopy.
The Perithecium
The ascocarp (perithecium) in the Laboulbeniales is almost invariably
an outgrowth of the receptacle (the only known exceptions are
occasional young perithecia in Herpomyces [Tavares, 1966]). Perithecia
are almost always produced by cell // (for exceptions, see Amorpho-
myces, Rhizopodomyces, Nanomyces, and most genera in the Dimor-
phomyceteae).
Typically the perithecium consists of stalk cells and a procarp or its
derivative cells surrounded and surmounted by one outer and one inner
TAVARES: PERITHECIUM
67
layer of wall cells (the inner layer may be restricted to the perithecial
neck). Each layer consists of 4 vertical rows of cells. Perithecia develop
in two ways — by the upgrowth of wall cells around the carpogonium
(characteristic of the Laboulbeniineae) or by the intrusion of the
carpogonium upward between the rows of wall cells (characteristic of the
Herpomycetineae).
The perithecia of all genera in the Laboulbeniineae probably develop
like those of Laboulbenia except those in which inner walls occur only in
the neck or possibly are lacking altogether. However, only a small
number of unstained specimens have been studied in most genera; well-
stained material of various early developmental stages or the use of
potassium hydroxide (Thaxter, 1896) should reveal cell division
sequences. Nevertheless, the configurations of the cells in the other genera
of the suborder are similar to those found in Laboulbenia.
In Thaumasiomyces and Tettigomyces, it is possible that extra cells
are formed in the outer walls of the venter of the perithecium, as in
Helodiomyces; however, such cells might have a different origin in these
two genera. In the Hydrophilomyceteae, the presence of 4 cells in the
basal cell tier above cells VI and VII, rather than 3 cells, may either
indicate lateral production of a wall cell by one of the basal cells (cf. cell
configuration in Hydraeomyces [PI. 34,e]) or a significant difference in
the sequence of development of the perithecium in this tribe. Possibly,
the two basal wall cells on the outer (anterior) side of the perithecium
are lobes of cell m; if so, the two basal cells on the inner side would be
cells n and n'.
A slightly different arrangement of basal cells was also reported in
Euzodiomyces by Benjamin and Shanor (1951); they believed there were
2 secondary stalk cells (the equivalent of VII and m), each of which gave
rise to 2 rows of perithecial wall cells. My own observations show that
this is one of the genera in which there is a slight lateral displacement
in the basal cell region, with cell n lying at the side, rather than over the
dorsal or ventral surface, so that it is sometimes difficult to see the
actual source of the second row from cell n.
Although the hyphal growth pattern typical of filamentous fungi is
readily apparent only in the appendages in the Laboulbeniales, the
growth of the entire thallus ordinarily is accompanied by the formation
of a transverse septum in the terminal cell of a shortened hypha, the
subterminal cell dividing after producing a lateral outgrowth. One
daughter nucleus migrates into the outgrowth; after the lateral branch
has been cut off by a septum, it may divide again in the manner of a
terminal cell (exceptions are the longitudinal or intercalary transverse
divisions in receptacles in Zodiomyces, Euzodiomyces, and some other
genera; cf. the formation of the basal wall cells of the perithecium of
Herpomyces).

68
MYCOLOGIA MEMOIR NO. 9
The divisions of the carpogenic cell of Laboulbenia, like the initial
divisions within the germinating spore and that of cell h, do not follow
the usual pattern of cross-wall formation. First, a transverse septum is
formed after nuclear division of the intercalary carpogenic cell. Later,
the supporting cells of the ascogonium are cut off at both ends of the
carpogenic cell by transverse or oblique septa.
The basic difference in the perithecia of the Laboulbeniineae is in the
relationship of the stalk cell VI and the secondary stalk cell VII to the
primary axis of the thallus. Cell m remains on the outside of the stalk
axis whether it is borne on the primary axis or on a secondary axis
terminating in a stalk appendage (see Pis. 17-23 and section on phylogeny).
There are four types of variation in basal and stalk cells in the
Laboulbeniineae besides the location with respect to the primary axis:
1. Rotation of basal cells and secondary stalk cell in the Laboulbenia-
ceae, as in Sphaleromyces, in which it is 180°, with cell m on the outside
(in Acallomyces, there is only a slight amount of displacement).
2. Amount of cell wall thickening in basal cells and secondary stalk
cell; in the Dimorphomyceteae, Euphoriomycetinae, and Aporomy-
cetinae, these cells are often destroyed early by growth of the ascogenic
cell mass downward. (Cf. Asaphomyces, Dermapteromyces, and Benja-
minella.)
3. Position on a uniseriate row of cells — in Kainomyces, Pselaphi-
domyces, Histeridomyces, and Trochoideomyces. (Cf. the Asaphomy-
cetinae, Dixomyces.)
4.Extra basal, stalk, and secondary stalk cells. There is an extra stalk
cell (VI1) in the more highly developed genera of the Ceratomyceteae
(such as Eusynaptomyces, Helodiomyces, Phurmomyces) which is
absent in Plectomyces, Drepanomyces, Thaumasiomyces, and Thripo-
myces (cf. Tettigomyces). Cell VI' is formed laterally from VI below the
level of n and n'\ it subtends m (VI' and m are distinguished by their
position in relation to the other basal cells of the perithecium).
In Ceratomyces mirabilis, there is an extra cell formed from VII
(VW) toward the inside of the thallus (toward the appendage); it
directly subtends the carpogonial cell. There is also an n" cell that
subtends a short-celled outer wall cell row and is paired with n from the
VII cell. Cell n' arises from VII'. Were it not for its position in the
primary axis of the thallus, VII might be called n here. Thus, the 1:3 cell
arrangement typical of the Laboulbeniineae has been altered here;
instead of there being two basal cells from the original cell VII (one
producing one row of outer wall cells and one giving rise to two rows), there
are actually three basal cells. (The basal cell designation has been given
TAVARES: PERITHECIUM
69
to the cells at the same level as rri). Possibly, an increase in the number
of cells in the thallus is accompanied by an increase in the number of
basal and stalk cells (cf. Tettigomyces).
From a phylogenetic standpoint, a primitive perithecium in the La-
boulbeniales is one in which each of the four vertical rows of outer wall
cells consists of many cells approximately equal in height (see
arrangement of genera in key; cf. Fig. 21). Genera of the more advanced groups
tend to have four vertical rows with a reduced number of outer wall
cells, usually with the lower cells much taller than the upper cells. In Co-
lonomyces and Zodiomyces, the cell walls are poorly defined as they are
in Drepanomyces and Thripomyces. The failure to develop distinct cell
walls is undoubtedly a highly modified characteristic and is often
accompanied by a reduction in the number of cells (cf. the more highly
evolved genera of the Dimorphomyceteae and Euphoriomyceteae).
There is a tendency for upper cells to be shorter than lower cells in the
outer wall of the perithecium in genera having equal or subequal cells
(for example, Cryptandromyces, Mimeomyces, Peyerimhoffiella, Ce-
ratomycetaceae), as well as in those having distinctly unequal cells. On
the other hand, in Kainomyces, the uppermost cells are taller, the
additional wall cells being cut off at the base of the apical cells. The most
recently formed septa may be too thin to detect easily, particularly in
species lacking constrictions at the septa (careful focussing should reveal
the exact levels of the inner and outer wall septa when they occur in
approximately the same plane).
Like the multiple-tiered perithecia of Herpomyces, the Euceratomy-
cetaceae, and the Ceratomycetaceae, those of Kainomyces, Pselaphido-
myces, Zodiomyces, and the Compsomycetinae may be considered to be
primitive. It is possible that the perithecium of Cryptandromyces
represents the ancestral type in the Laboulbenieae. The nature of the peri-
thecial wall cells, together with the presence of secondary development
of the receptacle in the Compsomycetinae, suggests that it is a relatively
old group. On the other hand, Scaphidiomyces and Clonophoromyces,
with a more complex secondary receptacle and unequal wall cells, are
more highly evolved. The 5 outer wall cell tiers typically occurring in
Autophagomyces, which tend to be subequal in the lower part of the
perithecium, suggest a relationship with Tetrandromyces and Dioi-
comyces; this is borne out by the form of the primary appendage. The
perithecia of Stigmatomyces, Laboulbenia, and many other genera have
a reduced number of tiers (4), which are decidedly unequal in height.
Genera in the Teratomycetinae tend to have very tall lower cells in the
vertical rows of outer wall cells; here there seems to be a well-developed
perithecium and retention of what may be considered a primitive
character — multiple perithecia, as well as a multicellular receptacle.

70 MYCOLOGIA MEMOIR NO. 9
Multiple perithecia appear in many genera that otherwise appear to be
relatively advanced phylogenetically — for example, most Dimorpho-
myceteae, Euphoriomyces, Balazucia, and Hydrophilomyces. On the
other hand, only one perithecium is present in thalli of the Ceratomy-
cetaceae under normal conditions, as well as in most genera in the Eu-
ceratomycetaceae.
Although the relationship between Hydraeomyces and Chitonomyces
seems clear because of the similar appendages and antheridia, there is a
great difference in the outer wall cells. The compound antheridia
occurring in some species of Mimeomyces demonstrate its affinity to other
genera of the Peyritschielleae; however, it is the only genus in the Pey-
ritschielloideae having outer wall cells that may be quite similar in
height.
In the Asaphomycetinae and the Euphoriomyceteae, the lowermost
(venter) cell in each vertical row of perithecial outer wall cells is very
tall. The second cell is very short and is at the level of the trichogyne
stump in many species in the Euphoriomycetinae. A very tall venter cell
is also characteristic of Distolomyces.
Although most genera in the Teratomyceteae and the Laboulbenieae
apparently have only 4 cells in each vertical row of perithecial outer wall
cells, there are a number of genera in which there are 5 cells: Apatelo-
myces, Cryptandromyces, Diclonomyces, Dipodomyces, Peyerimhof-
fiella, Rhizopodomyces, Sphaleromyces, Stemmatomyces, Sym-
podomyces, Synandromyces, and a few other genera, some taxa of
which (like Autophqgomyces) may have more than 5 cells. There are
probably late divisions in subterminal outer wall cells in a number of
genera having 4 or 5 cells in each vertical row (for example, some of the
genera in the Haplomyceteae and Acallomyces, possibly also
Mimeomyces). In some genera there are cells of different heights in different
rows near the apex of the perithecium — for example, Apatomyces and
Mimeomyces.
In Ceratomyces, Eusynaptomyces, Synaptomyces, Phurmomyces,
Botryandromyces, and Dixomyces, all vertical rows of outer wall cells
do not have the same number of cells. There are reduced numbers of
cells in some outer wall cell rows in certain species, such as Ecteino-
myces trichopterophilus, in which a difference of level of the basal cells
is responsible. In Plectomyces, one of the cell rows from n has a short
cell, comparable in height to n' and the cell above m, whereas the other
row from n has a tall lower cell. In this genus, n' is higher in position i
than m and n.
Very few observations have been published on inner walls; it is I
usually necessary to study well-stained material in order to determine
i
TAVARES: PERITHECIUM
71
the origin of the inner wall cell rows and the number of cells in each
row. There appear to be 4 inner wall cell rows arising from the basal
cells in the Euceratomycetaceae (PI. 17,f) and Ceratomycetaceae (PI.
18,e), as in most Laboulbeniaceae that have been studied, but their
structure, origin, and number have not been determined definitely.
In Herpomyces, the inner wall cell rows start at the third or fourth tier
level from the outer wall cells and are thus the equivalent of periphyses.
It is possible that in species of reduced size (PI. 43,e), inner walls often
may be lacking — at maturity outer walls are detectible only at the apex
of the perithecium. However, in Phaulomyces octotemni there appears
to be an inner cell layer (possibly consisting of fewer than 4 cells) arising
from the second tier of outer wall cells. The same is true in Dimero-
myces, in which there apparently are inner rows of 2 cells each. It is
probable that the inner walls function during spore discharge.
The apex of the perithecium sometimes is armed with lobes or
outgrowths. In genera in which some species have ornamented perithecia
and some have simple (entire) perithecial apices (for example,
Autophagomyces, Chitonomyces, Corethromyces, Diphymyces, Dixomyces,
Laboulbenia, Mimeomyces, Misgomyces, Rhizomyces, Sphaleromyces,
and Synandromyces), the exact structure of the outgrowths should be
determined. It may be possible to detect very reduced ornamentation
that shows the relationship between species clearly. Species of
Laboulbenia that parasitize Gyrinidae typically bear apical outgrowths (see Bala-
zuc, 1971c, 1973d, 1975d). Some genera having apical outgrowths are
Acallomyces, Dipodomyces, Distolomyces, Enarthromyces, Helodio-
myces, Ilyomyces, Ilytheomyces, Kruphaiomyces, Polyandromyces,
Prolixandromyces, Smeringomyces, and Stemmatomyces. Within a
single species such as Hesperomyces virescens, there may be
considerable variation in lobe development. Outgrowths that arise subapically
are found in Acompsomyces, Chitonomyces, Dioicomyces, Hetodio-
myces, Kyphomyces, Misgomyces, Smeringomyces, Triceromyces, and
Zodiomyces (see also Ceratomyces and Autoicomyces, as well as
Herpomyces). The apex of the perithecium in Kainomyces and Euceratomyces
ends in a rostrum or narrowed apical region including all four outer wall
cell rows, la Euceratomyces, the protoplasts within the rostrum
deteriorate and the cells below form broad, rounded lobes (PI. 17,d).
Conspicuous apical and subapical outgrowths may function as trigger
organs to initiate spore discharge (Thaxter, 1924, p. 388). When thalli
grow on tarsi of beetles, the lack of these organs in the Ceratomycetinae
may be unimportant because movements of the hosts control the
opening of the ostiole. Thick cell walls on simple apices and what
appear to be narrow channels from the protoplasts to the surface (as in
species of Laboulbenia [Pi. 48,d] and Corethromyces [PI. 48,a])
probably are part of the mechanism for opening the ostiole.

72
MYCOLOGIA MEMOIR NO. 9
The trichogyne is very frequently similar in appearance to the
appendage branches (cf. Laboulbenia gyrinidarum, Diplomyces, and Sander-
soniomyces with Diaphoromyces, Hydraeomyces, and Chitonomyces;
see also the lobes of the trichogyne in Dermapteromyces ctenophorus).
The lowest septum in a series of black septa may persist after the
trichogyne breaks off. In Rhizomyces crispatus Thaxter, the trichogyne is
abundantly branched and darkens like the appendage.
Trichogynes sometimes produce many slender branches or broad
apical lobes, depending on the species, but this probably occurs after
fertilization has taken place in many genera. In Acompsomyces ato-
mariae, for example, the tip of the trichogyne is close to the neck of the
phialide before outer wall cells are formed; later, the trichogyne
develops large, broad lobes (Thaxter, 1908, PI. XLII, Fig. 8) at a time
when fertilization may have already occurred. In Euceratomyces, the
extended trichogyne may bear short branchlets that resemble
exogenous spermatia (cf. Euzodiomyces [Benjamin and Shanor, 1951]).
There are several species in which a very long trichogyne seems to
grow directly downward (or upward) to an antheridium (for example,
Euphoriomyces cioideus [PL 42,b], Kainomyces isomali [PI. 26,e],
Zodiomyces vorticellarius [PI. 25,b], Scelophoromyces osorianus [PI.
33,c], and Scaphidiomyces scaphicomae [PI. 30,d]). In Euphoriomyces
pterogenii, trichogynes grow either upward or downward toward an
antheridium. In other taxa, such as Ecteinomyces trichopterophilus and
Asaphomyces cholevae (PI. 42,c), the receptive trichogyne is very short
and originates close to the mouth of the antheridium. An intricately
coiled trichogyne extends between the perithecium and the antheridium
in Smeringomyces trinitatis (PI. 31,a). The trichogyne is usually
terminal at the end of the trichophoric cell, but in some taxa, such as
Sphaleromyces lathrobii (PI. 47,a) and Neohaplomyces (Benjamin,
1955), it is lateral in position. It may also be somewhat lateral in Synan-
dromyces.
In the Laboulbeniaceae, the trichogyne, which is usually at the third
wall cell level, is much higher in relation to the wall cell tiers than it is in
most genera of the Ceratomycetaceae (exceptions are Helodiomyces and
Thaumasiomyces). In most genera, the stump of the trichogyne is visible
on the inner side of the perithecium (see Pis. 4,a,e; 39,e), often as a dark
or slightly raised scar. In the Laboulbeniaceae, septa are usually formed
in the outer wall of the perithecium at or just below the base of the
trichogyne, where the perithecium continues its upward growth beside it
(see Laboulbenia [PI. 4,b], Kainomyces [PI. 26,a,b]). In Rhachomyces,
the stump of the trichogyne may be just below the perithecial apex,
which indicates that outer wall growth is all intercalary (considerable
wall elongation takes place).
TAVARES: PERITHECIUM
73
A persistent trichogyne base occurs in Acrogynomyces and Aporo-
myces (PI. 41,h). The bases of these trichogynes become thick-walled
and the upper parts usually break off; the perithecia grow very little, if
any, beyond the trichogyne base.
In the Ceratomycetaceae, the trichogyne is strictly basal only in Tetti-
gomyces and in the Ceratomycetinae. In Plectomyces it emerges from
the side of the perithecium at the level of the second tier of outer wall
cells, with the trichophoric cell lying along the posterior margin of the
lowermost wall cells. The trichogyne arises from the apex of the two-
tiered perithecium in Drepanomyces and Thripomyces (Tavares,
unpublished), in which the walls of the perithecium grow up around the
carpogonial cell row as they do in the Laboulbeniaceae and Eucerato-
mycetaceae (cf. Benjamin, 1955, Benjamin and Shanor, 1951, Tavares,
1980).
Although it is possible that in some genera the number of ascogenic
cells is constant in all species, in others such as Stigmatomyces, the
number appears to vary according to the size of the perithecium. In
Acallomyces there may be 2, 4, or perhaps more, depending on the age
of the thallus (Tavares, 1973). Polyascomyces, Helodiomyces, Microso-
myces, Thaumasiomyces (PI. 19,c), and Compsomyces insignis have a
large number of ascogenic cells, at least in old perithecia. Thaxter (1896)
reported eight ascogenic cells in Haplomyces and Cantharomyces,
although I have detected no more than four groups of asci. There are
many genera in which there is only one ascogenic cell — for example,
Euphoriomyces, Rhipidiomyces, and probably Kainomyces. In many
species, such as Euphoriomyces cybocephali, the perithecium is so small
that only one ascus is formed (PI. 43,d).
It is possible that reduced perithecial size may also have an effect on
the number of spores in the ascus. Thaxter (1931) reported that there are
only 2 spores in the ascus of Kruphaiomyces. My own observations
showed that although this is true, some spores may have been forced out
of the asci very early by pressure resulting from growth. Spegazzini
(1917) thought there might be 2-spored asci in Cucujomyces; the
appearance of the asci in C. cylindrocarpus Spegazzini suggests that this
may be true. Although there are 8-spored asci in Herpomyces in Her-
pomycetineaerthe usual number of spores in an ascus in the Laboul-
beniineae is 4, but four of the eight nuclei in the ascus deteriorate and
the remaining four divide once after the spore walls are formed. As a
result, each spore is 2-celled (including those of Amorphomyces). I have
observed 8-spored asci only in some species of Compsomyces
(Laboulbeniaceae; PI. 28,b) and in Euceratomyces (Euceratomycetaceae;
Tavares, 1980). Although the spore septum is median in Herpomyces
and nearly median in Compsomyces verticillatus, it is near the base in

74
MYCOLOGIA MEMOIR NO. 9
Euceratomyces terrestris (cf. PI. 15,d; Tavares, 1980, Fig. 6). Benjamin
and Shanor (1951) reported that Euzodiomyces (Euceratomycetaceae)
has 8-spored asci (Cepede and Picard's illustration [1908, PI. IV]
showed three or four spores at each level in the perithecium). Autoi-
comyces (Ceratomycetaceae) appears to have 4-spored asci. The two
cells of each spore in the 8-spored asci of Compsomyces verticillatus and
Euceratomyces terrestris are unequal in length, whereas those of
Herpomyces species are equal.
The number of spores in an ascus can be detected accurately only by
pressing whole asci out of the perithecium; the count is best made on
slightly immature asci from which spores have not been discharged.
In 8-spored asci, the spores tend to be small and they are usually
arranged more or less in two tiers of more than two spores each. In 4-
spored asci, the spores are usually arranged in two pairs, the pair on the
lower level extending slightly above or below the uppermost pair; the
spores are usually quite long in relation to the length of the perithecium.
The Appendages
The upper cell of the germinating spore in the Laboulbeniales
produces the primary appendage system only, except in Herpomyces, in
which perithecia may arise from extensions of the subapical appendage
cell in the female (Tavares, 1966) or from a daughter cell of the sub-
apical cell (see Fig. 16,b). Primary appendages may consist of only one
or two cells, as in Filariomyces, Dioicomyces, Tetrandromyces, Rhizo-
podomyces, Rhipidiomyces, and some species of Herpomyces. Very
simple primary appendages also occur in Diandromyces and Chae-
tarthriomyces, as well as in the genera having inflated appendages like
those of Peyritschiella. The primary appendage becomes aborted in
Dipodomyces monstruosus and probably also in Chaetarthriomyces
flexatus (cf. Amorphomyces). Extensive primary appendage systems
occur in many species of Laboulbenia and Corethromyces, as well as in
Rhizomyces and some other genera.
Any sterile or antheridial branches from the lower spore segment are
designated secondary appendages. When secondary appendages are
close to the primary appendage (as in Amphimyces [PI. 42,k], Helo-
diomyces [PI. 20,d], Kainomyces [PI. 26,e], andMeionomyces [PI. 44,e]
[see also Apatelomyces]), they may be hard to distinguish. The
abundant, narrow appendages visible in the mature thalli of Zodiomyces (PI.
25,c) are all secondary in origin. The primary appendage of Thauma-
siomyces apparently undergoes little development and secondary
appendage branches predominate. The secondary appendages are also
intermingled with the primary appendage in most genera of the
TAVARES: APPENDAGES
75
Teratomycetinae and in Euphoriomyces bilateralis. In Smeringomyces,
the secondary appendages may obscure the primary appendage (see PI.
31,a,b), which in S. trinitatis appears to be a two-celled structure on the
lower surface rising from a broad, rounded III cell (Fig. 19,e). Although
similar to the secondary appendages, the primary appendages of
Trochoideomyces (PI. 27,a,b) and Clematomyces are located close to
the normal position; however, there may be some divisions in the
subtending cells of the suprabasal cell complex. In Rhachomyces, the
primary appendage is borne in the normal position — on cell 777, and
the secondary receptacle arises at the side of 77 in the same manner in
which perithecia are formed in other genera. In this genus, the primary
septum may be black and constricted, whereas it is colorless in Rickia,
Peyritschiella, Chitonomyces, and related genera, in which the
conspicuous black septum is at the top of the basal cell of the appendage as
it is in Botryandromyces ornatus. The primary appendage in Rickia
is at or near the apex of the thallus beside the perithecium, whereas in
Peyritschiella, it is usually centrally located and may be difficult to
distinguish from the secondary appendages.
Secondary appendages may arise only above the perithecia (see
Asaphomyces, Dermapteromyces, Kruphaiomyces, Ormomyces, and
Peyerimhoffiella, as well as two species of Rhadinomyces, some species
of Autophagomyces and Stichomyces, and Kyphomyces bicornis
[PI. 32,b] [cf. also Scelophoromyces]) or they may occur both above
and below the perithecium (see Amphimyces, Filariomyces, and
Homaromyces; cf. Chaetomyces). In Euphoriomyces, the whole range
of variation in position of the appendages occurs in a single genus; in E.
pterogenii, the upper branches may be either primary or secondary in
origin (PI. 43,j,k). From a taxonomic standpoint, whether or not a
branch arises from 777 or from the basal cell of the primary appendage
has minor significance, especially if similar branches occur on both
cells, as they do in Rhadinomyces. The presence of secondary
appendages below the perithecium is considered to be a primitive
characteristic.
Many genera, such as Stigmatomyces, seem to have no capacity for
producing branches from cell 777. As a rule, Corethromyces produces no
secondary appendages; however, in C. circinellus and C. laminifer, a
small appendage may arise from a cell between cells 77 and VI.
(Compare Autophagomyces and Stichomyces, as well as Laboulbenia proli-
ferans&ndL. variabilis.)
Accessory appendages bearing a precise relationship to the perithecia
occur in Scaphidiomyces and Clonophoromyces (PI. 30). Similar
secondary appendages that are somewhat different in nature are those paired
with the perithecia in Histeridomyces (PI. 29,a).

76
MYCOLOGIA MEMOIR NO. 9
The primary appendages of Chitonomyces and Hydraeomyces seem
to break off very early just above the constricted black septum (see PI.
34,g). In Peyritschiella, the broken appendage may proliferate,
producing two cells above the constricted black septum. Broken-off
primary appendages are difficult to detect in some genera (for example,
Rhizopodomyces), but in Scaphidiomyces, Histeridomyces, and Clono-
phoromyces, they are easily recognized beside the secondary
receptacles. The primary appendage of Coreomyces is destroyed as the peri-
thecium develops within the upper part of the receptacle (Thaxter,
1908).
The outer (main) axis of the primary appendage of Laboulbenia may
break off as the thallus develops, leaving only the inner appendage,
together with a few young branches from the inner side of the outer
appendage (Pis. 49,a; 52,a,h). Outer branches of the appendage of Core-
thromyces may also break off. Breakage completely alters the
appearance of the appendage in Aphanandromyces. Proliferation of new
hyphae through broken cells occurs regularly in the main axis of the
primary appendage of Laboulbenia flagellata, in its upper branches,
and in the inner antheridial branches (PI. 2,a); it also occurs in many
species of Dimeromyces. When breakage of the primary appendage of
L. gyrinidarum takes place, proliferation may sometimes occur;
however, lateral branches may be formed, instead, below the black cross-
walls.
The identity of appendages — whether primary or secondary — and
the nature of breaks in branches in mature thalli should be taken into
consideration when descriptions of species and genera are written. It is
necessary to distinguish between an unbroken appendage system and
one in which secondary proliferation has taken place. This is
particularly true in a genus like Laboulbenia, in which the appendage system is
of considerable taxonomic importance.
The basal cell of the primary appendage of Laboulbenia is the
insertion cell (e), from which the inner and outer appendages arise. Ce-
pede (1914) assumed that the insertion cell of L. blanchardii Cepede was
the darkened, thickened upper wall of the uppermost receptacle cell.
Cell e was mistaken by Istvanffi (1895a) for a black ring encircling the
base of the appendages of L. gigantea Istvanffi. However, he clearly
showed the origin of cell g from an undifferentiated cell e in a very
young thallus (Tab. II, Fig. 6); he believed that these two cells
subtended the insertion cell (probably he thought them to be cells IV
and Fbecause he lacked sufficient developmental stages).
In Laboulbenia and Stichomyces, the upper cell of the spore elongates
directly upward and no trace of the spore tip remains at maturity. By
contrast, in many taxa (for example, Zeugandromyces australis [see
TAVARES: APPENDAGES
77
Thaxter, 1931, Pi. XXI, Fig. 13], Euzodiomyces lathrobiae [Benjamin
and Shanor, 1951], Sphaleromyces lathrobiae [PI. 47,a,b], and Herpo-
myces periplanetae and H. stylopygae [male thalli; PI. 14,c, above the
upper black disk; also Tavares, 1966]) extension of cytoplasm into
appendage branches takes place just below the spore apex, which is spi-
nose, and the spine takes a lateral position (see also Microsomyces).
The failure of the spore apex in these genera to elongate directly upward
suggests that the spinose tip has a wall incapable of expansion. A spine
may also occur on the compound antheridia of Haplomyces, Poro-
phoromyces, Neohaplomyces, and Tettigomyces africanus, as well as on
the sterile primary appendage of Trenomyces (Thaxter, 1926).
Another form of persistent spore apex is found in Corethromyces
cryptobii, in which it is blunt and rounded (cf. C. cornutus and its allies,
Thaxter, 1931, pi. XXXV). In C. catalinae, the persistent spore apex is a
terminal phialide that becomes an inflated vesicle when it ceases to
function.
Similar vesicles terminate appendage branches of Colonomyces and
Euceratomyces, which are reminiscent of the inflated appendage cells of
Thaumasiomyces scabellularius. Pointed cells terminating branchlets in
some genera of the Teratomycetinae probably are specialized phialides
that cease functioning early (see Sandersoniomyces).
Aside from lateral outgrowths just below a persistent spore apex,
growth of appendages is either apical or by means of lateral extension of
intercalary cells from the upper angles of the cells (Pis. 2,a; 33,c; 36,f;
44,c; 46,e; 47,a). Corner cells do not have any particular significance;
they merely represent a particular placement of the septum following
cell division (that is, small cells triangular in optical section formed
from the upper angles of cells of the appendages). They seem to be
characteristic of certain genera (for example, Hydrophilomyces and
Chaetarthriomyces) and occur in others, such as Corethromyces, in old
thalli (PI. 48,a). They also occur in Diphymyces curvatus (PI. 46,e).
In Hydrophilomyces species on Phaenonotum they function as
phialides, probably growing out into appendages after the cessation of sper-
matia formation. Appendages develop from the corner cells in other
genera, as well (for example, Ormomyces, Stichomyces, and Euphorio-
myces). Cells similar to corner cells, but cut off the lower angles of
appendage and receptacle cells, occur in Dermapteromyces and Eu-
phoriomyces cioideus.

78
MYCOLOGIA MEMOIR NO. 9
The Antheridia
Although both Herpomyces and Laboulbenia have simple
endogenous antheridia (phialides), those of Herpomyces differ primarily by
their greater length in contrast to the length of the underlying cells from
which they arise (cf. Prolixandromyces). In Laboulbenia, Scaphidio-
myces, Stigmatomyces, and many other genera, the phialides are
relatively short, flask-shaped structures with slightly narrowed necks,
perhaps with one or two spermatia free within the broad cavity (cf. PI.
ll,c). In other genera, such as Euphoriomyces, and in Hydrophilo-
myces coneglianensis, the antheridia are wide at the apex and the
spermatia appear to be produced at the tip of the protoplast, although there
is actually a very short collar (formed by a thin cell wall) protruding
beyond the protoplast. In Histeridomyces, the empty outer wall
remaining after the discharge of the first spermatium encloses a subspherical
cavity into which subsequent spermatia are extruded (cf. PI. 29,a; Fig.
19,a; cf. also Homaromyces).
Unlike the spherical spermatia of Histeridomyces, those of
Laboulbenia and Herpomyces are somewhat cylindrical. They have relatively
large nuclei and a narrow peripheral layer of cytoplasm, as well as a thin
cell wall. Although the spermatia of Scelophoromyces are thin-walled
and spherical at first, apparently those formed later by the antheridia
are thick-walled, with smaller protoplasts (PI. 33 a,b). In Cochliomyces
(PI. 18 a-c), the spermatia borne in the antheridium terminating the
stalk appendage may be quite long, and the apex of the primary
appendage produces spermatia-like cells that are considerably larger in size
than the spermatia themselves. It is possible that some taxa only
produce spermatia-like stuctures lacking nuclei (cf. Pi. 3,c; see Thax-
ter's statement [1896, p. 217] that he was unable to detect nuclei by
staining) and never form functional spermatia (cf. also the abnormal
nucleate cells produced by Laboulbenia gyrinidarum or L. borealis).
Phialides on unstained thalli are being assumed, possibly in error, to be
producing nucleate cells.
Thaxter (1896) referred to spermatia as naked cells, although he
suggested that spermatial walls are probably present on small spermatia as
well as on the large ones of Enarthromyces, even though they may not
be visible. He apparently believed that a wall is secreted after the
spermatium is formed. As has been shown in Laboulbenia flagellata (PI. 3,
0, the spermatial wall is continuous with that of the antheridial
protoplast and is conspicuous from the beginning.
The diaphragm separating the neck cavity of the phialide appeared to
Thaxter (1896) to consist of wall material perforated in the middle. In
Laboulbenia flagellata, the diaphragm material is deposited as a layer
TAVARES: ANTHERIDIA
79
lining the phialide wall after the first spermatium is formed and the apex
of the phialide becomes perforated (PI. 3,0- It lacks the birefringence of
the adjacent wall layers and must differ from them in structure. The
collar it forms around the base of the neck of the phialide thickens as
consecutive septa are produced. A similar collar was shown in Neuro-
spora crassa Shear & Dodge by Lowry et al. (1967). In Laboulbenia,
there is an outer clear area surrounding the collar similar to those shown
in Aspergillus giganteus Wehmer by Trinci et al. (1968) and in Metar-
rhizium anisopliae (Metchn.) Sorokin by Hammill (1972). This clear
area may result from the pulling away of the diaphragm material. In all
of these fungi, the illustrations showed that the narrow wall
surrounding the developing microconidium is continuous with a narrow wall on
the inside of the dark collar, although this was not mentioned by any of
these authors. The continuity of the inner wall of the phialide protoplast
and spore was recognized by Roquebert and Abadie (1973) in Stil-
bothamnium nudipes Haum.
Although there are conidia formed in flask-shaped phialides in
Coniochaeta ligniaria (Grev.) Massee (Rogers, 1965), conidia are also
produced from pores in intercalary cells as in Euceratomyces. In Phialo-
myces, the phialides resemble those of Laboulbenia; the first spore is
terminal and the second is cut off just below the apex of the phialide
(Misra and Talbot, 1964).
The phialides reported by Dodge (1932) in Neurospora are basically
like those of Coreomyces in their extrusion of phialospores from a
lateral opening in an intercalary cell. Exogenous antheridia of Rhyncho-
phoromyces and Zodiomyces probably develop in the same way, the
distinction between these antheridia and the intercalary antheridia of
Corethromyces being the large size of the spermatia with respect to the
initiating cell (cf. the suggestionof Backus [1934] that exogenous
spermatia should be regarded as reduced antherids). The long, rodlike
spermatia that are borne laterally in Rhynchophoromyces and
terminally in Zodiomyces were believed sufficiently distinct by Thaxter (1896)
to warrant putting the organisms possessing them in a separate group
from genera with endogenous antheridia. Included with them were
species of Ceratomyces in which the spermatia were thought to be
formed from indistinct branchlets that broke up into segments. Thaxter
(1908) was forced to admit that a clear distinction could not be drawn
between endogenous and exogenous antheridia when he was faced with
a transitional type such as the undifferentiated intercalary antheridia of
Coreomyces, which have short lateral necks. However, he continued to
segregate the Laboulbeniales into families on the basis of antheridial
characters. Thaxter asked (1908, p. 234) whether the unspecialized
antheridial cells of Zodiomyces and Rhynchophoromyces had become
the specialized cells of the Laboulbeniaceae or whether the reverse were

80
MYCOLOGIA MEMOIR NO. 9
true, resulting in an adaptation that would facilitate fertilization in the
aquatic environment. He suggested Ceratomyces as the culmination of
this tendency. However, it is probable that the spermatia of
Ceratomyces are formed in the same manner as those of Rhynchophoromyces
and are present only on very young thalli. In Autoicomyces minusculus
there is a large solitary spermatium produced by the terminal phialide
(PI. 23,d-e); the neck appears to be fragile and easily broken by the large
spermatium. Much more study is needed on spermatium formation in
the Ceratomycetaceae, as well as in Zodiomyces, in which the spermatia
are terminal in position (PI. 25,a).
It is possible that only one or two spermatia are formed in each
antheridium in genera in which they fragile, evanescent, and rarely
observed. The position of the trichogyne in young thalli or the direction of
growth of its tip should indicate the position of such antheridia.
Phialides may be borne on the primary appendage (PI. 2,a), on
secondary appendages below the perithecium {Chaetomyces [PI. 43,h],
Amphimyces [PI. 42,m], and some species of Euphoriomyces [PI.
43,a]), or on secondary appendages from cell /// (Asaphomyces
[PI. 42,c], Dermapteromyces, and some thalli of Autophagomyces and
Stichomyces). They may be embedded within the outer cell layer of the
receptacle with their necks protruding at the surface (Rhipidiomyces
[PI. 42,e], Rickia, and Benjaminella). Lateral phialides on appendages
are sessile or have short (examples are Troglomyces and Stigmatomyces)
or long stalks (Fanniomyces). Although in most genera the necks of the
phialides extend toward the perithecium, in Stemmatomyces they are on
the outer side of the appendage.
Phialides may be solitary and terminal in Mimeomyces (PI. 36,g),
Smeringomyces (PI. 31,a), Kyphomyces (PI. 32,b) and Hydrophilo-
myces coneglianensis (PI. 32,a); they may be borne in clusters, as in
Clonophoromyces (PI. 30,a), Dipodomyces (PI. 29,b), Laboulbenia
(PI. 3,b), Rhadinomyces, Filariomyces, and Gloeandromyces; or they
may be seriate. Sessile or subsessile phialides may form a series on an
appendage, as in Trochoideomyces (PI. 27,a), Chaetarthriomyces (PI.
31,d), and Tavaresiella. In Corethromyces (PI. 47,d) and several other
genera, including Chaetomyces (PI. 43,h), Symplectromyces, and
Coreomyces, the seriate antheridia are undifferentiated intercalary
appendage cells; the series is usually terminated by a free phialide (the
pointed terminal structure in Sandersoniomyces is probably a phialide
in which the protoplast is used up in the production of spermatia before
the seriate antheridial branchlets develop). In the antheridial series in
Euceratomyces, a thick-walled, inflated, free phialide terminates an
appendage consisting of spermatia-producing intercalary cells (PI. 17,a;
cf. PI. 48,b).
TAVARES: ANTHERIDIA
81
The large phialides clustered around the spore septum in Botryandro-
myces are reminiscent of the compound antheridia of Mimeomyces;
both genera have slender phialides on the appendages. The stalk
appendage of Euzodiomyces may bear a cluster of phialides (Benjamin
and Shanor, 1951) (cf. Cochliomyces [PI. 18,a]).
Proliferation of appendages from old antheridia occurs in Misgo-
myces (PI. 38,0, Hydrophilomyces, and possibly also in Phaulomyces
(cf. PI. 41,d) and Euphoriomyces pterogenii, in which the first cell of
some appendages probably functions at first as a phialide. By contrast,
the antheridia of Chitonomyces (PI. 35,e) and Hydraeomyces (PI. 34,d)
apparently become functional after the apices of the short, inflated
secondary branchlets deteriorate. It is possible that phialides may arise
below deteriorating branchlets in Homaromyces, also.
Faull (1911) suggested that compound antheridia originated more
than once. The character of the perithecium of Tettigomyces shows that
this is true. The basic difference between the antheridial structure in the
Monoicomycetoideae and the Peyritschielloideae is in the manner in
which the spermatia-producing cells are formed. In Monoicomyces the
fertile cells of the antheridium produce cells from their inner surface;
each of these cells presumably forms spermatia. The cavity of the
antheridium into which the spermatia are discharged is an intercellular
space. In the Dimorphomyceteae and Haplomyceteae (and presumably in
the Peyritschielleae, as well) the initial fertile cells become subdivided,
the daughter cells producing spermatia-forming cells. Sterile cells break
down to form a discharge tube (which is intracellular with respect to the
initial fertile cell).
In other groups of fungi, there seems to be no counterpart of the
compound antheridium of the Dimeromyces type (see diagram of D.
adventiosus Thaxter, 1924, p. 332; cf. also PI. 38,d). The intercalary
position of the functional cells with respect to the cells terminating the
branchlet in early stages indicates that these compound antheridia are
not modified from simple phialides by an increase in the number of
spermatia-producing cells. The rare antheridia of Botryandromyces
having two fertile cells do not have subtending stalk cells within the
body of the antheridium; such stalk cells subtend one or more of the
fertile cells in well developed compound antheridia. It is probable that
proliferation of cells in the antheridium results in increasing numbers of
fertile cells with age in Mimeomyces. From an evolutionary standpoint,
there has probably been both an increase in number of spermatia-
forming cells (possibly the massive antheridium of Polyandromyces is
the result of polyploidy) and a reduction in number in the
Peyritschielloideae. This may be true in the Haplomyceteae; a reduction in number
certainly occurs in Dimeromyces. In Mimeomyces there is evidence of

82
MYCOLOGIA MEMOIR NO. 9
an initial neck cell (unpublished); the antheridium is borne on a stalk
cell (PI. 36,f). In Enarthromyces there is no external stalk cell; usually
the spermatia-producing cells form a vertical row along the side of the
primary axis of the thaljus.
The simpler antheridia in the Haplomyceteae are not very different in
structure from those of the Peyritschielleae; however, the fertile cells are
not arranged in a basal layer — their positions are more irregular. None
of the genera of the Haplomyceteae have simple phialides, such as those
occurring in Dimeromyces (PI. 38,a,c), Diaphoromyces, Rickia, and
Mimeomyces. Those in Mimeomyces are terminal on the fusiform
appendages, whereas in the other genera, their positions are similar to
those of the compound antheridia (if any are present).
The massive antheridia of Tettigomyces (PI. 18,d-f) and those of
some of the Haplomyceteae are somewhat reminiscent of spermagonia
of other fungi in the mature stages. This is also true of the compound
antheridium of Eumonoicomyces. Early developmental stages in Neo-
haplomyces (Benjamin, 1955) show that very regular septations take
place in the terminal cell of the thallus, resulting in a lateral
arrangement of antheridial cells. These discharge spermatia into an
anterior cavity. The arrangement of fertile cells in rows recalls the linear
cell rows in what is probably the antheridial region of Kainomyces,
as indicated by the direction of growth of the trichogyne (PI. 26,e); in
Kainomyces, there is a single cell row in each tier, rather than two rows
with an intervening cavity.
In Tettigomyces, the spermatia-producing cells may divide in the
plane of the cell row axis, so that there are double rows of cells, at least
near the axial cell of the antheridium (PI. 18,d); in Neohaplomyces, the
cell rows are single. In species having a highly organized antheridium,
such as Tettigomyces africanus and T gryllotalpae Thaxter, fertile cells
discharge spermatia into a central cavity, starting at the anterior edge.
In T. confusus and some other species, each cell row consists of a single
series of cells (PI. 18,h); although there is an upper and lower cell layer
with some intervening cells, there does not seem to be a distinct central
cavity in the antheridium.
In Polyandromyces and most of the Haplomycetinae, the primary
appendage becomes transformed into a compound antheridium.
However, its poorly developed perithecial basal cells indicate that there
is no close relationship between the dioecious genus Polyandromyces
and the monoecious Haplomycetinae.
Although Mimeomyces also has a compound antheridium formed
from the primary appendage, it constitutes (together with its stalk cell) a
lateral branch. The antheridia in Misgomyces result from division of the
TAVARES: FERTILIZATION
83
lowest cells of the primary appendage; however, the receptacle is
multicellular, whereas those of the Haplomycetinae consist of only three
cells. The only genera in the Peyritschielleae having multicellular
receptacles are characterized by thick, black constricted septa on the
appendages.
Although the compound antheridia of Kleidiomyces are borne on
appendages arising from cell 77, somewhat like those of Monoicomyces,
the manner in which the antheridial initial divides seems to ally it with
the Haplomycetinae, rather than with the Monoicomycetoideae. Even
though the development of the antheridia is different, there is a great
deal of similarity between those of Polyandromyces, Haplomyces, and
Eumonoicomyces. In fact, the similarity between the perithecia of the
Monoicomycetoideae and the Haplomyceteae suggests a closer
relationship than is indicated by the structure of the simpler antheridia.
Fertilization and Sexuality
Moller's proposal (1901) that the simple endogenous spermatia are
conidia similar to those of some other Ascomycetes was criticized by
Thaxter (1908) chiefly because it was clear to him that thalli never arose
from germination of spermatia. Numerous reports of microconidial
fertilization of trichogynes have been published since that time and the
distinction between micro- and macroconidia has become clearer.
Dodge (1935) mentioned that microconidia in Neurospora germinated
to form normal mycelia.
Because he had observed spermatia of Zodiomyces and other genera
attached to trichogynes, Thaxter (1896) assumed that fertilization
occurs. In his illustrations of Stigmatomyces baeri (Thaxter, 1896), he
indicated the presence of spermatia on the trichogyne (these had been
reported by Karsten [1869], although they were not observed by Pey-
ritsch [1873]). However, Thaxter (1931) later concluded that these
objects were outgrowths of the trichogyne. (Cf. the protrusions on the
elongating trichogyne of Herpomyces ectobiae.)
Thaxter's arguments (1908) in favor of the sexual function of the
spermatia and trichogynes seem to be a reflection of the prevailing
notion of red algal origin of the Ascomycetes, which he himself actually
did not share. He maintained that the sexual apparatus must be
operative because so much evidence was in favor of this view. No cytological
demonstration of fertilization has been presented in the numerous
publications that have appeared since Thaxter's observations were made.
However, most authors have assumed, together with Thaxter, that

84
MYCOLOGIA MEMOIR NO. 9
penetration of the extended trichogyne by a spermatium nucleus occurs,
followed by fusion of this nucleus with that of the carpogenic cell.
Bistis (1956) showed that the trichogyne in Ascobolus stercorarius
(Bull.) Schroet. developed in response to the presence of an oidium
placed nearby; at a certain stage of development it grew rapidly toward
it (cf. Thaxter's discussion [1896] of Zodiomyces and Pis. 25,b, 42,b).
In Neurospora sitophila Shear & Dodge, Backus (1939) observed that
some hours after fusion with a conidium, the adjacent wall of the
trichogyne tip might be swollen as if gelatinized. Migration of the sper-
matial protoplast into the trichogyne was reported, but total evacuation
was not seen. It was assumed that nuclear migration was rapid,
although nuclei were not observed. In Collema fragrans (Sm.) Ach. em.
Degel. (as C. microphyllum), Stahl (1877) observed an opening between
spermatium and trichogyne, as well as gelatinization of septa below (see
Taf. II, Figs, 4,5). In Gelasinospora, the densely stained trichogyne
became vacuolate, which suggested to Goos (1959) that a nucleus from
an empty, adherent spermatium had passed through it.
In Mycosphaerella tulipiferae (Schw.) Higgins, Higgins (1936) found
that empty spermatia were adherent to the receptive area of the
trichogyne (cf. also Shoemaker's observations [1955] on Cochliobolus).
Higgins reported an extra nucleus in the trichogyne, as well as one in the
base of the ascogonium at a later stage; however, he definitely stated
that he had not observed actual passage of a spermatial nucleus.
The initiation of ascocarp development after spermatization has been
demonstrated in various genera, such as Sclerotinia (see Drayton, 1932),
Pleurage (Dodge, 1936), and Neurospora (Dodge, 1932). However,
nuclear migration of the spermatial nucleus down through the
trichogyne was never definitely shown. That migration occurs was verified
experimentally by Goos (1959), who obtained a 1:1 ratio of mating types
in Gelasinospora from ascospores resulting from spermatized cultures.
This indicated that the spermatial nucleus contributed genetic material
at karyogamy.
In the antheridium-bearing species Laboulbenia flagellata, no
evidence has been found of spermatia fusing with the trichogyne or of
penetration of the trichogyne by a male nucleus. A second nucleus in the
trichophoric cell may be a daughter of the original nucleus of the cell;
unlike the deteriorating nucleus, it remains in active condition.
However, it is possible that the active nucleus is a male nucleus that has
migrated down through the trichogyne, which tends to deteriorate at
this time (cf. sequence reported by Faull [1912] for taxa in which the
presence of functional antheridia has not been confirmed).
TAVARES: FERTILIZATION
85
The ease with which appendage nuclei can pass through pores
indicates that the migration of a male nucleus could be rapid and could
occur without leaving any clear evidence of the process (PI. 3,a). There is
no breakdown of septa until the general deterioration of the trichogyne
takes place. If fertilization occurs in Laboulbenia, it must be just before
the wall breaks down between the trichophoric and carpogenic cells.
Benjamin and Shanor (1951) suggested that although fertilization
probably takes place at the time of early wall cell formation and
trichogyne extension, it might take place much earlier. The proximity of
trichogynes to antheridia in very young thalli suggests this in Mimeo-
myces deplanatus (PI. 36,f), Chaetomyces (PI. 43,g), Euphoriomyces
pterogenii (PI. 43,k), Dermapteromyces ctenophorus (PI. 44,b), and
Smeringomyces anomalus (PI. 31,a). Trichogynes that are growing
toward an antheridium are probably attached to perithecia that have not
yet been spermatized.
In Herpomyces ectobiae a second nucleus appears in the central cell
before it has even extended to the apex of the perithecium. No other
nucleus was seen in the centrum until after trichogyne breakdown. The
trichogyne nucleus probably is a sister of the upper centrum nucleus. No
spermatia were observed on or in the trichogyne. The nucleated
trichogyne is soon cut off by a septum and deterioration of the protoplast
begins, possibly following migration downward of a spermatial nucleus,
which would be hard to see in the heterogeneous cytoplasm of the
centrum. The size and position of the carpogonial nucleus shown in Fig.
13,c indicate that it probably arose from the ci nucleus.
If there is a delay in the initiation of the conjugate divisions, as there
seems to be in Laboulbenia (conjugate divisions may not occur until the
secondary inferior supporting cell is formed), perhaps this delay also
occurs in Herpomyces, where it is not certain that there is conjugate
division until the formation of the asci (Tavares, 1965,1966; see
discussion of Herpomyces). Possibly a spermatial nucleus enters the centrum
(either through the trichogyne or before it is formed), remaining there
and growing in size until a very late stage without dividing. Perhaps it
begins to divide when upgrowths are being formed from the carpogonial
initial cell. The small cells at the apices of some of these upgrowths
might be comparable to the small terminal cell of a crozier. More
knowledge is needed of the stages of development between those illustrated by
Hill (1977) in Figs. 3 and 4.
The occurrence of diakinesis in Herpomyces indicates that fusion in
the ascus is of dicaryotic nuclei and that meiosis does occur. One of the
nuclei in the dicaryon is probably from a male thallus.

86
MYCOLOGIA MEMOIR NO. 9
Although it is possible that monoecious species of the Laboulbeniales
produce asci in the absence of spermatial fertilization, it seems
improbable that spermatia are nonfunctional in dioecious species,
particularly those with two very distinct growth forms (however, see
discussion of Herpomyces above). Raper (1959) suggested that male-
female pairs may represent a physiological accommodation that is
possible between two thalli that develop together. Undoubtedly, he was
thinking primarily about Laboulbenia formicarum, which is obviously a
recently developed dioecious species, not about such genera as Dioico-
myces, in which large female and small male spores may be
differentiated within the perithecium (see photograph, p. 45, Benjamin, 1965,
of recently discharged spores). Furthermore, the males of Herpomyces
periplanetae and H. ectobiae produce antheridial branches very soon
after germination (Fig. 16, b; PI. 11, a), indicating that this character is
inherent in the thallus and not the result of slower growth or adjustment
to the greater development of the females. As Benjamin pointed out
(1971), males do not necessarily grow more slowly in the
Laboulbeniales. Olive's proposal (1966) that the more vigorous thallus of a pair is
the female and the slow-growing one is the male, therefore, has limited
application. Slower development of one member of a pair, resulting in
the growth of a small male beside a monoecious thallus, was reported by
Thaxter (1908) to occur commonly in some species, although it is
relatively rare in the order as a whole (cf. PI. 31,f with PI. XLIX, Figs. 16,
17, 1908).
In a species such as Laboulbenia formicarum, in which male and
female thalli are quite similar in appearance, it is possible that slower
development of one member of a pair could result in the suppression of
perithecium initiation. However, spermatia have never been seen on the
supposed female (Benjamin and Shanor, 1950b). If there is a
physiological advantage gained by the female which is accompanied by
loss of antheridia, there may be a mechanism that results in the absence
of antheridia except under adverse conditions (compare production of
antheridia by females in Herpomyces, Thaxter, 1908, Tavares, 1965).
In Hypomyces solani f. cucurbitae Snyder & Hansen, El-Ani (1956)
found that sex determination was a function of two loci, one controlling
protoperithecium formation, whereas the other was concerned with
spermatium production. Hermaphrodites and neuter thalli arose by
crossing-over of the two loci, which were on the same chromosome;
compatibility factors were independently inherited.
Herpomyces is an archaic dioecious genus. Thripomyces, possibly
dioecious, is relatively primitive. Other dioecious taxa are those in the
Dimorphomyceteae and Amorphomycetinae and some species of Cryp-
tandromyces, Euphoriomyces, Laboulbenia, and Aporomyces. Picar-
TAVARES: FERTILIZATION
87
della and Laboulbenia vignae W. Rossi (1978b) (which may not be a
Laboulbenia) have small males, unlike those of L. formicarum. Apato-
myces has small male thalli and may be dioecious. These genera are
relatively advanced phylogenetically. Just as heterothallism appears to
have arisen repeatedly in the fungi, according to Olive (1958), dioecism
seems to have arisen several times in the Laboulbeniales, as indicated by
the diverse groups to which the above genera belong.
If the Laboulbeniales were initially a dioecious order, then the
hermaphroditic genera could have arisen as in Hypomyces solani f.
cucurbitae. However, the absence of spermatium formation in the normal
female of Herpomyces and its occurrence in aborted females of H. para-
nensis (Tavares, 1965) suggest that the genus arose from a monoecious
progenitor (see Olive, 1966). In monoecious genera such as Laboulbenia
and Plectomyces (PI. 22,a), there is a similar tendency for aborted peri-
thecia to become filled with appendages or for appendages to grow
out from the base of the perithecium (Thaxter, 1896, PI. II, Fig. 8).
It is possible that in Laboulbenia formicarum there were two
mutations of hermaphrodites to unisexual thalli, the first mutation
producing the male, the other producing the female by suppression of the
antheridia. Perhaps delayed development of the perithecium in the
"male" thallus was succeeded by a genetic loss of ability to produce a
perithecium. A continuation of this evolutionary process might result in
a divergence of form with two entirely different kinds of thalli resulting,
as in Amorphomyces and Herpomyces. On the other hand, the thalli
might remain similar in form, as in Trenomyces, but they would bear
either perithecia or antheridia.
Laboulbenia formicarum should be examined carefully to determine
whether abnormality in development of the perithecium is accompanied
by antheridium formation; perhaps experimental work could be done
on ants in culture. More highly differentiated dioecious genera that
could be studied in this manner are Dimeromyces on termites and
Trenomyces on bird lice.
There may be both heterothallic and homothallic monoecious genera
in the Laboulbeniales (Olive, 1966). Experimental studies could be done
on Laboulbenia species on ants (Baumgartner, 1934; Blum, 1924),
beetles, or flies, as well as on Stigmatomyces and other genera occurring
on flies. Careful examination might reveal unusual morphological
characteristics, such as extra receptacle cells that appear to result from
mutation (Picard, 1913b, p. 533); such mutants could be crossed with
normal thalli. Hill's population (1977) of Herpomyces might be a
suitable mutant. Large species of Laboulbenia could easily be studied.

88
MYCOLOGIA MEMOIR NO. 9
It is my belief that true dimorphism is controlled by genetic factors,
although sometimes there are retarded or entirely undeveloped thalli in
the monoecious genera. Even though spermatial fertilization has not
been demonstrated conclusively, the relative positions of the tricho-
gynes and spermatia certainly suggest that they are functional in the
Laboulbeniineae, although there may be species in which their
reproductive functions have been lost. Although Olive (1966) mentioned
imperfect insect parasites having microconidial thalli only, most of
these taxa have spores quite different in shape from the microconidia or
spermatia of the Laboulbeniales (Thaxter, 1914b, 1920a). Therefore it is
probable that they are colonizers of the insect exoskeleton from other
taxonomic groups of fungi. Those with conidia similar to the spermatia
of the Laboulbeniales should be carefully studied in order to determine
whether they may be asexually reproducing mutants.
THE CLASSIFICATION OF THE LABOULBENIALES
Introduction
Although species of Laboulbenia tend to be remarkably similar in
their basic structure, occasionally there are thalli in which unusual
multiplication of perithecia and appendages has taken place (Picard,
1913b; Colla, 1933; Balazuc, 1976; see also PI. 2,c). If, as Middelhoek
(1957) suggested, such thalli simulate an earlier state in which a constant
number of receptacle cells has not yet been reached, the indeterminate
type of growth characteristic of Herpomyces and some other genera
must represent the primitive condition.
Thaxter (1908) apparently found it difficult to accept the concept of
simplified structure as indicative of an advanced condition, because he
said of Herpomyces: "The structure of the perithecium and the
relatively greater complication in the general structure of both sexes might
be assumed to place it higher in the scale than either Amorphomyces or
Dioicomyces, although the occurrence of a series of forms on Blattidae,
which are supposed to be representatives of one of the most ancient
types of true insects, might perhaps have been expected to be correlated
with a more primitive type in the parasite. But although the unisexual
forms with simple antheridia might for some reasons be assumed to be
the more primitive, the present genus is distinguished by a far more
complicated structure than the other unisexual forms of the same type,
and A morphomyces still remains... by far the simplest in structure of all
the Laboulbeniales."
In any study of phylogeny in the Laboulbeniales, the concept of the
basic taxonomic importance of total thallus structure (Chatton and
Picard, 1909; Picard, 1913b) deserves serious consideration. The Ce-
ratomycetaceae were accepted as primitive by Picard (1913b); however,
he placed more emphasis on the structure of the thallus as a whole than
on the male reproductive organs as an indication of relationship. His
use of Herpomyces and Trenomyces as examples of morphologically
similar genera shows that if thallus structure is to be considered, it must
be in conjunction with other characters.
Observations were made in 1959 and 1960 on all genera in Thaxter's
collection to determine the extent of receptacle development, the
position of the spore septum in the mature thallus, and the number of
spores in the ascus. Compsomyces, a genus on Coleoptera having a
89

90
MYCOLOGIA MEMOIR NO. 9
secondary receptacle of a variable number of cells and eight-spored asci
in some species, bears a strong resemblance to Herpomyces in its habit
of growth and its perithecia with short outer wall cells equal in height.
Clearly, this genus can be considered to be primitive. However, a
comparison of perithecial development in Compsomyces and Herpomyces
with that of Laboulbenia and Ceratomyces showed that the perithecia
of Compsomyces and Laboulbenia developed in the same manner,
whereas perithecium formation in Herpomyces was totally different.
The perithecium in Ceratomyces differed from that of Compsomyces
primarily in the position of the stalk cell and secondary stalk cell with
respect to the primary axis of the thallus. A comparison of
Compsomyces with Euceratomyces and Cochliomyces revealed that the position
of these two cells with respect to the primary axis was identical in all
three genera. However, a stalk appendage extended beyond the
secondary stalk cell in Euceratomyces and Cochliomyces. A complete
survey of all genera in Thaxter's collection followed, and it was
discovered that the formation of the perithecium was similar to that of
Laboulbenia in all genera except Herpomyces.
Because of the difficulty of observing structures in unstained
specimens, it was not possible to be certain about the septa in the perithecial
walls or of the outlines of the basal and stalk cells of the perithecia.
Therefore, further studies may require some minor changes in the
arrangement of taxa. In addition, more knowledge of the structure
of compound antheridia may necessitate additional readjustments.
A Comparison of Classification Systems
Although Rouget (1850) recognized that Laboulbenia rougetii was a
living organism, it was Robin (1852) who placed it among the fungi.
Robin (1853) included Laboulbenia in the division Clinosporei Lindl.
(which also included Puccinia), subdivision Endoclinei Lindl.
(having spores borne inside a receptacle), section Sphaeronemei
Lindl.; in the genus description, he called Laboulbenia a member of the
family Pyrenomycetes. Montagne (1856) put Laboulbenia into the
family Pyrenomycetes, order Dichaenacei, with Sphaeronaema (now in
the Sphaeropsidales) and Ostropa (now in the Discomycetes). The
Dichaenacei were placed between the Sphaeriacei and the Perisporiacei,
which are still included in the Pyrenomycetes.
Karsten (1869) failed to recognize the ascomycetous nature of
Stigmatomyces and placed the genus in the Mucorinee. Brauer (1870)
TAVARES: CLASSIFICATION SYSTEMS 91
realized that Arthrorhynchus, which had been described as an
acanthocephalan worm, was related to Stigmatomyces. Soon afterward,
Hoffman (1871) recognized the relationship between Stigmatomyces
and Laboulbenia. In 1873, Peyritsch established the family Laboul-
beniaceae with five genera. Karsten (1880-1883; 1895) continued to
deny the presence of asci; he established a new group, the Stigmato-
mycetes, which he placed between the Zygomycetes and the
Pyrenomycetes.
Confirmation by Thaxter (1891) of the formation of ascospores was
followed in 1894 by his arrangement of 23 genera according to antheri-
dial characters. In the first volume of his monograph of the Laboul-
beniaceae Thaxter (1896) established two groups, the Exogenae, having
spermatia formed exogenously (one "order," the Zodiomyceteae,
was included), and the Endogenae, with spermatia formed endo-
genously (this included the Peyritschielleae with compound antheridia
and the Laboulbenieae with simple antheridia; each of these "orders"
was divided into one dioecious and one monoecious group).
The order Laboulbeniineae, consisting of a single family, was
established by Lindau (1897a). The following year, Engler (1898)
established the class Laboulbeniomycetes; other classes were Ascomy-
cetes, Basidiomycetes, and Phycomycetes. The class included only one
order, the Laboulbeniales. A similar classification was published by
Schaffner (1909), who established the class Laboulbenieae in the
phylum Nematophyta, together with the Ascomycetae (Nematophyta
included fungi and algae). In 1908, Thaxter elevated the two
"orders" in the Endogenae to family status, created two suborders (the
Laboulbeniineae and the Ceratomycetineae), and introduced several
tribes within his key to the genera. See Table I.
Thaxter (1924) placed some genera with compound antheridia in the
Dimorphomyceteae, which he called a family. A Latin diagnosis for the
family Ceratomycetaceae was published in 1934 by Colla; she
considered the Laboulbeniaceae to consist of two families — the
Laboulbeniaceae homothallicae and the Laboulbeniaceae hetero-
thallicae. She placed dioecious genera with compound antheridia in the
Dimorphomycetaceae. Nannizzi (1934) established the family Zodio-
mycetaceae, which like the Ceratomycetaceae, included the genera with
exogenous spermatia.
The Laboulbeniales were included in the subclass Euascomycetidae
by Martin (1961) in his key to the families of fungi. However, Ains-
worth (1973) followed Engler's example in regarding the Laboulbeniales
as a separate class, but in his classification, the class Ascomycetes was
elevated to the subdivision Ascomycotina (cf. the inclusion by Alexo-

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MYCOLOGIA MEMOIR NO. 9
TABLE I
COMPARISON OF THE PRESENT CLASSIFICATION
WITH THAXTER'S CLASSIFICATION (1908)
THAXTER'S CLASSIFICATION PRESENT CLASSIFICATION
Laboulbeniineae
Peyritschiellaceae
Dimorphomyceteae
Rickieae
Peyritschielleae
Enarthromyceteae
Monoicomyceteae
Haplomyceteae
Laboulbeniaceae
Herpomyceteae
Amorphomyceteae
Stigmatomyceteae
Idiomyceteae
Teratomyceteae
Corethro my ceteae
Laboulbenieae
Rhachomyceteae
Clematomyceteae
Compsomyceteae
Chaetomyceteae
Ecteinomyceteae
Misgomyceteae
Ceratomycetineae
Ceratomyceteae (Thaxter
omitted the family,
Ceratomycetaceae)
Zodiomyceteae
suborder (but including some other taxa)
subfamily in Laboulbeniaceae
tribe in Peyritschielloideae
put with Peyritschiellinae, the subtribe
tribe (Chitonomyces and Hydraeomyces removed)
subtribe in Peyritschielleae with two new sub-
tribes, Diandromycetinae and Mimeomycetinae
subfamily in Laboulbeniaceae
tribe in Peyritschielloideae, with Polyascomyces
removed; new subtribe Kleidiomycetinae
family
suborder HERPOMYCETINEAE
subtribe of Laboulbenieae (Smeringomyces
removed to subtribe of Teratomyceteae)
subtribe of Laboulbenieae
put into Teratomyceteae
tribe, seven new subtribes added: Scelophoromyce-
tinae, Filariomycetinae, Smeringomycetinae, Asa-
phomycetinae, Rhipidiomycetinae, Amphimy-
cetinae, Histeridomycetinae
put into Stigmatomycetinae
subfamily, tribe, with two new subtribes
added: Chitonomycetinae, Chaetarthriomycetinae
subtribe in Teratomyceteae
put with Compsomyceteae
tribe, one new subtribe added: Kainomycetinae
subtribe in Teratomyceteae
put into Laboulbeniinae
subtribe of Laboulbenieae
Three new tribes added to Laboulbenioideae:
Euphoriomyceteae (none of genera were in
Thaxter's key [1908]; includes Aporomycetinae);
Hydrophilomyceteae (Hydrophilomyces was in
Ceratomyceteae); Coreomyceteae (was tribe in
Ceratomycetineae)
family now in suborder Laboulbeniineae
tribe (Euceratomyces later removed from
Ceratomyces by Thaxter [1931] — now is in
separate family EUCERATOMYCETACEAE)
(Hydrophilomyces removed to new tribe in
Laboulbenioideae)
family Ceratomycetaceae now includes two
subfamilies: Tettigomycetoideae, Ceratomycetoideae.
Ceratomycetoideae has a new tribe Drepanomyce-
teae; Ceratomyceteae has a new subtribe
Helodiomycetinae
subfamily in Laboulbeniaceae; Euzodiomyces now
in Euceratomycetaceae, Kainomyces now in sub-
tribe of Compsomyceteae
TAVARES: CLASSIFICATION, DIAGNOSES
93
poulos and Mims [1979] of subclass Laboulbeniomycetidae in the
Ascomycetes). The Pyrenomycetes were also included by Ainsworth
(1973) as a class in the Ascomycotina, together with the Loculoasco-
mycetes, Plectomycetes, Hemiascomycetes, and Discomycetes. Aside
from the habitat, the characters that distinguished the Laboulbeniomy-
cetes from all other Ascomycetes with unitunicate asci were not made
clear. Their basic structure does not seem to warrant such a separation.
It is probable that some of the taxa included in the Pyrenomycetes are
more distantly related to one another than they are to the Laboulbeni-
ales.
The classification proposed by Locquin (1974) seems to be based
primarily on erroneous interpretation of the literature. The subdivision
Cladomycotina was established, which included three separate classes.
One class, the Rhynchophoromycetes, was segregated on the basis of a
supposed coenocytic pseudotheciutti, containing indistinct spores (the
appendage characters were also considered). Another class, the
Futuromycetes, included only the order Amorphomycetales. This class
was distinguished by the supposed one-celled spores, unequal in size, as
well as by "zoospores" formed in the phialides.
Separation into families on the basis of a single character — the
manner in which spermatia are formed — has failed to provide a sound
classification; this is shown by the presence of similar antheridia in
genera in which the formation of the perithecium is entirely different.
Perithecial initiation is a characteristic that is little influenced by
environment.
Classification, Diagnoses, and Comparisons
Ordo Laboulbeniales Lindau in Englerand Prantl, 1897a, p. 491.
Subordo Herpomycetineae (Thaxt.) I. Tavares, 1981b, p. 469.
subordo Herpomycetineae (Thaxt.) I. Tavares, 1980, p. 485 (nom. nud.).
Famiiia Herpomycetaceae (Thaxt.) I. Tavares, 1981b, p. 469.
tribus Herpomyceteae Thaxter, 1908, p. 237.
Subordo Laboulbeniineae Thaxter, 1908, p. 236.
Famiiia Ceratomycetaceae Colla, 1934, p. 134.
subordo Ceratomycetineae Thaxter, 1908, p. 239 (see Cooke and Hawksworth,
1970).
famiiia Ceratomycetaceae Maire, 1916a, p. 32 (nom. nud.).

94
MYCOLOGIA MEMOIR NO. 9
Subfamilia Tettigomycetoideae nom. nov. Antheridium ex mole
cellularum numerosarum parvarum congestarum constans, in
latere interiore axis primarii thalli admodum supra perithecium
situm. Perithecii paries exterior ex seriebus quattuor verticalibus
cellularum constans, serie quaque ex cellulis numerosis brevibus
applanatis altitudine subaequis composita, cellulis serierum
omnium altitudine similibus. Perithecii cellula pedicellaris et cellula
pedicellaris secondaria sunt cellulae superpositae axis primarii
thalli. Trichogyna supra perithecium in angulo inter perithecii
basim et receptaculum superius emergens. Perithecium admodum
infra apicem abrupte angustatum. Typus: Tettigomyces Thaxter,
1915, p. 20.
Subfamilia Ceratomycetoideae
Tribus Thaumasiomyceteae nom. nov. Perithecii paries exterior
ex seriebus quattuor verticalibus cellularum, serie quaque ex
cellulis numerosis brevibus altitudine subaequis composita. Tricho-
gynae vestigium apice seriei horizontalis tertiae cellularum parie-
tis exterioris emergens. Axis thalli admodum supra perithecii cel-
lulam pedicellarem et cellulam pedicellarem secondariam, quae
cellulae superpositae sunt axis primarii thalli, discedens, fasci-
culum densum appendicum, quae cellulas amplificatas interdum
gerunt faciens. Pes niger typicus nullus, thalli basi quasi in
pedem dilatata vel ramificante. Typus: Thaumasiomyces
Thaxter, 1931, p. 332.
Tribus Ceratomyceteae Thaxter, 1908, p. 239.
Subtribus Helodiomycetinae nom. nov. Perithecium latum, ra-
mos laterales in serie verticali in latere exteriore partis inferioris
perithecii gerens. Perithecii paries exterior ex seriebus quattuor
verticalibus cellularum, serie quaque ex cellulis numerosis
brevibus altitudine subaequis composita. Cellula pedicellaris et
cellula pedicellaris secondaria perithecii sunt cellulae
superpositae axis primarii thalli. Ostiolum perithecii processibus
erectis filamentosis cinctum. Typus: Helodiomyces Picard,
1913b, p. 557.
Subtribus Ceratomycetinae
Tribus Drepanomyceteae nom. nov. Perithecium longum, an-
gustum; parietes cellularum parietis exterioris perithecii tam
macri ut maturitate non facile discerneri possint. Trichogyna
apice seriei horizontalis secundae cellularum parietis exterioris
emergens. Perithecii cellula pedicellaris et cellula pedicellaris
secondaria sunt cellulae superpositae axis thalli. Spermatia
lateraliter extrusa ex antheridiis quae cellulae intercalares appen-
dicis non speciales sunt vel antheridia ignota. Typus: Drepano-
myces Thaxter, 1931, p. 339.
TAVARES: CLASSIFICATION, DIAGNOSES
95
Familia Euceratomycetaceae I. Tavares, 1980, p. 488.
Familia Laboulbeniaceae Peyr., 1873, p. 246.
Subfamilia Zodiomycetoideae (Thaxt.) stat. nov.
tribus Zodiomyceteae Thaxter, 1908, p. 239 (listed as subfamily by Cooke and
Hawksworth, 1970).
familia Zodiomycetaceae (Thaxt.) Nannizzi, 1934, p. 263 (see also Colla, 1934,
p. 134).
Subfamilia Laboulbenioideae
Tribus Compsomyceteae Thaxter, 1908, p. 239.
Subtribus Compsomycetinae
Subtribus Kainomycetinae nom. nov. Perithecium stipiti uni-
seriato multicellulari appendices laterales carenti insidens
terminate, stipite in axe primario ipso thalli portato. Perithecii
paries exterior ex seriebus quattuor verticalibus cellularum
constans, serie quaque ex cellulis brevibus quinque vel pluribus
altitudine subaequis composita. Phialides in latere interior
axis primarii thalli productae aut cellulae parvae (forsitan
antheridiales) in latere interiore axis primarii thalli admodum
supra perithecium productae. Typus: Kainomyces Thaxter,
1901b, p. 44.
Tribus Hydrophilomyceteae nom. nov. Cellula accessoria elon-
gata in latere perithecii exteriore ad cellulas altas seriei ventralis
parallela interdum cornus lateralis instar; paries perithecii
exterior ex quattuor seriebus cellularum horizontalibus altitudine
inaequis constans supra seriem infimam ex quattuor ut videtur
non tribus (ut solet) cellulis basalibus constantem. Typus: Hy-
drophilomyces Thaxter, 1908, p. 431.
Tribus Coreomyceteae Thaxter, 1908, p. 239.
Tribus Teratomyceteae Thaxter, 1908, p. 238.
Subtribus Teratomycetinae
Subtribus Rhachomycetinae (Thaxt.) stat. nov.
tribus Rhachomyceteae Thaxter, 1908, p. 239.
Subtribus Chaetomycetinae (Thaxt.) stat. nov.
tribus Chaetomyceteae Thaxter, 1908, p. 239.
Subtribus Filariomycetinae nom. nov. Thallus elongatus,
gracilis, uniseriatus. Perithecii cellula pedicellaris in axe
primario lateralis. Perithecii parietes exteriores ex seriebus
quattuor verticalibus cellularum, serie quaque ex cellulis
altitudine distincte inaequis composita. Antheridia sunt phialides
in appendicibus brevibus lateralibus secus axem primarium
thalli supra infraque perithecia portatae. Septa crassa nigra
constricta nulla. Typus: Filariomyces Shanor, 1952, p. 499.

MYCOLOGIA MEMOIR NO. 9
Subtribus Smeringomycetinae nom. nov. Receptaculum breve,
angustum; receptaculum inferius infra perithecium ex cellulis
plus quam duabus constans; septa verticaliter singula vel plura
in receptaculo inferiore praesentia aut grex cellularum parvus
ad planitiem cellulae pedicellaris perithecii repertus.
Perithecium, appendix primaria, et appendices secondariae sin-
gulae vel plures ad apicem receptaculi aggregata; appendicum
bases cum cellula pedicellari brevi perithecii fere complanae.
Perithecium ab receptaculo superiore et appendicibus late-
raliter liberum. Appendices simplices vel sparse ramosae,
phialides ad apices portantes. Typus: Smeringomyces Thaxter,
1908, p. 296.
Subtribus Scelophoromycetinae nom. nov. Perithecia axibus
secondariis uniseriatis multicellularibus non ramosis in utro-
que latere axis primarii terminaliter vel axe primario aut axibus
secondariis lateraliter insidentia. Processus rhizoidales
deorsum crescentes ad thalli basin adsunt, nonnunquam
axibus secondariis non ramosis insidentes. Antheridia sunt
phialides in appendicibus lateralibus brevibus in axe primario vel
in axibus secondariis portatae. Perithecii paries exterior ex
seriebus quattuor verticalibus cellularum constans, serie qua-
que ex cellulis altitudine distincte inaequis composita. Typus:
Scelophoromyces Thaxter, 1912a, p. 210.
Subtribus Histeridomycetinae nom. nov. Thalli axis primarius
axes uniseriatos secondarios laterales singulos vel plures ex
cellula suprabasali producens, axe quoque perithecia singula
vel plura praebente. Perithecium quidque stipiti uniseriato
multicellular! cellulam pedicellarem perithecialem sustinenti
insidens terminale, stipite appendices antheridiales uniseriatas
multicellulars singulas vel plures basi gerente. Antheridia
phialides terminales sunt vel cellulae intercalares prope apices
appendicum antheridialium. Typus: Histeridomyces Thaxter,
1931, p. 181.
Subtribus Rhipidiomycetinae nom. nov. Receptaculum latum,
multiseriatum aut biseriatum, ex cellulis verticaliter elongatis
vel cubicis constans. Perithecii paries exterior ex seriebus
quattuor verticalibus cellularum, serie quaque ex cellulis altitudine
distincte inaequis composita. Antheridia sunt phialides in mar-
gine exteriore receptaculi partim inclusae. Septa crassa nigra
constricta nulla. Typus: Rhipidiomyces Thaxter, 1926, p. 509.
Subtribus Amphimycetinae nom. nov. Receptaculum magnum,
multiseriatum. Perithecii paries exterior ex seriebus quattuor
verticalibus cellularum, serie quaque ex cellulis altitudine
TAVARES: CLASSIFICATION, DIAGNOSES
97
distincte inaequis composita. Antheridia sunt phialides
terminales vel cellulae intercalares appendicum lateralium
receptaculo infra perithecium insidentes, forsitan etiam
appendicibus supra perithecium insidentia. Septa crassa nigra
constricta nulla. Typus: Amphimyces Thaxter, 1931, p. 305.
Subtribus Asaphomycetinae nom. nov. Monoeciae. Perithecii
paries ex seriebus quattuor verticalibus cellularum constans,
serie quaque ex cellula una infima altissima et cellulis tribus
(vel duabus) superioribus brevibus composita; cellula infima
quam quaevis cellula superior 6-plo altior. Perithecium
longum, angustum. Cellulis una duabusve quae parietes cras-
sos assequantur exceptis, parietes cellularum basalium
perithecii et cellulae pedicellaris secondariae tenuissimi manentes,
maturitate itaque invisibles. Antheridia sunt phialides vel
cellulae intercalares appendici brevi supra perithecium
insidentes; cellula pedicellaris perithecialis cellula lateraliter ex
una cellularum superpositarum receptaculi producta subtenta.
Perithecia plus quam unum adesse possunt, appendicibus inter
perithecia. Typus: Asaphomyces Thaxter, 1931, p. 310.
Tribus Laboulbenieae Thaxter, 1908, p. 238.
Subtribus Laboulbeniinae
Subtribus Misgomycetinae (Thaxt.) stat. nov.
tribus Misgomyceteae Thaxter, 1908, p. 239.
Subtribus Chitonomycetinae nom. nov. Appendix primaria bi-
cellularis, cellulis septo crasso nigro constricto separatis.
Antheridia (si adsunt) simplicia, ampulliformia, vel in margine
receptaculi superioris inclusa vel externe in receptaculo
superiore supra perithecium portata; antheridium septo crasso
nigro constricto superatum. Receptaculum uniseriatum infra
perithecium vel septum unum verticale diagonaleve praebens.
Typus: Chitonomyces Peyritsch, 1873, p. 250.
Subtribus Chaetarthriomycetinae nom. nov. Receptaculum
inferius infra perithecium ex cellulis duabus vel pluribus
constans. Thalli axis uniseriatus, non ramosus, phialides sessiles
vel partim inclusas supra perithecium lateraliter portans.
Cellulae parvae ab angulis superioribus cellularum in parte
superiore thalli interdum divisae. Perithecii paries exterior ex
seriebus quattuor verticalibus cellularum, serie quaque ex
cellulis altitudine distincte inaequis composita; cellula acces-
soria nulla. Typus: Chaetarthriomyces Thaxter, 1931, p. 319.
Subtribus Stigmatomycetinae (Thaxt.) stat. nov.
tribus Stigmatomyceteae Thaxter, 1908, p. 237.

98
MYCOLOGIA MEMOIR NO. 9
Subtribus Amorphomycetinae (Thaxt.) stat. nov.
tribus Amorphomyceteae Thaxter, 1908, p. 237.
Tribus Euphoriomyceteae nom. nov. Perithecii paries exterior ex
seriebus quattuor verticalibus cellularum constans, serie quaque
ex cellula una infima longissima et cellulis brevibus duabus tri-
busve superioribus composita. Parietes cellularum basalium
perithecii et cellulae pedicellaris secondariae tenuissimi ma-
nentes, maturitate itaque invisibiles. Trichogyna terminalis
manet vel paries perithecii exterior ex seriebus tribus cellularum
ternarum altitudine inaequarum contiguis verticalibus et ex serie
una cellularum quattuor constantur, cellula quarta apicaliter
protrudente. Monoeciae vel dioeciae; in plantis dioeciis,
perithecii cellula pedicellaris brevis et cellula basalis receptaculi non
se dividens. Antheridia sunt phialides. Typus: Euphoriomyces
Thaxter, 1931, p. 307.
Subtribus Euphoriomycetinae. Monoeciae vel dioeciae. Paries
perithecii exterior ex seriebus tribus cellularum ternarum
altitudine inaequarum contiguis verticalibus et ex serie una
cellularum quattuor constantur, cellula quarta apicaliter
protrudente.
Subtribus Aporomycetinae nom. nov. Dioeciae vel monoeciae.
Masculae simplices. Femineae vel hermaphroditae: parietes
cellularum basalium perithecii et cellulae pedicellaris
secondariae tenuissimi manentes, maturitate itaque invisibiles; ob
incrementum partis inferioris perithecii partes receptaculi
superiores et inferiores discedentes, maturitate itaque
appendix primaria ad latus perithecii affigeri visa; perithecii apex
non praeteriens sursum supra basim trichogynae, quae
maturitate terminalis manet. Typus: Aporomyces Thaxter, 1931,
p. 74.
Subfamilia Peyritschielloideae (Thaxt.) stat. nov.
familia Peyritschiellaceae Thaxter, 1908, p. 236.
Tribus Peyritschielleae Thaxter, 1908, p. 236.
Subtribus Peyritschiellinae
Subtribus Mimeomycetinae nom. nov. Antheridia sunt
phialides rami appendicis primaria insidentes terminales, interdum
antheridiis quoque multiplicibus ampulliformibus in axe pri-
mario thalli admodum supra perithecium portatis. Appendix
primaria ramum unum vel ramos plures ex latere interiore
cuiusque cellulae superpositae producens. Receptaculum in-
ferius plerumque fuscatum, ex cellulis superpositis duabus
perithecium subtendentibus constans, cellula basali longa,
TAVARES: CLASSIFICATION, DIAGNOSES
99
obconica, cellula suprabasali compressa, lata. Cellula
receptaculi suprema plerumque in cellulas superpositas duas se
dividens, harum superiore appendicem lateralem vel antheridium
complex fere semper producente. Perithecii paries exterior ex
seriebus quattuor verticalibus cellularum, serie quaque ex
cellulis quinque altitudine distincte inaequis composita.
Perithecium ab appendicibus et receptaculo lateraliter liberum.
Typus: Mimeomyces Thaxter, 1912a, p. 163.
Subtribus Enarthromycetinae (Thaxt.) stat. nov.
tribus Enarthromyceteae Thaxter, 1908, p. 236.
Subtribus Diandromycetinae nom. nov. Antheridia complicia,
ampulliformia, admodum infra perithecium portata. Septa
nigra nulla. Appendix primaria brevis, simplex. Typus: Di-
andromyces Thaxter, 1918b, p. 208.
Tribus DimorphomyceteaeThaxt., 1908, p. 236.
familia Dimorphomyceteae Colla, 1934, p; 50 (Thaxter, 1924, p. 315, referred
to the tribe as a family)
Tribus Haplomyceteae Thaxter, 1908, p. 237.
Subtribus Haplomycetinae.
Subtribus Kleidiomycetinae nom. nov. Monoeciae.
Antheridium complex, intercalare in appendice secondaria
cellulae suprabasali indivisae thalli insidente, cum peritheciis
cellulis pedicellaribus longis praeditis; cellulae fertiles antheridii
spermatia poro laterali emittentes. Typus: Kleidiomyces
Thaxter, 1908, p. 280.
Subfamilia Monoicomycetoideae (Thaxt.) stat. nov.
tribus Monoicomyceteae Thaxter, 1908, p. 236.
The characteristics of the taxa above the rank of genus are indicated
in the key. Characters occurring throughout the order are discussed in
sections on parts of the thallus. See Tables I-III.
Although the position of stalk and secondary stalk cells is not always
easy to determine, the presence of numerous, short perithecial wall cells
often indicates that the taxon belongs to the Ceratomycetaceae or the
Euceratomycetaceae. Careful focussing will reveal the angle at which
the wall cell rows emerge from the underlying cells and the direction of
their growth with respect to the axis of the carpogonium, enabling
recognition of the Ceratomyceteae. The stalk appendage cells can be
traced in the Euceratomycetaceae.
Both the antheridial and the sterile appendages in the Chitonomy-
cetinae have thick black septa. The subtribe is readily distinguishable

SUMMARY OF CHARACTERISTICS OF FAMILIES AND SUBFAMILIES OF THE LABOULENIALES
FAMILY, SUBFAMILY
EUCERATOMYCETACEAE
CERATOMYCETACEAE
Tribes separated in Cer-
atomycetoideae by
thickness of perithecial cell
walls, trichogyne posir-
tion, branching of primary
axis; subtribes by
perithecial structure
LABOULBENIACEAE
Primitive to advanced
ZODIOMYCETOIDEAE
Distinguished by
position of perithecia
LABOULBEN10IDEAE
Subtribes based on
receptacle structure, position
of antheridia and
appendages; dioecism, short
appendages characterize
Amorphomycetinae; sub-
tribes in Euphoriomyce-
teae based on perithecial
differences
PEYRITSCHIELLOIDEAE
Subtribes based on form
of receptacle and
position of antheridia,
which become septate
(compound)
MONOICOMYCETOIDEAE
Antheridial cells
from surface, extend
into area between cells
OUTER WALL CELLS OF PERITHECIUM
More or less equal in height;
no modification except reduction
in number and change in cell
shape in Colonomyces
Little difference in height of
outer wall cells in vertical
row; advanced genera may have
different heights in
alternating rows; increased wall cell
number in Rhynchophoromyces,
Thaumasiomyceteae. Thin cell
walls in Drepanomyceteae
Heights of outer wall cells in
vertical rows equal to unequal
Reduced cell wall thickness;
outer cells equal in heigth;
unusual outgrowths
Reduction in wall cell number;
cell heights equal in Compsomy-
ceteae, rarely so in other
tribes. In Euphoriomyceteae,
basal cell walls fail to thicken
'(partial tendency in Rhipidiomy-
cetinae, Asaphomycetinae, Amphi-
mycetinae)
Reduced cell number in vertical
rows, strongly unequal wall
cell heights except in Mimeo-
mycetinae. Failure of cell
walls to thicken in Dimor-
phomyceteae
As in Haplomyceteae
ANTHERIDIA
Simple phial-
ides or sper-
matia
laterally borne
Exogenous
spermatia if
known in Cer-
atomycetoideae ;
compound
anther idium in
Tettigomycetoi-
deae
Exogenous to
compound
Exogenous
(terminal)
Laterally
borne
spermatia in Compso-
myceteae, Cor-
eomyceteae,
some Stigmato-
mycetinae.
Primitive
compound antheri-
dium in Mis-
gomycetinae
Flask-shaped
in Peyritschi-
elleae;
various, on
secondary or primary
appendages in
Haplomyceteae
Flask-shaped
or paired
fertile cells
RECEPTACLE
Minor generic
differences.
Primary axis only
Primary axis only
(except secondary
in
Drepanomyceteae upon loss of
primary). Mostly
horizontal septa
Primary and
secondary axes
Many vertical
septa; internal
core
Primary and
secondary axes, many-
celled to 3-celled
receptacle (2 in
Amorphomyces)
Secondary axes in
some Peyritschiel-
leae, Dimorphomy-
eteae; 3-celled
in Haplomycetinae;
few to
many-celled in Pey-
ritschielleae
Secondary axes;
short primary
axis
GENERA SEPARATED ON BASIS OF
Extent of appendages; shape of
wall cells; number of perithecia;
vertical septa in receptacle;
antheridial arrangement
In Ceratomycetinae separated on
basis of number of cells
subtending perithecium
(segregating closely related genera) or
on kind of vertical wall cell
rows {Synaptomyces and Eusynap-
tomyces distinguished by
perithecial outgrowths)
Tribes based on equal vs. unequal
wall tiers combined with number
of receptacle cells; extra lateral
cell on perithecium; internal
perithecium; lack of cell wall
thickening; antheridia above or below
perithecium, combined with number
of receptacle cells. Genera based
on arrangement of receptacle cells,
antheridia, appendages, number of
wall cell tiers
Genera based on cell arrangement
in antheridium in Haplomycetinae;
on presence or absence of
secondary appendages in Haplomyceteae;
on receptacle in Peyritschielleae
and Dimorphomyceteae
Complexity of antheridia
TABLE III
BASES FOR SEPARATION INTO TAX0N0MIC CATEGORIES IN THE LABOULBENIALES
SPECIES
APICAL Some genera in Stig-
OUTGROWTHS matomycetinae, Lab-
OF PERI- oulbeniinae, Cerato-
THECIUM mycetinae; Smevingo-
myoee
APPENDAGE
STRUCTURE
Separates species
in Ldboulbenia
GENUS
Separates
Synaptomyoes,
Eusynaptomyaes
APPENDAGE POSITION Segregates Ormomyces in
Laboulbeniinae, Peyer-im-
hoffi-ella, etc., in
Stigmatomycetinae
ANTHERIDIUM TYPE
Separates genera of
Haplomyceteae;
Botryandromyoes
MANNER OF THALLUS DEVELOPMENT
POSITION OF STALK AND SECONDARY
STALK CELL WITH RELATION TO
PRIMARY AXIS
SUBTRIBE
Segregates Chitonomycetinae,
Chaetarthriomycetinae;
Peyrit sch iellinae
Separates Misgomycetinae
in Laboulbenieae
Zodiomycetoideae
Families in
Laboulbeni-
ineae
MANNER IN WHICH PERITHECIAL WALLS
FORM WITH REGARD TO CARPOGONIAL
CELL ROW
TRIBE
Monoicomyceteae
Haplomyceteae
distinguished
Distinguishes
the suborders
SUBFAMILY
Peyritschielloi-
deae, Tetti-
gomycetoideae
Ceratomyceta-
ceae generally
di st ingu i she d
RELATIVE HEIGHT
OF PERITHECIAL
OUTER WALL CELLS
IN VERTICAL ROW
Cryptandromyoee,
Peyein-mhoff-Ce I la
distinguished
Mimeomycetinae cells more
subequal than those in
other tribes of
Peyritschielleae
Separate Compso-
myceteae, Tera-
tomyceteae
generally
NUMBER OF OUTER
WALL CELLS IN
PERITHECIUM
Separates many in Laboulbeniinae,
Stigmatomycetinae; Colonomyces
Separates Aporomycetinae from
Euphoriomycetinae
TRIBE
All in Eucera-
tomycetaceae,
Ceratomyceta-
ceae — equal
Separates Euphoriomyceteae from
Teratomyceteae, Compsomyceteae from
other Laboulbeniaceae
THICKENING OF
CELL WALLS IN
PERITHECIUM
Separate Asaphomycesy
Dermapteromyces
Distinguishes Drepanomyceteae;
separate some subtribes in
Teratomyceteae
Distinguishes Euphoriomyceteae (La-
boulbenioideae), Dimorphomyceteae
(Peyritschielloideae)
SPECIAL
MODIFICATION OF
PERITHECIAL WALLS
Separates some genera in
Ceratomycetinae
Separates Helodiomycetinae
from Ceratomycetinae
Separates Coreomyceteae, Hydrophilo-
myceteae (Laboulbenioideae)
RECEPTACLE
CHARACTERS
Used in Ceratomycetinae,
Laboulbeniinae , Stigmatomycetinae
Used in Compsomyceteae,
Laboulbenieae , Teratomyceteae,
Peyritschielleae, Haplomyceteae
Separate Teratomyceteae, Laboulbenieae
MANNER APPENDAGES
EMERGE FROM
RECEPTACLE
Separates Rhynchophoromyces in
Ceratomycetinae, genera in Hydro-
philomyceteae, Laboulbenioideae
Separates Thaumasiomyceteae
POSITION OF
ANTHERIDIA
Separates Mimeomycetinae in
Peyritschielleae, Kleidiomy-
cetinae from Haplomycetinae
Teratomyceteae mostly distinguished
from Laboulbenieae, Haplomyceteae
from others

102
MYCOLOGIA MEMOIR NO. 9
from the Peyritschielleae by the uniseriate receptacle, although there
may be one diagonal or vertical septum at some levels.
The absence of thick, black, constricted septa readily distinguishes
the Rhipidiomycetinae and the Amphimycetinae from the Pey-
ritschiellinae. Thalli of Rickia (Peyritschiellinae) may bear simple, sub-
sessile antheridia that are somewhat similar in appearance to those
occurring in the Rhipidiomycetinae.
The perithecia of the Asaphomycetinae are more strongly modified
than those of Benjaminella and Amphimyces, but less so than those of
the Euphoriomyceteae. Although the lowest tier of outer wall cells in
these taxa may be proportionally almost the same in height, a more
slender perithecium is characteristic of the Asaphomycetinae. Although
there may be only two cells visible between the perithecial stalk cell and
the wall cells in Dermapteromyces (Asaphomycetinae), as well as in
the Rhipidiomycetinae and Amphimycetinae, only one thick-walled cell
can be detected there in Asaphomyces. Although the receptacle of
Dermapteromyces, as well as its antheridia, may bear a resemblance
to those of Benjaminella (Rhipidiomycetinae), the number and
arrangement of receptacle cells is small and precise and there are distinctive
lateral branchlets on the appendage. The perithecia in the Euphorio-
mycetinae bear a resemblance to those of the Dimorphomyceteae;
however, the stalk cell tends to be shorter and there is no evidence of
division of cell /.
Taxa in the Mimeomycetinae having simple antheridia are difficult to
distinguish from some genera in the Laboulbenioideae, although they
have the characteristic thallus form of Mimeomyces. The perithecium is
always free laterally from the receptacle and appendages, as contrasted
to the adnate perithecia of Apatomyces and most species of Laboul-
benia.
Generic Key to the Laboulbeniales
(* = taxon not seen)
I. Dioecious, on cockroaches (Blattaria); spore becomes 4-celled primary axis (in some
species, two middle cells of female may subdivide); in females, and in male of Herpo-
myces ectobiae, one or more secondary receptacular axes grow out from suprabasal cell
and produce reproductive structures; perithecia with many short outer wall cells in each
vertical row, these cells equal in height; carpogonial upgrowth grows inward and
upward from one cell of first tier of outer wall cells; spores 8 in ascus, with median
septum; antheridia long phialides
HERPOMYCETINEAE — HERPOMYCETACEAE — Herpomyces
TAVARES: GENERIC KEY
103
P. Mostly monoecious; on various hosts (only one monoecious Laboulbenia known
from Blattaria); thallus form and antheridia varied; perithecial walls grow up around
carpogonial upgrowth (in Ceratomycetaceae having axillary trichogynes, walls grow
up at angle, so that trichophoric cell is embedded in perithecial wall just above
junction with upper receptacle); spores 4 in ascus (except 8 in some primitive species), with
septum usually near lower end in ascus; mostly without secondary receptacle
LABOULBENIINEAE
A. Stalk cell (VI) and secondary stalk cell (VII) of perithecium are consecutive
intercalary cells of primary axis (or principal axis in Drepanomyceteae) (outer wall cell
row on outer [anterior] side of perithecium grows up from VI or its sister cell VI' and
the other 3 rows arise from VII); perithecium with many short equal or subequal cells
in outer wall cell rows (in some genera, cells in different vertical rows are not same
height) CERATOMYCETACEAE
1. Many-celled compound antheridium usually present above perithecium on
primary axis; trichogyne axillary; usually long, mostly uniseriate receptacle; on
Gryllotalpa (Orthoptera, Gryllotalpidae — mole crickets); terrestrial
TETTIGOMYCETOIDEAE — Tettigomyces
V. Antheridia undifferentiated intercalary cells or terminal cells of appendage,
producing exogenous spermatia singly (antheridia not seen in all genera);
trichogyne axillary or emerging from interior of perithecium above the base; aquatic
or semiaquatic; chiefly on Hydrophilidae CERATOMYCETOIDEAE
2. Primary axis subdivided at level of perithecium, much-branched, with branches
similar in size to axis; appendages in dense cluster, some of them specialized,
with enlarged cells; receptacle short, developing secondary feet or flattened
attachment zone; typical black feet absent; perithecium has long neck; outer wall
cells are approximately equal in height, the tiers of cells numerous; trichogyne
emerges from internal carpogonium at upper end of third tier of outer wall
cells; on Amphiops THAUMASIOMYCETEAE — Thaumasiomyces
2'. Primary axis simple or bearing narrow branches laterally or terminally above
perithecium; no enlarged appendage cells; receptacle short or long; perithecium
has long neck only in Rhynchophoromyces; no secondary foot development. If
outer wall cells distinct, outer walls of perithecium grow up at angle to
carpogonial upgrowth, so that trichogyne is axillary or slightly higher (if perithecial
wall cells indistinct, cf. Aporomyces, 88).
3. Outer wall cells of perithecium distinct; trichogyne axillary or nearly so
CERATOMYCETEAE
4. Perithecium has 4 slender filaments surrounding ostiole; lateral branches
arise from lower outer wall cells on outer (anterior) side of perithecium;
extra short row of cells on inner (posterior) side of perithecium (probably
part of carpogonial cell row); on Dryopidae:.
HELODIOMYC'ETINAE — Helodiomyces
4'. Perithecium simple or having 1 or more subapical horns (or lateral
outgrowths on neck in some species of Rhynchophoromyces); trichogyne
axillary; on Hydrophilidae (one report on Dytiscidae).CERATOMYCETINAE
5. Perithecium with long, slender neck; inflated venter usually appearing
innate (attached laterally to primary axis above perithecial base) and
surmounted by cluster of slender appendage branches; wall cell tiers ca.
25 or more, with short cells equal in height in all 4 rows (note that R.
unguicola is reduced in size); receptacle rarely with as few as 3 cells (R.
minor Spegazzini); perithecium usually without outgrowths except on
neck Rhynchophoromyces

ORIGIN AND DISTRIBUTION
A Brief Outline of Paleontological History
Primitive wingless hexapods (insects in a broad sense) were already
well developed by mid-Devonian, as indicated by the remains of
Rhyniella (apparently a coUembolan), found in the Rhynie fossil beds in
Scotland (Jeannel, 1960). During the early part of the period, the psilo-
phytes, plants of comparatively small size, were the predominant land
vegetation. Forests of pteridosperms, lycopods, and horsetails
(sphenopsids) appeared in late Devonian (Stokes, 1960). Possibly fungi
attacked these plants, but present-day records give no indication of any
group, such as the fern-conifer rusts, that may have arisen on the
vascular plants of the Devonian. As Savile (1955) pointed out, early fungi,
being very primitive and poorly adapted, were unlikely to survive (see
also discussion of the position of the Laboulbeniales among fungi).
During the Upper Carboniferous period, abundant members of two
groups of insects lived in the warm humid forests of the equatorial belt
of Laurentia, the European-North American continent — one of these
groups included the extinct superorder Palaeodictyoptera, an extinct
order in the superorder Ephemeroptera (mayflies), and one in the super-
order Odonatoptera (dragonflies) — all in the section Palaeoptera
(insects having primitive wings that were never folded backward) (Jeannel,
1960). The other group included some extinct families of roaches in the
superorder Blattopteroidea, as well as families in the superorder
Orthopteroidea (locusts, crickets, and walking sticks or phasmids) — all
in the section Polyneoptera, possessing primitive wings that fold
backward and undergoing gradual metamorphosis (hemimetabolous or
heterometabolous). In the coal-forming forests, fairly large trees
flourished, belonging to the lycopods and sphenopsids among the
pteridophytes and to the primitive gymnosperms; smaller plants
occurred on the forest floor (Stokes, 1960). Blattids (roaches) lived in
vegetative debris (Jeannel, 1960). The records of fungi for this period
(Meschinelli in Saccardo, 1892; Wolf and Wolf, 1947) are inadequate.
However, there were structures that resemble hysterioid fungi on leaves,
which were probably ascomycetous fungi (see Magdefrau, 1956, Lang-
ford, 1958). The occurrence of such fungi during this period is certainly
351

352
MYCOLOGIA MEMOIR NO. 9
compatible both with the theories of Savile (1955) and Buller (1950) on
the concurrent development of Ascomycetes and higher plants and with
the existence of well-authenticated specimens from later periods
(Suzuki, 1910, Dilcher, 1965).
During the Carboniferous, there was considerable glaciation in
southern Gondwanaland, the large continent of the southern
hemisphere (Stokes, 1960) and it was presumably in the adjacent
temperate areas, such as southern Africa, that the holometabolous insects
and the hemimetabolous Homoptera evolved (Jeannel, 1960). Holo-
metaboly (complete metamorphosis with pupal stage) was an adaptation
to a temperate climate with cold winters (Jeannel, 1960). With the
coming of the Permian in Laurentia, conditions became unfavorable for
many organisms, which consequently became extinct. The climate
became cooler and more arid everywhere. Upland and desert types of
vegetation replaced the swamp vegetation of the Carboniferous period
(Stokes, 1960). Retreat of the Tethys Sea (the primitive Mediterranean)
permitted a great increase in northward movement of Gondwanian
insects along with the conifers, reptiles, and primitive ancestors of the
mammals (there had been limited migration across this sea during
earlier periods). Some Laurentian insect lines passed to Gondwanaland
(Jeannel, 1949b), where they evolved during the Mesozoic on the
fragments of Gondwanaland (see MacKerras, 1974, for times of
separation of segments of the continent in terms of millions of years).
Among the insects, some Paleozoic groups, such as the Palaeodicty-
optera, died out entirely; in others, new lines replaced the old. Of the
orders occurring in the Carboniferous, only the roaches (Blattaria;
suborder Blattodea of Dictyoptera according to Jeannel's classification,
1949b) remain. The present-day roaches belong to the recent families,
which are close to Mesozoic roaches (Jeannel, 1949b). The most
significant new Permian group was the Coleoptera (beetles), which suddenly
appeared in the lower Permian (Tshekardocoleus) in the Urals (Jeannel,
1960). Permocupes, with elytra venation very similar to that of modern
Cupedidae, has been found in Upper Permian Russian fossil beds
(Jeannel, 1949b). Several extinct beetle-like groups also existed (Brues et al.,
1954), in which the humeral callus permitting the elytra to be held rigid
as stabilizers during flight had not yet appeared; they also had primitive
wing venation (Jeannel, 1949b). Beetle-like fossils occur in sub-
antarctic Australian Upper Permian and Triassic beds, as well as in the
U.S.S.R. (Jeannel, 1942b; Riek, 1970). The immediate fossil
progenitors of the Coleoptera are not known and the origin of the order is
not clear. Coleoptera belong to the section Oligoneoptera, which are
holometabolous; they have reduced wing venation and wings that fold
back. Some members of the superorder Mecopteroidea, also in the
Oligoneoptera, appeared in the Permian, although the order with which we
TAVARES: PALEONTOLOGICAL HISTORY
353
are concerned — the Diptera — was first in evidence in Australia in the
Triassic (Jeannel, 1960). In the section Paraneoptera (hemimetabolous,
with wings folding backward and reduced venation), the order
Homoptera was abundant in the Permian. It migrated into Laurentia from
Gondwanaland together with the Oligoneoptera (Jeannel, 1949b). It
appears that evolution was rather rapid during this period.
Gryllidae (crickets) existed during the Triassic in South Africa and the
Jurassic in Europe and Turkestan (Jeannel, 1949b). During the Triassic,
Heteroptera (bugs) appeared, undoubtedly evolving from Homoptera;
the first fossils of the superorder Hymenopteroidea (Oligoneoptera),
which includes ants, and of Dermaptera (Dermapteroidea, Poly-
neoptera) did not appear until the Jurassic (Jeannel, 1960). All
insects of the Mesozoic except roaches belonged to Gondwanian lines
(Jeannel, 1949b). Probably a few of these (such as Ephemeroptera —
may flies) arose in Gondwanaland from Laurentian roots that migrated
before the Permian (Jeannel, 1960).
The predominant higher plants of the early Mesozoic were true ferns,
cycads, Ginkgo, and primitive conifers such as araucarias, pines, and
sequoias (Stokes, 1960). (Modern ferns are parasitized by the simple
Ascomycete Taphrina and primitive rusts having alternate stages on
Abies.)
In the Cretaceous, the angiosperms appeared and became the most
abundant group of plants; as they spread, they were accompanied by
flies and flower beetles (Buller, 1950). Angaran (Asian) lines spread
through the northern hemisphere in mid-Cretaceous (Jeannel, 1949b).
By the end of the period, Africa and South America had separated,
although there were probably residual land masses between the two
continents which facilitated migration of organisms for some time (see
Jeannel, 1942b). North America and Europe were still adjacent to one
another in the far north, permitting interchange of organisms.
The Tertiary was a period of extensive migration of plants and
animals. Both the land masses and the living organisms took on a modern
appearance. The higher plant vegetation was similar to that of the
Cretaceous. All of the insect groups had already appeared. Fossil
insects such as those in amber of the Oligocene are very close to modern
species (see Stokes, 1960; Jeannel, 1942b).

354
MYCOLOGIA MEMOIR NO. 9
Position of the Laboulbeniales among Fungi
Although the Laboulbeniales have prototunicate ascus walls (see
Alexopoulos and Mims, 1979), they may be considered to be primitive
members of the unitunicate line of Pyrenomycetes (see Luttrell, 1955).
They have simple, thin-walled asci with no apical pore and the ascocarps
have neither paraphyses nor pseudoparenchyma in the usual sense.
Inner wall cell rows arising from the basal cells of the perithecium can
be compared with paraphyses. Those that arise from the upper, inner
angles of outer wall cells are comparable to periphyses. The close
relationship of the Laboulbeniales to filamentous members of the
subclass Euascomycetidae was confirmed by Hill (1977), when he
demonstrated the presence in Herpomyces of flattened spindle pole bodies,
Woronin bodies associated with simple septal pores, and ascus vesicles.
Phialides occur not only in the Laboulbeniales, but they are found
sporadically at various evolutionary levels in the unitunicate
Pyrenomycetes. They may be present in the Helotiales among the orders having
true paraphyses and the Melanosporaceae among the families with the
Diaporthe type of centrum (Luttrell, 1955).
The centrum of the Laboulbeniales is comparable to that of the
Chaetomiaceae (having the Xylaria type of centrum), in which
paraphyses arise only from the lateral walls of the perithecium and the thin
walls of the asci are deliquescent. As in Laboulbenia, there is in the
Chaetomiaceae a true perithecial wall arising from cells underlying the
ascogonium (or from nearby vegetative hyphae [Luttrell, 1951]). In
Laboulbenia, however, there are no internal cells other than derivatives
of the carpogonial cell row (the equivalent of the ascogonium and
trichogyne of the Chaetomiaceae). In the Melanosporaceae (having the
Diaporthe type of centrum, in which asci expand into an
undifferentiated pseudoparenchymatous tissue), the asci have thin walls. Cory-
neliales, which have been reported to be ascostromatic (Luttrell, 1951),
also have thin-walled asci. It is possible that the ascostromatic,
unitunicate genera have evolved from an ancestral form (now extinct or not
yet recognized among living genera — see Savile, 1955, on the probable
nature of primitive fungi) in which deliquescing, thin-walled, clavate
asci developed within a true perithecial wall consisting of cells produced
by those subtending the ascogonium (cf. LuttrelPs discussions, 1951).
Although there is an ascostroma into which fertile cells grow in
Herpomyces, its structure is essentially the same as that of the true
perithecial wall of Laboulbenia, and at maturity it appears as if the
perithecial wall has developed from the stalk of the fertile portion as it does
in Laboulbenia. It is difficult to comprehend how the Laboulbeniales
could be polyphyletic, despite the basic difference in the pattern of
TAVARES: POSITION AMONG FUNGI
355
development. All genera have two-celled spores and perithecia
consisting of four rows of outer and probably four rows of inner wall cells
(although the latter may be limited to the neck). A consideration of the
difference in the development of the perithecial wall in the two
suborders may enable investigators to understand the relationships
between genera in other groups of Ascomycetes in which taxa with true
perithecial walls and those that are ascostromatic have similar centrum
development, ascus structure, and spores.
In the ancestor of the Laboulbeniales, the thallus may have developed
from a spore in the manner of Herpomyces ectobiae, the perithecium
being produced like that of Laboulbenia and having a random number
of wall cell rows growing out from the upper cells of the perithecium-
bearing branch. There may have been more than two wall cell layers; the
present inner wall layer may actually be the remnant of a rudimentary
system of periphyses or paraphyses (or both). The nature of the
upgrowths within the centrum in Herpomyces suggests that perhaps
there may have been a form of crozier in the ancestral organisms, at
least when certain conditions favored their development.
Many other Ascomycetes have two-celled spores; they include a large
percentage of marine Pyrenomycetes, as well as terrestrial genera in the
bitunicate and unitunicate series (Denison and Carroll, 1966). Chade-
faud (1960) suggested that the two-celled spore is the ancestral form.
Spathulospora, a parasite producing a perithecium on the exterior
surface of a red alga, was proposed as a connecting link between the
Laboulbeniales and the Pyrenomycetes (Kohlmeyer, 1973). Although
Spathulospora produces spermatia from phialides, lacks paraphyses,
and has a trichogyne and deliquescing, simple, unitunicate asci, its asco-
spores are one-celled and bear appendages. The spermatia, at least in
some species, bear a filiform, coiled appendage. It was believed that the
thallus of Spathulospora consisted of a compact surface structure with
peglike penetrating cells. However, Walker et al. (1979) have shown that
the perithecium is produced by hyphae that emerge from an internal
crust reminiscent of parts of lichen thalli.
Despite the differences between the two orders, the Laboulbeniales
and the Spathulosporales probably both arose from a group of primitive
Ascomycetes having undifferentiated spermatia and asci. There seems
to be no justification for placing them together in a separate class or
subclass unless other taxa with simple perithecial structure and
prototunicate asci are also included. Probably, the reduced thallus of the
Laboulbeniales is an adaptation to growth conditions. It is doubtful,
however, that their simple asci have resulted from the modification of
asci of more complex structure.

356
MYCOLOGIA MEMOIR NO. 9
The thalli of some ancestral Ascomycetes might have been lichenlike
in form, as suggested by Cain (1972), with the parasitic groups
undergoing some reduction in size and becoming more compact. Cain
postulated that the earliest Ascomycetes were autotrophic, with a discrete
haploid thallus bearing a parasitic dikaryotic stage; he suggested that
they colonized soil in a tropical, moist climate and formed the major
land flora. Some of these organisms became lichenized, whereas others
became epiphytic on vascular plants, which began to appear. Cain
believed that asci developed mechanisms for aerial dispersal in terrestrial
habitats.
Like Cain, Denison and Carroll (1966) proposed that the ancestors of
Ascomycetes were autotrophs, and they explained briefly, in their
reexamination of the question of red algal origin, how autotrophic
nutrition may account for the differences in food reserves and cell wall
composition between the Ascomycetes and the red algae. They also
suggested that extensive mycelial growth developed as bulky plant
tissues became available that could be penetrated for nutrients.
Demoulin (1975) believed that parasitism was the first step in
heterotrophic life taken by red algal ancestors of fungi, with saprophytism
evolving later from necrotrophic parasitism; he proposed that adelpho-
parasitic (i.e., parasites that are related to their hosts) red algae were the
first step in the development of the Ascomycetes. Kohlmeyer (1973) also
thought that the Ascomycetes developed from parasitic ancestors living
on algae.
Recent ultrastructural, biochemical, and developmental studies of red
algae support the concept of a common ancestry for the Rhodophyco-
phyta and the Ascomycetes, in the opinion of Demoulin (1975).
However, Hill (1977) pointed out differences in red algal pit connections and
spindle pole bodies. That the polar ring of Membranoptera (Ceramiales,
a highly evolved order) to which Hill referred (McDonald, 1972) is not
characteristic of all algae was shown clearly in illustrations of the
complex spindle pole body of Porphyridium (Bangiales, a more
primitive order) (Schornstein and Scott, 1980). The proximal portion of
the latter organelle somewhat resembles spindle pole bodies of
Ascomycetes (cf. Hill, 1977).
The concept of a common ancestry between Rhodophycophyta and
Ascomycetes is compatible with the theories of Cain (1972) and
Kohlmeyer (1973, 1975). Some heterotrophs (as suggested by Denison and
Carroll, 1966) may have become saprophytes on the increasingly
abundant terrestrial plant debris floating in the sea. Eventually, such
saprophytes would be deposited on the shore, where interrelationships
with terrestrial insects could be initiated. Possibly, association with
green algae to form littoral perithecial lichens also took place. If entry
TAVARES: PROPOSED PHYLOGENY
357
into the terrestrial environment has taken place more than once, by
fungi that have reached different evolutionary levels before emerging
from the sea, traces of these levels may be detectable by means of
anatomical and physiological comparisons.
Proposed Phytogeny and Host Relationships
of Ancestral Laboulbeniales
Herpomyces occurs on a primitive host, is phylogenetically isolated,
and its thallus morphology may represent the ancestral form in the
Laboulbeniales. However, the ascogonial development does not have a
counterpart among known genera of Ascomycetes. On the other hand,
the carpogenic cell row of Laboulbenia is similar to the ascogonium
and trichogyne of some other Ascomycetes. Thus, it might be proposed
that the ancestral form bore a resemblance in structure to a primitive
member of the Laboulbeniineae, although it possessed an extensive
thallus similar to that of Herpomyces ectobiae. Possibly, the
Herpomyces perithecium arose after abortion of the normal carpo-
gonium, with a secondary fertile branch from the side substituting for
it. Perhaps the primordial cell that divides into the first wall cells
represents a procarp that divided abnormally, instead of developing
into the carpogonial cell row. The subsequent growth of a substitute
ascogonial branch from one of the basal cells would have compensated
for this failure to develop normally (despite the normal precision of cell
division, there is considerable plasticity in the Laboulbeniales, which
is expressed in abnormal growth). (See Fig. 20.)
If it is assumed that the Laboulbeniineae are closest to the ancestral
form, it might also be supposed that the original host group was the
precursor of the true beetles (Coleoptera), parasitized perhaps during the
Carboniferous. However, beetles were thought by Jeannel (1942b) to
have arisen in Gondwanaland under the influence of cold temperature,
whereas roaches (of extinct groups) were abundant in the warm
northern continent Laurentia, where the climate presumably was more
favorable for primitive fungi. Precursors of beetles and roaches
presumably diverged in the Devonian (Jeannel, 1949b). It seems
unlikely that the fungi have evolved with both of these insect orders.
Probably one of the two groups was secondarily infected. The original
host could have belonged to another order, instead. Perhaps, as
Kohlmeyer (1973) suggested, precursors of the Laboulbeniales parasitized
marine arthropods. If this is true, it is doubtful that the particular
precision of cell numbers which characterizes this order arose before the
terrestrial environment was invaded; this would have been necessary
had the initial host been a common ancestor of both roaches and

358
MYCOLOGIA MEMOIR NO. 9
beetles. Although a reduction in size may have accompanied growth on
the integument, probably a drier environment and the physical nature of
the habitat were responsible for the development of a reduced thallus
resistant to desiccation.
An ancestor close to other primitive perithecial Ascomycetes may
have parasitized roaches in the tropics of Laurentia during the
Carboniferous, undergoing reduction in the number of perithecial wall cells. On
the other hand, debris in the developing coniferous forests of Gond-
wanaland may have harbored the progenitors of both the Laboulbeni-
ales and the beetles, particularly if the vegetation was already quite
varied (Kuschel, 1969, pointed out that slow decomposition of forest
litter in temperate climates provides a good habitat for beetles, although
this may not be true in a forest of uniform composition). During the
Permian there was an equalizing of temperatures and a general
exchange of flora and fauna (Jeannel, 1942b). Possibly, cross-infection
took place between roaches and beetles at that time.
Coleoptera appeared in large numbers in Europe during the
Jurassic, when that area became suitable for colonization. Presumably they
came from Angara Land (Jeannel, 1942b), where they were elaborated
from Gondwanaland roots. Meanwhile, roaches were evolving in
Gondwanaland that probably gave rise to Tertiary groups (Tillyard,
1919, 1926; Jeannel, 1960). Jeannel (1942b) based his conclusions
about the origin and distribution of beetles largely on present-day
distribution of primitive groups and on a study of routes taken by particular
genera as indicated by relationships between species; the knowledge we
now have about continental drift (see Hurley, 1968) shows that his
assumptions were quite well founded.
Savile (1955) proposed an evolutionary parallelism between higher
plants and rusts, whereas Buller (1950) made suggestions about the
possible relationship between the development of flies and fungi.
Probably, as the beetles evolved, so did the Laboulbeniineae. If cross-
infection between beetles and roaches took place during the Permian,
it might be postulated that as Herpomycetineae developed on
roaches, the Laboulbeniineae began to evolve on beetles during the
Triassic. The precise 1:3 arrangement of the outer wall cell rows of the
perithecium (one row from the stalk cell VI, with m as its basal cell,
and three rows from the secondary stalk cell VII, two arising from n and
one from n") must have developed after beetles became parasitized.
This arrangement is merely the result of the condensation of the hyphae
so that a lateral branch arises from each cell (see Fig. 7).
After this modification of the perithecium had taken place,
alterations appeared in the position of the stalk cells in the thallus.
Assuming that a certain amount of time was necessary for each stage of
TAVARES: PROPOSED PHYLOGENY
359
development, one may postulate that the perithecial position (on a
lateral branch) characteristic of the Euceratomycetaceae appeared in the
Jurassic and probably a little later, the Ceratomycetaceae, with its peri-
thecia borne directly on the primary axis, began to differentiate. The
Euceratomycetaceae were probably ancestral; it is difficult to
comprehend, however, how the stalk cells could have been transferred from
a lateral axis to the primary axis (or vice versa), because the orientation
with respect to the primary axis is different. The Ceratomycetaceae
seem to represent a strong departure from the probable ancestral form;
no perithecia are formed on the primary axis in Herpomyces. Perhaps
the ancestral form bore perithecia both on the primary axis and on
lateral axes, the primary perithecia being lost in the Euceratomycetaceae
and the lateral perithecia disappearing in the Ceratomycetaceae. The
Drepanomyceteae may have evolved separately from the
Euceratomycetaceae; if so, they should be placed in a separate family, rather than be
regarded as a subfamily in the Ceratomycetaceae.
Tettigomyces occurs on a primitive terrestrial host in the Orthoptera.
Although it has an entirely different kind of antheridium, its trichogyne
is axillary like those of the Ceratomycetinae, parasitic on aquatic
Coleoptera. This trichogyne position undoubtedly developed from the
more elevated one characteristic of the Euceratomycetaceae, Thau-
masiomyceteae, and Drepanomyceteae. Perhaps Tettigomyces diverged
from the Ceratomycetinae after the axillary trichogyne developed in
that subtribe. On the other hand, the axillary position in Tettigomyces
may have arisen independently, possibly because of the massive
character of the thallus. The discovery of closely related genera and the
development of a better understanding of thallus structure in the
Ceratomycetaceae might reveal whether the ancestral taxa were aquatic or
terrestrial.
Eventually the stalk appendage was lost in an ancestral member of the
Euceratomycetaceae and the Laboulbeniaceae began to differentiate
(see Fig. 21). Pseudoecteinomyces may represent a step in the
development of the Laboulbeniaceae. Interestingly, this genus occurs in
southeast Asia, which appears to be the source of many plants and animals.
The arrangement of the antheridia is reminiscent of that of Pselaphi-
domyces. Because the Jurassic was the period in which many modern
beetle families were first in evidence, it is possible that all of the major
groups of the Laboulbeniaceae arose then — just as it was apparently a
time of rapid development in the beetles, it also may have been among
their fungi. Apparent Caraboidea and Hydrophilidae (now host groups
of Cochliomyces and Thaumasiomyces — probably two of the most
primitive genera) are known from Jurassic deposits in Europe (Jeannel,
1942b); Elateridae (hosts today of Stemmatomyces and Laboulbenia)
and Nitidulidae (present hosts of Rickia and Aphanandromyces)
have also been reported.

360
MYCOLOGIA MEMOIR NO. 9
Possibly, ancestral forms of Cochliomyces and Thaumasiomyces
parasitized carabids and hydrophilids during the Jurassic.
Cochliomyces is closely related only to Pseudoecteinomyces, not to the other
genera of the Euceratomycetaceae, which occur on Staphylinidae and
Colonidae. Thaumasiomyces is isolated from other genera in the
Ceratomycetoideae.
The wide range of Euphoriomyces and the variety of form exhibited
in the genus suggest Cretaceous origin and Tertiary distribution, when
European temperatures were warmer (see section on distribution
patterns). Carpophoromyces, which, like Euphoriomyces, occurs in Sri
Lanka, is either an exceptionally well-developed variant of
Euphoriomyces, in which secondary receptacular axes have been formed, or it is a
primitive relict. The absence of secondary axes in Asaphomyces, which
may be considered ancestral to the Euphoriomyceteae, indicates that
these axes are probably a recent modification in Carpophoromyces.
(For characters of the perithecium and receptacle indicative of phylo-
genetic relationships, see section on parts of the thallus; see genera; cf.
Fig. 20, Tables II, III).
The Cretaceous was probably the period when all widespread genera
came into being, as well as small genera that have primitive characters
and are quite isolated. Genera such as Diplopodomyces and Fannio-
myces have undoubtedly evolved quite recently (see discussion of
geographical distribution).
Distribution of the Laboulbeniales
Introduction
Thalli of a species of the Laboulbeniales may be randomly dispersed
on the bodies of individual hosts or they may appear to be limited to a
single location (Benjamin [1971] has discussed in detail position
specificity and sex-of-host specificity). Often thalli that are anatomically
similar in many ways may differ in general form when they occur in
different positions on hosts of the same or different sexes. These thalli
may be described as different species if their position and the habits of
the hosts are not considered (see Scheloske, 1976a, 1976b). Thalli of
Eusynaptomyces benjaminii Scheloske (1976a) growing on claws of
males have short receptacles, whereas those growing on the underside of
the pronotum of the female have elongate receptacles and produce peri-
thecial outgrowths. Scheloske (1976b) doubted the existence of sex-of-
host specificity in the Laboulbeniales.
Strict position specificity of species of Chitonomyces on Orectochilus
(see Benjamin, 1971) might be altered by accidental infection. A new
TAVARES: DISTRIBUTION ON HOSTS
361
position of growth for a taxon could affect its morphology. A different
pattern of distribution of species might develop in a host population if
some species germinate in unusual places.
Whereas dispersal of thalli on individual hosts is related to present
living habits, geographical dispersion and distribution on different
host groups reflect the habitats and host interrelationships of the
past. Co-evolution of hosts and parasites has undoubtedly occurred,
although Frank (1982) believes that parasite-specific reactions by the
hosts should be evident.
Distribution on Host Groups
There are two basic patterns of distribution of Laboulbeniales on the
families and orders of hosts:
1. Unrelated genera of fungi parasitize a particular group of hosts;
this results from recurrent waves of infection. The Staphylinidae bear
genera from almost every tribe in the Laboulbeniales, with the exception
of the Ceratomyceteae, which are limited to aquatic beetles
(particularly Hydrophilidae), and the tribes consisting of only one genus, such
as Coreomyceteae.
2. Many closely related genera parasitize genera in a single family of
hosts, indicating diversification of a common ancestral form and
suggesting a certain degree of parallel evolution of hosts and
parasites — on Staphylinidae, there occur four genera in the
Haplomyceteae, five in the Teratomyceteae, and several in the Stig-
matomycetinae.
Two of the genera parasitizing Staphylinidae — Euceratomyces and
Euzodiomyces — may be considered relict genera. Other genera on
these beetles obviously have evolved more recently (for example, the
genera in the Haplomyceteae). Probably, the Staphylinidae have been
parasitized by Laboulbeniales at many different times since the Meso-
zoic. The Staphylinidae undoubtedly arose quite early, the absence of
an extensive fossil record probably being the result of their fragility
(Paulian, 1943). Their habits and habitats seem to be favorable for
parasitism by the Laboulbeniales, because this family bears the greatest
number of genera of all the families of hosts. This large family cannot
be compared with the numerous small families in this respect; however,
the large family Chrysomelidae is host only to Laboulbenia and Di-
meromyces, two of the ubiquitous genera that occur on almost all host
groups. By contrast, 47 genera have been reported on Staphylinidae (see
discussion, Frank, 1982). The other large group parasitized by several
genera is the Caraboidea (often called the Carabidae); only 16 genera
have been reported on these beetles.

362
MYCOLOGIA MEMOIR NO. 9
One of the largest families of beetles — the Scarabaeidae, with over
12,000 species — is parasitized only by Laboulbenia and Rickia.
Another large family, the Tenebrionidae, with over 14,000 species,
is parasitized by six genera, including Laboulbenia and Dimeromyces,
but Rickia has not been reported as a parasite.
Herpomyces obviously has evolved on Blattaria; it occurs on no other
order of hosts and is an isolated genus. The habitats of cockroaches in
nature are similar to those frequented by beetles and other arthropods;
despite this, the genus apparently has not cross-infected any other
group. On the other hand, Laboulbenia has been found on a roach in
West Africa, and probably other collections may be made on roaches
living under stones and in other natural habitats. Most collections of
Herpomyces have probably been obtained from taxa that live in and
around buildings. The stable environment of these roaches may be
conducive to the retention of an archaic parasite. Even among the
genera of Blattaria, it is unlikely that much cross-infection takes place
(see Richards and Smith, 1954). Roaches in all stages of development
frequently are found living together in large numbers. The heaviest
infections are usually found in crowded, old colonies.
On Isoptera, another primitive order of insects, only Laboulbenia
and Dimeromyces have been found. There appears to be no genus of
Laboulbeniales that has evolved with the Isoptera. Isoptera serve as
hosts for a number of ectoparasitic anamorphs (Blackwell and Kim-
brough, 1976). Two of the species of Laboulbenia on termites (L.
hagenii Thaxter and L. geminata Buchli, both from East Africa on Ter-
mitidae, Macrotermitinae) have profusely branched appendages and a
cubical cell VI, whereas the third species (L. felicis-caprae W. Rossi, on
Hodotermitidae from North Africa) has an appendage that is little-
branched and a flat cell VI. The two species of Dimeromyces belong to
the typical structural group. Termites in many areas should be examined
in order to determine whether evolution has taken place on the termites,
whether there are any distinctive distribution patterns, and what role
associated insects have in the dissemination of the fungi.
Gryllotalpa (Orthoptera) is parasitized by Tettigomyces, an isolated
genus most closely related to the Ceratomyceteae on aquatic beetles; it
is also parasitized by Rickia. Only Laboulbenia and Dimeromyces
are known to parasitize other Orthoptera (Gryllidae — crickets, which
probably encounter other insects more frequently than do the
burrowing mole crickets [Gryllotalpidae]). Hosts are likely to contact
unrelated species, even though the contacts may be of shorter duration
than those between conspecific individuals of opposite sex (Frank,
1982).
TAVARES: DISTRIBUTION ON HOSTS
363
The species of Laboulbenia on different groups of ants (Hymenop-
tera, Formicidae) are not closely related. Laboulbenia formicarum
occurs only in North America, where it is widespread on Formicinae
(Formica, Lasius, Myrmecocystus, Prenolepis, and probably Poly-
ergus). The appendages in this species have abundant constricted black
septa. The Brazilian species, L. ecitonis Blum (1924), from Eciton
(Dorylinae), belongs to the L. galeritae group, whereas L. camponoti S.
T. Batra (1963), on Camponotus (Formicinae) in India, belongs to the
L. vulgaris group (cf. PI. 50,a,b,c). The structure of L. napoleonis Bac-
carini is not clear from Baccarini's figures (1904); this taxon was found
on Echinomegistus foreli (Wasmann), a mite occurring on Lasius
alienus (Foerster), a European ant (see Balazuc, 1971f). However, only
Rickia has been reported on ants (Myrmica [Myrmicinae]; possibly it
also occurs on Leptothorax [Pseudomyrmicinae]) in Europe. Dimor-
phomyces occurs on Paratrechina (Formicinae) in Argentina. It is not
known whether closely related species of Laboulbenia occur on ants on
a continent or whether only one widespread species such as L.
formicarum is present in each area. It should be determined whether
monoecious and dioecious species differ in the amount of variation that
occurs on different host genera.
In addition to Rickia, Diplopoda are parasitized by two other genera,
which are unknown on any other hosts — Diplopodomyces and Troglo-
myces. These two genera are closely related.
Rickia also occurs on Coelostoma, a terrestrial genus in the Hydro-
philidae that is also parasitized by Blasticomyces and Chaetarthrio-
myces. Rickia parasitizes genera in the Staphylininae, Omaliinae,
Osoriinae, and Tachyporinae in Asia, the Philippines, and Indonesia.
It occurs on several families that bear only one or two genera of
Laboulbeniales; these families may characteristically frequent fungi (Eroty-
lidae, Scaphidiidae) or rotten wood and detritus (Passalidae).
Phoretic mites are probably the means of transmission of
Dimeromyces and Laboulbenia, as well as Rickia, to a number of host groups.
These mites generally use the insect hosts merely as a means of transport
from one feeding ground (such as dung) to another (Hughes, 1959);
however, some mites live more regularly on beetles, such as under the
elytra. Dimeromyces and Rickia, which parasitize many host
groups, 'show much structural diversity. By contrast, Trenomyces,
which occurs only on Mallophaga and Hippoboscidae (bird and
mammal ectoparasites) is a very uniform genus, with little variation among
species. The Mallophaga on American land birds were found by
Kellogg (1896b) to be identical or very similar to those occurring on
Old World birds that were closely related. Kellogg (1896a) found that
although uniformity of living conditions tended to encourage little

364
MYCOLOGIA MEMOIR NO. 9
change, groups of Mallophaga on individual host birds were isolated
and close breeding occurred. Perpetuation of slight variations insured
that the members of the community would soon differ from the parental
type. Kellogg (1896a) compared life on the birds to island life. He
reported that lice of maritime birds leave hosts when they die; they may go
from one host to another in places where birds congregate, but without
the warmth of the body of a bird, the lice die. The Mallophaga may thus
be found on host species that are not customarily inhabited; however,
Kellogg (1896a) reported that despite such accidental transfers, there
does not seem to be any long-term change in the host species parasitized
by a particular species of louse (cf. Blackwell, 1980a, on bat parasites).
Chitonomyces, which is confined to water beetles, is found on several
families in the Adephaga; its primary host family is probably the Dytis-
cidae. Apparently it does not parasitize the Hydrophilidae (in the
Polyphaga). This suggests that Chitonomyces evolved with the water
beetles of the Adephaga; perhaps differences in living habits have
discouraged transfer to the Hydrophilidae. Species of Chitonomyces seem,
with few exceptions, to be quite restricted in range. The related genus
Hydraeomyces apparently is limited to Haliplidae (in the Adephaga),
which also is parasitized by Chitonomyces.
Among aquatic hosts, the greatest diversity of fungi occurs on the
Hydrophilidae, on which there is a group of closely related fungi in a
single tribe — the Ceratomyceteae. Obviously, the fungi and beetles
have evolved together. There have also been several waves of infection
on the Hydrophilidae. If the most primitive fungi constituted the first
wave (the sequence could have been quite different), then the
Hydrophilidae would have been infected first by the Ceratomycetaceae,
then by Zodiomyces, with Chaetarthriomyces, Hydrophilomyces,
and Blasticomyces appearing later. Most species in the latter group of
genera do not occur on aquatic beetles, however.
There are various factors to take into account when trying to
determine the time at which the different host groups probably became
infected, the area where the transfer may have been made, and the host
group that could have been the source of the infection. One is the time
that the area became suitable for colonization because of the receding of
water and establishment of land connections or because of temperature
changes and resultant changes in the vegetation. The amount of phylo-
genetic development that had been achieved by the host and the fungus
groups at the time that the initial infection took place may be reflected
in the relationships between the fungi inhabiting a particular host
group. The habitats that were occupied by the hosts at different times
during their evolution may be indicated by the interrelationships of the
parasites.
TAVARES: GEOGRAPHIC DISTRIBUTION
365
Geographic Patterns of Distribution
Distribution of species in the Laboulbeniales is not a matter of
dispersal of spores by air currents. The spores are transmitted by
contact — either with an infected host or with spores deposited on the
substrate. Possibly dispersion has occurred as follows: The infected host
species carried the fungus with it on its exoskeleton as it extended its
range; the infection may have died out in some populations or it may
have been transmitted to another species of host; eventually, many of
the fungi underwent structural changes, sometimes resulting in a
complex of closely related species on a single host genus (as in
Chitonomyces), while other fungi remained unchanged. When a fungus infected
a new host genus, small structural differences usually arose.
The interrelationships of the species in the host genus, as well as their
ranges, might indicate the route along which the fungus was
transported. A tropical insect could have passed westward into Europe and
then into North America during the warm, humid period of the early
Tertiary; as the temperature grew colder in the north, it may have
migrated south. Tropical insects of African origin might have moved
into tropical America before the Cretaceous separation of Africa and
South America. Most widespread beetles probably were dispersed
before the glacial period in the northern hemisphere, although there
has been a certain amount of dispersal since then, particularly of those
that repopulated previously glaciated areas.
Data gathered on distribution of water beetles in Great Britain (Bal-
four-Browne, 1953, 1960) suggest that active flight over short
distances is an important means of dispersal, particularly^when aided by
a steady breeze. Regular migrations into Britain from the south,
southeast, northeast, and west show that they may take place
independently of the direction of the prevailing wind (Balfour-Browne,
1953). Balfour-Browne (1960) pointed out that environmental
changes and migrations cause the fauna of a particular place to change
continually, although lakes are more stable than smaller bodies of
water. The order of arrival of a species or its strength in numbers at a
particular time affects the later composition of the fauna. Temporary
changes of weather or lack of suitable food, especially when a new
generation appears, may induce beetle movement. The survival of a
species depends upon its power of adjustment to altered conditions and
its ability to locate another suitable habitat (Balfour-Browne, 1960).
Many beetles probably were transported to America from Europe
with ballast rock (Lindroth, 1957); distribution of pests with
agricultural crops has also occurred (Price, 1975). The habitats of beetles
and other insects, such as cockroaches, should be fair indicators of

366
MYCOLOGIA MEMOIR NO. 9
their possible relations to the activities of man (see Price, 1975, chapter
21). Lindroth (1957) also suggested that there might have been some
movement in the Gulf Stream eastward. Oceanic transport in driftwood
is possible when living habits permit it (Kuschel, 1969). Although wind
dispersal is considered to be the most important agent for dispersal of
insects, Udvardy (1969) pointed out that cold and desiccation at higher
altitudes results in the death of most insects transported long distances.
Although migrating birds might carry some insects or their eggs,
the movement would involve only those of appropriate habitats and the
distance they were carried would depend on the persistence of their
attachment (bird lice or Mallophaga would be least likely to become
detached from the carrier en route, although they may move from one
bird to another at roosting places). Long-distance distribution by air
drift or similar means would result in the introduction of a very small
sample of the gene pool of the parent population (Price, 1975); under
the influence of different environmental conditions, a rapid divergence
of genetic stock between the new and parent populations would
probably result in large differences between species occurring in the two
areas.
Thaxter (1908) studied a large number of species of Caraboidea from
Hawaii, including Bembidion bearing Laboulbenia vulgaris. The
remaining hosts belong to two groups, according to Sharp's disposition
(Sharp, 1913); Mecyclothorax was referred to Harpalini, Pterostichides
(Jeannel, 1941, placed it in Mecyclothoracinae in the Trechidae) and the
remaining genera were placed in Harpalini, Anchomenides (Jeannel
thought they should be carefully studied). Four species and two varieties
of Laboulbenia were reported besides L. vulgaris. The appendage
systems of these four species are all similar, indicating a close
relationship. The basal cells of the outer and inner appendages are subequal,
and the abundant branches darken. The receptacle is normal in L. dise-
nochi Thaxter, but in L. sphyriopsidis Thaxter, cell Kis free from the
perithecium, which curves outward; in L. cauliculata Thaxter, the
perithecium is stalked; and in L. hawaiiensis Thaxter, the perithecium is
adnate to the receptacle for much of its length and cell Kis broadened so
that the perithecium is forced outward. Laboulbenia hawaiiensis
parasitizes several genera, including Mecyclothorax (but it does not occur on
Brosconymus and Metromenus); L. sphyriopsidis parasitizes
Metromenus; L. cauliculata and its varieties occur on Colpocaccus, Ate-
lothrus, Mesothriscus, and Metromenus; andi,. disenochi was reported
on Disenochus and Brosconymus. Colla (1926b) also reported L.
disenochi on Abropus in Colombia and on an undetermined carabid in
the Canary Islands, but she included no illustrations, so it is impossible
to evaluate her determination.
TAVARES: GEOGRAPHIC DISTRIBUTION
367
As Jeannel pointed out (1942b), there are a great number of endemics
in Hawaii; evidently, colonizers underwent considerable evolution while
invading a great variety of habitats; striking patterns of adaptive
radiation have been reported. The relationships of the genera of cara-
bids listed above are not well known, but most of them probably are
quite closely related. Yet the species of Laboulbenia that parasitize them
are few and they appear to be descended from a single ancestor, with
the exception of L. vulgaris. The ancestral fungus probably invaded the
area with the initial hosts and has undergone little change by
comparison to the evolutionary modifications that have occurred in the hosts.
Morion (Caraboidea, Pterostichidae) is a very old, primitive genus
(Jeannel, 1942b), with a range extending from South America through
Africa to Indonesia. Its range and morphological characteristics suggest
that Morion occupied the continuous area that was broken up by the
separation of the African-Brazilian continent in the Cretaceous period.
There is little difference between species on the two sides of the Atlantic
(Jeannel, 1942b, p. 251). Beetle populations that have migrated long
distances, encountering various barriers interfering with gene flow and
stimulating diversification, should have undergone considerable
differentiation (Price, 1975). Consequently, it is improbable that
Morion moved through Europe and North America to reach the
neotropical region.
Morion is the host of Laboulbenia morionis Thaxter (1893), L. papu-
ana Thaxter (1899), and L. barbata Thaxter (1899). Although a thallus
of L. morionis from Java (PI. 49,b) has an unmodified cell Kand thus
differs considerably from the type from Mexico (Thaxter, 1896, PL
XXI, Fig. 19), each has a perithecium that is almost entirely adnate to
the receptacle and a thick cell //, so that the thallus is roughly
cylindrical in shape. Thaxter obtained a specimen from New Guinea (PL
49,c) with an enlarged free V separating the perithecial apex from the
appendages, but to a lesser extent than in the type. This suggests very
early evolution of L. morionis, before Morion extended its range,
although it appears that the enlargement of Kis greater in western thalli.
It is possible, of course, that the New Guinea thallus is a secondary
modification of the ancestral form, which is probably represented by
the Javan specimen. Laboulbenia papuana, from New Guinea, has a
normal V and a perithecium that extends farther beyond the insertion
cell. It is not very different from the Javan specimen, otherwise.
Laboulbenia barbata, known only from the Western Hemisphere, has
an enlarged V that extends up the side of the perithecium. However,
its unmodified appendage system appears to be somewhat similar to
those of the other species on Morion and they are all clearly related.

368
MYCOLOGIA MEMOIR NO. 9
The cooling of Africa during the early Tertiary (when the South Pole
was near South Africa and Europe was on the equator) resulted in
the displacement of the African-Brazilian fauna, which was able,
however, to flourish in South America (Jeannel, 1942b). In the Paleocene,
India and Africa were joined to Asia Minor, providing a place of refuge
for tropical faunas and a route by which they could enter southern
Europe. Galerita s. lat. has been found in Tertiary amber in northern
Europe, according to Jeannel (1942b, p. 192). Although it possibly
migrated northward, it could have come from America or from the east;
the structural characteristics of fossil beetles (and of any Laboulbeniales
that might occur on them), as well as specimens of Laboulbenia on
Asian genera allied to Galerita (Balazuc [unpublished host index]
lists a collection on Galeritella from Indonesia), should be useful in
determining the probable distribution route.
Galerita (conserved, Bull. Zool. Nomen. 25: 98. 1968; synonyms:
Galeritula Strand, 1936, Galeritina Jeannel, 1949a [Caraboidea, Dryp-
tidae, Galeritinae]) has been divided into several genera (see Jeannel,
1949a). Laboulbenia longicollis Thaxter, which occurs in Africa on
Galeritiola and Galeritella, is quite different from L. galeritae Thaxter
and its allies, which parasitize Galerita and Progaleritina in the Western
Hemisphere. Other taxa, unrelated to L. galeritae, that occur both on
Galerita and Progaleritina are L. mexicana Thaxter (apparently allied to
L. rougetii) and L. melanotheca Thaxter (with a stalked perithecium,
having an appendage system close to that of L. flagellata).
Progaleritina occurs in North and Central America; Galerita, which is related,
but more highly evolved (see Jeannel, 1949a), occurs in Central and
South America.
Among Laboulbenia species on Chrysomelidae that have an
undivided cell /// (Ceraiomyces group), Laboulbenia dislocata (Thaxt.)
Thaxter (1915) (PI. 54,c) on Chaetocnema and Epitrix (Halticinae) and
L. trinidadensis (Thaxt.) Thaxter (1915) (PI. 54,e) on Epitrix have a
short insertion cell and a few-celled short appendage; these species occur
in the West Indies. *Laboulbenia temperei Balazuc (1973a) (in France
on Chaetocnema) is similar, although an insertion cell was not detected;
it resembles more closely L. minuscula (Thaxt.) Thaxter (1915) on
Chaetocnema and L. obesa (Thaxt.) Thaxter (1915) on Epitrix, also
from the West Indies; they have indistinct insertion cells. On the other
hand, L. nisotrae (Thaxt.) Thaxter (1915) on Nisotra (Halticinae) (PI.
54,a,b), L. monoleptae Balazuc (1975f) on Monolepta (Galerucinae),
and L. motasii Balazuc (1975f) on Podagrica (Halticinae) (in Africa and
Madagascar) have a taller insertion cell and a longer appendage,
although they are similar in structure. Possibly, L. temperei and the
West Indian species are descended from a taxon whose range extended
. TAVARES: GEOGRAPHIC DISTRIBUTION
369
from Europe to North America during the warm Eocene period and
whose progeny subsequently moved southward into tropical America.
Brachycerous Diptera (which include the hosts of Stigmatomyces,
Fanniomyces, Rhizomyces and Ilytheomyces), together with associated
beetles, came into Europe from the east with the flowering plants in the
Cretaceous (Jeannel, 1949b). Stigmatomyces must have evolved with
these flies, probably arising from a beetle parasite such as
Autophagomyces or Zeugandromyces, and is now dispersed worldwide;
it exhibits a high degree of host specificity, each species usually
occurring on a single host genus. Fanniomyces, on the other hand, is a
segregate of Stigmatomyces. The occurrence of F. ceratophorus from
Europe to North America (also Central America, according to Whisler,
1968) suggests a Tertiary distribution, although proximity of its host to
human habitations probably indicates more recent dispersal. Only very
small genera with limited distribution that are only slightly
differentiated from their relatives (or that represent mutants of distinctive
appearance) are probably of very recent origin. Rhizomyces is known only
from Diopsidae in Africa, whereas Ilytheomyces (on Ephydridae), a
genus with simpler structure, ranges from the East Indies through
Africa to tropical America. Possibly Ilytheomyces was dispersed
throughout this area before the Cretaceous separation of Africa and
South America, whereas Rhizomyces may have evolved from some
African species of Ilytheomyces more recently. It should be ascertained
whether Ilythea (host of the latter) may have been dispersed by air or
other means over long distances, or whether it underwent a gradual
extension of its range over a long period of time.
Probably, Stigmatomyces and Arthrorhynchus arose from a single
source as the Diptera evolved. Its distribution pattern indicates that
Arthrorhynchus, like Stigmatomyces, may have originated in Asia. By
contrast, Gloeandromyces is known only from Streblidae (less highly
modified dipteran bat parasites than those bearing Arthrorhynchus)
in Venezuela and the West Indies; it may have evolved from a poorly
differentiated species of Stigmatomyces or from a related taxon on
beetles (possibly scavengers on bat guano). Zeugandromyces has a
perithecium that is simpler in form than that of Stigmatomyces and it
occurs on beetles. However, its distribution pattern suggests a center of
dispersion in southeast Asia. It should be determined whether Streblidae
in other areas bear Gloeandromyces and Nycteromyces or whether these
are secondary infections that took place in the neotropical area;
Nycteromyces may have evolved from Dimeromyces living on insects in
bat caves.
A paleantarctic dispersal is suggested by the occurrence of Diphy-
myces in Argentina, Chile, Europe, Java, and New Zealand.

370
MYCOLOGIA MEMOIR NO. 9
Jeannel (1962) considered the Chilean Catopidae to be of paleantarctic
origin, with closely related genera in Australia and New Zealand. He
believed that Ptomaphagus, a holarctic genus, was derived from
ancestors that had migrated north from South America and spread
widely in the Tertiary. The European species Diphymyces niger, which
parasitizes Ptomaphagus, is quite different in form from the species of
the Southern Hemisphere.
The New Zealand taxa of Cucujomyces parasitize Cryptophagidae,
whereas those in Chile and Argentina occur on Leiodidae, Cucujidae,
and Biphyllidae. The European taxon occurs on Silvanidae, which is
closely related to Cucujidae. More collections of Cucujomyces are
needed in order to determine the probable manner of distribution. Of
the two tribes treated by Jeannel (1962) in his study of South American
Leiodidae, Neopelatopini occurs in New Zealand and South America,
whereas Hydnobiini occurs both in the holarctic and in the paleantarctic
region. Leiodidae in New Zealand should be examined for
Laboulbeniales.
Darlington (1965) pointed out that many or most of the invertebrates
and plants common to South America and Tasmania are confined to the
narrow wet forest zone or subantarctic moors on the western edges
of these land masses. Nothofagus, a prominent tree in this zone, also
occurs in New Zealand, where it has a slightly different distribution
pattern. In the highlands of New Guinea it is associated with a different
fauna, derived from that of the surrounding lowlands. It is important
that the beetles of the Nothofagus forest of Chile, Tasmania, and New
Zealand be examined for Laboulbeniales, as well as the areas of
southwestern Australia that harbor primitive relict beetles. The oldest
elements of the Australian fauna are in the southwest (Jeannel, 1942b,
p. 220); they show affinities with organisms in India, New Guinea,
South Africa, and Madagascar.
The Laboulbeniales Parasitizing Trechinae
(Including the Cave Beetles of Europe and North America)
At the beginning of the Upper Jurassic, land connections between
Asia (Angara Land) and the Australian-New Zealand region (eastern
part of the southern paleantarctic continent separated from Gondwana-
land) enabled the ancestors of the tribe Trechini (Caraboidea, Trechi-
dae, subfamily Trechinae) to enter Angara Land (Jeannel, 1942b).
Presumably the Trechini evolved in southeast Asia; the tribe is represented
today in New Zealand, Asia, Europe, North America, and Africa.
Meanwhile, other tribes of the Trechinae evolved in the paleantarctic —
TAVARES: LABOULBENIALES ON TRECHINAE
371
Homaloderini (primarily living in the mountains) and Aepini (chiefly
living in the marine intertidal zone). Trechodini is a montane tribe of
earlier origin, with close relatives occurring at present both in Africa
and Australia — that is, East Gondwanaland; Perileptini also originated
in this region — they are ripicolous (Jeannel, 1942b; see also Jeannel,
1930). Aepini occur in temperate South America, the subantarctic
islands, and New Zealand, whereas Homaloderini occur in
southeastern Australia and in temperate South America, ranging north in the
Andes. There are five genera in Europe belonging to these southern
hemisphere tribes — Aepus and Aepopsis (Aepini, intertidal along the
Atlantic coast), Iberotrechus (Homaloderini, cavernicolous in the
mountains of northern Spain), Thalassophilus (Trechodini, with a wide
distribution across central Europe, extending also to the Canary
Islands), and Perileptus (Perileptini, with an even more extensive
range). Jeannel (1960) believed that the paleantarctic fauna moved into
Europe through North America; on the other hand, Trechodini
undoubtedly entered from Asia.
Laboulbeniales have been reported from each of these genera except
Aepus. Thalassophilus is parasitized by Rhachomyces, whereas the
other three genera bear Laboulbenia. Infraspecific taxa within L.
vulgaris occur on Perileptus in Yugoslavia (Banhegyi, 1960) and on
Amblystogenium (Trechodini) in the subantarctic Crozet Islands,
southeast of South Africa (Lepesme, 1947), whereas a species closely
related to L. vulgaris (*L. francoisiana Lepesme, 1942c, nom. nud.)
parasitizes Pachydesus (syn. Plocamotrechus) (Trechodini) in South Africa.
This distribution pattern suggests that there was a connection between
these islands and South Africa, as proposed by Jeannel (1942b), which
enabled dispersal of a temperate fauna differentiated in southern Africa
during early Tertiary.
The species occurring on Aepini are unrelated: in *L. jeannelii
Lepesme (1947, nom. nud.) on Temnostega, Crozet Islands, there is a
well-developed inner appendage and an enlarged, wedge-shaped cell
V, resembling that of L. morionis; on the other hand, in *L. marina
Picard (1908a) on Aepopsis, cells IVand Fare rectangular and of equal
height — there are constricted black septa on the appendage. In *L.
bolivarii Gonzalez Fragoso (1924) on Iberotrechus, an extra cell
apparently is cut off above cell IV in the receptacle; the species was
thought by Gonzalez Fragoso to be allied to L. proliferans. In *L.
chiliensis Lepesme (1942c, nom. nud.) on Trechisibus
(Homaloderini) in Chile, cells IV and V are of equal height, but there are no
constricted black septa on the appendages. These four species are closer
to taxa parasitizing subfamilies other than Trechinae among the
Caraboidea. By contrast, all species of Laboulbenia on Trechini —

372
MYCOLOGIA MEMOIR NO. 9
L. vulgaris and its varieties, *L. shanorii Banhegyi (1960), L. sub-
terranea Thaxter (1893), and *L. heimii Lepesme (1942b, nom. nud.) on
Madeira) — are very closely related. Balazuc (personal communication)
believes that Siemaszko and Siemaszko (1928) probably mistook L.
vulgaris on Trechus for L. polyphaga.
There were two principal migrations of Trechini into Europe — one
at the beginning of the Tertiary, when land connections permitted mass
migrations from the east, and the other in the Oligocene (Jeannel,
1942b). Some beetles of the first migration reached the Pyrenees
(progenitors of Aphaenops and Geotrechus, today parasitized by
Rhachomyces); others, also now parasitized by Rhachomyces —
Pheggomisetes and Typhlotrechus, are remnants of genera that
colonized eastern Europe. Other beetles of the first migration extended into
North America — Trechus and relatives of Trechoblemus, from which
most North American cave beetles (including Pseudanophthalmus
[Krekeler, 1958]) were derived. Paratrechus probably became isolated
in the Eocene (Jeannel, 1928).
Beetles of the second migration populated Asia Minor, southeastern
Europe, and North Africa. Duvalius subgenus Duvalius occupied Asia
Minor until Upper Miocene, according to Jeannel (1942b), when the
disappearance of a water barrier in the vicinity of Mount Olympus enabled
it to spread through western Europe, displacing Allegrettia, Para-
phaenops, and Speotrechus from the first migration; the latter relicts
remain today in isolated caves (Allegrettia in Italy and Speotrechus in
France are parasitized by different taxa of Rhachomyces). Amero-
duvalius in caves of southeastern United States is related to Duvalius
(Barr, 1965). Laboulbenia and Rhachomyces occur on Duvalius and
Geotrechus (Balazuc, personal communication); Aphaenops bears
Rhachomyces only. In the United States, it is possible that
Pseudanophthalmus is the only cave beetle parasitized by Rhachomyces, although
Laboulbenia occurs on other cave beetles (unpublished) and on Trechus
(Thaxter, 1896; Lepesme, 1947). However, Rhachomyces occurs on
Paratrechus in Mexico (Balazuc, 1975e).
As the Atlantic Ocean opened up (Hurley, 1968; Jeannel, 1942b;
Demoulin, 1973) and finally joined with the Arctic, interchange of
organisms between North America and Europe became progressively
more restricted, with the connection in the far north remaining until
rather late in the Tertiary, permitting migration of cool temperature
taxa. On the other hand, taxa of warm climates became isolated
earlier on the two continents (see section on geographical distribution).
Evidently, with glaciation in the Western Hemisphere, Trechus
moved south, returning to more northern areas after the retreat of the
TAVARES: LABOULBENIALES ON TRECHINAE 373
glaciers, whereas the cave beetles went into the caves at the edge of the
glacier in southern Indiana, Tennessee, and the Kentucky region during
the glacial epoch. Barr (1965) thought that the ancestors of
Pseudanophthalmus, being preadapted to cave life, colonized caverns during
interglacial periods; otherwise, they were epigean in wetter, colder
environments (compare habitat variations of Duvalius [Neoduvalius] in
East Europe).
As the climate of Europe cooled, beetles of the second migration in
the mountains of eastern Europe adjusted to the increasing cold by
going underground (Jeannel, 1942b). Surface-dwelling Trechus moved to
lower altitudes in order to escape the glaciers of the mountains, then
returned to their higher altitude forest habitats after the glaciers
retreated. Most Duvalius species adapted to an underground
existence and did not change their ranges after the glacial period.
Aphaenops and Geotrechus are found in caves along the line of the
furthest extension of glaciation of the Pyrenees (Jeannel, 1942b). Most
are known from France. Aphaenops is very highly modified in form,
having long legs and lacking wings and eyes; it inhabits walls of
stalagmite caverns. Geotrechus is less modified in shape and inhabits a
different ecological niche, often being found near entrances to caves.
According to Jeannel (1942b), the extreme form of Aphaenops
indicates that it probably did not pass through an endogeous stage like
Duvalius and therefore lived on the surface before the glacial period.
Duvalius species have a more normal shape and there is a rudimentary
eye in some species (see Jeannel, 1941). Many beetles that live under
leaves and rocks may be apterous and have reduced eyes. Thus, many
species of Trechus have these characters, whereas others have normal
eyes or wings or both. Jeannel (1942b) proposed that the surface-
dwelling Aphaenops was nivicolous like Duvalius (Trechopsis) lapiei
(Peyerimhoff) of North Africa. It was suggested by Vandel (1964) that
nivicoles entered caves at the end of the Pleistocene in order to escape
the warmer, drier climate.
Most of the specimens of Laboulbenia found on European Trechini
were identified as L. subterranea Thaxter (Baumgartner, 1923, Cepede
and Picard, 1908, Banhegyi, 1940, 1949, 1960, Colla, 1926b, 1934,
Gonzalez Fragoso, 1924, Maire, 1916a, Lepesme, 1944, Picard,
1913b, Siemaszko and Siemaszko, 1928, Spegazzini, 1914b, 1915b).
The type (from a North American Pseudanophthalmus) (Thaxter, 1896,
PI. XIII, Fig. 9) has extremely short VI, III, and IV cells and two
darkened outer appendage cells; the base of the perithecium is unin-
flated. In the thallus in Thaxter's Fig. 10, PI. XIII (1896), cells ///, IV,
and Fare longer and the upper receptacle wider, as in Balazuc's thalli
(1974) from North American Pseudanophthalmus; lengthening of / and

374
MYCOLOGIA MEMOIR NO. 9
// seems to be accompanied by elongation of /// and IV in this species.
Thaxter (1908) placed in L. subterranea a European specimen on
Trechus (PI. LIII, Fig. 15) that corresponds to L. vulgaris var. trechi-
phila Spegazzini (1914b); another European specimen on Trechoblemus
(PI. LIII, Fig. 14) has a long IV and VI, with no swelling of the
perithecial base — it is L. subterranea subsp. lecoareri Balazuc (1974).
As Balazuc pointed out, this subspecies has a taller insertion cell than
the typical subspecies and a small wedge-shaped basal cell of the inner
appendage, rather than a cylindrical cell. Colla's illustrations (1934) of
L. subterranea show great variation; the hosts were Duvalius and
Trechus.
Laboulbenia shanorii Banhegyi (1960), which occurs on Neotrechus
in eastern Europe, is a short, stout species, characterized by darkness on
the upper part of cell / and the lower part of the perithecium and
outer appendage. Some of the hosts of these species of Laboulbenia
are cavernicolous; others are not. There is no clear distinction between
L. vulgaris and L. vulgaris var. trechiphila, according to Balazuc (1974).
On the other hand, Lepesme (1947) observed that thalli of L. vulgaris
are never the same on different hosts or in different localities.
Laboulbenia vulgaris was believed by Lepesme (1947) to be the oldest
species in the genus on the basis of its distribution pattern. Its structure
supports his view — the poorly developed inner appendage probably
represents the first step in the process of appendage elaboration
following modification of the basal cell of the primary appendage into a
flat, dark insertion cell.
Because of the large number of species of Peryphus (Trechidae, Bem-
bidiinae, often treated as a subgenus of Bembidion) parasitized by L.
vulgaris, Lepesme (1947) believed it to be'the primary host. This fungus
occurs on Bembidion in India (Batra, 1963) and in temperate South
America (Thaxter, 1912a), as well as in North Africa, Europe, western
Asia, Siberia, and Central America (Lepesme, 1947, as-Plataphus,
Peryphus). Darlington (1965) indicated that Bembidion was more recently
dispersed in South America than the Trechinae of that continent. It
undoubtedly carried L. vulgaris with it as it extended its range. Balazuc
(unpublished) has obtained/,, vulgaris var. trechiphila from Trechisibus
in Chile. Careful comparison of thallus morphology might indicate
whether the latter populations were derived from fungi parasitizing the
ancestral beetles or whether they arose through cross-infection with
Bembidion.
The widespread occurrence of L. vulgaris on the ripicolous genus
Bembidion, as well as on Trechinae occupying various habitats, suggests
that it may have parasitized the ancestral stock of these two subfamilies.
TAVARES: LABOULBENIALES ON TRECHINAE
375
On the other hand, cross-infection could have occurred in the riparian
habitat between Perileptus and Bembidion.
It is unlikely that the taxa in the Crozet Islands and in South Africa
became infected with L. vulgaris through contact between Trechus,
which migrated very late along the high mountain chains into East
Africa, and Pachydesus, which migrated north in the mountains
from South Africa (see Jeannel, 1942b). The two genera do not occur in
the same mountains at present. Moreover, Laboulbenia has not been
reported from this subgenus of Trechus (Elgonotrechus) (Balazuc, 1970,
1972, found Rhachomyces instead). Although Perileptus became
dispersed in areas around the Indian Ocean, Laboulbenia has been
reported on this host only in the Mediterranean region (Banhegyi, 1960;
Maire, 1916b). Laboulbenia vulgaris var. perilepti Banhegyi resembles
some of the specimens found on Amblystogenium (cf. Lepesme,
1947, Fig. 1, right) in the Crozet Islands.
The variation in thallus length that occurs in Laboulbenia vulgaris
living in caves, as contrasted to the lack of any conspicuous changes
in thalli of Rhachomyces, led Lepesme (1942a) to believe that the
introduction of the latter into the subterranean habitat was more recent than
the entrance of Laboulbenia. However, the two genera seem to respond
differently to environmental influences. Rhachomyces shows a greater
degree of host specificity than Laboulbenia vulgaris. Laboulbenia
species often vary in the length of the receptacle, perhaps because of the
position on the host. In their study of Laboulbenia on Bembidion,
Benjamin and Shanor (1952) found that the short species L. truncata
Thaxter occurred on the tarsi, whereas the larger taxon L. perpendi-
cularis Thaxter grew on the prosternum, presumably a place with a
better supply of nutrients.
When species of Laboulbenia and Rhachomyces entered caves
depends on the identity of the host. Both genera occur on cavernicolous
species of Trechus in Spain. In southeastern Europe, both genera occur
on surface-dwelling species of Trechus. Some species of Duvalius
parasitized by Laboulbenia in the Maritime Alps live in caves, whereas
others live under stones. In Tuscany, the hosts are cave dwellers
(Jeannel, 1927,1928).
Among the hosts of Rhachomyces, Anophthalmus, host of R. anoph-
thalmi, lives in caves at low altitudes and under stones in humid places
in the mountains. Jeannel (1928) believed that Orotrechus (host of R.
bucciarellii) colonized caves earlier than Neotrechus (host of L.
shanorii), which probably lived in the superficial fauna during the Tertiary.
On the other hand, Duvalius baldensis Putzeys (host of jR. maublancii)
is a surface dweller (see discussion at end of section on Rhachomyces).

TAVARES: ACKNOWLEDGMENTS
377
ACKNOWLEDGMENTS
The author is deeply indebted to the late Dr. Betty S. Davis, Hastings
Natural History Reservation, University of California, for her work on
the drawings of Laboulbenia and Herpomyces ectobiae. She is grateful
to Mrs. Charlotte M. Hannan for assistance with drawings of
Herpomyces periplanetae and other genera, as well as with maps and charts.
She wishes also to acknowledge the work of the following
photographers of the Scientific Photographic Laboratory, University of
California, Berkeley: Mr. Victor Duran, for Pis. 1, a-e, 2, 3, a-c, 4,
a, e, 5, b-c, e, 6, a-b, 7, c-e, 8, 9, 10, b-c, 11, 12, a-d, 14, b, f, 15, 38, d;
Mr. SamEhrlich, for Pis. 24, b-c, 31, e, 32, a, f. 37, a-c, 39, b, 51, g, 56,
a-c, e; Mr. Gene Groppetti, for Pis. 24, a, d-e, 32, g, 42, d, 44,1, 46, f-g,
48, a-b; Mr. James Hendel, for Pis. 17, a, c, e, 18, a-c, 19, a-b, 20, a-c,
21, a, 24, f, 26, a-b, 27, c, 29, 30, 31, g, 32, c-e, 34, c, 36, a, 37, d-f, 38,
a-b, 39, a, c-e, 40, a, c-e, 42, c, j-k, 44, c, k, 45, a-c, 46, a-d, h-j, 48, d,
49, a, d, 50, c, d, 51, a-d, f, 52, a, c-i, 53, b, 56, d, f-h; and Mr. Alfred
Blaker, for Pis. 1, f, 3, d-f, 4, b-d, 5, a, d, 6, c, 7, a-b, 10, a, 12, e, 13,
14, a, c-e, 16, 17, b, d, f, 18, d-i, 19, c, 20, d, 21, b-c, 22, 23, 25, 26, c-e,
27, a-b, d-e, 28, 31, a-d, f, 32, b, 33, 34, a-b, d-g, 35, 36, b-i, 38, c, e-h,
40, b, 41, 42, a-b, e-i, 43, 44, a-b, d-j, 45, d-g, 46, e, 47, 48, c, e-g, 49, b-
c, e-f, 50, a-b, e, 51, e, 52, b, 53, a, c, 54, 55; as well as that of Dr. Mel-
vin Fuller, for PI. 6, d.
The author is also indebted to Mr. Miklos Nagy for translation of
Istvanffi's paper (1895a) and to the Curt Dietz Fund for most
photographs of Pis. 1-15. Determinations of beetle hosts for ontogenetic
studies of Laboulbenia flagellata, L. gyrinidarum, andZ. borealis were
made by Mr. H. B. Leech, California Academy of Sciences, San
Francisco; sections of Hesperomyces were prepared by Miss Nel Rem from
material sent by Dr. N. H. Hussey and Miss Barbara Gurney,
Glasshouse Crops Research Institute, Rustington, Littlehampton, Sussex,
England; specimens of Herpomyces periplanetae were obtained from
Insect Toxicology, Department of Entomology, University of
California, Berkeley, from the Insectary, Department of Entomology,
Cornell University, and from the collections of Dr. Barbara Stay, Harvard
University; and slides of various species were lent by Drs. Jean Balazuc,
Tomasz Majewski, R. K. Benjamin, Lars Huggert, and Walter Rossi —
the author is grateful for all of this assistance. The primary source of
material, however, for study of the many genera of the Laboulbeniales
was the Thaxter collection at the Farlow Herbarium, without which this
publication could not have been written; an important secondary source
has been Spegazzini's collection, from which only a few specimens have
been observed so far. The author is deeply indebted to Dr. I. M. Lamb,

378
MYCOLOGIA MEMOIR NO. 9
Dr. Reed Rollins, Dr. Donald Pfister, and the many helpful staff
members of the Farlow Herbarium who have assisted her through many
years of study, as well as Drs. Irma Gamundi de Amos, Instituto de
Botanica "Spegazzini," La Plata, Argentina, for sending specimens on
loan, including the type of Laboulbenia borealis. Preserved material
from the Museum of Comparative Zoology, Harvard University, must
also be acknowledged.
Finally, bibliographic assistance and advice from Dr. John Ingram
and Dr. William J. Dress, Bailey Hortorium, Cornell University, as well
as advice from Dr. Paul Silva, University of California, Berkeley,
has been much appreciated. In addition, Dr. William Dress has kindly
provided the Latin diagnoses. Dr. Jean Balazuc has provided much
assistance on insect names and Dr. Paul J. Spangler, United States
National Museum, has kindly checked names of aquatic beetles. The
author was also assisted by Miss Alta Pray, who helped to prepare an
index to the Laboulbeniales and their hosts in 1965. The Louise Kellogg
Fund has been the source of valued financial assistance.
BIBLIOGRAPHY
with annotated entries for the Laboulbeniales
*indicates reference not seen;** indicates reference not cited in text
(Of the references listed only in text,
only the CMI Index of Fungi mentions Laboulbeniales.)
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York. London.
Alexoponlos, C. J., and C. W. Mims. 1979. Introductory Mycology. Third edition.
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Applebaum, S. W., R. Kfir, U. Gerson, and U. Tadmor. 1971. Studies on the summer
decline of Chilocorus bipustulatus in citrus groves of Israel. Entomophaga 16: 433-
444. Conclude that Hesperomyces had no appreciable effect on host.
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Backus, M. P. 1934. Initiation of the ascocarp and associated phenomena in Coccomyces
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Mem. 2. 158 pp., 112 Figs., map. Reports of Laboulbeniales on pp. 18, 41, 58, 63,
77,104,126-7 (Arthrorhynchus, Rhachomyces).
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379

380
MYCOLOGIA MEMOIR NO. 9
—. 1971a. Laboulbeniales inedites, parasites de carabiques. Nouv. Rev. Entomol. 1:
107-117. New taxa in Laboulbenia (including L. benjaminii, L. coiffaitii) (see 1974,
1975b) and Rhachomyces (nomina nuda).
—. 1971b. Bibliographic des Laboulbeniales (Ascomycetes).. Bull. Mens. Soc. Linn.
Soc. Bot. Lyon 40: 134-149. Entries numbered; these numbers used in subsequent
papers.
—. 1971c. Notes sur les Laboulbeniales. II. — Laboulbenia parasites des Gyrinus (plus
particulierement europeens et nord-africains). Bull. Mens. Soc. Linn. Soc. Bot. Lyon
40:160-168. Discussion of North American species L. borealis included.
—. 1971d. Laboulbeniales inedites, parasites de carabiques et d'un catopide. Nouv.
Rev. Entomol. 1: 245-254. New species in Corethromyces, Eucantharomyces,
Laboulbenia, including L. crolandii, L. paulianii, L. vadonii (see 1975b) (nomina nuda).
Discusses species on Catopidae.
—. 1971e. Notes sur les Laboulbeniales. III. Rectifications, synonymies et mises au point.
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corrections in species names in Laboulbenia; suggests correct host genera; lists
Laboulbeniales on bat parasites, their hosts and localities. Additional notes on Rhachomyces
on Trechidae.
—. 1971 f. Plaidoyer pour une flore francaise des Laboulbeniales parasites. L'Entomo-
logiste 27: 113-119, 158 (correction). Popular article. Lists hosts which may bear La-^
boulbeniales in France and should be collected.
—. 1972. Laboulbeniales nouvelles, parasites d'Insectes. Bull. Soc. Entomol. France
76 (1971): 226-235. New species in Dimeromyces, Dioicomyces (first report of
Laboulbeniales from Thysanoptera), Laboulbenia (as Misgomyces), Rhachomyces
(nomina nuda) (see 1975b). Discusses genera with wide range of unrelated hosts. Makes
some corrections in host names.
—. 1973a. Une Laboulbenia nouvelle (Ascomycetes), parasite d'un Altise (Coleopteres,
Chrysomelidae). Bull. Soc. Linn. Bordeaux 3 (2): 27-28.
—. 1973b. Ebauche d'une flore des Laboulbeniales de Roumanie (Ascomycetes). Pp. 463-
477. In: Instit. Speol. "Emile Racovitza," Livre Cinquant., Colloque Natl. Speol.
Chief Ed. T. Orghidan. Acad. Republ. Social. Roman. Bucharest. Reports
Arthrorhynchus, Laboulbenia, Misgomyces, Peyritschiella, Rhachomyces, Treno-
myces. Mentions specimens found on Diplopoda including undescribed Rickia (as
Rhachomyces).
—. 1973c. Laboulbeniales de France. Bull. Mens. Soc. Linn. Soc. Bot. Lyon 42: 244-
256, 280-285. New taxa in Corethromyces, Rhachomyces. Reports Asaphomyces,
Autoicomyces (as Ceratomyces), Compsomyces, Coreomyces, Corethromyces
(including Diphymyces, Peyerimhoffiella, Rhadinomyces), Euzodiomyces, Helodiomyces,
Misgomyces (including Botryandromyces, Hydrophilomyces), Rhachomyces, Rhyncho-
phoromyces, Zodiomyces. Introduction to Laboulbenia.
—. 1973d. Recherches sur les Laboulbeniomycetes. I. Trois especes nouvelles et une mal
connue. Rev. Mycol. [Paris] 37 (1972): 253-262. New taxa in Laboulbenia, including
L. chopardii, Polyandromyces (as Eudimeromyces); includes description of Rickia
coleopterophagi Paoli.
—. 1974. Laboulbeniales de France (suite). Bull. Mens. Soc. Linn. Soc. Bot. Lyon 43:
12-21, 57-64, 73-79, 253-262, 295-315, 346-368. New taxa in Laboulbenia, including L.
parriaudii. Reports Arthrorhynchus, Autophagomyces, Cantharomyces, Chitono-
TAVARES BIBLIOGRAPHY
381-
myces, Dimeromyces, Haplomyces, Herpomyces, Hesperomyces, Hydraeomyces,
Idiomyces, Ilyomyces, Monoicomyces, Peyerimhoffiella (as Corethromyces),
Peyritschiella, Picardella (as Dioicomyces), Rhachomyces, Rhynchophoromyces, Rickia,
Stemmatomyces? (as Stigmatomyces), Stigmatomyces, Symplectromyces, Teratomyces,
Trenomyces, Laboulbenia. Includes extensive discussion of L. flagellata, giving list of
hosts in France; discussion of many other species, including L. nebriae, L. polyphaga,
L. subterranea, L. vulgaris. Extralimital reports: Asaphomyces, Compsomyces,
Euzodiomyces, Hydrophilomyces (as Misgomyces), Laboulbenia, Misgomyces,
Rhachomyces.
**—. 1975a. Sur les Laboulbenia (Ascomycetes) parasites d'Odontonyx (Coleoptera
Caraboidea, Pterostichidae, Anchomenini). Nouv. Rev. Entomol. 5: 97-100.
Discussion, report of L. olisthopi in France and Madeira.
—. 1975b. Diagnoses nonnullorum Laboulbenialium nuper francogallice descriptorum.
Acta Mycol. 11: 49-57. New species in Dimeromyces, Dioicomyces, Eucantharomyces,
Laboulbenia (in part as Misgomyces), Rhachomyces (see 1970,1971a, d, 1972).
**—. 1975c. Description de 4 especes no[u]velles de Laboulbenia (Ascomycetes),
parasites de Coleopteres. Acta Mycol. 11:67-76.
—. 1975d. Laboulbeniales nouvelles (Ascomycetes), parasites de Coleopteres exotiques.
Bull. Mus. Hist. Nat. ser. 3 Bot. [Paris] 22: 177-200. New taxa in Dimeromyces,
Laboulbenia, including L. blumii, L. darlingtonii, L. minetii, L. peyrierasii,
Misgomyces.
—. 1975e. Recherches sur les Laboulbeniomycetes. II. Description de cine especes
nouvelles de Rhachomyces, parasites de Coleopteres Carabiques. Rev. Mycol.
[Paris] 38(1973): 218-227.
—. 1975f. Recherches sur les Laboulbeniomycetes. III. Laboulbeniales parasites de
Coleopteres Chrysomelides, avec description d'especes nouvelles. Rev. Mycol [Paris]
39 (1974-1975): 189-211. New species in Laboulbenia, including L. dorstii, L. motasii.
Discussion and catalogue arranged according to subfamily of host.
—. 1976. Quelques cas teratologiques observes chez des Laboulbeniales (Ascomycetes).
Rev. Mycol. [Paris] 40: 51-55. Gives teratological examples of Laboulbenia and
Ormomyces.
—. 1977a. Laboulbeniales nouvelles (Ascomycetes), parasites de Coleopteres exotiques
(suite). Bull. Mus. Hist. Nat. ser. 3 Bot. [Paris] 29: 1-14. New species in Laboulbenia,
including!,, ardoinil, Rhachomyces.
—. 1977b. Mission biospeologique cubano-roumaine a Cuba, 1969. Laboulbeniales
(Ascomycetes) parasites de Coleopteres. Pp. 407-411. In: Resultats des Expeditions Bio-
spSologiques Cubano-Roumaines a Cuba. Vol. 2. Eds. T. Orghidan et al. Acad.
Republ. Social. Romania, Bucharest. New subspecies of Cochliomyces. Reports
Autoicomyces (in part as Ceratomyces), Laboulbenia, Neohaplomyces, Rhachomyces;
includes list of all records for Cuba.
**—. 1977c. Deuxieme mission biospfeologique cubano-roumaine a Cuba (1973).
Laboulbeniales (Ascomycetes) parasites de Coleopteres Carabiques. Pp. 413-415. In:
Resultats des Expeditions Biospeologiques Cubano-Roumaines a Cuba. Vol. 2.
Eds. T. Orghidan et al. Acad. Republ. Social. Romania, Bucharest. New species of
Laboulbenia. Reports Laboulbenia, Rhachomyces.
—. 1978. Laboulbeniales (Ascomycetes) de la region francaise Antilles-Guyane. Bull.
Mens. Soc. Linn. Soc. Bot. Lyon 47: 488-500. New species in Laboulbenia. Reports
Ceratomyces, Dimeromyces, Herpomyces, Laboulbenia, Rhachomyces.

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MYCOLOGIA MEMOIR NO. 9
—. 1979. Recherches sur les Laboulbeniomycetes. IV. Descriptions de cinq especes de
Laboulbeniales. Rev. Mycol. 43:393-404. New species in Fanniomyces, Laboulbenia.
—. 1980. Laboulbeniales nouvelles (Ascomycetes), parasites de Coleopteres et de
Dipteres. Bull. Mus. Hist. Nat. ser. 4 Bot. [Paris] 2: 209-219. New species in
Laboulbenia, including L. cyrtonatis, and in Rickia, including R. huggertii.
—. 1982a. Laboulbeniales (Ascomycetes) de Madagascar, des Comores et des Mas-
careignes. Bull. Mens. Soc. Linn. Soc. Bot. Lyon 51: 6-27. Reports Dimeromyces,
Dioicomyces, Dixomyces, Enarthromyces, Eucantharomyces, Herpomyces,
Laboulbenia, Ormomyces, Peyritschiella, Polyandromyces, Rhachomyces, Rickia,
Trenomyces.
—. 1982b. Description de trois Laboulbeniales nouvelles (Ascomycetes). Int. J. Myc.
Lich. 1: 39-48. New species in Laboulbenia, Stigmatomyces.
Balazuc, J., and J. Demaux. 1973. Une nouvelle espece de Laboulbenia (Ascomycetes,
Laboulbeniales), parasite de Coleopteres Chrysomelides Hispinae. Bull. Mens. Soc.
Linn. Soc. Bot. Lyon num. spec. (150e ann.) 42: 7-9.
Balazuc, J., X. Espadaler, and J. Girbal. 1982. Laboulbenials (Ascomicets) iberiques.
Collectanea Bot. 13: 403-421. New species of Laboulbenia. Reports Arthrorhynchus,
Hydraeomyces, Laboulbenia, Rhachomyces, Rickia, Symplectromyces, taxon on
Caraboidea allied to Laboulbenia (as ? Dioicomyces).
Balcells Rocamora, E. 1954. Quiropteros de cuevas Catalans: Campana de 1952-
53. SpeleonS: 105-110. Reports Arthrorhynchus (p. 108).
—. 1955. Datos para el estudio de la fauna pupipara de los quiropteros en Espaiia.
Speleon 6: 287-312. Mentions Laboulbeniales on Nycteribia, Penicillidia, and Basilia
in Spain (pp. 307-309).
—. 1968. Revision faunistica de nicteribidos y estrgblidos de quiropteros espafioles y su
especificidad. Rev. Iber. Parasitol. 28: 19-31. Mentions that summer abundance of
Nycteribiidae is correlated with major frequency of Laboulbeniales.
Balfour-Browne, F. 1953. The aquatic Coleoptera of the western Scottish islands with a
discussion on their sources of origin and means of arrival. Entomol. Gaz. 4: 79-127.
—. 1960. The aquatic Coleoptera of Scotland and their routes of arrival. Entomol.
Gaz. 11:69-106.
Banhegyi, J. 1940. Elomunkalatok a magyarorszagi Laboulbenia-felek monografia-
jahoz. Etudes preliminaires sur les Laboulbeniales de la Hongrie. Index Horti Bot.
Univ. Budapest. 4: 39-59. Tab. VII-X. New species of Dimorphomyces (as
Dimeromyces); reports Arthrorhynchus, Idiomyces, Laboulbenia, Misgomyces, Peyritschiella
(as Dichomyces), Symplectromyces, Teratomyces.
—. 1944. A Balaton kornyekinek Laboulbenia-feUi (Les Laboulbeniales aux environs
du lac de Balaton). Bot. Kozlem. 41:49-61. Tab. I-II. New species: Aporomyces szaboi;
reports Cantharomyces, Dioicomyces, Laboulbenia, Misgomyces, Peyritschiella.
—. 1949. Les Laboulbeniales de la Transylvanie. Index Horti Bot. Univ. Budapest.
7:93-101. Tab. VII-VIII. Reports Laboulbenia, Peyritschiella, Rhachomyces.
—. 1950. Ritka Laboulbeniak a Karpatmedencebol (Laboulbeniales rares du Bassin
Carpatique). Ann. Biol. Univ. Budapest. (Budapesti Tudomanyegyetem Biol,
intezeteinek EvkOnyve) 1: 189-196. Reports Asaphomyces (as Barbariella),
Hydraeomyces, Laboulbenia, Rhachomyces.
—. 1960. Contributions a la connaissance des Laboulbeniales de la peninsule des Balkans.
Ann. Univ. Sci. Budapest. Rolando Eotvos, Sect. Biol. 3: 49-67. New taxa in Laboul-
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benia, Rhachomyces; reports Chitonomyces, Corethromyces, Euzodiomyces,
Idiomyces, Laboulbenia, Rhachomyces.
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Rolando EOtvos, Sect. Biol. 7: 19-27. Reports Rhachomyces tenenbaumii; taxonomic
discussion of Laboulbenia elaphricola and other species on Elaphrus, L. leisti.
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Bary, A. de. 1884. Vergleichende Morphologie und Biologie der Pilze, Mycetozoen,
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Batra, Suzanne W. T. 1963. Some Laboulbeniaceae (Ascomycetes) on insects from India
and Indonesia. Amer. J. Bot. 50: 986-992. New species in Laboulbenia,
Stigmatomyces.
Baumgartner, R. 1923. Contribution a l'etude des Laboulbeniales de la Suisse. Jahrb.
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Corethromyces, Euzodiomyces, Idiomyces, Laboulbenia, Misgomyces; general discussion
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Emile G. Racovitza 1868-1968. Chief. Ed. T. Orghidan. Editions Acad. Republ.
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—. 1968a. Sandersoniomyces, a new genus of Laboulbeniales allied to Diplomyces, Sym-
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—. 1968b. Balazucia, a new genus of Laboulbeniales allied to Cucujomyces Spegazzini.
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compares with related genera.
—. 1971. Introduction and supplement to Roland Thaxter's contribution towards a
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—. 1979. Laboulbeniales on semiaquatic Hemiptera. III. Rhizopodomyces. Aliso 9:
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—, and —. 1951. Morphology of immature stages of Euzodiomyces lathrobii Thaxter
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Laboulbenia on Bembidion picipes. Amer. J. Bot. 39: 125-131. New species in
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Blagoveshtchensky, D.I. 1950. Mallophaga s ptit Barabinskih ozer (II). Parazit.
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cette espece. Bull. Soc. Acad. Arrondissement Boulogne-sur-Mer 11(1922-1928): 636-
640. L. flagellata literature reports; use of name Phycascomycetes justified.
Cepede, C, and F. Picard. 1907. Observations biologiques sur les Laboulbeniacees
et diagnoses sommaires de quelques especes nouvelles. Compt. Rend. 36me Session
Assoc. Franc. Avancem. Sci., Reims, 1907(part 2): 778-784. New genus: Rheophila
(syn. of Peyritschiella), new species in Euzodiomyces, Laboulbenia; infection
experiments on Laboulbenia.
—, and —. 1908. Contribution a la biologie et a la systematique des Laboulbeniacees de
flore francaise. Bull. Sci. France Belgique 42: 247-268. Pis. Ill, IV. New species in
Laboulbenia, including L. giardii, Rhachomyces; list of species of Europe; discussion
of nutrition, etc.
Chadefaud, M. 1960. Les Vegetaux non Vasculaires (Cryptogamie). (vol. 1, Chadefaud,
M., and L. Emberger, Traite de Botanique Systematique). Masson et Cie, Paris. 1018 p.
Discusses structural patterns in Laboulbeniales based on arrangement of cladomes and
pleuridia (see pp. 487-496 on Laboulbeniales).
Chagnon, G., and A. Robert. 1962. Principaux Coleopteres de la Province de Quebec.
Second edition. Universite de Montreal Press, Montreal. 440 p.
Chatton, E., and F. Picard. 1908. Sur une Laboulbeniaciee: Trenomyces histophtorus n.
g., n. sp., endoparasite des poux (Menopon pallidum Nitzsch et Goniocotes abdomi-
nalis P.) de la poule domestique. Compt. Rend. Hebd. Seances Acad. Sci. [Paris]
146: 201-203. New genus on chicken lice.
—, and —. 1909. Contribution a l'etude systematique et biologique des Laboulbeniacees;
Trenomyces histophtorus Chatton et Picard, endoparasite des poux de la poule
domestique. Bull. Soc. Mycol. France 25: 147-170. Pis. VII, VIII. Discussion of nutrition,
effect on host, place in classification; comparison with Dimeromyces, Dimorphomyces,
Herpomyces.
China, W. E. [Editor]. 1966. Official List of Specific Names in Zoology. Second
Instalment: names 1526-2045. Pp. 207-286. International Trust for Zoological Nomenclature,
London.
China, W. E., and R. L. Usinger, 1949. Classification of the Veliidae (Hemiptera) with
a new genus from South Africa. Ann. Mag. Nat. Hist. (ser. 12) 2:343-354.
Chopard, L. 1967. Gryllides. Fam. Gryllidae: Subfamily Gryllinae (Trib. Gymnogryllini,
Gryllini, Gryllomorphini, Nemobiini). Pars 10. Pp. 1-211. In: Orthopterorum Cata-
logus. Ed. M. Beier. W. Junk, The Hague.
Clements, F. E., and C. L. Shear. 1931. The Genera of Fungi. H. W. Wilson Co., New
York. 496 p. Pis. 1-58. Note spelling of generic names in Laboulbeniales.
Coiffait, H. 1958. Les coleopteres du sol. Vie et Milieu, Suppl. 7 (Actualites Scienti-
fiques et Industrielles 1260). Hermann & Cie, Edit., Paris. 204 p. Points out rarity of
Laboulbeniales on endogeous beetles; reports Laboulbenia and Rhachomyces on
Geotrechus and Picardella (as Dioicomyces) on Anillus (see pp. 117-118).
Coiffait, H., and F. Saiz. 1966. Les Quediini du Chili [Col. Staphylinidae]. Ann. Soc.
Entomol. France (n. s.) 2:385-414.
Cole, A. C, Jr. 1935. Laboulbenia formicarum Thaxter, a fungus infesting some Idaho
ants, and a list of its known North American hosts (Hym.: Formicidae). Entomol. News
46: 24. Host list for North America.
—. 1949. New ant hosts of the fungus, Laboulbenia formicarum Thaxter. Entomol.
News 60:17. Reports two hosts in Formica and Acanthomyops (as Lasius).
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Colla, Silvia. 1925. Contributo alia conoscenza dei Laboulbeniali piemontesi. Atti
Acad. Sci. Torino 60: 250-269. New taxa in Laboulbenia, Monoicomyces; reports
Amorphomyces, Chitonomyces, Laboulbenia, Peyritschiella (as Dichomyces).
—. 1926a. Sull'organo d'assorbimento della specie del gen. "Laboulbenia" Rob. Atti
Accad. Sci. Torino 61: 277-280. Shows penetration of haustorium into host cuticle from
foot.
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Torino. Atti Accad. Reale Naz. Lincei Mem. CI. Sci. Fis. [Roma] (ser. 6) 2: 153-193. T.
I-IV. (Preceded by relazione letta dal Socio Mattirolo, pp. 151-152, referring to this
paper). New taxa in Chaetomyces (as Dimeromyces), Laboulbenia, Rhachomyces;
reports Laboulbenia, Monoicomyces, Peyritschiella (as Dichomyces), Rhachomyces.
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Sci. Nat. Milano 65: 136-147. Reports Laboulbenia; includes chart of species in the
area.
—. 1932a. "Troglomyces Manfredii" n. gen. et n. sp.: nuova Laboulbeniacea sopra un
miriapode. Nuovo Giorn. Bot. Ital. (n. s.) 39: 450-453. New genus: Troglomyces.
—. 1932b. Una Laboulbeniale nuova per l'ltalia: Rachomyces [sic] aphaenopsis Th.
Nuovo Giorn. Bot. Ital. (n. s.) 39: 512. Reports Rhachomyces.
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sistematico. Mem. Reale Accad. Sci. Torino Sci. Fis. (ser. 2) 67 (6): 1-14. Description of
different thallus forms and taxonomic discussion; mentions teratological forms.
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Heterothallicae, Laboulbeniaceae Homothallicae, Ceratomycetaceae. Fasc. 16: 1-157.
In: Flora Italica Cryptogama, pars I: Fungi. Eds. P. A. Saccardo and H. Dalla Costa.
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Soc. Entomol. Belg. 83: 21-35. Historical account; general discussion of host groups.
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Cooke, R. 1977. The Biology of Symbiotic Fungi. John Wiley and Sons, London, New
York, Sydney, Toronto. 282 p. Proposes different ways in which the Laboulbeniales
might obtain nourishment, such as slow degradation of cuticle, leaving microscopic
lacunae over a wide area, rather than severe localized erosion.
Cooke, W. B., and D. L. Hawksworth. 1970. A preliminary list of the families proposed
for fungi (including the lichens). Mycol. Pap. no. 121: 1-86. Include Ceratomycetaceae
Thaxter (1908), Zodiomycetaceae (Thaxt.) Nannizzi (1934); indicate illegitimacy of
Laboulbeniaceae heterothallicae S. Colla (1934) and Laboulbeniaceae homothallicae
S. Colla (1934).
**Costantin, J. 1888. Les Mucedinees Simples. Libraire P. Klincksieck, Paris.
210 p. Brief mention of Laboulbeniales (pp. 23, 171-172); key; places in group with
Saccharomyces, Speira, etc.
Dainat, Hannelore, 1970. Presence en France d'une nouvelle Laboulbeniale du genre
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Comp. 45: 129-137. New taxon: S. harantii. Lists parasites of brachycerous Diptera
in Europe, also Stigmatomyces (s. I.) on Coleoptera. Discusses specificity of Laboul-

390
MYCOLOGIA MEMOIR NO. 9
beniales on different host groups. Lists groups of Diptera on which Stigmatomyces are
found. Compares species with similar species.
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Mentions related species; lists some species found both in Europe and on other
continents.
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drides et de Drosophilides recoltees dans le Midi Mediterraneen de la France (these). Pp.
I-IV, 1-144, I-IX. Pis. I-XXXIII. Academie Montpellier, U.S.T.L. Discussion of
habitats, description of species, notes on hosts, relation to hosts, biogeography, maps,
photographs, drawings.
Dainat, Hannelore, and J. Dainat. 1973. Sur dix especes du genre Stigmatomyces
(Laboulbeniales) parasites de Dipteres Acalypteres dans le Sud de la France. Bull. Soc.
Mycol. France 89:337-352. New taxa in Stigmatomyces. Reports other species.
Dainat, Hannelore, Delage, A., and M.-C. Lauraire. 1971. Contribution a l'etude des
Dipteres Acalypteres de la region de Montpellier. Nouv. Rev. Ent. 1: 89-106. Diptera
belonging to 24 families reported; Laboulbeniales seen only on Ephydridae.
Dainat, Hannelore, and J.-F. Manier. 1974. Haustoria de Stigmatomyces scaptomyzae
Thaxter (Laboulbeniale) parasite de Scaptomyza graminum Fallen (Diptere, Droso-
philide). Bull. Soc. Mycol. France 90: 217-221. Penetration of integument of host is
demonstrated.
Dainat, Hannelore, Manier, J.-F., and J. Balazuc. 1974. Stigmatomyces majewskii n.
sp., Stigmatomyces papuanus Thaxter 1901, Laboulbeniales parasites de Dipteres
Acalypteres. Bull. Soc. Mycol. France 90: 171-178. Comparison with S. entomophilus.
Discussion of possible synonyms and host specificity of S. papuanus.
Darlington, P. J., Jr. 1965. Biogeography of the Southern End of the World. (Second
printing, 1969). Harvard University Press, Cambridge, Massachusetts. 236 p.
DeLamater, E. D. 1948. Basic fuchsin as a nuclear stain for fungi. Mycologia 40:423-429.
Delfinado, Mercedes D., and D. E. Hardy. 1973. A Catalog of the Diptera of the Oriental
Region. Vol. I. Suborder Nematocera. University Press of Hawaii, Honolulu. 618 p.
—, and —. 1975. A Catalog of the Diptera of the Oriental Region. Vol. II. Suborder
Brachycera through Division Aschiza, Suborder Cyclorrhapha. University Press of
Hawaii, Honolulu. 459 p.
—, and —. 1977. A Catalog of the Diptera of the Oriental Region. Vol. III. Suborder
Cyclorrhapha (excluding Division Aschiza). University Press of Hawaii, Honolulu.
854 p.
Demoulin, V. 1973. Phytogeography of the fungal genus Lycoperdon in relation to the
opening of the Atlantic. Nature 242:123-125.
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Denison, W. C, and G. C. Carroll. 1966. The primitive Ascomycete: a new look at an old
problem. Mycologia 58:249-269.
Deonier, D. L. 1971. A systematic and ecological study of nearctic Hydrellia (Diptera,
Ephydridae). Smithsonian Contr. Zool. no. 68: 1-147. Describes trap for collecting
specimens; reports Stigmatomyces on adults and larvae; concludes that Stigmatomyces
may have killed several larvae (pp. 20,45-46).
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genus. Mentions supposed fungus on human louse.

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MYCOLOGIA MEMOIR NO. 9
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**Fassiatova, Olga, and M. Fassati. 1956. Prlspevek k poznani nasich zastupcu
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Outlines sequence of division in the Laboulbeniaceae; compares it with other Ascomy-
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Laboulbenia; discusses relation to host and moisture relations.
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Laboulbenia to subspecies. Reports Corethromyces (in part as Stigmatomyces), Dimeromyces,
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Lotsy, J. P. 1907. Vortrage iiber botanische Stammesgeschichte. Band 1: Algen und
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Corethromyces (in part as Sphaleromyces, Eucorethromyces), Dioicomyces, Euzodiomyces,
Helodiomyces, Herpomyces, Hydraeomyces (in part as Parahydraeomyces),
Laboulbenia, Misgomyces, Monoicomyces, Peyritschiella (as Dichomyces), Rhachomyces.
Host index.
—. 1916b. Sur quelques Laboulbeniales. Bull. Soc. Hist. Nat. Afrique N. 7: 100-
104. New species: Cantharomyces thaxteri; name change: Dimeromyces thaxteri for D.
falcatus Thaxt. non Paoli: reports Haplomyces, Laboulbenia, Misgomyces,
Peyerimhoffiella, Peyritschiella (as Dichomyces).
—. 1916c. Sur une nouvelle laboulbeniale parasite des Scaphidiidae. Bull. Sci. France
Belgique 49: 290-296. New species: Rickiapeyerimhoffii.
—. 1920. Troisieme contribution a l'etude des Laboulbeniales de l'Afrique du Nord. Bull.
Soc. Hist. Nat. Afrique N. 11: 123-138, 143-170. New taxa in Autophagomyces
(as Cryptandromyces), Botryandromyces (as Misgomyces), Cantharomyces,
Dimeromyces (see Balazuc, 1971f)> Dioicomyces, Euphoriomyces (as Ecteinomyces),
Laboulbenia, Teratomyces. Reports Amorphomyces, Chitonomyces, Corethromyces,
Dimeromyces, Dioicomyces, Eumonoicomyces, Herpomyces, Laboulbenia,
Misgomyces, Monoicomyces, Peyritschiella, Rhachomyces, Rickia, Stichomyces,
Stigmatomyces. Discusses Mimeomyces (as Corethromyces).

402
MYCOLOGIA MEMOIR NO. 9
**—. 1945. Etudes mycologiques. Fasc. 5. Bull. Soc. Hist. Nat. Afrique N. 36: 24-42.
Name change for Laboulbeniapicardii Lepesme, 1940: L. lepesmei (p. 40).
—. 1948. Contributions a l'etude de la flore de 1'Afrique du Nord. Fasc. 35. Bull. Soc.
Hist. Nat. Afrique N. 39:129-137. ReportsDioicomyces.
Maire, R., and R. G. Werner. 1937. Fungi maroccani. Catalogue raisonne des
champignons connus jusqui'ici au Maroc. Mem. Soc. Sci. Nat. Maroc 45: 1-147. Lists species of
Laboulbeniales in Morocco.
Majewski, T. 1972a. Rare and new Laboulbeniales from Poland. Acta Mycol. 7: 269-277.
New species in Eusynaptomyces (as Rhynchophoromyces). Reports Asaphomyces,
Botryandromyces (as Misgomyces), Chaetarthriomyces, Coreomyces, Ecteinomyces
(as Misgomyces), Fanniomyces (as Stigmatomyces), Hydrophilomyces (as
Misgomyces), Laboulbenia, Rhynchophoromyces.
—. 1972b. Rare and new Laboulbeniales from Poland. II. Acta Mycol. 8: 229-237. New
genus Fanniomyces. New species, reports in Stigmatomyces.
—. 1973a. Rare and new Laboulbeniales from Poland. III. Acta Mycol. 9: 111-124. New
species in Dioicomyces, Diphymyces (as Corethromyces), Siemaszkoa (as Misgomyces),
Stichomyces. Reports Asaphomyces, Laboulbenia, Rhachomyces.
—. 1973b. The genus Coreomyces Thaxter (Laboulbeniales) in Poland. Acta Mycol. 9:
217-228. Discusses species in Poland; includes key.
—. 1973c. Rare and new Laboulbeniales from Poland. IV. Acta Mycol. 9: 229-238. New
species in Autophagomyces, Phaulomyces (as Euphoriomyces). Reports Acomp-
somyces (as Stigmatomyces), Compsomyces, Euzodiomyces, Peyritschiella (as Dicho-
myces), Rhachomyces.
—. 1974. Rare and new Laboulbeniales from Poland. V. Acta Mycol. 10: 267-282. New
species in Cucujomyces, Distolomyces (as Hesperomyces), Hydrophilomyces, Rickia.
Reports Herpomyces, Rickia, Stigmatomyces, Symplectromyces.
—. 1981. Rare and new Laboulbeniales from Poland. VI. Acta Mycol. 16(1980): 141-153.
New genera: Tavaresiella, Triceromyces; new species in Euphoriomyces. Reports
Corethromyces, Dimeromyces, Euphoriomyces (as Asaphomyces), Herpomyces,
Laboulbenia.
—. 1982. Rare and new Laboulbeniales from Poland. VII. Acta Mycol. 17(1981): 53-62.
New species in Dipodomyces, Rickia. Reports Chitonomyces, Dimeromyces,
Laboulbenia, Sphaleromyces, Stigmatomyces.
Manfredi, Paoia. 1931. Un nuovo miriapodo cavernicolo italiano Trogloiulus mirus
n. gen., n. sp. Atti Soc. Hal. Sci. Nat. [Milano] 70: 181-189. Mentions frequency of
Troglomyces infection and location on host.
Martin, G. W. 1961. Key to the families of fungi. Pp. 497-517. In: Ainsworth, G. C,
Ainsworth & Bisby's Dictionary of the Fungi. Fifth edition. Commonwealth Myco-
logical Institute, Kew, Surrey.
**Matruchot, L. 1899. Revue des travaux sur les champignons publies en 1894, 1895,
1896, et 1897 (suite). Ascomycetes. I. — Laboulbeniacees. Rev. Gen. Bot. 11: 471-
484. Review of Thaxter's work on Laboulbeniales.
**—. 1900. Revue des travaux sur les champignons publies en 1894, 1895, 1896, et
1897 (suite). Ascomycetes. I. — Laboulbeniacees. Rev. Gen. Bot. 12: 25-31.
Mayer, K. 1934. Die Metamorphose der Ceratopogonidae (Dipt.). Ein Beitrag zur
Morphologie, Systematik. Okologie und Biologie der Jugendstadien dieser Dipteren-
familie. Arch. Naturgesch. (n. f.) 3: 205-288. Reports possible Laboulbeniales on
Forcipomyia, a midge (suborder Nematocera) (p. 286).
TAVARES: BIBLIOGRAPHY
403
Mayr, G. 1853. Abnorme Haargebilde an Nebrien und einige Pflanzen Krains. Verh.
Zool.-Bot. Ges. Wien 2: 75-77. One of first records of Laboulbeniales; not recognized
as an organism.
McDonald, K. 1972. The ultrastructure of mitosis in the marine red alga Membranoptera
platyphylla. J. Phycol. 8: 156-166.
McKittrick, Frances A. 1964. Evolutionary studies of cockroaches. Mem. Cornell Univ.
Agric. Exp. Sta. No. 389:1-197.
**Meijer, J. 1975. Carabid (Coleoptera, Carabidae) migration studied with
Laboulbeniales (Ascomycetes) as biological tags. Oecologia 19: 99-103. Reports: Laboulbenia,
Misgomyces. Compared % of infected beetles in newly reclaimed polder with % in
adjacent areas; results support view that migration usually occurs in early adult life.
Meola, Shirlee, and Joyce DeVaney. 1976. Parasitism of Mallophaga by Trenomyces
histophtorus. J. Invert. Pathol. 28: 195-201. Describe structure of haustoria, their
effect on fat body, muscles; color photographs of stained section of thallus on host.
Meola, Shirlee, and Isabelle I. Tavares. 1982. Ultrastructure of the haustorium of
Trenomyces histophthorus and adjacent host cells. J. Invert. Pathol. 40:205-215.
Mercier, L., and R. Poisson. 1927. Une Laboulbeniale, Stigmatomyces ephydrae n. sp.,
parasite d'Ephydra riparia Fall. (Dipt. Ephydridae). Bull. Soc. Zool. France 52:
225-231. Discuss effect on host, Laboulbeniales on European Diptera.
Merisuo, A. K. 1944. Notulae mallophagologicae. I. Ann. Entomol. Fenn. 10: 198-
226. Mentions presence of Trenomyces (p. 212).
Merola, A. 1952. Interessante ritrovamento di labulbeniologia cavernicola: Arth-
rorhynchus acrandros n. sp. (con considerazioni sul gen. Arthrorhynchus). Bol. Soc.
Naturalisti Napoli 60 (1951), Suppl. (Stud. Speleol. Faunist. Ital. Merid.) no. 16: 1-
30. Tav. I. Compares with other species; includes key to species of genus: discusses
geographical distribution, habitats, haustoria, spore discharge.
—. 1953. Unisessualita di una Labulbeniacea omotallica e suoi rapporti con il dioicismo
della Labulbeniacee eterotalliche. Delpinoa (Bull. Orto Bot. Univ. Napoli, n.s.) 6:
61-92. Concludes that absence of appendage on thallus of Arthrorhynchus acrandros
resulted from atrophy of upper cell of receptacle; proposes that dioecious taxa are
derived from monoecious taxa and are more closely related to them than they are to
one another.
Meschinelli, A. 1892. Fungi fossiles. Pp. 741-808. In: P. A. Saccardo. Sylloge Fungorum.
Vol. X. Supplementum Universale Pars. II. — Discomyceteae — Hyphomyceteae.
Published by the author. Padua.
Middelhoek, A. 1941. Dichomycesprinceps Thaxter. Fungus 12: 56-57. Report in
Netherlands (now in Peyritschiella).
—. 1942. Een nieuwe Laboulbeniacae voor ons land. Fungus 13: 52-53. Report of
Rhachomyces from Netherlands.
—. 1943a. Enige nieuwe Laboulbeniales voor ons land. Fungus 14: 57-59. Report of
Peyritschiella (as Dichomyces), Monoicomyces, Symplectromyces from Netherlands.
—. 1943b. Parasitaire Keverschimmels uit Zuid-Limburg. Natuurhist. Maandbl.
(Maandbl. Natuurhist. Genootsch. Limburg) 32: 58-60. Reports Monoicomyces
(as Eumonoicomyces), Rhadinomyces, Symplectromyces; includes list of species found
in Netherlands.
—. 1943c. Enige nieuwe Laboulbeniales voor ons land (Vervolg.) Fungus 14: 71-72.
Reports Euzodiomyces, Rhachomyces.

404
MYCOLOGIA MEMOIR NO. 9
—. 1943d. Laboulbeniaceae in Nederland. Ned. Kruidk. Arch. 53: 86-115. New species
in Laboulbenia, Mimeomyces; reports Corethromyces, Euzodiomyces, Haplomyces,
Laboulbenia, Misgomyces, Monoicomyces, Peyritschiella (in part as Dichomyces),
Rhachomyces, Rhadinomyces, Symplectromyces, Teratomyces.
—. 1945. Twee keverschimmels op een gastheer. Fungus 16: 6-8. Reports Corethromyces,
Laboulbenia on Stilicus (syn. of Rugilus).
—. 1947a. Laboulbeniaceae in Nederland. II. Ned. Kruidk. Arch. 54: 232-239.
Reports Laboulbenia, Idiomyces, Monoicomyces, Peyritschiella (in part as
Dichomyces'), Rhachomyces.
**—. 1947b. Wij en de keverschimmels. Natura 44: 89-93. Popular discussion, including
comments on trichogynes and spermatia, structure, relation to host, examining
specimens; short host list.
—. 1949. Laboulbeniaceae in Nederland. III. Ned. Kruidk. Arch. 56: 249-260. New
genus: Barbariella (now Asaphomyces); reports Cantharomyces, Laboulbenia.
—. 1951. About some interesting variations in the genus Laboulbenia. Biol. Jaarb.
(Dodonaea) 18: 122-129. Discussion of abnormal development in Laboulbenia proli-
ferans.
—. 1957. Eine neue Gattung der Laboulbeniales. Fungus 27: 72-75. New genus: Schizo-
laboulbenia (syn. of Laboulbenia).
Mirande, M. 1905. Sur une nouvelle fonction du tegument des Arthropodes considere
comme organe producteur de sucre. Arch. Anat. Microscop. 7: 232-238.
Misra, P. C. and P. H. B. Talbot. 1964. Phialomyces, a new genus of Hyphomycetes.
Canad. J. Bot. 42:1287-1290. PI. I.
Moesz, G. 1931. Mykolbgiai kozlemenyek VIII. Kozlemeny. (A Laboulbeniaceae
csalad ket faja Magyarorszagban.) Bot. Kozlem. 28: 161-174. Reports Arthrorhynchus
(Helminthophana), Teratomyces from Hungary. (German translation, pp. 170-174.)
Moller, A. 1901. Phycomyceten und Ascomyceten. Heft 9. Pp. 1-319. In: Botanische
Mittheilungen aus den Tropen. Ed. A. F. W. Schimper. Gustav Fischer, Jena.
Discusses lack of proof of sexual function in Laboulbeniales, mentions other Ascomycetes
with conidia similar to spermatia of Laboulbeniales (pp. 45-46).
Montagne, J. F. C. 1856. Sylloge Generum Specierumque Cryptogamarum. J.-B. Bail-
liere, Paris. 498 p. Places Laboulbenia in Pyrenomycetes.
Miiller, T. 1927. Beobachtungen iiber die Mallophagen der Frischen Nehrung. Ber. Ver-
samml. Westpreuss. Bot.-Zool. Vereins Danzig 49:1-43. Mentions discoloration of host
that could have resulted from the presence of Laboulbeniaceae (see pp. 15-17).
—. 1932. Erganzungen zu den Beobachtungen tiber die Mallophagen der Frischen
Nehrung mit Berilcksichtigung ihrer Parasiten. Ber. Versamml. Westpreuss. Bot.-Zool.
Vereins Danzig 54: 17-37. Brief discussion of presence of Laboulbeniales on Mallo-
phaga; new records (pp. 35-37).
Nannizzi, A. 1934. Repertorio Sistematico dei Miceti dell'Uomo e degli Animali.
Pp. 1-557. Vol. IV. In: Trattato di Micopatologia Umana. Ed. G. Pollacci. S. A.
Poligrafica Meini. Siena. Establishes family Zodiomycetaceae.
Nowasad, A. 1973. Arthrorhynchus nycteribiae (Peyritsch) Thaxter (Ascomycetes,
Laboulbeniales) w Polsce. Arthrorhynchus nycteribiae (Peyritsch) Thaxter
(Ascomycetes, Laboulbeniales) in Poland. Polskie Pismo Entomol. 43: 423-430. Includes
photograph.
TAVARES: BIBLIOGRAPHY
405
Olive, L. S. 1958. On the evolution of heterothallism in fungi. Amer. Naturalist 92: 233-
251. Points out lack of evidence that dimorphism in Laboulbeniales is genetically
determined; suggests it may have a physiological basis.
—. 1966. Sexual dimorphism in the Laboulbeniales. Mycologia 58: 478-479. Suggests
that dimorphic species could have been derived from heterothallic forms with
monoecious thalli and that there may be both homothallic and heterothallic monoecious
species.
Otani, Y. 1957. How to collect the Laboulbeniales. Trans. Mycol. Soc. Japan 1(5): 8-10.
Paper in Japanese. Lists undescribed species of Laboulbenia and Rhachomyces
(species illustrated by Ishikawa, 1948). Mentions Zodiomyces.
Pacheco M., F. 1964. Sistematica, Filogenia y Distribucibn de los Heteroceridos
de America (Coleoptera: Heteroceridae). Monogr. Col. Post.-grad. no. 1. Escuela
Nacional de Agricultura, Colegio de Post-graduados, Chapingo, Mexico. 155 p.
Figs. 1-501.
Paoli, G. 1911. Nuovi Laboulbeniomiceti parassiti di Acari. Redia 7: 283-295. Tav. XII.
New species in Dimeromyces, Rickia (?minuta based on immature material); transfers
Rhachomyces berlesianus to Rickia.
—. 1912. Nuovi Laboulbeniomiceti parassiti di Acari. Malpighia 24: 329-340. Tav. V.
Same as 1911 reference.
Parriaud, H. 1964. Laboulbeniales de la zone littorale du Bassin d'Arcachon. Proces-
verb. Soc. Linn. Bordeaux 101: 73-77. Reports Haplomyces, Laboulbenia,
Peyritschiella on hosts in saline soil of littoral zone.
**Pasquet, O. 1909. Nouvelles especes de Laboulbeniacees. Bull. Soc. Sci. Med. Ouest
[Rennes] 18: 166-169. List of species found in northwestern France (no new records).
Patton, R. L. 1963. Introductory Insect Physiology. W. B. Saunders, Philadelphia.
245 p.
Paulian, R. 1943. Les Coleopteres, formes — moeurs — role. (Bibliographie Scienti-
fique.) Payot, Paris. 396 p.
Pearse, A. S., Marguerite T. Patterson, J. S. Rankin, and G. W. Wharton. 1936. The
ecology of Passalus cornutus Fabricius, a beetle which lives in rotting logs. Ecol.
Monogr. 6:455-490.
Peck, C. H. 1885. Report of the botanist. New York State Bot. Rep. 38: 77-138. Pis.
1-3. New genus (as Deuteromycete): Appendicularia (syn. of Stigmatomyces) (pp.
95-96, pi. 3).
**Petch, T. 1944. Notes on entomogenous fungi. Trans. Brit. Mycol. Soc. 27: 81-93.
Comments on British records of Laboulbeniales.
Peyerimhoff, P., de. 1910. Nouveaux Coleopteres du Nord-Africain (onzieme note:
faune cavernicole du Djurdjura). Bull. Soc. Entomol. France. 1910: 149-154. Reports
Rhachomyces.
Peyritsch, J. 1871. ttber einige Pilze aus der Familie der Laboulbenien. Sitzungsber.
Kaiserl. Akad. Wiss., Math.-Naturwiss. CI., Abt. 1 [Wien] 64: 441-458. Taf. .1-11.
New taxa in Laboulbenia. Discusses relationship of L. muscae to host and sexuality of
the fungus. Includes discussion of Kolenati's species of Arthrorhynchus. English
summary: On some fungi belonging to the family Laboulbeniae. Ann. Mag. Nat. Hist,
(ser. 4)8:440-441. 1871.
—. 1873. Beitrage zur Kenntniss der Laboulbenien. Sitzungsber. Kaiserl. Akad.
Wiss., Math-Naturwiss. CI., Abt. 1 [Wien] 68: 227-254. Taf. I-III. Places taxa in

406 MYCOLOGIA MEMOIR NO. 9
Laboulbeniaceae. New genera: Chitonomyces, Heimatomyces (now Chitonomyces), <
Helminthophana (now Arthrorhynchus). Includes Stigmatomyces. New species of
Laboulbenia. All known species listed except L. pilosella (now Rhachomyces).
General discussion of relationship to host, distribution, sexuality.
—. 1875. Uber Vorkommen und Biologie von Laboulbeniaceen. Sitzungsber. Kaiserl.
Akad. Wiss., Math.-Naturwiss. CI., Abt. 1 [Wien] 72: 377-385. Extensive discussion i
of biology. Infection experiments described; information about collecting. Lists
parasitized beetles collected. Mentions undescribed species: Laboulbenia gracilis.
Considers fungi on Haliplus and Hydrobius to be possible new genera. J
Picard, F. 1908a. Sur une Laboulbeniacee marine {Laboulbenia marina n. sp.), parasite
d'Aepus Robini Laboulbene. Compt.-Rend. Hebd. Seances Mem. Soc. Biol. [Paris]
65:484-486. Discusses possible manner of obtaining nutrients.
—. 1908b. Les Laboulbeniacees et leur parasitisme chez les insectes. Feuille Naturalistes
39: 29-34. PI. III. Popular account; includes comment on nutrition. J
—. 1909. Sur une Laboulbeniacee nouvelle {Hydrophilomyces digitatus n. sp.) parasite
d'Ochtebius marlnus Paykull. Bull. Soc. Mycol. France 25: 245-249.
—. 1912. Description de deux Laboulbeniacees nouvelles, parasites de Coleopteres.
Bull. Soc. Entomol. France 1912: 178-181. New species in Cantharomyces, Dtoico-
myces (now in Picardella).
—. 1913a. Sur une Laboulbeniacee nouvelle, parasite de Stenus aceris Steph. Bull. Soc.
Entomol. France 1913:462-465. New species: Acallomyces lavagnei (now Ilyomyces). <
—. 1913b. Contribution a 1'etude des Laboulbeniacees d'Europe et du nord de 1'Afrique.
Bull. Soc. Mycol. France 29: 503-571. Pis. XXIX-XXXII. New genus: Helodlomyces.
New species in Autoicomyces (as Ceratomyces), Laboulbenia, including L. pasquetii,
Misgomyces. Reports Arthrorhynchus, Cantharomyces, Chitonomyces, Compsomyces,
Coreomyces, Corethromyces (in part as Eucorethromyces), Dimeromyces, Euhaplo- '
myces, Euzodiomyces, Haplomyces, Herpomyces, Hydraeomyces, Hydrophilomyces,
Idiomyces, Laboulbenia, Misgomyces, Monoicomyces, Peyritschiella (in part as Dicho-
myces), Picardella (as Dioicomyces), Polyascomyces, Rhachomyces, Rhadinomyces,
Rickia, Sphaleromyces, Stigmatomyces, Symplectromyces, Teratomyces, Trenomyces,
Zodlomyces. Corrects orthography of Laboulbenia pseudomasei Thaxt., 1899.
Discusses abnormalities in L. proliferans Thaxt., 1893. Host index. '
—. 1917. Sur quelques Laboulbeniales d'Europe. Bull. Sci. France Belgique 50: 440-460.
PI. VI. New genus: Ilyomyces; new species in Cantharomyces, Laboulbenia (in part
now in Scalenomyces), including L. lichtensteinii, Rhachomyces (in part as
Dimeromyces), Stigmatomyces (now StemmatomycesT). Reports Cantharomyces,
Compsomyces, Euzodiomyces, Helodiomyces, Laboulbenia, Monoicomyces, Peyritschiella
(inpart as Dichomyces), Rickia, Teratomyces.
Poelt, J. 1952a. Laboulbeniales aus Sudbayern. Mitt. Bot. Staatssamml. Munchen 1: |
115-118. New species: Laboulbenia buehlmannii; reports Euzodiomyces, Laboulbenia,
Monoicomyces, ?Peyerimhoffiella (genus not named); Peyritschiella (as Dichomyces).
—. 1952b. Laboulbenien und ihr Vorkommen in Sudbayern. Nachrichtenbl. Bayer.
Entomol. 1: 33-36. General account with host index (gives name and locality of host of
unnamed genus of 1952a). .
Poisson, R. 1929. Contribution a la connaissance des Laboulbeniales parasites des insectes
hemipteres hydrocorises. Paracoreomyces Thaxteri gen. nov., sp. nov., laboulbeniale
parasite de Stenocorixa protrusa Horv. Compt. Rend. Hebd. Seances Acad. Sci.
[Paris] 188: 824-826.
f
TAVARES: BIBLIOGRAPHY
407
—. 1930. A propos de l'insertion superficielle de certaines Laboulbeniales sur leurs hfltes
et sur la presence en Normandie de Laboulbenia fasciculata Peyrit[s]ch (= brachiata
Thaxter) parasite de Leistus fulvibarbis Dej. (Coleopt. Carabidae). Bull. Soc. Linn.
Normandie (ser. 8) 2 (1929) (Trav. orig.): 65-68. Reports haustoria in Laboulbenia;
general discussion of occurrence in Laboulbeniales.
—. 1954. Sur une Laboulbeniale ectoparasite de Velia osborniana Kirkaldy (Hemipt.
Veliidae) Laboulbenia (s. g. Veliomyces nov.) titschacki n. sp. Pp. 81-82. In: Beitrftge
zur Fauna Perus. Vol. 4. Ed. E. Titschack. Gustav Fischer, Jena.
—. 1957. Faune de France. 61. Heteropteres aquatiques. P. Lechevalier, Paris. 263 p.
Reports Autophagomyces, Coreomyces (in part as Paracoreomyces); new taxon (nom.
nud.): Autophagomyces mesoveliae (mention only, p. 16).
Pont, A. C. 1972. Family Muscidae. no. 97: 1-111. In: A Catalogue of the Diptera of
the Americas south of the United States. Museu de Zoologia, Univ. SaO Paulo, Brazil.
**Pope, F. M. 1891. Micro-organisms in their relations to the higher animals. Trans.
Leicester Lit. Soc. (ser. 3) 2: 256-262. Effect of Stigmatomyces on host confused with
that of Empusa muscae (Fr.) Cohn (p. 259).
Popham, E. J. and A. Brindle. 1966. Genera and species of the Dermaptera 3. Carci-
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—, and —. 1967. Genera and species of the Dermaptera 5. Spongiphorinae and
Labiinae. Entomologist 100: 255-262.
Price, P. W. 1975. Insect Ecology. John Wiley, New York. 514 p. (See Chapt. 21, Paleo-
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Princis, K. 1969. Blattariae: Subordo Epilamproidea. Fam.: Blattellidae. Pars 13.
Pp. 713-1038. In: Orthopterorum Catalogus. Ed. M. Beier. W. Junk, The Hague.
Racovitza, E. G. 1908. IX. Isopodes terrestres (seconde serie). Biospeologica. Arch.
Zool. Exp. Gen. 39: 239-415. Pis. IV-XXIII. Specimens on isopod possibly
Laboulbeniales, according to author (cf. Amphoromorpha, Thaxter, 1920a) (pp. 271-272).
Radford, C. D. 1950. Systematic Check List of Mite Genera and Type Species. Union
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Raper, J. R. 1959. Sexual versatility and evolutionary processes in fungi. Mycologia
51:107-124. Mentions heterothallism in Laboulbeniales.
Richards, A. G. 1951. The Integument of Arthropods, the Chemical Components
and their Properties, the Anatomy and Development, and the Permeability. Univ.
Minnesota Press, Minneapolis. 411 p.
—. 1954. Similarities in histochemical differentiation of insect cuticle and the walls of
parasitic fungi. Science 120:761-762.
Observations on Herpomyces.
Richards, A. G., and Myrtle N. Smith. 1954. Infection of cockroaches with Herpomyces
(Laboulbeniales). III. Experimental studies on host specificity. Bot. Gaz. [Crawfords-
ville] 116: 195-198. Cross-inoculation by mixing of host species resulted in new host
records; species shown to be stable.
—, and —. 1955a. Infection of cockroaches with Herpomyces (Laboulbeniales). I.
Life history studies. Biol. Bull. Mar. Biol. Lab. Woods Hole 108:206-218. Herpomyces
stylopygae on Blatta orientalis: distribution on host, spore discharge, volume
increase of thallus during growth. Comments on H. ectobiae and H. paranensis (as H.
tricuspidatus).

408
MYCOLOGIA MEMOIR NO. 9
—, and —. 1955b. Infection of cockroaches with Herpomyces (Laboulbeniales). IV.
Development of H. stylopygae Spegazzini. Biol. Bull. Marine Biol. Lab. Woods Hole
109:306-315. Development of male and female thalli.
—, and —. 1956. Infection of cockroaches with Herpomyces (Laboulbeniales). II.
Histology and histopathology. Ann. Entomol. Soc. Amer. 49: 85-93. Herpomyces stylopygae
on Blatta orientalis: fungus cell wall structure, characteristics of haustoria. Brief
comments on haustoria of H ectobiae and H. paranensis (misidentified as H. tricus-
pidatus). Effect of infection on host; loss at molting.
Rick, J. 1903. Zur Pilzkunde Vorarlbergs. V. Oesterr. Bot. Z. 53: 159-164. Reports
, Rickia wasmannii; comments on possible symbiotic relationship.
Riek, E. F. 1970. Fossil History. Pp. 168-186. In: The Insects of Australia, a Textbook
for Students and Research Workers. Div. Entomol., Commonw. Sci. Industr. Res.
Organ., Canberra. (Melbourne Univ. Press.)
Robin, C. P. 1852. Vegetaux parasites sur un insecte du genre Brachynus. Compt. Rend.
Hebd. Seances Mem. Soc. Biol. [Paris] (ser. 1) 4: 11. One of earliest records of
Laboulbenia.
—. 1853. Histoire Naturelle des Vegetaux Parasites qui Croissent sur l'Homme
et sur les Animaux Vivants. J.-B. Bailliere, Paris, One vol., 702 p. Atlas, 24 p., pis.
I—XV. New genus: Laboulbenia. Illustrations; discussion, which includes some
errors in interpretation (pp. 622-639, Pis. VIII-X).
—. 1871. Traite du Microscope. J. B. Bailliere et fils, Paris. 1028 p. New taxon:
Laboulbenia pilosella (now Rhachomyces); brief comment accompanying excellent
illustration, pp. 912-913.
Rogers, J. D. 1965. The conidial stage of Coniochaeta ligniaria: morphology and
cytology. Mycologia 57: 368-378.
Rogers, W. P. 1962. The Nature of Parasitism, the Relationship of some Metazoan
Paiasites to their Hosts. (Vol. 2, Theoretical and Experimental Biology, an
International Series of Monographs, ed. J. F. Danielli.) Academic Press, New York,
London. 287 p.
Roquebert, Marie-France, and M. Abadie. 1973. Etude ultrastructurale de la sporogenese
chez un micromycete: Stilbothamnium nudipes Haum. Compt. Rend. Hebd. Seances
Acad. Sci. [Paris] (ser. D) 276:2883-2885.
Rossi, W. 1974. Una nuova specie di Laboulbenia (Ascomycetes, Laboulbeniales)
parassita di termiti. Doriana [Suppl. Ann. Mus. Civ. Storia Nat. "G. Doria" Genova]
5(215): 1-5. New taxon: Laboulbenia felicis-caprae (correction made in CMI Index of
Fungi.Pt. 11, Jan. 1976).
—. 1975. Su alcune Laboulbeniali (Ascomycetes) nuove per L'ltalia. Giorn. Bot.
Ital. (Nuovo Giorn. Bot. Ital. n. s.) 109: 71-85. Reports Asaphomyces, Corethro-
myces, Euzodiomyces, Helodiomyces, Laboulbenia, Misgomyces, Monoicomyces,
Peyritschiella, Rhachomyces, Rhadinomyces (as Corethromyces). Host index.
Photographs.
—. 1977. Pseudoecteinomyces, a new genus of Laboulbeniales (Ascomycetes). Mycologia
69:1073-1076.
—. 1978a. Due nuove Laboulbeniali della Sierra Leone (Ascomycetes). Natura; Rivista
Sci. Nat. [Milano] 69: 17-22. New taxa: Laboulbenia schulleri (probably Dixomyces),
Rickia leonis. Reports hyperparasite (near Monilid) on Rickia.
—. 1978b. Sulle Laboulbeniali (Ascomycetes) parassite dei Trechini di Turchia (Coleop-
tera, Carabidae). Pp. 1-8. In: Fauna Ipogea di Turchia. Eds. V. Sbordoni, A.
TAVARES: BIBLIOGRAPHY
409
Vigna Taglianti. Quad. Speleol., Circolo Speleol. Romano. Vol. 3. New taxa:
Laboulbenia vignae, Rhachomyces gratiellae. Reports Laboulbenia, Rhachomyces.
—. 1978c. Une espece inedite de Rhachomyces (Ascomycetes, Laboulbeniales) parasite
du Carabique troglobie, Speagonum mirabile Moore de la Nouvelle-Guinee (resultats
zoologiques de l'expedition sp&eologique britannique en Papouasie-Houvelle Guinee,
1975, 3). Int. J. Speleol. 9: 365-368.
—. 1979. Sui Rhachomyces (Ascomycetes, Laboulbeniales) Darassiti dei Duvalius
italiani (Coleoptera, Carabidae, Trechini). Int. J. Speleol. 10(1978): 323-330. Validates
Rhachomyces maublancii; reports R. stipitatus. Discussion.
—. 1980. On two Laboulbeniales (Ascomycetes) parasitic on Histeridae (Insecta,
Coleoptera). Mycologia 72: 430-433. New taxon: Histeridomyces europaeus; reports Homa-
romyces.
—. 1981. Una nuova specie de Monoicomyces (Ascomycetes, Laboulbeniales). Candollea
36: 375-378.
—. 1982a. Aphanandromyces, a new genus of Laboulbeniales (Ascomycetes).
Mycologia 74: 520-523.
**_. 1982b. Two new species of Laboulbenia (Ascomycetes, Laboulbeniales) from
Asia. Kew Bull. 37:69-71. PI. 2.
—.1982c. New or interesting Laboulbeniales from China. Mycologia 74: 1023-1026.
New taxa in Laboulbenia, Rhachomyces. Photograph of L. manubriolata Thaxter.
Rossi, W., and J. Balazuc. 1977. Laboulbeniales parasites de Myriapndes. Rev. Mycol.
[Paris] 41: 525-535. New genus: Diplopodomyces. New species in Rickia. Extensive
discussion of Laboulbeniales found on Diplopoda.
Rossi, W., and M. Graziella Cesari. 1974. Due nuove specie di Rhachomyces
(Ascomycetes, Laboulbeniales), parassiti di Trechini italiani (Coleoptera, Carabidae).
Atti Soc. Ital. Sci. Nat. [Milano] 115: 175-180. New species: Rhachomyces bucciarellii,
R. vignae.
—, and —. 1976. Contributo alia conoscenza della Laboulbeniali (Ascomycetes) parassite
di Carabidi italiani (Insecta, Coleoptera). Giorn. Bot. Ital. 110: 145-153. New species
in Laboulbenia, Rhachomyces. Reports Corethromyces, Dimeromyces, Laboulbenia.
Photographs.
Rossi, W., and M. Graziella Cesari Rossi. 1977a. Due nuove specie di Dimeromyces
(Laboulbeniales). Rivista Parassitol. 38: 109-113. New species in Dimeromyces from
Italy, Africa.
—, and —. 1977b. Deux Laboulbeniales (Ascomycetes) nouvelles, parasites de Zuphiini
(Coleoptera, Carabidae). Canad. J. Bot. 55: 1575-1578. New species in Eucantharo-
myces, Laboulbenia from Africa. Photographs.
—, and —. 1977c. Sulle Laboulbeniali (Ascomycetes) parassite dei Trechinae del
Messico (Coleoptera, Carabidae). Pp. 373-376. Tav. I. In: Subterranean Fauna of
Mexico. Part III. Further results of the Italian zoological missions to Mexico,
sponsored by the National Academy of Lincei (1973 and 1975). Quaderno Accad.
Naz. Lincei (Probl. Attual. Sci. Cult., Sez.: Missioni Esplor. — I.) No. 171. New
species: Laboulbenia sbordonii; reports Laboulbenia, Rhachomyces. Photographs.
—, and —. 1978. Contributo alia conoscenza delle Laboulbeniali (Ascomycetes)
parassite di Stafilinidi italiani (Insecta, Coleoptera). Giorn. Bot. Ital. 112: 63-74.
New species in Corethromyces, Diplomyces, Rickia. Reports Cantharomyces,
Compsomyces, Idiomyces, Monoicomyces, Symplectromyces, Teratomyces.
Photographs.

410
MYCOLOGIA MEMOIR NO. 9
—, and —. 1979a. Trois Laboulbeniales (Ascomycetes) nouvelles, parasites de
Dipteres. Canad. J. Bot. 57:993-996. New species in Stigmatomyces from Africa.
**—, and —. 1979b. Due species nuove di Laboulbenia (Ascomycetes Laboulbeniales)
parassite di Chrysomelidae (Insecta Coleoptera). Natura; Rivista Sci. Nat.
[Milano] 70: 89-93. New species in Laboulbenia (Italy, Peru).
—, and —. 1979c. Su alcune specie di Stigmatomyces (Ascomycetes, Laboulbeniales)
parassite di Ditteri italiani. Boll. Mus. Civico Storia Nat. Venezia 30: 13-17.
Figs. 1-7.
—, and —. 1979d. Tre nuove specie di Stigmatomyces (Ascomycetes, Laboulbeniales)
parassite di Ditteri italiani. Giorn. Bot. Ital. 113:379-385.
—, and —. 1980a. Su alcune Laboulbeniali (Ascomycetes) parassite di insetti
acquatici italiani. Rivista Idrobiol. 19:147-152. Report Autoicomyces, Cantharomyces,
Helodiomyees, Hydrophilomyces, Rhynchophoromyces, Zodiomyces.
—, and —. 1980b. Nuovo contributo alia conoscenza delle Laboulbeniali (Ascomycetes)
parassite di Stafilinidi italiani (Insecta, Coleoptera). Giorn. Bot. Ital. 114: 187-192.
New taxa in Camptomyces, Corethromyces. Reports Compsomyces, Haplomyces,
Smeringomyces, Teratomyces.
—, and —. 1981. Two new species of Rhachomyces (Ascomycetes, Laboulbeniales)
parasitic on Orotrechus (Coleoptera, Carabidae, Trechini). Mycologia 73:554-559.
—, and —. 1982. Quelques especes nouvelles de Laboulbeniales (Ascomycetes)
parasites de Carabiques. Canad. J. Bot. 60: 306-309. New species in Laboulbenia,
Rhachomyces.
Rossi, W., and A. Vigna Taglianti. 1979. Considerazioni sulle Laboulbeniali (Ascomy-
cets) [sic] parassite dei Duvalius italiani (Coleoptera, Carabidae, Trechini). Fragm.
Entomol. [Roma] IS: 7-15. Discussion of Laboulbenia vulgaris, Rhachomyces stipi-
tatus, R. maublancii — distribution of species, relationships of hosts.
Rostrup, O. 1916. Bidrag til Danmarks Svampeflora. I. Dansk. Bot. Ark. 2: 1-56.
Tav. I-III. First records for Denmark; reports Eumonoicomyces, Laboulbenia (pp.
10-11, tav. I, Figs. 2-3). (Cf. note [Bot. Not. 1933: 615] that O. Ryberg gave first
report of Laboulbeniales for Sweden and Denmark.)
Roth, L. M. and E. R. Willis. 1960. The biotic associations of cockroaches.
Smithsonian Misc. Collect. Vol. 141 (Smithsonian Inst. Publ. 4422). Washington, D. C.
470 p. Brief discussion of fungi; list of Herpomyces species (see pp. 127-129, 134-
138), including some new records.
Rouget, A. 1850. Notice sur une production parasite orservee [sic] sur le Brachinus
crepitans. Ann. Soc. Entomol. France (ser. 2) 8: 21-24. One of first reports of
occurrence of Laboulbeniales. (See also vol. 7 for mention of reading of paper by Desmarest,
p. LI; examination of specimens by Laboulbene, etc., pp. LV-LVI, LXIII at meeting.)
Ruffieux, L. 1904. I. Contribution a l'etude de la flore cryptogamique fribourgeoise.
Les champignons observes dans le Canton de Fribourg. Mem. Soc. Fribourg. Sci.
Nat. 1:165-214. Reports Laboulbenia, Stigmatomyces (pp. 202-203).
Ryberg, O. 1947. Studies on Bats and Bat Parasites, Especially with Regard to Sweden
and Other Neighbouring Countries of the North. I. Bokforlaget Svensk Natur,
Stockholm. 330 p. Reports Arthrorhynchus in Sweden and Denmark.
Samsinakova, Anna. 1960a. Novy nalez Rickia berlesiana (Bacc.) Paoli (Laboulbeniales).
Ein neuer Fund des Pilzes Rickia berlesiana (Bacc.) Paoli (Laboulbeniales). Ceska
Mykol. 14:49-52. Reports Rickia from India.
TAV ARES: BIBLIOGRAPHY
411
—. 1960b. Prispevek k poznani entomofytnlch hub na muchulovitych (Nycteribiidae).
Beitrage zur Kenntnis der entomophagen Pilze auf den Nycteribiiden. Zool. Listy
(ser. 2) 9: 237-238. Reports Arthrorhynchus (as Stigmatomyces nycteribiidarum) in
Poland.
—. 1968. Nalez houby Dimeromyces falcatus Paoli (Laboulbeniales) na novem hostiteli.
Fund des Pilzes Dimeromyces falcatus Paoli (Laboulbeniales) auf einem neuen
Wirt. Ceska Mykol. 22:225-228. Report on mite on carabid.
Sarna, Anna, and Jolanta Milewska. 1977. Laboulbeniales from Poland parasitizing
on semi-aquatic insects. Acta Mycol. 13: 301-311. New taxa in Hydrophilomyces,
Rhynchophoromyces. Report Autoicomyces, Cantharomyces, Chaetarthriomyces,
Chitonomyces, Helodiomyees, Hydraeomyces, Hydrophilomyces, Laboulbenia,
Rhynchophoromyces, Zodiomyces.
Savile, D. B. 0.1955. Aphylogeny of theBasidiomycetes. Canad. J. Bot. 33: 60-104.
Schaffner, J. H. 1909. The classification of plants, IV. Ohio Nat. 9: 446-455. Laboul-
benieae a class of phylum Nematophyta (algae and fungi).
Schellenberg, G. 1923. Ober die Laboulbeniaceen. Verh. Internat. Verein. Theor.
Angew. Limnol. 1: 311-313 (reviewed by H. Gams, Centralbl. Bakteriol. Abt. 2,
62: 575. 1927). Popular account; mentions collection on millepedes.
Scheloske, H.-W. 1969. Beitrage zur Biologie, Okologie und Systematik der
Laboulbeniales (Ascomycetes) unter besonderer Beriicksichtigung des Parasit-Wirt-Verhalt-
nisses. Parasitol. Schriftenreihe Heft 19: 1-176. New species in Chaetarthriomyces,
Dichomyces (now Peyritschiella), Eucantharomyces, Eusynaptomyces, Laboulbenia,
Misgomyces (now Siemaszkoa), Monoicomyces, Rhynchophoromyces, Rickia
(one on millepede), Stigmatomyces (now Acompsomyces). Reports Asaphomyces,
Autoicomyces (as Ceratomyces), Botryandromyces (as Misgomyces), Camptomyces,
Cantharomyces, Chitonomyces, Colonomyces, Compsomyces, Coreomyces,
Corethromyces, Ecteinomyces (as Misgomyces), Euzodiomyces, Helodiomyees,
Hydraeomyces, Hydrophilomyces (as Misgomyces), Laboulbenia, Misgomyces,
Monoicomyces, ?Peyerimhoffiella (as Corethromyces), Peyritschiella (in part as
Dichomyces), Rhachomyces, Rhynchophoromyces, Symplectromyces, Teratomyces,
Zodiomyces, and undetermined genus on Cercyon. Intensive ecological survey,
observations on relation to hosts, etc.
—. 1976a. Eusynaptomyces benjaminii, spec, nova, (Ascomycetes, laboulbeniales)
und seine Anpassungen an das Fortpflanzungsverhalten seines Wirtes Enochrus testa-
ceus (Coleoptera, Hydrophilidae). Plant Syst. Evol. 126: 267-285. Describes growth
form A on lower side of pronotum of both sexes and reduced growth form B on claws
of males; compares with E. enochri; discusses mating behavior; rejects concept of
sex-of-host specificity.
—. 1976b. Morphologische Anpassungen eines ektoparasitischen Pilzes (Ascomycetes:
Laboulbeniales: Misgomyces coneglanensis) an Korperbau und
Fortpflanzungsverhalten seines Wirtes (Coleoptera: Hydrophilidae: Laccobius minutus). Entomo-
logica Germ. 3: 227-241. Discusses two morphological forms that occupy different
positions on host, transfer of spores during mating.
Scheloske, H.-W., and D. Matthes. 1966. Ein parasitischer Pilz. Mikrokosmos 55: 314-
315. Reports Zodiomyces. Photograph.
Schornstein, Kathleen L., and J. Scott. 1980. Reevaluation of mitosis in the red alga
Porphyridiumpurpureum. Nature 283:409-410.
Schrantz, J.-P. 1967. Presence d'un aster au cours des mitoses de l'asque et de la
formation des ascospores chez l'Ascomycete Pustularia cupularis (L.) Fuck. Compt.
Rend. Hebd. Seances Acad. Sci. [Paris] (ser. D) 264:1274-1277. Pis. I-IV.

412
MYCOLOGIA MEMOIR NO. 9
Schubart, O. 1934. Tausendfussler oder Myriapoda. I. Diplopoda. 318 p. Teil 28. In:
Dahl, F. Die Tierwelt Deutschlands und der angrenzenden Meeresteile nach ihrem
Merkmalen und nach ihrer Lebensweise. Eds. Maria Dahl and H. Bischoff. Gustav
Fischer, Jena. Mentions Verhoeff's records.
—. 1945. Os Proterospermophora do Distrito Federal (Myriapoda, Diplopoda).
Arq. Mus. Nac. Rio de Janeiro 38: 1-156. Reports possible Laboulbeniales
(figure resembles imperfect fungus) (pp. 133-135).
Scott, H. 1914a. H. Sauter's Formosa = Ausbeute. Nycteribiidae. Arch. Naturgesch.
69(1913) (Abt. A) (8): 92-103. Mentions Laboulbeniaceae on Nycteribiidae from
Taiwan and Ceylon (pp. 94, 96).
—. 1914b. On some oriental Nycteribiidae [Diptera Pupipara]. Ann. Mag. Nat. Hist,
(ser. 8) 14: 209-235. Pis. X-XII. Mentions occurrence of Laboulbeniales on Nycteribia
parilis, etc. (pp. 213, 214, 234).
—. 1925. Zoo-geographical and systematic notes on the Nycteribiidae (Diptera Pupipara)
of India, Ceylon and Burma. Records Indian Mus. 27: 351-384. Includes table of
Nycteribiidae of India; mentions Laboulbeniales (pp. 362, 365), report by Falcoz
(p. 369).
—. 1936. Descriptions and records of Nycteribiidae (Diptera Pupipara), with a
discussion of the genus Basilia. J. Linn. Soc. Zool. 39: 479-505. Reports Laboulbeniales
on Nycteribia euxesta from Ceylon (p. 487).
Seevers, C. H. 1978. A generic and tribal revision of the North American Aleocharinae
(Coleoptera: Staphylinidae). Fieldiana: Zool. 71:1-VI, 1-289. (Publication 1282.)
Seguy, E. 1951. Ordre des Dipteres. Pp. 449-744. In: Traite de Zoologie, Anatomie,
Systematique, Biologie. Ed. P.-P. Grasse. Vol. 10. Insectes Superieurs et Hemipter-
oides. Premiere Fasc. NeVropteYoides, Mecopteroides, Hymenopteroides (Symphytes
et Ter^brants). Masson et Cie, Paris.
Shanor, L. 1952. The characteristics and morphology of a new genus of the
Laboulbeniales on an earwig. Amer. J. Bot. 39:498-504. New taxon: Filariomyces.
—. 1955. Some observations and comments on the Laboulbeniales. Mycologia 47: 1-12.
Extensive discussion of the Laboulbeniales.
Sharp, D. 1913. Coleoptera. III. Coleoptera Caraboidea. Pp. 175-292, Pis. VI-VII.
In: Fauna Hawaiiensis, being the Land Fauna of the Hawaiian Islands. Vol. 3.
Cambridge Univ. Press, Cambridge.
Shaw, S. 1952a. Some records of Laboulbeniales (Fungi) on Coleoptera. J. Soc.
Brit. Entomol. 4:116-117. Reports Laboulbenia, Rhachomyces.
—. 1952b. The entomology of Spurn Peninsula. 8. Coleoptera, Carabidae. Naturalist
[London] No. 843:170-173. Reports Laboulbenia.
Shoemaker, R. A. 1955. Biology, cytology, and taxonomy of Cochliobolus sativus.
Canad. J. Bot. 33: 562-576.
Sibilia, C. 1927. Alcune Laboulbeniaceae parassite di Acari. Redia 16: 79-88. New species
in Rickia. Reports Dimeromyces, Rickia; also Rhynchophoromyces, Herpomyces,
Stigmatomyces, which may have been misidentifications. Discusses teratological forms
of Rickia.
Siemaszko, Janina, and W. Siemaszko. 1928. Owadorosty polskie i palearktyczne.
(Laboulbeniales polonici et palaearctici.) Polskie Pismo Entomol. 6: 188-211.
Tav. VII. New taxa in Cryptandromyces, Laboulbenia, Rhachomyces. Reports Can-
tharomyces, Chitonomyces, Helodiomyces, Laboulbenia, Misgomyces, Rhachomyces
(in part as Rickia), Zodiomyces. Discussion of Laboulbeniales. Host index.
TAVARES: BIBLIOGRAPHY
413
—, and —. 1932. Owadorosty polskie i palearktyczne. (Laboulbeniales polonici et
palaearctici.). II. Polskie Pismo Entomol. 10:149-188. Tab. VII-X. Reports Cantharo-
myces, Corethromyces, Idiomyces, Laboulbenia, Monoicomyces, Peyritschiella
(in part as Dichomyces), Rhachomyces (in part as Rickia), Rhadinomyces (as
Corethromyces) Symplectromyces. Refers collections on Pselaphidae to Corethromyces and Au-
tophagomyces. Includes host index.
—, and —. 1933. Owadorosty polskie i palearktyczne. (Laboulbeniales polonici et
palaearctici.). III. Polskie Pismo Entomol. 12: 115-138. Tab. IX-X. Reports Autoi-
comyces (as Ceratomyces), Cantharomyces, Chitonomyces, Helodiomyces, Hydraeo-
myces, Laboulbenia, Zodiomyces. Discusses Laboulbenia gyrinicola and puts into
synonymy with L.funeralis Spegazzini, in part. Includes host index.
Smith, M. R. 1917. An infestation of Lasius niger L. var. americana with Laboulbenia
formicarium [sic]Thaxter. J. Econ. Entomol. 10:447.
—. 1928. Remarks concerning the distribution and hosts of the parasitic ant fungus,
Laboulbenia formicarium Thaxter. Bull. Brooklyn Entomol. Soc. 23: 104-106.
Reports new hosts and localities.
—. 1946. Ant hosts of the fungus, Laboulbenia formicarum Thaxter. Proc. Entomol.
Soc. Wash. 48: 29-31. List of hosts in United States; comments on occurrence.
—. 1961. Another ant genus host of the parasitic fungus Laboulbenia Robin (Hymenop-
tera: Fdrmicidae). Proc. Entomol. Soc. Wash. 63: 58. Reports new host genus for
L. formicarum.
Sorokin, N. 1871. Mikologicheskie Ocherki. Trudy Obsc. Isp. Prir. Imp. Har'kovsk.
Univ. 3(3): 1-48. Tab. I-IV. New taxon: Laboulbenia pitraeana; suggested as possible
synonym: L. baeri. Gives brief account of thallus development; suggests that mycelium
probably develops inside host and reemerges to form pedicel and perithecium (p. 39, PI.
IV). (See review by Janczewsky.)
—. 1883. Rastitel'nye parazity chelovieka i zhivotnykh, kak prichina zaraznikh bolez-
nei. Vol. II. St. Petersburg. 544 p. Tab. I-XXXIII. Includes brief historical
discussions; outlines development, discussing Peyritsch's work, etc.; describes known
species; mentions Laboulbenia pitraeana as synonym of Stigmatomyces baeri
(pp. 406-425; pis. XXXI-XXXIII).
Spegazzini, C. 1902. Mycetes argentinenses (series II). Anales Mus. Nac. Hist. Nat.
Buenos Aires (ser. 2) 8:49-89. New species in Laboulbenia (pp. 79-80).
**—. 1910a. Laboulbeniaceas nuevas chilenas. Rev. Chilena Hist. Nat. 14: 71-72. New
species in Laboulbenia.
**—. 1910b. Fungi Chilenses. Contribuci6n al Estudio de los Hongos Chilenos.
Libreria Nacional. J. Lajouane & Cia., Buenos Aires. 205 p. Descriptions of the
taxa of 1910a; also reports other Laboulbenia species.
—. 1912. Contribuci6n al estudio de las Laboulbeniomicetas argentinas. Anales Mus.
Nac. Hist. Nat. Buenos Aires 23: 167-244. New genera: Cochliomyces, Laboulbeniella
(syn. of Laboulbenia). New species in Cantharomyces (C. bruchii), Corethromyces,
Dichomyces (now Peyritschiella), Dimorphomyces, Eumonoicomyces, Laboulbenia
(including L. leathsii), Monoicomyces, Sphaleromyces (later Laboulbenia). Reports
Amorphomyces, Compsomyces, Corethromyces, Dixomyces (as Laboulbenia),
Laboulbenia, Monoicomyces, Peyritschiella (as Dichomyces), Rhachomyces.
Includes key to genera.
—. 1914a. Fungi nonnulli Senegalenses et Canarienses. Anales Mus. Nac. Hist. Nat.
Buenos Aires 26:117-134. Reports Dioicomyces (p. 129).

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MYCOLOGIA MEMOIR NO. 9
—. 1914b. Primo contributo alia conoscenza della Laboulbeniali italiani. Redia 10:
21-75. T. I-IX. New species in Laboulbenia, including L. coneglanensis. Reports Di-
meromyces, Herpomyces, Laboulbenia, Peyritschiella (as Dichomyces), Rickia.
Includes key to his subgenera, sections, subsections in Laboulbenia (of these, only Apsal-
lia, Ceratotheca, Macromastiga, Monomastiga, Oligomastiga, Psalliophora, Schizo-
soma are included among taxa listed; the remaining names suggested as subdivisions of
genera serve only to distinguish dichotomies in the key).
—. 1915a. Laboulbeniali ritrovate nelle collezioni di alcuni musei italiani. Anales Mus.
Nac. Hist. Nat. Buenos Aires 26: 451-511. New species in Dichomyces (now
Peyritschiella), Ecteinomyces (now Pseudoecteinomyces), Eumisgomyces (E. dohrnii, syn. of
Laboulbenia partita), Laboulbenia, including L. desgodinsii (see Balazuc, 1971e), L.
lagarii, L. langsbergii, L. leonardii, Laboulbeniella (now Laboulbenia), Rickia,
including R. copriphis, R. jacobsonii. Reports Corethromyces, Dioicomyces, Eucan-
tharomyces, Herpomyces, Laboulbenia, Peyritschiella (as Dichomyces), Rhachomyces.
Includes additional subdivisions of Laboulbenia and makes some changes in their
spelling.
—. 1915b. Segunda contribuci6n al conocimiento de las Laboulbeniales italianas.
Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 37-74. New genera: Parahydraeomyces
(syn. of Hydraeomyces), Thripomyces. New species in Amorphomyces, Autoicomyces,
Cantharomyces, Chitonomyces, including C. aculeifer, C. ensifer, Dichomyces (now
Peyritschiella), Dioicomyces, Hydraeomyces, Hydrophilomyces, Laboulbenia,
Monoicomyces, Stigmatomyces. Reports Chitonomyces, Coreomyces, Ecteinomyces,
Laboulbenia (some new forms), Monoicomyces, Peyritschiella (as Dichomyces), Treno-
myces.
—. 1917. Revision de las Laboulbeniales argentinas. Anales Mus. Nac. Hist. Nat. Buenos
Aires 29: 445-688. New genera: Cucujomyces, Pselaphidomyces, Stephanomyces
(syn. of Cucujomyces). New taxa in Acompsomyces (now Autophagomyces),
Amorphomyces, Autoicomyces (later Ceratomyces), Cantharomyces, Ceratomyces,
Chitonomyces, including C. bruchii, Clematomyces (position uncertain), Compso-
myces (now Scaphidiomyces), Coreomyces, Corethromyces (including ?C. andicola,
later Eudimeromyces, now Dimeromyces; ?C. subsigmoideus, probably Cryptandro-
myces) (in part, now Cryptandromyces, Mimeomyces), Dimorphomyces, Dioicomyces
(including forms of D. formicillae), Ecteinomyces (now Aporomyces,
Hydrophilomyces, Kyphomyces, Siemaszkoa), Herpomyces, Laboulbenia, including L. bergii,
L. brachinicola, Monoicomyces, Rickia (in part now Benjaminella, Dimorphomyces),
Sphaleromyces (now Mimeomyces), Stigmatomyces (now Corethromyces,
Mimeomyces, and probably Sphaleromyces, in part), Trenomyces. Reports
Acompsomyces, Amorphomyces, Autoicomyces, Botryandromyces (as Laboulbenia), Camp-
tomyces, Cantharomyces, Ceratomyces, Chaetomyces, Compsomyces, Cochliomyces,
Corethromyces, Cryptandromyces, Dimeromyces, Dimorphomyces, Dioicomyces,
Ecteinomyces, Eumonoicomyces, Herpomyces, Hesperomyces (as Stigmatomyces),
Hydraeomyces, Hydrophilomyces and Kyphomyces (as Ecteinomyces), Laboulbenia,
Mimeomyces (as Corethromyces), Monoicomyces, Peyritschiella (in part as
Dichomyces), Rhachomyces, Rhadinomyces (as Corethromyces), Rhynchophoromyces,
Rickia, Scaphidiomyces, Sphaleromyces (as Corethromyces), Stigmatomyces,
Trenomyces, Includes key to his sections of Laboulbenia and extensive discussion (in Spanish)
of collection methods, morphology, transmission, etc.
—. 1918. Observaciones microbiologicas. Anales Soc. Ci. Argentina 85: 311-323.
New species in Coreomyces; new name proposed for Laboulbenia paupercula Speg.
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Discusses Autoicomyces, Laboulbenia, Parahydraeomyces (pp. 320-323).
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**—. 1919. Les Hongos de Tucuman. Soc. Argentina Ci. Nat. Pjimera Reunion Nac.
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—. 1924. "Corethromyces Bruchi" nueva Laboulbenial argentina. Physis [Buenos
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Ceratomyces), Cantharomyces, Chitonomyces, Coreomyces, Corethromyces,
Cryptandromyces? (in part as Corethromyces), Dimeromyces, Dioicomyces, Ecteinomyces
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Hydraeomyces, Hydrophilomyces (as Misgomyces), Idiomyces, Laboulbenia, Mimeomyces,
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Reports Laboulbenia.

416
MYCOLOG1A MEMOIR NO. 9
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Cantharomyces, Chitonomyces, Cryptandromyces (as Corethromyces), Dioicomyces,
Enarthromyces, Euzodiomyces, Filariomyces, Hydraeomyces, Kainomyces,
Laboulbenia, Misgomyces, Peyritschiella (in part as Dichomyces), Rhachomyces, Rickia.
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**Sugiyama, K. 1981a. Notes on Laboulbeniomycetes of Formosa III. Trans. Mycol. Soc.
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**Sugiyama, K., and E. Shazawa. 1977. Notes on Laboulbeniomycetes of Formosa.
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(Herpomycetineae, nom. nud.).

418
MYCOLOGIA MEMOIR NO. 9
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Photographs of Colonomyces, Eusynaptomyces, Herpomyces, Mimeomyces.
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Tavares, Isabelle I., and T. Majewski. 1976. Siemaszkoa and Botryandromyces,
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*—. 1947. Rhachomyces girardii Lepesme et Tempere, Laboulbeniacee cavernicole
nouvelle des Pyrenees basques. Science et Montagne (Trav. Comm. Sci. Com. Med. Sect. S.
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year; position on host.
Terada, K. 1976. Some species of the Laboulbeniales from Taiwan. Trans. Mycol. Soc.
Japan 17: 23-24. Reports Blasticomyces (as Misgomyces), Corethromyces, Dimero-
myces, Dixomyces (as Misgomyces), Filariomyces, Laboulbenia, Peyritschiella (as
Dichomyces).
—. 1977. Some species of the Laboulbeniales newly recorded from Japan. Hikobia 8:
124-131. Reports Cantharomyces, Idiomyces, Monoicomyces, Rickia, Smeringomyces,
Stichomyces.
—. 1978. Additions to the Laboulbeniales of Taiwan, with descriptions of two new
species. Trans. Mycol. Soc. Japan 19: 55-64. New species of Kainomyces, Rickia.
Reports Dimeromyces, Laboulbenia, Rhachomyces, Rickia.
—. 1980. New or interesting species of the Laboulbeniales found on some coleopterous
insects of Japan. Trans. Mycol. Soc. Japan 21: 193-203. New species of Balazucia,
Dichomyces (now Peyritschiella), Laboulbenia, Rickia. Reports Dimeromyces, Rickia.
—. 1981. Osoriomyces, a new genus of the Laboulbeniales from Taiwan. Mycotaxon 13:
412-418.
Thaxter, R. 1890. On some North American species of Laboulbeniaceae. Proc. Amer.
Acad. Arts Sci. 25: 5-14. New genera: Cantharomyces, Peyritschiella. New species in
Laboulbenia. Reports Laboulbenia; transfers Appendicularia, Appendiculina to Stig-
matomyces. Includes brief introductory discussion of group.
—. 1891. Supplementary note on North American Laboulbeniaceae. Proc. Amer. Acad.
Arts Sci. 25: 261-270. New genera: Hesperomyces, Zodiomyces. New species in
Laboulbenia, Peyritschiella.
—. 1892. Further additions to the North American species of Laboulbeniaceae. Proc.
Amer. Acad. Arts Sci. 27: 29-45. New genera: Acanthomyces (syn. of Rhachomyces),
Ceratomyces, Corethromyces. New species in Heimatomyces (later placed in Chito-
nomyces and Hydraeomyces), Laboulbenia. Reports Chitonomyces, Laboulbenia.
TAVARES: BIBLIOGRAPHY
419
—. 1893. New species of Laboulbeniaceae from various localities. Proc. Amer. Acad.
Arts Sci. 28: 156-188. New genera: Amorphomyces, Chaetomyces, Dichomyces
(now Peyritschiella), Dimorphomyces, Haplomyces, Idiomyces, Rhadinomyces,
Teratomyces. New species in Cantharomyces, Peyritschiella, Acanthomyces,
Ceratomyces (in part now Autoicomyces, Rhynchophoromyces), Corethromyces, Dixomyces
(as Laboulbenia), Heimatomyces, Laboulbenia. Reports Laboulbenia. Cantharomyces
emended to exclude C. verticillatus.
—. 1894. New genera and species of Laboulbeniaceae, with a synopsis of the known
species. Proc. Amer. Acad. Arts Sci. 29: 92-111. New genera: Camptomyes, Compso-
myces, Moschomyces (now Compsomyces), Sphaleromyces. New species in
Cantharomyces, Ceratomyces (now Autoicomyces, Euceratomyces), Dichomyces,
Dimorphomyces, Heimatomyces, Peyritschiella, Teratomyces. Includes key to known species,
brief discussion of sexual structures, etc.
—. 1895. Notes on Laboulbeniaceae, with descriptions of new species. Proc. Amer.
Acad. Arts Sci. 30: 467-481. New genera: Diplomyces, Eucantharomyces,
Rhachomyces. New species in Ceratomyces, Dichomyces, Heimatomyces, Laboulbenia,
Rhachomyces, Sphaleromyces. Places Thaxteria in synonymy with Laboulbenia.
—. 1896. Contribution towards a monograph of the Laboulbeniaceae. Mem. Amer.
Acad. Arts Sci. 12: 187-429. Pis. I-XXVI. Includes historical account and extensive
discussion of group. New genera: Dimeromyces, Enarthromyces, Hydraeomyces,
Rhizomyces (all genera of previous papers included). New species in Laboulbenia,
Rhachomyces, Teratomyces.
—. 1899. Preliminary diagnoses of new species of Laboulbeniaceae. — I. Proc. Amer.
Acad. Arts Sci. 35:151-209. New species of Laboulbenia.
—. 1900. Preliminary diagnoses of new species of Laboulbeniaceae. — II. Proc.
Amer. Acad. Arts Sci. 35: 407-450. New genera: Clematomyces, Eucorethromyces
(now Corethromyces), Euzodiomyces, Limnaiomyces, Misgomyces, Monoicomyces,
Polyascomyces. New species in Amorphomyces, Autoicomyces (as Ceratomyces),
Cantharomyces, Ceratomyces, Chitonomyces, Compsomyces, Corethromyces (in part
as Sphaleromyces), Dimeromyces, Dimorphomyces, Dixomyces (as Misgomyces),
Eucantharomyces, Hydrophilomyces (as Ceratomyces), Mimeomyces (as
Sphaleromyces), Peyritschiella (in part as Dichomyces), Rhachomyces, Rhizomyces,
Rhynchophoromyces (as Ceratomyces), Symplectromyces (as Teratomyces), Teratomyces.
—. 1901a. Preliminary diagnoses of new species of Laboulbeniaceae. — III. Proc.
Amer. Acad. Arts Sci. 36: 395-414. New genus: Ceraiomyces (now Laboulbenia). New
species in Arthrorhynchus, Dimeromyces, Rhizomyces, Stigmatomyces.
—. 1901b. Preliminary diagnoses of new species of Laboulbeniaceae. — IV. Proc. Amer.
Acad. Arts Sci. 37: 19-45. New genera: Acompsomyces, Dioicomyces, Euhaplomyces,
Eumonoicomyces (in part now Monoicomyces), Kainomyces, Stichomyces. New species
in Ceratomyces, Chitonomyces, Corethromyces, Eucantharomyces, Kleidiomyces
(as Monoicomyces), Mimeomyces (as Corethromyces, Sphaleromyces),
Monoicomyces, Peyritschiella (in part as Dichomyces), Rhachomyces, Sphaleromyces,
Teratomyces.
—. 1902. Preliminary diagnoses of new species of Laboulbeniaceae. — V. Proc. Amer.
Acad. Arts Sci. 38: 7-57. New genera: Acallomyces, Coreomyces, Ecteinomyces,
Herpomyces. New species in Acompsomyces, Corethromyces (in part as Stichomyces),
Dimeromyces, Dixomyces (as Laboulbenia), Laboulbenia, Monoicomyces,
Smeringomyces (as Rhachomyces).
**—. 1903. Notes on the genus Herpomyces. Science (ser. 2) 17:463. [Abstract.]

420
MYCOLOGIA MEMOIR NO. 9
—. 1905. Preliminary diagnoses of new species of Laboulbeniaceae. — VI. Proc. Amer,
Acad. Arts Sri. 41: 301-318. New genus: Distichomyces (now Rickia). New species in
Acompsomyces, Autoicomyces (as Ceratomyces), Chitonomyces, Coreomyces,
Dimeromyces, Eucantharomyces, Herpomyces, Laboulbenia, Monoicomyces,
Rhachomyces, Stigmatomyces.
—. 1908. Contribution toward a monograph of the Laboulbeniaceae. Part II. Mem.
Amer. Acad. Arts Sci. 13: 217-469. Pis. XXVIII-LXXI. New genera: Autoicomyces,
Hydrophilomyces, Kleidiomyces, Rhynchophoromyces, Smeringomyces, Sym-
plectromyces. New species in Laboulbenia, Ceratomyces. Extensive discussion of
group; key to genera.
—. 1912a. New or critical Laboulbeniales from the Argentine. Proc. Amer. Acad. Arts
Sci. 48: 153-223. New genera: Autophagomyces, Cryptandromyces, Mimeomyces,
Scaphidiomyces, Scelophoromyces, Synandromyces, Synaptomyces, Tetrandromyces,
Zeugandromyces. New species in Amorphomyces, Autoicomyces (in part as
Ceratomyces), Benjaminella (as Rickia), Botryandromyces (as Laboulbenia), Cantharomyces,
Ceratomyces, Corethromyces (in part as Stichomyces) Diaphoromyces (as Rickia),
Dimeromyces, Dimorphomyces, Dioicomyces, Kyphomyces (as Ecteinomyces),
Laboulbenia, Mimeomyces (as Corethromyces), Monoicomyces, Rhachomyces, Sphalero-
myces (as Corethromyces), Stemmatomyces (as Stigmatomyces). Brief notes on
Acompsomyces, Camptomyces, Chaetomyces, Eumonoicomyces, Herpomyces, Hesperomyces
(as Stigmatomyces), Hydrophilomyces (as Ecteinomyces), Kleidiomyces, Peyritschiella
(as Dichomyces), Rhynchophoromyces, Zodiomyces.
—. 1912b. Preliminary descriptions of new species of Rickia and Trenomyces. Proc.
Amer. Acad. Arts Sci. 48:363-386.
—. 1914a. Laboulbeniales parasitic on Chrysomelidae. Proc. Amer. Acad. Arts Sci.
50: 15-50. New species in Dimeromyces, Ceraiomyces (syn. of Laboulbenia), including
C. minusculus, Laboulbenia, including L. bruchii (Sphaleromyces bruchii Speg.), L.
disonychae (Laboulbeniella disonychae Speg.), L. homophoetae (Laboulbeniella
homophoetae Speg.), L. papuana (syn. of L. macarthuri Balazuc, 1971d; proposed
also by Benjamin, 1971,,p. 132).
—. 1914b. On certain peculiar fungus-parasites of living insects. Bot. Gaz. [Craw-
fordsville] 58: 235-253. Pis. XVI-XIX. Describes imperfect fungi; discusses briefly
relationship of Laboulbeniales to hosts.
—. 1915. New Indo-Malayan Laboulbeniales. Proc. Amer. Acad. Arts Sci. 51: 1-51. New
genus: Tettigomyces. New species in Blasticomyces (as Misgomyces), Corethromyces,
Cryptandromyces, Diclonomyces (as Cryptandromyces), Dimeromyces, Diphymyces
(as Corethromyces), Dixomyces (as Misgomyces), Euphoriomyces (as Stichomyces),
Herpomyces, Laboulbenia, Monoicomyces, Peyritschiella (as Dichomyces),
Rhachomyces, Rickia, Synandromyces, Zeugandromyces (as Stigmatomyces). Reports Arth-
rorhynchus, Rickia; reduces Ceraiomyces to synonymy with Laboulbenia.
—. 1916. New or critical species of Chitonomyces and Rickia. Proc. Amer. Acad. Arts
Sci. 52:1-54.
—. 1917. New Laboulbeniales, chiefly dipterophilous American species. Proc. Amer.
Acad. Arts Sci. 52: 647-721. New genera: Ilytheomyces, Nycteromyces. New taxa
in Acompsomyces, Gloeandromyces and Hesperomyces (as Stigmatomyces),
Laboulbenia, Rickia, Stigmatomyces.
—. 1918a. Extra-American dipterophilous Laboulbeniales. Proc. Amer. Acad. Arts
Sci. 53: 695-749. New species in Dimeromyces, Hesperomyces (as Stigmatomyces),
Ilytheomyces, Laboulbenia, Rhizomyces, Stigmatomyces.
TAVARES: BIBLIOGRAPHY
421
—. 1918b. New Laboulbeniales from Chile and New Zealand. Proc. Amer. Acad.
Arts Sci. 54: 205-232. New genera: Diandromyces, Eudimeromyces (now
Dimeromyces). New species in Cantharomyces, Coreomyces, Cucujomyces, Diphymyces
(as Corethromyces), Herpomyces, Mimeomyces (as Corethromyces), Laboulbenia,
Monoicomyces, Peyritschiella (as Dichomyces).
—. 1920a. Second note on certain peculiar fungus-parasites of living insects. Bot. Gaz.
[Crawfordsville] 69: 1-27. Pis. I-V. Describes imperfect fungi; compares with
Laboulbeniales.
—. 1920b. New Dimorphomyceteae. Proc. Amer. Acad. Arts Sci. 55: 209-282. New
genus: Polyandromyces. New species in Dimeromyces, Dimorphomyces. Reports
Dimeromyces (in part as Eudimeromyces).
—. 1924. Contribution towards a monograph of the Laboulbeniaceae. Part III. Mem.
Amer. Acad. Arts Sci. 14: 309-426. Pis. I-XII. New taxa: nine species of Chitonomyces.
Includes descriptions from recent papers for Dimeromyces, Dimorphomyces,
Nycteromyces, Polyandromyces (Dimorphomyceteae); recent descriptions of
Chitonomyces. Mentions other species in these genera, as well as Hydraeomyces, Parahy-
draeomyces (syn. of Hydraeomyces). Extensive discussion of morphology.
—. 1926. Contribution towards a monograph of the Laboulbeniaceae. Part IV. Mem.
Amer. Acad. Arts Sci. 15: 427-580. Pis. I-XXIV. New genera: Diaphoromyces,
Porophoromyces, Rhipidiomyces. New species in Camptomyces, Chitonomyces,
Dimeromyces, Dimorphomyces, Eucantharomyces, Limnaiomyces, Rickia,
Tettigomyces. Morphological discussion includes Trenomyces. Some descriptions from recent
papers. Excludes four species from Rickia (now in Benjaminella).
—. 1931. Contribution towards a monograph of the Laboulbeniaceae. Part V. Mem.
Amer. Acad. Arts Sci. 16: 1-435. Pis. I-LX. New genera: Acrogynomyces,
Adelomyces (now Phaulomyces), Amphimyces, Apatelomyces, Apatomyces,
Aporomyces, Asaphomyces, Carpophoromyces, Chaetarthriomyces, Clonophoro-
myces, Dermapteromyces, Diclonomyces, Dicrandromyces (now Tetrandromyces),
Dipodomyces, Distolomyces, Drepanomyces, Euceratomyces, Euphoriomyces,
Eusynaptomyces, Gloeandromyces, Histeridomyces, Kruphaiomyces, Meionomyces,
Microsomyces, Nanomyces, Phaulomyces, Phurmomyces, Plectomyces, Rhizopodo-
myces, Schizomeromyces (now Clematomyces), Stemmatomyces, Thaumasiomyces,
Triandromyces (now Tetrandromyces), Trochoideomyces. Hesperomyces reestablished.
Includes descriptions from recent papers on genera not included in parts III, IV,
but Laboulbenia omitted.
Theodor, O. 1954. 66A. Nycteribiidae. Pp. 1-44, Taf. I-XVIII. In: Lindner, E.,
Die Fliegen der Palaearktischen Region, Lief. 174. E. Schweizerbart'sche Ver-
lagsbuchhandlung (Erwin Nagele), Stuttgart.
—. 1967. An illustrated catalogue of the Rothschild Collection of Nycteribiidae (Diptera)
in the British Museum (Natural History) with keys and short descriptions for the
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* Theodorides, J. 1952. Contribution a I'etude ecologique des parasites et commensaux
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Includes table showing Laboulbeniales on Ceuthosphodrus (Caraboidea, Ptero-
stichidae, Sphodrini).
*—. 1954. Parasites et phoretiques des Coleopteres des grottes de l'Ariege et autres cas
connus de parasitisme chez les Coleopteres cavernicoles. Publ. Mus. Hist. Nat.
[Paris] (Notes Biosptol.) 9: 187-197. Reports Laboulbenia, Rhachomyces.

422
MYCOLOGIA MEMOIR NO. 9
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H. Bischoff. Gustav Fischer, Jena.
**Tieghem, P. van. 1901. L'oeuf des plantes considere comme base de leur classification.
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**Trinchieri, G. 1910. Intorno a una Laboulbeniacea nuova per l'ltalia (Trenomyces
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TAVARES: BIBLIOGRAPHY
423
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(P. 43).

TAVARES: GLOSSARY
425
Glossary
accessory cell — cell growing on outer side of perithecium, on exterior of outer wall
cells of lower part of perithecium — occurs in Hydrophilomyceteae.
andropodium — used by Spegazzini (1915b) for basal cell of inner (anterior) appendage
in Laboulbenia (cell g).
anterior — in the direction of the side of the perithecium away from the appendage; on
the appendage, it is the inner side, toward the perithecium.
antheridia
simple — one-celled phialide, producing spermatium within neck, which may be
extremely short, but is usually moderately long and narrow; the spermatia are
said to be endogenous,
compound — massive, with variously arranged cells, or shaped like a phialide and
having a basal layer of spermatia-producing cells (the latter is characteristic
of the Peyritschielleae and Dimorphomyceteae).
intercalary — a cell of an appendage branch that produces spermatia laterally,
usually through a distinct lateral neck. When the spermatium is borne at the apex
of the neck and is relatively large, the spermatium is said to be exogenous —
it appears to be borne exteriorly (in Zodiomyces, they are borne terminally
on short appendages),
seriate — a series of consecutive intercalary cells functioning as antheridia;
usually the terminal cell of the series is a phialide of normal form,
antheridial tuft — cluster of antheridial branches growing from a group of cells; these
tufts are produced in a series on the male of Herpomyces ectobiae.
antheridiophore — the inner (anterior) appendage of Laboulbenia (Spegazzini, 1917).
appendage
primary — cells derived from upper spore segment, the upper of the two cells
of the germinating spore; the cells above the spore septum, which is often
conspicuously thicker or darker than those formed later,
secondary — appendages derived from lower spore segment (not used for those
growing on the perithecium itself); they are on the receptacle above or below the
perithecium. (Note: Benjamin [1973b] called the branches of the primary ap-
endage secondary appendages.)
stalk appendage — distal part of appendage that includes stalk cell and secondary
stalk cell of perithecium as intercalary cells — occurs in Euceratomycetaceae.
see inner and outer appendages below,
armed, armament — outgrowths on the perithecium, primarily at or near the apex
(as contrasted to unarmed or simple),
ascogonium — the fertile cell in row of cells formed from trichophoric cell and carpogenic
cell; tnis binucleate ceil usually divides to form more than one cell that functions
as an ascogenic (ascogenous) cell,
axil, axillary — pertaining to the angle between the primary axis and the perithecium,
toward the apex of the thallus (refers to position of trichogyne in most genera of the
Ceratomycetaceae).
axis
primary — principal axis of thallus, roughly coinciding with axis of spore from
which thallus develops,
secondary — axis of thallus growing out from receptacle of thallus at an angle to the
primary axis; it may take the appearance of the principal axis of the thallus, with
the primary axis becoming relatively small and inconspicuous.

426
MYCOLOG1A MEMOIR NO. 9
basal cell of thallus — cell /— foot cell, part of which usually turns black.
basal cells of perithecium — the lowest cells in the perithecium, just above the stalk and
secondary stalk cells; usually 3 in number — m arises from VI; n and n' arise
from VII; n gives rise to 2 wall cell rows,
beak — an outgrowth of apex of the perithecium — pointed, so that apex resembles a
bird's beak (occurs in Kruphaiomyces).
biseriate — double row of cells in receptacle,
buffer cells, buffer outgrowths — short outgrowths just above foot, presumably
concerned with keeping thallus in suitable position on host,
carpogonial cell — fertile cell subtending trichophoric cell; the central cell just above
cell VII in the Laboulbeniineae, before binucleate cells are formed in the central
row of cells of the perithecium.
carpogonial upgrowth — upgrowth of one of the lower wall cells in Herpomyces,
passing through the interior of the perithecium to its apex; later refers to secondary
upgrowths in the carpogonium (centrum),
claw form — simplified thallus growing on tarsi or claws of insect hosts of the Ceratomy-
cetinae — lacks horns and differs in number of thallus cells from typical form,
corner cells — small cells formed from upper angles (lower angles in Euphoriomyces cioi-
deus) of cells in primary axis of thallus, usually above perithecium.
corticate — layer of small cells cover surface of a larger cell,
determinate — refers to receptacle in which there is a fixed number of cells in a particular
taxon.
exogenous — large spermatium borne at apex of lateral neck of intercalary antheridium
or apically on antheridial cell; appears to be borne externally,
fertile cells — cells that will give rise to antheridia, spermatia, or perithecia.
filaments — slender upgrowths around ostiole of perithecium.
flask or flask-shaped — used for phialides or for a compound antheridium of similar
shape having a basal layer of fertile cells,
foot
primary — basal point of attachment of thallus; usually blackened.
secondary — a secondary point of attachment of thallus to host integument, taking
the place of the primary foot functionally (not used in Herpomycetineae).
haustorium — extension of cytoplasm of lowermost cell of thallus, penetrating into
integument of host (from secondary foot or secondary receptacle in some genera);
usually from primary foot (called rhizomycelium by Benjamin, 1971).
horn (cornu) — apical or subapical outgrowth on perithecia in Ceratomycetaceae
(usually multicellular, but in some species short and unicellular); also used for
broad outgrowth borne in a lower position on perithecia.
indeterminate — refers to receptacle of species in which number of receptacle cells varies,
as contrasted to the fixed number in a determinate receptacle,
inferior supporting cell — lowermost cell in 5-celled carpogonial cell row in
Laboulbeniineae.
innate — refers to perithecium or antheridium that is embedded within the outer margin
of the thallus and opens out at its edge (or protrudes slightly),
inner — inner side of perithecium or primary appendage, as contrasted to the side on the
outside of the thallus (of the appendage, the inner side is the anterior side; of the
perithecium, the inner side is the posterior side),
inner appendage — in Laboulbenia; arises from cell g; called antheridiophore by Spegaz-
zini (1914b).
inner appendage basal cell — lowermost cell of inner appendage (cell g); called andro-
podium by Spegazzini (1914b).
insertion cell — lowermost cell of primary appendage in Laboulbenia; usually flat
and blackened (cell e); called psallium by Spegazzini (1914b), but referred to as
an annulus.
TAVARES: GLOSSARY
427
ligula — strap-shaped outgrowth borne laterally on perithecium (used in Zodiomyces).
lip — cells surrounding aperture, as of compound antheridium.
lobes — usually refer to short, relatively broad outgrowths at or near apex of perithecium.
multicellular perithecial stalk — uniseriate row of cells directly subtending perithecial
stalk cell (cell VI), forming a straight row of cells which includes VI.
multiseriate — row of cells in receptacle in which there are more than two cells at each
level (sometimes used to include biseriate).
neck — narrowed apex of antheridium (also used for apex of antheridium above point
where septa of spermatia are formed); upper part of perithecium (usually narrowed)
above the wider venter, which contains the asci.
outer (posterior) appendage — in Laboulbenia; arises from cell/; see paraphysis.
outer (posterior) appendage basal cell — lowermost cell of outer appendage (/); see
paraphysopodium.
paraphysis — main axis of appendage system (Spegazzini, 1914b).
paraphysopodium — outer appendage basal cell (Spegazzini, 1914b).
perithecium (perithecium proper of Thaxter, 1896, included the wall and stalk cells)
primary — borne on primary axis of thallus.
secondary — borne on secondary axis of thallus.
posterior — in the direction of the side of the perithecium toward the appendage; on the
appendage, it is the outer side, away from the perithecium.
primary septum — original spore septum. For detection, see section on receptacles,
procarp — the central cell row borne on the secondary stalk cell of the perithecium
(formed from cell i); later develops into trichogyne, ascogonium, and associated
cells (called procarpe by Thaxter in 1896).
prospore — one-celled ascospore before division (see Hill, 1977).
pseudoreceptacle — upper part of receptacle in Laboulbenia (cells ///, IV, and V);
sometimes considered to be the stalk of the appendage; called androstichum by
Spegazzini (1914b).
pseudoperithecium — perithecium in Coreomyces which is formed within the outer wall of
the receptacle (Thaxter, 1908).
receptacular cavity — shallow cavity in top of thallus in Zodiomyces.
receptacular cell complex — receptacle of numerous cells so arranged that it is not certain
whether perithecia are borne on the primary or a secondary axis (called a suprabasal
complex when it appears to consist chiefly of cells resulting from division of cell II).
receptacular stalk cell — single cell in axis of receptacle subtending perithecial stalk cell;
a cell that has been produced laterally(as in Dermapteromyces, Asaphomyces).
receptacle
primary — cells in primary axis that were derived from lower spore segment.
secondary — cells produced by primary receptacle to form a secondary axis
bearing one or more perithecia; also used for receptacular cell complex,
rhizoidlike — refers to irregular, elongate downgrowths in lower part of thallus (as in
Scelophoromyces).
rhizomycelium — haustorium, particularly applicable to slender haustoria, such as those
of Hesperomyces virescens (see Benjamin, 1971).
rostrum — extremely narrow perithecial apex which includes all four outer wall cell rows
(as in Kainomyces, Synandromycesfloriformis).
secondary inferior supporting cell — cell just above inferior supporting cell,
secondary receptacular axis — an axis of the thallus other than the primary axis,
bearing one or more perithecia.
secondary stalk cell — cell VII, between cell VI and the procarp.
shield — group of secondary receptacle cells subtending perithecium in Herpomyces;
sometimes obscuring young perithecia.
spine — a more or less pointed, elongate outgrowth at or near the perithecial apex
(cf. auricles of Peyritschiella, used by Thaxter, 1896, 1908, for paired outgrowths);
not multicellular, as the horn in Ceratomycetaceae.

428
MYCOLOG1A MEMOIR NO. 9
spore segment — upper spore segment is the cell of the two-celled spore that is distal in the
germinated position and proximal within the perithecium; lower spore segment is the
other cell (attaching to the host),
spore septum — original septum of spore.
stalk appendage — appendage extending beyond cell K//in the Euceratomycetaceae.
stalk cell — intercalary cell of the primary axis in the Ceratomycetaceae and of a
secondary axis (the stalk appendage) in the Euceratomycetaceae; cell VI, which subtends
all cells of the perithecium (together with cell VII forms the gynostichum [Spegaz-
zini, 1914b] or perithecial primordium [Benjamin, 1971]).
stigmata — small rounded structures on cells of the secondary receptacle in Rhachomyces
(see discussion by Thaxter, 1908).
subsessile — lateral phialides that are partially embedded, not borne free on surface,
superior supporting cell — cell distal to ascogonium in carpogonial cell row in Laboul-
beniineae.
suprabasal cell (called subbasal cell later by Thaxter) — cell //, the cell just above the
basal cell of the thallus (not a cell secondarily produced in age),
suprabasal cell complex — a complex of cells formed from the suprabasal cell; a form
of secondary receptacle in which the primary axis is not readily identifiable,
teeth — very minute perithecial outgrowths; apical or subapical in position (see Diphy-
myces).
trichogyne stump — the persistent base of the trichogyne, borne laterally on the
perithecium (called the residuum by Spegazzini, 1914b).
trichophoric cell — cell separating carpogonial cell and trichogyne; septum breaks down,
causing fusion with carpogonial cell; a new septum is later formed,
trigger organs — spines, horns, etc., of the perithecium that presumably cause the
ostiole to open when they are pressed (see Thaxter, 1924, p. 388).
umbo — short to tall, rounded, erect dark apical outgrowth of perithecium in species
of Laboulbenia on Gyrinidae.
unarmed — simple perithecium without any outgrowths,
uncus — paired hook-like outgrowths on apex of perithecium in Laboulbenia species
on Gyrinidae (may be reduced to low protrusions); associated with umbo,
uniseriate — a single row of receptacle cells,
venter — lower, enlarged part of perithecium, which encloses ascogenic cells and young
asci.
wall cells
outer — outer layer of wall cells of perithecium.
inner — inner layer of wall cells of perithecium; Thaxter (1896) called the lower
tier parietal cells (Laboulbeniineae only), whereas the cells within the neck were
called canal cells (those at the base of the neck in Stigmatomyces were called
guard cells).
equal — equal in height in the different tiers.
subequal — nearly equal in height.
unequal — markedly unequal in height.
TAVARES: FIGURE INTERPRETATION
429
Interpretation of the Figures
Upper layer of cells (outer wall cells in perithecium) — heavy lines,
with outlined stippled nuclei or nucleoli (in central cells, sometimes
outline of nucleoli only because of lack of space).
Lower layer of cells (lower layer of outer wall cells in perithecium) —
heavy outline of dashes, with dashes forming the outlines of nuclei
and nucleoli.
Second layer from upper surface (upper row of inner wall cells in
perithecium) — solid, thin outline with no stippling.
Second layer from lower surface (lower row of inner wall cells in
perithecium) — dotted outlines of cell, nucleus, and nucleolus.
Outer cells of perithecium that are overlaid by other cells have narrower
outlines or "dash" outlines.
Central cell area — "zipatone," with nuclei stippled heavily but not
outlined.
Septation between central cells is represented by white lines which are
solid at upper levels and broken at lower levels. Broken white
lines also may indicate probable septa or constrictions.

TAVARES: ABBREVIATIONS
431
Alphabetical Guide to Abbreviations used (based on
those used by Benjamin and Shanor, 1950b)
a original septum of spore
ac ascogenic cell (ascogenous cell)
ace accessory cell on outer side of perithecium in Hydrophilo-
myceteae
acg accessory appendage associated with a perithecium in a
definite pattern
am ascogonium
an ahtheridium
ap apical cell of carpogonial upgrowth in Herpomyces
ap'," - secondary and tertiary apical cells (from later upgrowths) in
Herpomyces
as ascus
b upper cell in lower spore segment — becomes cell ///
ba primordium of antheridial tuft of perithecium in Herpomyces
bb basal cell of perithecium or antheridial tuft in Herpomyces that
is slightly higher in position than cell be (these two cells
are formed from lower daughter cell of ba)
bb'," - secondary cells cut off laterally from bb
be lower of two daughter cells of initial basal cell formed by
division of ba
bd cell cut off below cell bb
bd, be sr cells
cells of secondary receptacle directly subtending cells bd
and be
c middle of three cells in lower spore segment — becomes cell //
ce centrum (in Herpomyces, after second upgrowth from ci has
been formed)
ci carpogonial initial cell in Herpomyces, initiating centrum
ci' a secondary cell formed adjacent to ci cell late in development
co cornu (horn) — multicellular (or one-celled) subapical
appendage of perithecium in Ceratomycetinae
cp carpogonial cell (early centrum in Herpomyces)
cp' second carpogonial cell, formed by division of fused tricho-
phoric-carpogonial cell; in Herpomyces, second upgrowth
from ci cell
cp" — third upgrowth from ci cell in Herpomyces
cp-tc fused trichophoric-carpogonial cell
cut — cuticle (integument) of arthropod host

432
MYCOLOGIA MEMOIR NO. 9
d perithecial initial cell; produced by cell c (cf. a', Thaxter,
1896, pi. I)
e first cell of primary appendage in Laboulbenia (insertion cell)
epi epidermal cell layer of arthropod host
/ second cell of primary appendage in Laboulbenia (basal cell of
outer appendage)
/// filament (ex. slender ostiolar appendages in Helodiomyces)
fo foot (point of attachment of thallus; usually darkened)
fo-2 — secondary foot
fo' modification of apex of primary axis of Herpomyces —
resembles foot
g basal cell of inner appendage in Laboulbenia
g' secondary cell forming an additional inner appendage
h lower cell formed by division of cell d (perithecial primordium of
Benjamin, 1971) (cf. c, Thaxter, 1896, pi. I)
ha haustorium
/ upper cell formed by division of cell d (primordial cell of pro-
carp, according to Benjamin, 1971) (cf. d, Thaxter, 1896,
Pl-I)
isc inferior supporting cell — lowermost cell of carpogonial cell
row
j middle cell in row of three forming young perithecium —
becomes cell VII
k lower cell in row of three — becomes cell VI
li ligula (tongue-like outgrowth) — near apex of perithecium of
Zodiomyces
m basal cell of perithecium formed from cell VI(o', Thaxter, 1896,
pi. II)
n basal cell of perithecium formed from cell VII — produces two
wall cell rows (d, Thaxter, 1896, pi. II)
n' second basal cell formed from cell VII — produces one wall
cell row (g, Thaxter, 1896, pi. II)
n " extra n cell in more complex genera of Ceratomycetaceae
o first outer wall formed by cell n; usually used for first outer wall
cell tier (e or/, Thaxter, 1896, pi. II)
o' outer wall cell formed by cell m (/, Thaxter, 1896, pi. II)
o" outer wall cell formed by cell n' (j, Thaxter, 1896, pi. II)
o'" — second outer wall cell formed by cell n (e or/, Thaxter, 1896,
Pi- ID
ost ostiole
p inner wall cells (parietal cells) — first tier of cells (sometimes
used for entire row of inner wall cells) of inner wall of
perithecium
pa primary appendage
per perithecium
TAVARES: ABBREVIATIONS
433
pr primary receptacle
px primary axis
ro rostrum (beaklike, prolonged apex of perithecium ex. Kai-
nomyces)
sa secondary appendage (appendage from lower spore segment)
se seta of arthropod host
sh shield (specialized portion of secondary receptacle in
Herpomyces)
sisc secondary inferior supporting cell — cell just above isc
sp spermatium
spi spine (tapered apical or subapical outgrowth of perithecium in
Laboulbeniaceae)
spo spore
sr secondary receptacle (secondary axis of thallus)
ssc superior supporting cell (cell between ascogonium and
reformed trichophoric cell)
st stalk appendage (in Euceratomycetaceae)
sx spore apex
tc trichophoric cell (cell between trichogyne and carpogonial cell)
tc' reformed trichophoric cell (formed from fused trichophoric-
carpogonial cell)
tr trichogyne
tr' secondary trichogyne, formed after first trichogyne deteriorates
um umbo; knoblike outgrowth on perithecial apex ( ex.
Laboulbenia gyrinidarum)
un uncus; hooklike (or straight) outgrowths associated with umbo
x inner wall cells — second tier
x' inner wall cells — third tier
x" inner wall cells — fourth tier
w outer wall cells — second tier
w' outer wall cells — third tier
w" outer wall cells — fourth tier
w-1, etc. outer wall cell tiers in Herpomyces
w3c' extra cell in third tier of outer wall cells, occurring in the
vertical row above cell ci in Herpomyces
w4c' extra cell in fourth tier of outer wall cells in the ci row in
Herpomyces
I basal cell of mature thallus
77 suprabasal cell of mature thallus
III upper cell of three-celled primary receptacle; may divide.
IV upper, outer cell of five-celled primary receptacle in
Laboulbenia (may divide to form secondary IV cells IVa and
IVb)
V upper, inner cell of five-celled primary receptacle in Laboulbenia
(may divide to form secondary V cells Va and Vb)

434 MYCOLOGIA MEMOIR NO. 9
VI stalk cell of perithecium (p, Thaxter, 1896, pi. II); gives rise to
m
VII secondary stalk cell of perithecium (h, Thaxter, 1896, pi. II);
subtends carpogonial cell and cells n and n'
(Note that Spegazzini [1914b] used Roman numerals for septa, not for
cells)
TAVARES: COLLECTION AND PREPARATION
435
Collecting, Mounting, and Staining Laboulbeniales
In any area, the examination of arthropod genera or families that
have been reported as hosts of the Laboulbeniales in other parts of the
world may reveal the presence of fungi previously unknown in that
area (for European examples, see Majewski, 1974, Rossi, 1980, and
Rossi and Cesari Rossi, 1978, 1980b [also see Balazuc, 1971f]; for
Japanese, see Sugiyama, 1973, and Terada, 1977, 1980). Genera or
higher taxa not known to be hosts may bear new genera of the
Laboulbeniales or new species in existing genera (for example, Hebridae
[Majewski, 1981]; see Benjamin, 1955, 1967, 1970, 1979, and Balazuc,
1972; see also Racovitza, 1908, Caroli, 1912, Folsom, 1916, Mayer,
1934).
Knowledge about the kinds of habitats in which infected hosts can be
found may be obtained from general works on particular groups, such
as Vitzthum's publication (1940-1943) on acarids (see also Baker and
Wharton, 1952; Hughes, 1959; Evans et al., 1961); for parasites of
bats and birds, publications on their habits should be consulted (see also
Kellogg, 1896a,b). Paulian (1943) provided information about habitats
of beetles; useful information was also included in Traite de Zoologie,
vol. 9, by Paulian (pp. 892-1026) and Jeannel (pp. 1026-1069) (see Jean-
nel, 1949b; also Blatchley, 1910). (See Giles [1970] on earwigs.)
Abundant populations of beetles, especially Caraboidea, may be
found along the margins of streams and ponds, particularly those with
sand, mud, or fine gravel. Beetles can often be found under large
stones imbedded in these substrates. The prevalence of beetles on
the surface is influenced by sun, shade, and temperature. In Scheloske's
study (1969), the greatest number of species of Laboulbeniales were
found on beetles in ponds, among accumulated reeds along pond
margins, and in swampy places among alders and associated trees.
Maire (1920) included helpful information about the living habits of
hosts.
Often large numbers of beetles can be found hibernating under logs,
in dead stems, etc., although Thaxter (1896) pointed out that their fungi
might be in poor condition. Infected beetles, particularly Staphylinidae,
may be found in mounds of fine grass clippings that are slightly damp.
Hincks (1960) believed that a combination of dampness and warmth
in heaps of debris favored a high rate of infection, although some
locations harbored more infected beetles than others. He referred
particularly to the uniform, relatively high temperature conditions of

436
MYCOLOG1A MEMOIR NO. 9
mole nests (a habitat examined also by Kossen, 1937). Termite and ant
nests might also harbor infected insects.
Many beetles can be found in decaying fungi, particularly polypores.
Spegazzini (1917) recommended searching in dead animals and dung,
particularly on cold mornings in the rainy season. He also suggested
collecting at an outdoor light at night. Dainat (1973) mentioned the use
of a trap of black muslin in which flies are attracted to a structure of
white muslin from which they fall into alcohol. She also suggested
(personal communication) setting out white dishes containing water with
detergent (see also Deonier, 1971). Masses of debris of various kinds can
be processed using a Berlese funnel (for instructions, see Benjamin,
1971). Benjamin (1971) gave directions on how to construct an aspirator
with which to catch rapidly moving terrestrial insects.
Benjamin (1971) recommended fixing specimens in 70% ethanol,
rather than drying collections, because the insects were less susceptible
to damage. However, many insect collectors routinely dry and pin their
specimens. Such collectors should take care not to discard specimens
that appear to be covered with excessive amounts of debris, because
these insects are most likely to bear many thalli of Laboulbeniales. For
cytological preparations, insects can be preserved in Carnoy's (1 part of
acetic acid to 3 parts of 95% ethanol). Glycerine is unsuitable for
preserving insects, although Dainat (1973) suggested that removal of fungi in
a viscid liquid facilitates detachment and manipulation. The substance
should then be removed in 70% ethanol.
Thalli on dried hosts can be removed with a dampened, clean needle
(formed into a flattened blade), a fine dissecting blade, a razor blade
fragment, or fine forceps (purchased or made from needles) (see Thax-
ter [1896], Spegazzini [1917], and Benjamin [1971] for detailed
instructions). Debris should be removed from the fungi with a hair
mounted in a matchstick or a fine camel's hair paintbrush trimmed so
that one hair protrudes beyond the others. Specimens may be placed in a
tiny drop of Hoyer's solution (glycerine, chloral hydrate, and gum ara-
bic). Then invert over the drop a coverslip bearing a drop of the
mounting medium (Benjamin, 1971). A precise arrangement of thalli can be
obtained by placing them in a drop of water, arranging them with a hair
mounted in a matchstick, removing the water, then covering them
with alcohol (Thaxter, 1896). Paper fibers under the coverslip prevent
flattening of the specimens (Thaxter, 1896).
If thalli are placed between two coverslips, as suggested by Spegazzini
(1917), a preliminary examination can be made under the microscope
TAVARES: COLLECTION AND PREPARATION
437
and a choice between the two sides is possible, depending on which
details are visible. There is also less likelihood that the specimens will
be lost if the slide is broken when inadequately packed for mailing.
The thalli are placed in a drop of mounting medium on a large
circular coverslip, which is then covered by a small circular coverslip.
By inverting the smaller coverslip, some of the sealing compound
goes under the edge of the upper, larger coverslip when they are
attached to a slide. However, the larger coverslip can be attached to the
slide directly with balsam or some other mounting medium and the
margin of the small coverslip sealed with a broad circle of sealing
material.
A thin coverslip should be used and the sealing ring should be thin
enough to permit the use of an oil immersion lens. When mounting
a fragment of host integument, its thickness should be reduced as much
as possible. Setae, appendages, or a small piece of integument cut with a
razor blade may permit study of very young thalli or haustoria.
Specimens of a collection should be dispersed on several slides,
particularly if they are to be stained with acetocarmine. This permits
staining different thalli for different lengths of time. The mounting
medium on an unstained slide can be softened to allow later staining.
For removal of fungi from living insects, the insect (etherized if
necessary) can be held between the thumb and forefinger on sticky tape
or some padding. The padded tips of forceps can also be used.
Glycerine is the best mounting medium, particularly for type
specimens. Thaxter (1896) added a small amount of a saturated alcoholic
solution of eosin and a trace of salt to the glycerine, which he
introduced slowly under the coverslip to replace the water. When glycerine
preparations dry out, additional glycerine can be introduced (the cover-
slip must be securely attached to the slide first — this can be done by
placing radiating lines of balsam across the deteriorated sealing ring).
The old seal is removed in one or two places; care must be taken
to insure that dark sealing material does not move under the coverslip.
Diaphane has proved to be a very satisfactory mounting medium,
both for stained and unstained preparations, although some drying can
occur; crystals develop rarely (for directions, see following techniques
and staining methods). Balsam is more difficult to use, because the
specimens must be dehydrated and placed in xylene; dark specimens are
cleared and internal details are easy to see.
Early observers of the Laboulbeniales did not report the use of stains

438
MYCOLOGIA MEMOIR NO. 9
in the preparation of their specimens. However, potassium hydroxide
was used by Peyritsch (1873, p. 240) to demonstrate the nature of the
cell wall layers. Later, Thaxter (1896) used it to trace the connections
between cells in the perithecium. Various reagents were also used for
microchemical tests. Thaxter (1896), using Schultze's chloriodide of
zinc, and iodine with sulphuric acid, found that the presence of cellulose
was not indicated (cf. von Wettstein, 1921).
In the first cytological studies of the Laboulbeniales, Faull (1911,
1912) relied primarily on the use of Flemming's weaker fixative,
Flemming's triple stain, and Heidenhain's iron haematoxylin. Material
was prepared by sectioning embedded insect parts in paraffin.
More recent investigators have used cotton blue in lactophenol, acid
fuchsin in lactophenol, and Hoyer's acid fuchsin for staining specimens
(Benjamin and Shanor, 1951; Shanor, 1952; Richards and Smith,
1955b). The latter authors also used Gram stain (Richards and Smith,
1955a).
Most of my material has been prepared as acetocarmine squashes.
Fixation was accomplished either by immersion in Carnoy's alcohol-
acetic acid fixative (3:1) for 5 minutes, followed by Randolph's
modified Navashin fixative (Craf; a chromic acid-formalin-acetic acid
mixture; see Johansen [1940]) for 24 hours, or by a modification of the
ferric hydroxide mordanting method of Hyde and Gardella (1953). In
the latter method, the material was fixed in a propionic acid-ferric
hydroxide solution (2 g. of ferric hydroxide gently heated in 500 cc. of
propionic acid), of which 1 part was mixed with 3 parts of 95% ethanol.
After rinsing, the specimens were stained in refluxed acetocarmine. The
material fixed in Craf was mordanted in an aqueous solution of ferric
acetate for 30 minutes, then-rinsed, placed in a drop of acetocarmine,
and heated. In preparing slides of Laboulbenia flagellata, it was
advantageous to harden the specimens in 70% alcohol for 2-7 days before
staining.
The thalli were removed from the host and placed in a drop of fluid,
then transferred to a small drop of acetocarmine small enough to
prevent the specimens from floating out from under the coverslip.
If mordant was used, a small amount was introduced under one side of
the coverslip during the heating process. The slide was heated slightly
over an alcohol lamp until it was warm to the touch on the lower side
(boiling caused a deposition of dark granules). The slide was then
examined under the microscope and the thalli pressed slightly to
break the outer envelope (they tend to adhere to the slide if this is
done; sparse material should be carefully watched, but not flattened
TAVARES: COLLECTION AND PREPARATION
439
until the end of the heating process). After the slide cooled slightly,
it was reheated gently, then cooled again. This process was repeated
until the desired amount of stain was obtained (up to 30 minutes or
more). During the heating process, additional acetocarmine was added
at one side of the coverslip. A heavy stain could be achieved more easily
by forcing (with the fluid) the darkest acetocarmine area to the position
of the specimens and maintaining it at that place. Additional mordant
could increase the concentration of stain. When a dark stain was
obtained, 20% aqueous acetic acid was introduced at one edge of the
coverslip and a narrow piece of cleansing tissue was placed at the other
to draw the fluid under the coverslip. (If material is scarce, destaining
should be observed under the compound microscope). When only the
thalli retained the stain, the slides were inserted in a 100% ethanol
chamber (if vertically inserted, all excess fluid was removed from the
mount so that the coverslip would not slip off); after 2-3 days, the
slides were placed at a slight angle in Petri dishes lined with filter paper
saturated with 100% ethanol; a drop of diaphane or Euparol was
placed at the upper end of the coverslip each day for about 3 days
before the slide was allowed to dry. If slides dried out later and required
the addition of more diaphane, they could be softened in 100% ethanol.
Slides stained too darkly in iron propionocarmine may be destained in
20-40% acetic acid, heated slightly if necessary. However, excessive
destaining should be avoided.
Sections of Herpomyces ectobiae were made of thalli attached to
appendages of the host. They were fixed in Karpechenko's fixative
(chromic acid, formalin, and acetic acid), then embedded in warm agar
to which eosin had been added. The agar was cut into very small blocks
when cool; these were infiltrated with paraffin after being run through
a toluene sequence. Sections were cut at 6 jim at cold temperature. They
were stained with iron haematoxylin, after being coated with a thin
solution of celloidin to prevent loss of sections. Sections of Laboulbenia
were prepared from detached thalli. Sections of Hesperomyces virescens
(prepared by Miss Nel Rem) were coated with a solution of parlodion
to prevent loss; they were stained with iron haematoxylin with an orange
counterstain. Richards and Smith (1956) used a contrasting stain in their
work (Mallory's triple stain, sometimes superimposed on a silver
deposit from the argentaffin reagent). They used Zenker's fixative
(mercuric chloride, potassium bichromate, and acetic acid), with
embedding in Tissuemat. They also used Carnoy's, with double
embedding in celloidin and paraffin. With Mallory's triple stain, the
fungus walls were usually stained blue or purple, the haustoria were
blue, and the epidermal cells of the cockroach were red.

440
Whole mounts were prepared from material of Herpomyces and
Laboulbenia fixed in Schaudinn's fixative (mercuric chloride,
potassium bichromate, acetic acid), hydrolyzed in hydrochloric acid,
mordanted in formalin solution, and stained with undecolorized basic
fuchsin (DeLamater, 1948).

Figs. 1-6. Laboulbenia flagellata, on Agonum ovipennis, San Mateo County, California,
November, 1956. Acetocarmine-stained specimens. Fig. 1 a. Initiation of perithecium
from upper part of cell //. Basal appendage cell has divided (its upper septum is dark in
this thallus); lateral branch has arisen from one cell of appendage; apex of appendage has
broken off; cytoplasm in lateral branch of appendage and in terminal cell is heavily
stained. LLf 7 (8). x 850. Fe(OH)3 fixative, b. Division of cell d into cells h and i. Cell b
has undergone nuclear division; division of cell e has resulted in the formation of cell g,
which is extending to form a lateral appendage branch; cytoplasm in g and in apex of
appendage stains heavily. Note that terminal cell of appendage has broken off and
contains no protoplasm; nucleus in subterminal cell has divided and cell is extending upward
through empty outer wall of cell above. LLf 3 (17). x 850. Craf fixative, c. Young
perithecium preceding formation of cells m, n, and carpogonial cell. Cell g has produced a
1-celled inner appendage; cell e is still hyaline. LLf 3 (11). x 1275. Craf.
442

f
e
EC
m
I
tr
cp
m
n
Fig. 2. Laboulbenia flagellata x 1275. a. Phialide formation. Protoplast of upper :
phialide has drawn away from the wall of the neck; it probably has not yet produced sper- '
matia; upper angle of subterminal cell of lowest antheridial branch at right is heavily
stained and is about to extend into a lateral branch; cell / has undergone nuclear division
but a septum has not yet been formed; cell n' has not yet been produced by celly. LLf 3 -m.
(4). Craf. b. Nuclear division has taken place in formation of first o cell, which arises from JU"
cell n; cell i has divided to form cells tr and cp; lower part of cell e is dark, but nucleus is
still visible. LLf 7 (16). Fe(OH)j. c. Perithecium after formation of cell «'. Two o cells
have been formed from cells m and n; trichogyne initial has elongated and trichophoric
cell has been formed; cell e is quite dark. LLf 7 (3). Fe(OH)j.
444

Fig. 3. Laboulbenia flagellata. x 1275. Fe(OH)j fixative, a. Thallus with four outer wall
cells in perithecium. Spermatia are being produced; the outer wall of cell e has darkened
completely. LLf 4 (4). b. Septum has been formed separating inner wall (parietal) cell
from cell m; trichogyne initial cell has divided to form a two-celled trichogyne; only one
cell V is present, which is typical; a division has occurred in the outer wall cell row from m
and in one of the rows from n prior to division of cell «'. LLf 4 (14). c. Inner wall cells
have been formed from cells n' and m (septum is indicated at base of that from cell n");
trichogyne (only basal cell shown) consists of four elongate cells with two-celled branch
from suprabasal cell and short branch from subterminal cell. LLf 4 (21).
446

Fig. 4. Laboulbenia flagellata. X 1275. Fe(OH)3 fixative, a. Three inner wall cells
in perithecium (their basal septa not visible in thallus); one of inner wall cells comes from
cell m, one from n, and one from n'\ cell/is unusually dark in color; there are two K cells
(or possibly one binucleate cell). LLf 4 (17). b. Perithecium prior to breakdown of septum
between carpogonial and trichophoric cells. Four inner wall cells are present — cell p at
right (from cell m) is longer than others; trichogyne is branched. LLf 4 (2). c. Perithecium
with reconstituted trichophoric cell (to"); three of nuclei from conjugate division are in
carpogonial cell, which has reformed. Trichogyne has deteriorated, but stump remains;
inner walls from n and m are bicellular; cell division has not occurred yet in other p cells —
their nuclei appear to be in prophase; nuclear division has taken place in posterior outer
wall cell at upper level (next to cell e), but septum has not yet been formed; nuclei of cells
K//and m appear to be undergoing deterioration. LLf 4(14).
448

Fig. 5. Laboulbenia flagellata. x 1275. Fe(OH)j fixative, a. Perithecium with inferior
supporting cell. Three nuclei, rather than four, visible in fused trichophoric-carpogonial
cell; inner wall cells have all undergone division (septum not indicated in inner wall cell
row at lowest level, which arises from cell«'); anterior and uppermost rows of inner wall
cells arise from cell n; perithecial walls will soon grow up beside trichogyne remnant,
which includes nucleus. Note cubical shape of cell VII at lower level (indicated by
stippling), as contrasted to angular shape at upper level. LLf 4 (10). b. Perithecium after
deterioration of trichogyne, showing growth Of its apex beyond trichogyne stump.
There are two outer wall cells (w, w") above the o cells and three cells (p, x, x") in each
inner wall cell row; central cell row consists of reformed trichophoric cell, binucleate asco-
gonium (before it has produced superior supporting cell), secondary inferior supporting
cell, and inferior supporting cell; supernumerary nuclei are present in subterminal outer
wall cells at left, as well as in subterminal inner wall cells at upper and lower levels; asco-
gonium stains more heavily than other cells in central row. LLf 4 (10). c. Central cells of
perithecium, including binucleate superior supporting cell and two binucleate ascogenic
cells (part of one at right actually lies over surface of ssc cell); secondary inferior
supporting cell has grown upward. LLf 4(17).
450

Fig. 6. Laboulbenia flagellata. Ascogenic cells at beginning of ascus production (two
superposed ascus nuclei at left; fused ascus nucleus at right); apical lobes are forming on
perithecium; outer walls are three cells in height; lower and posterior inner wall cell rows,
which are three-celled, arise from cell n; supernumerary nuclei are present in subterminal
outer wall cells, and in lower and posterior subterminal inner wall cells; superior
supporting cell, containing two large nuclei, lies below ascogenic cells and the cell cut off from
upper extension of secondary inferior supporting cell (one ssc nucleus lies between those in
central ascogenic cell and one lies just above that of sisc extension). LLf 4 (26). x 1275.
Fe(OH)3.
452

Fig. 7. Laboulbenia flagellata. Cell sequence (diagram), indicating letter designations,
etc., of cells derived from a four-celled sporeling. Lines not terminating in letters should
be compared with a completed sequence. Line above/indicates the upper part of the outer
appendage.
454
CELL SEQUENCE IN
LABOULBENIA FLAGELLATA
Upper cell of spore
/
lowermost celK e e
inner
.appendage
g
Lower cell of spore
b—III-
I 1
JV
ws
p o'"'
,o,M
,w'
/P- /W W
Jr ^n^n .o«—o»
.n'— n'<_
n'
-\ VII VII
-m;
-VI
-tr
cp-—tc
o'-
-m
.tc'
0'
ac
ssc
cp-tcCT / /
cp" ^cp'—cp1—am—am.
\. \. ac
ISC SISC

Figs. 8-10. Laboulbenia gyrinidarum, L. borealis (species not distinguished). On Gyrinus
plicifer, Tilden Regional Park, Contra Costa County, 1948-1955. Acetocarmine-stained
specimens. Fig. 8. a. Germinating spore having enlarged foot; pigmented ring lies just
outside darkened lateral protrusion of protoplast; narrow outer wall layer of spore
(enveloping membrane) is visible. LLg 13 (117). x 1275. Craf. b. First division has taken place
in receptacle of germinating spore, cutting off basal cell /; lower part of foot is quite dark;
nucleus of cell / is already deteriorating. LLg 13 (36). x 850. Craf. c. Sporeling; cell b has
been formed first (rather than /, as in 8,b); dark area extends across upper part of enlarged
portion of foot; first two septa of primary appendage have darkened. LLg 13 (109). x
850. Craf. d. Perithecial primordium (d) before division; foot is entirely black except
for middle of upper enlarged area. LLg 14(11) x 850. Fe(OH),.
456

Fig. 9. Laboulbenia gyrinidarum, L. borealis (species not distinguished). Craf fixative,
a. Thallus in which main axis of primary appendage has broken off and anterior branch
arises from cell g; Va cell is present, but septum has not yet been formed between cells ///
and IV, although nuclear divison has taken place; cell d has divided to produce cells h and
/'. LLg 13 (84). x 850. b. Perithecium in which cells /, j, and k have been produced;
nucleus of cell /// is elongated, with the nucleolus at upper end; cells Va and Vb have been
formed — the upper one has produced a one-celled secondary appendage, the septum of
which is already black; cell g has not yet initiated an appendage; all cells of primary
appendage above/are devoid of protoplasm; black septa of primary appendage represent
bases of two branches; pore of large septum is eccentric. LLg 13 (39). x 1275. c. Thallus
in which cell IVa has been formed, together with Va, Vb cells. Dark septum is at base of
lateral branch of broken primary appendage; wall of cell/is black; secondary appendages
have been formed from cells Va and Vb; two appendages arise from cellg. LLg 13 (37). x
1275. d. Perithecium in which cells m and Kf have been formed from cell k. LLg 13 (72).
X 1275.
458

Fig. 10. Laboulbenia gyrinidarum, L. borealis (species not distinguished), a. Formation of
m, n, and n' cells in base of perithecium; n is larger than n'\ two cells have been produced
at apex of cell IV; cell g', derived from cell e, gives rise to appendage; Vb cell has produced
two appendages; only the heavy, dark outer wall of cell/remains. LLg 13 (29). x 1275.
Craf. b. Perithecium, with trichophoric cell, carpogenic cell, and trichogyne; four o cells
are present; cell m is in mitotic metaphase prior to formation of first inner wall cell
(position of chromosomes is indicated). LLg 13 (130). x 1650. Craf. c. Perithecium with first
inner wall cell (here arising from n")\ division is taking place in o cell at right on lower
level; o cells at left have divided; darkening of outer wall of perithecium is indicated at
right. LLg 14 (19). x 1275. Fe(OH)j. d. Perithecium with fused trichophoric-carpogenic
cell containing three nuclei (two from division of carpogenic cell nucleus); four inner wall
cells are present (two at upper level are from cell n). Note small dark spot and nearby
heavily staining area at base of trichogyne. LLg 14(45). x 1275. Fe(OH)3.
460

Figs. 11-15. Herpomyces ectobiae. On Blattella germanica (L.), Life Sciences Building,
University of California, Berkeley, I. Tavares, 1949-1956. Acetocarmine-stained
specimens. Fe(OH)3 fixative. Fig. 11. a. Germinating spore detached from seta; primary
axis has black apex and black basal foot, in which hyaline aperture of haustorium is
visible; suprabasal cell extends downward. LHe 8 (4). x 2450. b. Male. Young antheridial
tuft primordium arising from terminal cell of secondary receptacle; nuclear division has
taken place, but septum has not yet been formed. LHe 8 (22). x 1650. c. Male. Two-celled
tuft primordium; two nuclei are in underlying secondary receptacle cell, but a septum has
not yet been formed following nuclear division; this cell has produced two haustoria. LHe
8 (16). x 1650. d. Male. Four-celled tuft with be cell extending in direction of primary
receptacle. LHe 8 (22). x 1650. e. Male. Tuft with two unicellular branches; one-celled
tuft (ba) has been produced by terminal cell of secondary receptacle; secondary receptacle
cells extend downward into haustoria. LHe 8 (4). x 1650. f. Male with branched
antheridial tuft and two-celled tuft primordium (all branches of tufts are on side facing primary
appendage); haustorial aperture of foot is visible; foot is attached to outer surface of seta
(dark, dotted line in seta indicates inner surface of cuticle of seta). Protoplast is lacking in
terminal cell of primary appendage. LHe 8 (16). x 1650.
462

Fig. 12. Herpomyces ectobiae. x 2450. a. One-celled perithecial primordium; secondary
receptacle cells extend downward to form haustoria. LHe 8 (15). b. Two-celled perithecial
primordium. LHe 8 (5). c. Three-celled perithecial primordium. LHe 6 (16). d. Four-celled
perithecial primordium. LHe 6 (18). e. Perithecium with two basal cells and three wall
cells; wall cell at right has not divided yet. LHe 6 (41). f. Four-celled perithecium with two
basal cells below. LHe 6 (16). g. Eight-celled perithecium with two basal cells. LHe 6
(18).
464

Fig. 13. Herpomyces ectobiae. x 2450. a. Young perithecium with central upgrowth from
carpogonium-initiating wall cell (ci), in which nuclear division has taken place; there are
three cells in the outer wall cell row that includes this cell; division has not taken place yet
in the other three rows. LHe 8 (21). b. Beginning of trichogyne extension, prior to
formation of inner wall cells; centrum contains one nucleus above and one in ci cell area;
carpogonial initial has only a narrow connection, if any, with central cell here. Cell w3c'
arises as upgrowth from subtending w2 cell; it lies between the lower part of the apical cell
of the ci outer wall cell row and the centrum. Secondary receptacle cell below perithecium
has divided; an additional perithecial initial (ba) is present. LHe 12 (2). c. Perithecium
next to primary receptacle (at right, but not shown), with one secondary receptacle cell
intervening; perithecium curves toward primary receptacle. Trichogyne is nucleate, but
not yet walled off; two nuclei are in centrum (cp) (including ci nucleus); division of apical
outer wall cell rows at upper level precedes that in rows at lower level; w3c' cell not yet
formed. LHe 8 (16).
466

Fig. 14. Herpomyces ectobiae. x 2450. (Only nucleoli are shown in centrum.) a. Four-
tiered perithecium having septum at base of trichogyne (apex of trichogyne is dark-
staining and nucleus stains faintly); first upward extension of inner walls has begun from
w3 cell at lower level at left and from cell w3c'; nucleus appears to be lacking in secondary
upgrowth from ci. LHe 6 (33). b. Four-tiered perithecium with four inner wall cells (only
nucleoli are shown); trichogyne has deteriorated. Centrum may include nuclei other than
the two indicated, but none were clearly visible; second carpogonial upgrowth (cp^ has
extended upward at right to top of third tier of outer wall cells; centrum nucleus at left is
within earlier upgrowth. LHe 8 (20). c. Perithecium that has undergone further
development of centrum; there are two nuclei in secondary upgrowth (cp1) apparently; ci nucleus
appears to be dividing. The uppermost nucleus in centrum seems to be deteriorating; it
appears to be set off in a separate cytoplasmic mass (ap). Two inner wall cells arise from
the ci cell row; one arises from the uppermost outer wall cell row and one is produced by
the outer row at right. LHe 6 (22).
468

Fig. 15. Herpomyces ectobiae. x 2450. (Only nucleoli are shown in centrum in fig. a and
in most cells in figs, b, c.) a. Perithecium with four inner wall cells, which arise from all
outer wall cell rows except that opposite ci. Small apical cells (ap', ap") have been
separated from the upgrowths in the centrum. Two centrum nuclei (ce) are present in addition
to the ci nucleus (the two smaller, lower nucleoli probably are in sister nuclei). Inner wall
cell from w3c' reaches outer surface of perithecium. (Internal cytoplasmic masses not
indicated.) LHe 8 (5). b. Perithecium with seven nuclei in centrum (pair of small nucleoli at
right are probably included within separate cell) (internal protoplasts are not
distinguished). Inner wall cell on upper level at right is in position of w4c' cell of//, periplane-
tae; it reaches the surface. An inner wall cell is formed by the displaced w3c' cell. Dark-
staining remnant of trichogyne is visible at apex of perithecium. LHe 8 (12). c.
Perithecium with six outer wall cells in each vertical row. Inner walls are three cells in
height. The relationships of cells ap' and ap" to cells below have become obscured. Three
elongate asci and one short ascogenic cell (at left) are present in centrum. Carpogonial
initial cell is binucleate and separated from centrum by septum; two small uninucleate cells
lie above it. LHe 8 (20).
470

Figs. 16-18. Herpomyces periplanetae. Acetocarmine-stained specimens. Insect
Toxicology section, Department of Entomology, University of California, Berkeley, 1951, Fe
(OH)j fixative (Figs. 16, a, c, e); Harvard University, Dr. Barbara Stay collection, 1959,
Craf fixative (Figs. 16,b, 17,b,c, 18,b); Insectary, Department of Entomology, Cornell
University, 1960, Fe(OH)3 fixative (Figs. 16, d, 17,a, 18,a). Fig. 16. a. Four-celled thalli
attached to small seta by adhesive disks (fo, fo")\ male at left. Female (right) with
suprabasal cell beginning to grow downward past basal cell. LHp 5 (4). x 2450. b. Young
female (left) and male (right) on small seta. Two-celled perithecium at left is subtended by
two-celled secondary receptacle, arising probably from downgrowth of primary
appendage. Male bears one-celled antheridial appendage on subapical cell of primary
appendage. LHp. 15 (7). x 2044. c. Mature male, producing spermatia; intercellular
pores visible between some cells. Antheridium at right is binucleate; that at left contains
mass of nuclear material apparently remaining from a previous mitosis. LHp 8 (13). x
1650. d. Female with three-celled perithecium (left, subtended by two sr cells, which arise
from downgrowth of suprabasal cell of primary receptacle). Black disk is on wall of basal
cell. LHp 14 (1). x 1639. e. Female with one-celled perithecium at left, subtended by one-
celled secondary receptacle. At right is one-tiered perithecium, with two basal cells
(be, bb) and three sr cells. Another perithecium may have broken away. Suprabasal cell
has produced perithecium at left; two downward-growing cells above it are sister cells of
subapical cell (although these three cells may be primary appendage cells, it is possible that
the primary appendage cell did not divide and they are actually receptacle cells). LHp 5
(7). x 1650.
472

Fig. 17. Herpomyces periplanetae. a. Young perithecium subtended by three basal cells;
mitosis is taking place in one pair of tier 1 (w/) cells, preparatory to formation of tier 2.
Shield-forming secondary receptacle cell (lower level) has divided once. LHp 14 (23). x
2044. b. Two-tiered perithecium with central upgrowth from carpogonium-initiating cell
in wl tier. Cell bd has been cut off from cell bb. Haustorial apertures are visible below two
sr cells. LHp 15 (5). x 2869. c. Four-tiered perithecium with primary trichogyne (tr) and
nucleated secondary carpogonial upgrowth (cp'). Septum separates apical cell (ap) (which
is continuous with the trichogyne extension and has not undergone nuclear division) from
basal part of first carpogonial upgrowth. Cell bd has fused with subtending sr cell. Shield-
forming sr cell at left has produced two narrow shield cells. LHp 15 (4). x 3483.
474

Fig. 18. Herpomyces periplanetae. (Only nucleoli are shown in all upper cells [except asci
in Fig. b].) a. Four-tiered perithecium bearing stump of trichogyne. Septum separates
small cell (ap') from cp' (secondary upgrowth from a cell). Tertiary upgrowth from ci cell
is at left (cp"). The two lowermost inner wall cells arise from the lateral w3 cells; the inner
wall cell at upper level at left is formed by w3c in the ci cell row. The fourth inner wall
arises as an upgrowth (upper right) from w4c', the extra cell in tier 4 in the ci cell row.
LHp 14 (2). x 2869. b. Fully extended nine-tiered perithecium with six tiers of inner wall
cells. Extra cell (ci') has probably been cut off toward the interior by cell ci. Centrum is
filled with spindle-shaped asci, which have probably been formed by two ascogenic cells;
those at upper level appear to have fusion nuclei. Ascogenic cell at lower right has formed
binucleate ascus (above it at left) and uninucleate ascus (above it at right). Some cells have
undoubtedly been lost through break in perithecial wall. The residual ap' cell (possibly a
subsequently formed apical cell) is visible at top of central cavity. Lower outer wall cell
row is not shown above tier 4; only the two upper inner wall cell rows are indicated.
LHp 15 (5). x 2459.
476

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Fig. 19. Semi-diagrammatic drawings of portions of photographed specimens, a. Histeri-
domyces venezolanus — young perithecium, left side of specimen at right, pi. 29,b. b. .
Thaumasiomyces scaurus — PI. 19,b. c. Clonophoromyces grenadinus — PI. 30,b.
d. Smeringomyces anomalus — PI. 31,a. e. Smeringomyces trinitatis — PI. 31,b.
478
a
6
S
6

Fig. 20. Diagram - types of perithecia in the Laboulbeniales.
480
Types of Perithecia in the Laboulbeniales
Laboulbeniineae
Ceratomycetaceae
Laboulbeniaceae
Herpomycetineae

Eus
ynaptomyces
PROPOSED PHYLOGENETIC RELATIONSHIPS IN THE LABOULBENI ALES
APOROMYCETINAE
EUPHORIOMYCETINAE
DIANDROMYCETINAE
AMORPHOMYCETINAE

Fig. 22. Map — distribution patterns in Rhachomyces (based on appendage arrangement
and structure). R. perigonae group: few-celled secondary axis, few appendages irregularly
arranged. R. alluaudii, R. orientalis: similar except appendages regularly arranged.
Usual thallusform: longer axis, row of appendages on one side. R. aphaenopsis, R. gratiel-
lae: similar, but having short cells in appendages. R. velatus group: axis half-enveloped or
totally surrounded by appendages. R. tenuis, R. javanicus, R. beronii: long axis with
appendages along one side and forming dense collar around perithecium.
484

PI. 1. Laboulbenia flagellata. On Agonum ovipennis. a-b. University of California
campus, Berkeley, ca. 1952. Basic fuchsin. LLf 2 (1). x 182. a. Thallus in which ascus
formation has begun, b. Mature thallus. c-e. San Mateo County, California, November,
1956. Acetocarmine. c. Two germinating spores with feet inflated at base; black
coloration extends upward, forming two lines of dark granules along one surface; remainder
of lower half of foot is dull yellow in transmitted light; protoplast in swelling at upper end
of foot tapers downward into narrow cone of cytoplasm (sporeling at left). LLf 3 (16). x
1440. Craf. d. Young sporeling with light-colored area remaining within inflated zone of
foot (this area later becomes filled with dark granules); slight constriction at original
spore septum; appendage has broken off. LLf 7 (8). x 1440. Fe(OH)j. e. Young thallus
after formation of cell V. Septum separating cell IV from cell 77/ has not yet been formed;
division has begun in cell/. Compare figs. lb,c. LLf 7 (2). x 700. Fe(OH)j. f. Thalli on
body of host, x 9.2.
486

PI. 18. Cochliomyces and Tettigomyces. a-c. Cochliomyces trinitatis. FH 4106, Emperor
Valley, Port-of-Spain, Trinidad, West Indies, Thaxter, 2833c, inferior abdomen (slide
lost), x 395. Glycerine, well stained with eosin. a. Perithecium with deteriorating tricho-
gyne; spermatia present in upper part of stalk appendage; there is a constriction below
large protoplasmic mass inside apex of primary appendage, b. Perithecium with tricho-
gyne; spermatia are in neck of terminal antheridium on stalk appendage, c. Early stage of
ascus formation; there is large spermatium-like structure at apex of primary appendage, d-
g. Tettigomyces africanus. FH 3260. On Gryllotalpa, Wambugu, East Africa, Mearns, on
inferior abdomen, x 328. Glycerine, well stained, d. Young thallus, with unbranched
trichogyne bordering compound antheridium above, e. Elongating perithecium; dark-
staining internal fertile cells visible below base of branched trichogyne. f. Older thallus,
undergoing deterioration of upper part of antheridium; lateral rows of inner wall cells of
perithecium are visible, g. Early ascus formation; trichogyne has disappeared, h.
Tettigomyces confusus. Young perithecium with masses of antheridial cells arising from cells
forming upper part of primary axis (above perithecium); other thallus of pair has broken
off below. FH 3286, Gryllotalpa africana, Samarang, Java, Nov., 1909, Jacobson. x
350. Glycerine, i. Tettigomyces vulgaris Thaxter; mature thallus, with primary axis broken
above perithecium. FH 3348, forewings, Gryllotalpa, Zamboanga, Philippines, x 328.
Glycerine.
520
I

PI. 19. Thaumasiomyces. a-b. Thaumasiomyces scaurus. a. Two sporelings, feet upward.
Thallus at right has three cells below black septum (lower arrow); appendage, which
curves around and extends upward at left, has outgrowth (upper arrow) at side of third
cell from apex that will probably form a spermatium; at base of this cell is spermatium-
like structure (arrow at left). FH 7107, on Amphiops globus Erichson, Cameroun, G.
Schwab, 1925. x 510. Glycerine, well stained, b. Thallus with young perithecium; tricho-
gyne emerges just below apex (out of focus) (see Fig. 19b); just above VI is narrow, wedge-
shaped cell, then broad, rounded cell (v-line), from which two appendage branches arise
(arms of inverted y), with appendage on lower level (arrow at right) extending directly
upward (foot of inverted y); appendage at left is obscured by perithecium. until it reaches its
apex (arrow at left); foot spreads irregularly at base of thallus. FH 7111, on Amphiops
globus, Cameroun, West Africa, G. Schwab, 1925. x 540. Glycerine, c. Thaumasiomyces
scabellularius (type), having mature perithecium with coiled neck; intact black secondary
foot at left, broken one at right, with small, colorless primary foot in center (arrow);
inflated appendages at right, with terminal filaments broken off (stump indicated by arrow
at right). FH 7094, on Amphiops asperatus (undescribed?), Cameroun, G. Schwab,
1925. x 260. Glycerine, well stained.
522

PI. 40. Dixomyces. a. D. spiralis, mature thallus with long, spiralled appendages; upper
receptacle cells subdivide; basal cells of appendage are probably derived from one
appendage cell, not from subtending receptacle cells. FH 5293, presumably on Morion guineen-
sis Imhoff (see Thaxter, 1931), Cameroun, West Africa (Thaxter no. 2355). x 380. b. D.
perpendicularis, young thallus, with trichogyne emerging from apex of perithecium,
which has two tiers of outer wall cells; phialides terminate appendages. FH 5277 (type
slide), on Caelostomus (as Drimostomd), Madagascar (Thaxter no. 3126), on lower left
base of abdomen, x 540. Glycerine, c. D. ornatus, mature thallus; perithecium bears
lateral hump (left) and terminal spine (one-celled); appendage branch grows inward from
basal cell of primary appendage (arrow). FH 5268, on Perigona, Samarang, Java, Jacob-
son, 1910 (Thaxter no. 2081f), on middle margin, right elytron, x 700. Glycerine, d-e. D.
compacts. FH 5205, presumably on Caelostomus, Madagascar (Thaxter no. 3126c),
distal on anterior tibia, d. Mature thallus in which divisions have taken place in cell III;
line at left indicates septum at top of first tier of outer wall cells; arrow indicates extra
septum on inner side of perithecium. x 600. Glycerine, e. Young thallus before extension of
trichogyne; outer walls of perithecium are two cells in height, x 760. Glycerine.
564