Aliso: A Journal of Systematic and Floristic Botany Aliso: A Journal of Systematic and Floristic Botany Corylophomyces, A New Dioecious Genus of Laboulbeniales on Corylophomyces, A New Dioecious Genus of Laboulbeniales on Corylophidae (Coleoptera) Corylophidae (Coleoptera)

A new dioecious genus of Laboulbeniaceae (Laboulbeniales), Corylophomyces, was established to accommodate five species parasitizing Corylophidae (Coleoptera; Cucujoidea): C. peyerimhoffii (=Cryptandromyces peyerimhoffit); C. sericoderi (=Autophagomyces sericoderi); C. sarawakensis (""A. sarawakensis); and two new species, C. reflexus and C. weirii. A key to the taxa was given and all were illustrated with line drawings. Corylophomyces was placed in Amorphomycetinae sensu Ta vares. The other genera included in this subtribe by Tavares in 1985, i.e., Amorphomyces, Dioicomyces, Nanomyces, Rhizopodomyces, and Tetrandromyces, were compared with one another and with the new genus.


hidae (one sp
cies), as well as a species on a semiaquatic bug (Heteroptera; Veliidae).

As characterized by Thaxter (1931 ), Autophagomyces includes Laboulbeniales having a simple, twocelled receptacle with the suprabasal cell bearing a stalked perithecium and a usually simple, sometimes furcate, appendage consisting of usually three or four superposed cells terminated by an antheridial phialide.It should be noted that the terminal cell of a primary three-celled receptacle derived from the lower cell of a two-celled spore was termed the basal cell of the appendage by Thaxter.Thus, in Thaxter's descriptions of Autophagomyces spp., as well as other related genera, the lowermost cell of the appendage is in reality the terminal cell of the receptacle, i.e., cell III as it now is defined (Tavares 1985: 433).The genus is classified in subtribe Stigmatomycetinae (Laboulbeniaceae; Laboulbenieae) by Tavares (1985).

Several species have been added to Autophagomyces since 1931, i.e., A. poissonii R. K. Benj.(Benjamin 1970) ( = the monoecious morph of Triceromyces poissonii [R.K. Benj.]R. K. Benj.[Benjamin 1986]) on Mesoveliidae (Heteroptera); A. falcatus T. Majewski (1973) on Cryptophagidae (Coleoptera; Cucujoidea); A. castellinii W. Rossi (1982) on Pselaphidae; A. tiwaie sis W. Rossi ( 1990) on Pselaphidae; and A. sericoderi Santam.(Santamaria 1993) on Corylophidae.

Three of the species that have been placed in Autophagomyces, namely A. peyerimhoffii (Maire) Maire in Thaxt. (1931: 366) (syn.Cryptandromyces peyerimhoffii Maire [1920: l38]), A. sarawakensis Thaxt. (1931: 94), and A. sericoderi (Santamaria 1993: 412), which parasitize members of Corylophidae, prove not only to be dioecious but also to have other characteristics that clearly separate them from Autophagomyces sensu Thaxter (1931) and Tavares (1985).Accordingly, because these taxa cannot be accommodated in an existing genus, they are here assigned to a new genus to which are added two new species, one found on the host bearing A. sarawakensis.In addition, my purposes shall be to ( l) illustrate the species with line drawings, (2 provide a key to the species, (3) comment briefly on morphology and development, and ( 4) compare the genus with other possibly allied genera of Laboulbeniales.


MATERIALS AND METHODS

While at Harvard University as a National Research Council Fellow many years ago (1951)(1952), where I studied the Thaxter collection of Laboulbeniales at the Farlow Herbarium (FH), I was fortunate in locating, both at the Farlow and the Museum of Comparative Zoology (MCZ), du licates of a few of the insect hosts from which Thaxter had removed Laboulbeniales he later described.These insects originally had been preserved in small, cork-stoppered, glass vials containing alcohol and a small amount of glycerine; only the glycerine remained because of the evaporation of the alcohol.Thaxter had included in each vial a label with minimal data, often only the country of origin of the collection, but in many instances he included the accession number of the host as cited in his published works along with the description of a given taxon.The late Drs.W. Lawrence White, (Director, FH), and P. J. Darlington, Jr., (Curator of Insects, MCZ), permitted me to retain on permanent loan many of these insects and to keep for future reference any specimens of Laboulbeniales removed from them that duplicated species described by Thaxter.Through the years, several of the fungi recovere

supplemented in some
nstances by specimens borrowed from the Thaxter collection, have provided me with valuable information pertaining to my studies of Laboulbeniales.In connection with the present investigation, specimens of A. sarawakensis, as well as a related new species not detected by Thaxter, were found on individuals in two small lots of hosts among the above insects, including duplicates of the same lot from which A. sarawakensis had been taken.Dr. Michael C. Thomas, Florida State Collection of Arthropods, Gainesville, who has examined these insects, found that they belong to the family Corylophidae, not Phalacridae as given by Thaxter.

Through the courtesy of Dr. Donald H. Pfister, Curator of the Farlow Herbarium, I have received on loan Thaxter's slide mounts of A. sarawakensis, including the type.Dr. Sergio Santamaria, Universitat Autonoma de Barcelona, Bellaterra, Spain, kindly has provided me with a loan of mounted specimens of A. sericoderi, including the holotype, and two collections of A. peyerimhoffii from Spain.Finally, Mr. Alex Weir, Stockton-on-Tees, Cleveland, and The Natural History Museum, London, U. K., sent me an undescribed species, congeneric with the others being treated here, which was found on undetermined Cory1ophidae collected in Sulawesi, Indonesia.

All of these fungi were mounted on slides either in glycerine (Thaxter's original specimens and those I re-covered from his duplicate hosts) or Amann's fluid (Santamaria's and Weir's specimens).

A Leitz Dialux microscope equipped with differential interference contrast optics was used for making microscopic observations, and, with the aid of a camera Iucida, for preparing drawings.

Terms and abbreviations as defined in the text are essentially those of Tavares (1985: 431-434).The female ascomata of these fungi typically are positioned so that they can be observed only in lateral view; thus, the drawings and descriptions of these are based, for the most part, on observations of specimens as seen from one side or the other.With reference to the perithecium and receptacle of female individuals, anterior (outward) is in the direction away from the appendage whereas posterior (inward) is in the direction towards the appendage.With reference to cells of the receptacle, stalk and basal cells of the perithecium, and cells of the appendage, inner is in the direction of the axis of the ascoma, outer is in the direction away from the axis.Unless indicated otherwise, height preceds width in the measureme ts given in the descriptions.Dioecious.Male.-Receptacle consisting of two superposed cells; with a single, simple, terminal antheridiom.Female.-Receptacleconsisting of three cells; the basal cell (I) o liquely separated distally from the suprabasal cell (II); the terminal cell (III) united below with cell I and more or less united on the inside below with cell II and above with the base of the primary stalk cell (VI) of the perithecium.Primary appendage, subtended by cell III of the receptacle, consisting of two or three superposed sterile cells, apically rounded or acute.Perithecium, subtended by cell II of the receptacle, with well-defined stalk and basal cells, and four vertical rows of outer wall cells of four or five cells each; trichogyne strongly deflexed, consisting of two cells separated by a transverse cross wall, the proximal cell more or less basally constricted, the distal cell forming slender branchlets terminally.Ascospores 1-septate.Male.-Hyaline except for a partially opaque f<;>ot, elongate, slender; basal cell of receptacle, including foot, ca. 25 X 4 !J-ill, ca.five times longer than the suprabasal cell, which is quadrangular, nearly isodiametric, 4 X 4 !J-ill; antheridium ± abruptly attenuate distally, 10-13 X 4 !J-ill.Total length ca.40 !J-ill.

Female.-Ascoma nearly straight or slightly curved; pale yellowish brown.Total length from tip of foot to tip of perithecium 140-160 !J-ill.Receptacle: Triangular, 35-40 !J-ill high, 14-18 !J-ill wide distally, tapered downward to the acute hyaline tip of the otherwise opaque foot; the basal cell (I) 27-35 X 7-13 !J-ill, subtending the terminal cell (III) and suprabasal cell (II), from which it is separated by a strongly diagonal cross wall; cells II and III ± trapezoidal, united, in part, on the inside and ± parallel to one another; cell II up to twice as high as wide, 14-20 X 6-l 0 !J-ill, subtending the primary stalk cell of the perithecium; cell III l 0-15 X 6-10 !J.ID, subtending the primary appendage.

Appendage: Suffused with brown, consisting of two superposed cells in line with cells I and III of the receptacle, 18-20 !J-ill long; basal cell quadrangular, wider than high, 5-6 X 1 O-Il ~ , 13-15 X 9-11 !J-ill, apically rounded to somewhat acute, separated from the basal cell by a transverse cross wall that becomes darkly suffused with brown.Perit ecium: Primary stalk cell (VI) elongate, 15-30 X 10-15 !J-ill, widest distally, ± constricted immediately above the base, nearly free, slightly incurved; secondary stalk cell (VII) small, nearly isodiametric, lying above cell VI on the outside, partially enclosed distally by the basal cells (m, n, n'), which, along with cell VII, constitute ca.25% of the t tal length of the body of the perithecium above cell VI and envelop the base of the ascigerous cavity; body, including basal cells, 93-105 !J-ill long, slightly inflated, median width 30-40 !J-ill, the outer margin more strongly convex than the inner, abruptly narrowed distally, the apex bluntly rounded, bent slightly inward, more darkly suffused with brown on the outside and inside than laterally, with a narrow, dark, longitudinal subterminal, postero-lateral band on each side, one terminal outer wall cell forming a ± inconspicuous projection ca.2.5 X 2 !J-ill.Ascospores hyaline, 34-41 X 3.5-4 !J-ill.Notes.-Mydescription and drawings of are based on specimens received from Dr. Santamaria.This material includes one nearly mature and two fully mature females, two immature females with early stages of development of the trichogyne, and three males.In addition there are two immature, but aberrant, females, and a mature receptacle and perithecium, which are widely separated from one another.The mature females, however, match closely Maire's (1920) description and somewhat diagrammatic illustration of Cryptandromyces peyerimhoffii, found on Arthrolips obscurus J. Sahib.var.sancta-balmae Abeille (Corylophidae).Maire's description was based on five perithecium-bearing individuals, and certain of the dimensions he cites differ somewhat from those given above, i.e., total length, 125-135 !J-ill; perithecial stalk, 20-23 !J-ill; perithecium 85-95 X 33-35 !J-ill; and ascospores, 33-36 X 4 !J-ill.Balazuc (197 4) records a collection of four specimens of Corylophomyces peyerimhoffii (as Autophagomyces peyerimhojfil) taken from the elytra of Sacium nanum (Muls.)from France.Dr. Tavares kindly sent me a photograph of one of Balazuc's specimens, which shows an ascoma corresponding closely to the taxon as illustrated by Maire in having an acute appendage with a similarly pigmented cross wall separating its two cells.Santamaria (1993) reports a single specimen, also as A. peyerimhoffii, on the elytron of Arthrolips sp. from Spain.

In the course of my studies of Laboulbeniales as a graduate student at the University of Illinois, 1947-


I I

A B cp be 1951, I found single male and female individuals of a dioecious fungus congeneric with but specifically different from the taxa described in this paper.They parasitized a specimen of Molamba fasciata Say (Corylophidae) collected in Arkansas.When I studied the fungus, I suspected that it might be related to Maire's Cryptandromyces peyerimhoffii.Unfortunately, the mounted specimens are now in very poor condition and not suitable for description, and the fungus has not been encountered again.When it is rediscovered and can be properly described, characteristics of the male as given in the generic diagnosis of Corylophomyces will need emendation, for, in addition to a terminal antheridium, two lateral, secondary antheridia arise from cells cut off distally by the receptacle.

It was during my tenure at the Farlow Herbarium, where I first studied Autophagomyces sarawakensis and determined its dioecious character, that I suspected that this taxon and very likely A. peyerimhoffii were misplaced generically (Benjamin 1970).Santamaria's description of A. sericoderi and his report of new collections of A. peyerimhoffii revived my interest in Laboulbeniales parasitizing Corylophidae, and his willingness to share his material with me enabled me to complete a study I started so long ago.Male.-Hyaline except for a partially opaque foot, elongate, slender; basal ce l of receptacle, including foot, 23-25 X 4.5-5 ,_.,m, ca.five times longer than the suprabasal cell, which is slightly higher than wide, 5-6 X 4.5-5 ~J-m; antheridium gradually attenuate distally, 12-14 X 4-5 IJ-m.Total length 40--45 IJ-m.

Female.-Ascoma elongate, nearly straight; hyaline to pale yellow.Total length from tip of foot to tip of perithecium 143-170 ,_.,m.Receptacle: Triangular, 31-39 IJ-m high, 10-13 IJ-m wide distally, tapered downward to the acute, hyaline tip of the otherwise opaque foot; basal cell (I) ca.three times.longer than wide, nearly cylindrical, only slightly wider above than be-low, 23-31 X 6-9 IJ-m, subtending the terminal cell (III) and suprabasal cell (II), from which it is separated by a strongly diagonal cross wall; cells II and III ± trapezoidal, united, in part, on the inside and ± parallel to one another; slightly longer than wide; cell II 9 11 X 5-7 IJ-m, subtending the primary stalk cell of the perithecium; cell III 7-9 X 6-8 ,_.,m, subtending the primary appendage.Appendage: Consisting of two superposed cells in line with cells I and III of the receptacle, 20-24 j..tm long; basal cell quadrangular, with slightly concave margins, nearly isodiametric, 8-11 X 7-10 ~J-m; terminal cell elongate, ± acute, 12-15 IJ-m high, 6-9 j..tm wide at the base, separated from the basal cell by a transverse cross wall.Perithecium: Primary stalk cell (VI) elongate, 25-40 X 10-12 ~J-m, incurved, widest distally, nea ly free, slightly constricted at the base; secondary stalk cell (VII) small, ± triangular in lateral view, lying above cell VI on the outside, partially enclosed distally by the basal cells (m, n, and n'), which, along with cell VII, constitute ca.20 % of the total length of the body of the perithecium above cell VI; basal cells barely enveloping the base of the ascigerous cavity; basal cell m externally ± strongly convex, as are the outer wall cells of the first three tiers; body, including basal cells, 85-105 !J-m long, slightly inflated, broadest medianly, 24-28 j..tm wide, outer margin nearly straight, inner margin convex, gradually narrowed toward the apex; the terminal outer wall cell of the antero-lateral row formed by basal cell n forming an elongate, slightly curved protuberance 5-7 X 3 IJ-m.Ascospores hyaline, 20-22 X 3 j..tm.Notes.-Theslide bearing the holotype, BCB-SS-991, includes three mature females (one lacking the foot), one nearly mature female, one mature perithecium separated from its receptacle, four receptacles with appendages, and six males (one broken); slide BCB-SS-1707 includes five immature females, six males (two broken), and one receptacle with appendage.The feperithecium, gives rise to cells VII, n, and n'; m, perithecial basal cell derived from cell VI, gives rise to one vertical row of wall cells; n, one of two perithecial basal cells derived from cell VII, gives rise to two vertical rows of wall cells; n', the other perithecial basal cell derived from cell VII, gives rise to one vertical row of wall cells; o, a primordial outer wall cell; pa, primary appendage; tc, trichophoric cell, lies between carpogenic cell and trichogyne; tr, trichogyne (also trichogynic remnant); w 1 -,, outer wall cell tiers I (basal) to 5 (terminal); I, basal cell of receptacle; II, suprabasal cell of receptacle; III, terminal cell of receptacle; VI, primary stalk cell of perithecium; VII, secondary stalk cell of perithecium.male of Corylophomyces sericoderi is readily distinguished from that of C. peyerimhoffii, which also has a two-celled appendage, by its nearly straight habit, by its relatively slender acute appendage lacking a pigmented cross wall, and especially by its perithecial apex with its two short prominences, the longer of which is somewhat curved.Male.-Hyaline except for a partially opaque foot, elongate, slender, straight or slightly curved, 43-55 JJ.m long, ± gradually attenuate from near the base to the tip of the antheridium; basal cell of receptacle, 24-27 X 5-6 JJ.m, ca.five times longer then the suprabasal cell, which is nearly isodiametric, 5 X 5 J.lm; antheridiom ± uniformly narrowed toward the apex, 13-23 J.lm long, 4-5 JJ.m wide at the base.

Female.-Ascoma elongate, erect, pale brownish yellow.Total length from tip of foot to tip of perithecium 135~156 J.lm.Receptacle: Triangular, 33-40 J.lm high, 10-15(-20) J.lm wide distally, tapered downward to the acute, hyaline tip of the otherwise opaque foot; basal cell (I) 22-31 X 8-10(-15) JJ.m, in contact above with the terminal cell (III) and suprabasal cell (II), from which it is separated by an oblique cross wall; cells II and III ± trapezoidal, united with each other, in part, on the inside; suprabasal cell slightly higher than wide, ~12 X ~8(-10) J.lm, subtending the primary stalk cell (VI) of the perithecium; cell III nearly isodiametric, 7-12 X 7-12 J.lm, subtending the primary appendage.Appendage: Hyaline, consisting of three superposed cells in line with cells I and III of the receptacle, 24-38 JJ.m long; basal cell quadrangular, slightly wider than high, 9-15 X 6-9 J.lm; median cell about half again as high as wide, ± convex externally, narrowed distally, 12-18(-21) X 9-13(-15) J.lm; terminal cell small, triangular, only slightly higher than wide, 6-11 J.lm high, 5-7 JJ.m wide at the base, separat d from the median cell by a transverse cross w ll.Perithecium: Primary stalk cell (VI) 20-27 J.lm long, 8-14 J.lm wide distally; secondary stalk cell (VII) small, ± triangular in lateral view, lying above cell VI on the outside, partially enclosed distally by the basal cells (m, n, n'), which, along with cell VII constitute ca.20% of the total length of the body of the perithecium above cell VI; basal cells barely enclosing the base of the ascigerous cavity; body, including basal cells, 65-100 JJ.m long, nearly straight, narrowest at the base, inflated distal y, 25-29 JJ.m wide; the apex broad, asymmetrical, with a pair of short, fingerlike protuberances, 5-8 X 3-5 JJ.m, distally on the inside opposite a broad-based, divergent, pointed prominence on the outside; and with a second pair of short, distally rounded protuberances, 5-8 X ~8 J.lm, borne laterally at the level of the median tier of outer wall cells.Ascospores hyaline, 33-35 X 3-4 JJ.m.Notes.-Thethree slides from the Thaxter collection (Ace.Nos.4037-4039) include a total of eight mature and six immature females, one receptacle, and one male.Specimens found on the other hosts from Sarawak (RKB 1331B, 1438B) include four mature females (two in very poor condition), two nearly mature females, 13 immature females, 20 males, and four appendages.This material has been ample for preparing the above revised description of Thaxter's taxon, and for illustrating the male as well as several stages of early development of the ascoma of the female.Male.-Hyaline except for a partially opaque foot, elongate, slender, nearly straight, 35-37 JJ.m long, ± attenuate from near the base of the foot to the tip of the antheridium; basal cell of receptacle 17-20 X 5-6 J.lm, ca.five times longer than the suprabasal cell, which is nearly isodiametric, 5 X 5 J.lm; antheridium gradually attenuate, ± abrup ly constricted apically, 12-13 tJ-m long, 5 tJ-m wide at the base.


Corylophomyces reflexus

Female.-Ascoma strongly recurved inwardly, the perithecial tip reaching nearly to the level of the basal cell of the receptacle, brownish orange, the body of the perithecium darker below the tip and above the primary stalk cell (VI), Total distance from the tip of the foot to the most distant point on the anterior margin of the perithecial body 104--136 tJ-m.Receptacle: More or less triangular, 27-35 tJ-m high, 12-20 f.Lm wide distally, tapered downward to the rounde hyaline tip of the otherwise opaque foot; basal cell (1) mo e than twice as high as wide, 23-28 X 9-12 f.Lm, the inner margin convex, the outer margin ± concave; suprabasal cell ± trapezoidal, slightly less high than wide, 6-11 X 9-13 f.Lm, subtending the primary stalk cell (VI) of the perithecium, from which it is separated by a transverse septum; cell III nearly isodiametric, 6-7 X 8-10 f.Lm, somewhat divergent, the free outer margin strongly convex, united on the inside with cells I, II, and the base of cell VI, broadly united above with the basal cell of the appendage.Appendage: Consisting of three superposed cells, curved outward slightly, 18-24 f.Lm long; basal cell 6-8 f.Lm high, 10-12 f.Lm wide, outer and inner margins strongly convex; median cell 9-13 tJ-m high, 12-17 tJ-m wide, outer and inner margins very strongly convex; terminal cell small, triangular, 4--5 tJ-m high, 5-8 f.Lm wide at the base, separated from the median cell by a transverse cross wall.

Perithecium: Primary stalk cell (VI) 51--65 tJ-m long, broadest distally, median width ca.20 tJ-m, ± inwardly curved; secondary stalk cell relatively small, slightly longer than broad, 14--18 X 11-15 f.Lm, outer margin strongly convex; basal cells (m, n, n') enclosing only the base of the ascigerous cavity, constituting ca.20% of the total length of the body of the perithecium above cell VI; basal cell n strongly externally rounded below; body, including basal cells strongly curved, ca. 110- 1 16 X 46-53 f.Lm; the basal and supra asal outer wall cells derived from basal cell m much shorter than the relatively greatly elongated and somewhat thickened basal and suprabasal cells of the basal-cell-n-derived row of outer wall cells that constitute the prominent anterior margin of the perithecial body; the apex distinguished by the ± protrudent, rounded terminal outer wall cells and a short, divergent, subapical, lateral prominence, 9-11 X 6-8 tJ-m, near the ju cture of the upper two outer wall cells of the lateral row of wall cells derived from basal cell n.Ascospores hyaline, 31-36 X 4-5 f.Lm.

Etymology.-Fromreflexus (L.), reftexed.

Holotype  Male.-Not observed.

Female.-Ascoma strongly inclined inwardly, ± pale yellow brown; distance from tip of foot to tip of perithecium 140-160 J.Lm.Receptacle: Dark brown, triangular, 40-44 J.Lm high, 24-26 J.Lm wide distally, tapered downward to the hyaline tip of the otherwise opaque foot; basal cell (I) triangular, 33-38 J.Lm high, 17-23 J.Lm wide distally, with opaque, dendriform striae as much as 10 J.Lm long extending upward from the foot on the surface opposite the haustoria!opening; cells II and III ± trapezoidal, united on the inside; cell II wider than high, 7-9 X 12-14 j..Lm, subtending the primary stalk cell of the perithecium; cell III about twice as wide as high 8-9 X 15-19 J.Lm, broadly separated from the basal cell of the appendage by a transverse cross wall.quadrangular, about half as high as wide, 6-8 X 16-19 J.Lm; median cel1 dome shaped, 10-12 J.Lm high, 15-16 J.Lm wide at the base; terminal cell relatively small, acute, 7-8 J.Lm high, 4-6 J.Lm wide at the base.Perithecium: Primary stalk cell (VI), free, abruptly constricted near the base where it joins cell II of the receptacle, inflated above, strongly convex on the outside, nearly straight or slightly concave on the inside, upper one half or more dark brown, 30-36 J.Lm long, 23-28 J.Lm wide distally; secondary stalk cell (VII) lying above cell VI on the outside, together with the basal cells (m, n, n') enveloping the base of the ascigerous cavity and constituting ca.one third the total length of the body of the perithecium above cell VI; body, including basal cells, 1 14-153 J.Lm long, nearly uniformly inflated, median width 45-60 J.Lm, abruptly and uniformly narrowed distally, the apex gradually narrowed, rounded, with the terminal outer wall cells projecting only slightly.Ascospores hyaline, ca.45-50 X 3.5-4 J.Lm.

Etymology.-Named for Alex Weir, British student of Laboulbeniales.

Holotype Notes.-Thespecimens of Corylophomyces weirii received from Weir were mounted on four slides and included six mature, one nearly mature, and three immature females as well as three female receptacles with appendages.Weir also forwarded two of the hosts from which he had removed some of the fungi, and from these insects I recovered three more immature females, but no males.None of the immature females examined was at a stage of development showing the nature of the trichogyne, being either too young or too old.The host insects appear, according to Weir, to belong to a single species, and they await identification.They currently are referenced under the British Museum code number given above.

The female of Corylophomyces weirii, like that of C. sarawakens s and C. reflexus has a three-celled appendage.However, as can be seen from an examination of the accompanying figures of these species, they differer greatly from one another in the morphology of their perithecia.The distinctive dendritic striae radiating upward from the opaque foot along the surface of the basal cell of the receptacle of C. weirii appear very early in the development of the receptacle.


OBSERVATIONS


Ascospore

Spores did not appear to differ enough in size in any of the species of Corylophomyces, when observed inside or outside the perithecium, to suggest male-female ascospore dimorphism.The body of the ascospore was divided by a single, transverse cross wall (a) into long and sh rt segments having a ratio to one another of ca.2:1 (Fig. 8 , 9, 33-35, 0-42), with the longest segment uppermost in the perithecium prior to discharge 18,32,37,39,44,45).


Male

In all instances, the male consisted of a receptacle of but two cells, in which the basal cell was uniformly about five times longer than the suprabasal cell.The latter was nearly isodiametric and subtended a single, elongate, more or less flask-shaped or attenuate antheridial phialide (an) (Fig. 1,2,10,12,14,(20)(21)(22)29,36).When viewed face-on, the opaque part of the otherwise hyaline foot, which had been in direct contact with the host, was distinguished by a small, circular, more or less median, clear opening that coincided with the proximal end of the contents (plasmolyzed) of the basal cell (Fig. 10,12,14,22,36).In all instances but one, males and females had been dispersed when mounted.However, in one preparation of C. reflexus where single male and female individuals still were attached to a fragment of host integument, the feet of the two sexes were were in contact.


Female

Receptacle.-Early stages of development of the receptacle of females from the two-celled ascospore were not found.In the juveniles encountered, the receptacle already consisted of three cells (Fig. 3,4,11,13,(23)(24)(25)(26)(27)(28) in which the relatively elongate basal cell (I) was separated from the suprabasal and terminal cells (II and III) by more or less oblique cross walls.CeJls II and III were nearly transversely arranged, with III positioned slightly above the level of II (Fig. 3,4,11,13,15,[23][24][25][26][27][28], and the relationship of these cells remained essentially unchanged as the ascoma matured 16,18,(30)(31)(32).

Appendage.-Inall instances but one, the primary appendage (pa) of the juvenile individuals examined consisted of two or three cells and, depending on the species, was morphologicaly nearly like that of the mature ascoma 11,13,15,16,18,(24)(25)(26)(27)(28)(30)(31)(32).In a single individual of C. sarawakensis, the appendage consisted of a single, relatively large, domeshaped cell, presumably representing the initial stage of development of the appendage from the upper cell of the original ascospore (Fig. 23, pa).

Perithecium.-Theearliest stage of perithecial development was found only in two immature specimens of C. sarawakensis (Fig. 23,24) where the perithecial initial (d) had arisen from the suprabasal cell (II) of the receptacle.As seen in the individual depicted in Fig. 24, the primary appendage (pa) already had reached its three-cell limit before further development of the perithecium from cell d had begun.In the next earliest stage of perithecial development encountered (Fig. 25), four cells, k, j, cp (the carpogonial cell), and e, had arisen from cell d.In the individuals shown in Fig. 3 and 26, cell m, the first of three perithecial basal cells, had arisen distally on the inside from cell k, which had become the primary stalk cell (VI) of the perithecium.As shown in Fig. 3, 25, and 26, the distal part of of cell e had begun to recurve posteriorly.In the young perithecium pictured in Fig. 4, cell j had divided and formed the secondary stalk cell of the perithecium (V

n had cut
ff an elongate cell (here termed be) that had grown upward around the base of the carpogenic cell (cp); cell m had divided and formed a primordial wall cell (o) distally on the inside; and cell e had given rise terminally, to the reflexed trichogyne initial (tr), which had become constricted above its base, and the trichophoric cell (tc), which had begun to grow upward.A somewhat later stage of perithecial development is shown in Fig. 27 where the other

o per
thecial basal cells n and n', derived from what was now cell VII, had formed.From these and basal cell m, the first permanent tier of outer wall cells {w 1 ) had developed, the cells of which had, in tum, given rise to a tier of primordial wall cells (o); the enlarged distal part of the reftexed trichogyne (tr), now strongly constricted near its base, had been divided into two parts by a transverse cross wall.

A few stages of trichogynic development beyond the level of that shown in Fig. 27 were found in juveniles of C. sericoderi and C. sarawakensis.In C. sericoderi, the relatively elongate, strongly reftexed, two-celled trichogyne extended downwardly and inwardly alongside the body of the immature perithecium (Fig. 11,13,15).The lower part of the constricted basal cell formed a lateral, bulbous enlargement broadly attached to the perithecium below the tip.The upper cell gradually

oward t
e apex where it ramified and formed a complex of slender, coenocytic branchlets (Fig. 11,13,15).A mature trichogyne of C. sarawakensis (Fig. 28), which was in poor condition and may have begun to degenerate at the time the specimen bearing it was mounted, appeared to have the same basic structure as that of C. sericoderi.As development of the perithecium progressed, the trichogyne had disappeared except for a portion of its base, which remained in the form of a slightly elevated scar on the surface of the perithecium (Fig. 16,17,(30)(31)(32)(43)(44)(45).This relatively large, persistent remnant was located at the juncture between the hird and fourth outer wall cells of the vertical rows of cells derived from basal cell m and the adjacent row of the two rows derived from basal cell n.

The outer wall of the mature perithecium consisted of four vertical rows of cells derived from the three basal cells, one row each from cells m and n', and two rows from cell n, one row adjacent to the basal-cellm-derived row, the other adjacent to the basal-cell-n'derived row (Fig. 30,31).Only the upper two tiers of inner wall cells could be discerned with any ce tainty in mature perithecia, those lower down having been compressed and rendered ± unrecognizable as ascospores matured within the ascigerous cavity.In fully mature individuals of C. peyerimhoffii (Fig. 6), C. sericoderi (Fig. 18), and C. weirii (Fig. 43, 44) the outer wall appeared comprised of five tiers of cells (w 1 -5 ); however, in C. sarawakensis (Fig. 32) and C. rejlexus (Fig. 37, 39) I could not detect a fifth cell in the basalcell-n-derived row adjacent to the basal-cell-n'-derived row.

In C. peyerimhoffii, C. sericoderi, C. sarawakensis, and C. weirii 16,18,32,43,44), the longest outer wall cells were those comprising the basal tier, with those of the suprabasal tier being only slightly shorter.Together, these cells comprised one half or more of the total length of the perithecial body.In these species and C. rejlexus as well (Fig. 37,38), the shortest cells were those of the third and fourth tiers, which were often subequal.In the recurved body of C. rejlexus, the basal and suprabasal cells of the basalcell-m-derived row were relatively short compared to those of the other three rows (derived from basal cells nand n'), with those of the row comprising the outside of the arc being about twice as long as those comprising the inside of the arc (Fig. 37,38).


DISCUSSION

When Thaxter studied the fungus he described as Autophagomyces sarawakensis he must not have found the single male individual, which would have suggested dioecism, located some distance from three mature, perithecium-bearing individuals closely associated on one of his slides (Ace.No. 4038), otherwise he undoubtedly would have mentioned it-he was careful about such details.Thus, he did not recognize that the fungus possibly was dioecious and that the appendage of the ascoma might be sterile.The apices of the terminal cell of several (four) appendages in Thaxter's preparations contain fra ments of cytoplasm resembling spermatia, and the malformed tip of one appendage is perforated much like that of a simple antheridium (Thaxter 1931: Pl. XVII, fig. 20).Thaxter's observation of these features, which resemble the antheridium of Autophagomyces as he defined the genus (Thaxter 1931: 94 ), especially species on Pselaphidae, apparently is what led him to assign the taxon to Autophagomyces.However, from my studies, it is clear that Thaxter's interpretation of the distal cell of the appendage of A. sarawakensis as a simple, spermatium-forming antheridium was based on artifacts.

Thaxter's handwritten notations (in black ink) on the labels of his slides of A. sarawakensis suggest his early assessment of the fungus prior to its placement in Autophagomyces (Thaxter 1931).When Thaxter actually mounted the specimens after he received the hosts from his correspondent in Sumatra (i.e., Moulton) is unknown to me.On the upper margin of each label is written "n.g.," indicating that initially he thought the fungus might represent a new genus.However, immediately below is written the binomial "Bordea sarawakensis," showing that he considered assigning it to Bordea Maire (Maire 1916), a genus he later transferred to Autophagomyces (Thaxter 1931).There follows "on base left el.minute Nitidu id Phalacridae."Thus, the host first was thought to belong to the Nitidulidae, but "Nitidulid" subsequently was struck through and Phalacridae added.Finally, at the bottom of each label is listed the locality and date, "Kuching, Sarawak, 1912."On one slide (Ace.No. 4039) is noted, in pencil, "near Olibrus," the genus he named as being allied to the host of A. sarawakensis.The name he actually gave to the fungus was not added to the labels.

In his discussion of Cryptandromyces peyerimhoffii, Maire ( 1920) states that he hesitated to classify the fungus in the genus Cryptandromyces as defined by Thaxter (1912: 173), and he stresses the fact that in no instance did the appendage of any of the specimens he studied bear antheridia.He also notes the resemblance of the reduced appendage to that of certain dioecious species.His collection did include a single pair o individuals, which he figures (Maire 1920, Pl. 1, fig. 11 ), in which one is well developed, the other aborted but with the same type of appendage.The male of this taxon, here referred to Corylophomyces, proves, like that of C. sarawakensis, to be small and inconspicuous, so it is not surprising that Maire, who doubtless was not looking for males, did not detect them when he mounted his specimens.The associated pair he illustrates must have consisted of juxtaposed females from which the males had become separated, as often happens when one mounts dioecious species of Laboulbeniales.Later, in correspondence with Thaxter, Maire agreed that the species be transferred to Autophagomyces (Thaxter 1931: 366).

Perithecial ontogeny in Corylophomyces is like that of the perithecia of most genera of subfamily Laboulbenioideae (Laboulbeniaceae) (Tavares 1985) whose morphological development has been described in some detail, e. g., Acallomyces Thaxt.(Tavares 1973), Acompsomyces Thaxt.(Benjamin 1989), Amorphomyces Thaxt.(Tavares 1970), Cupulomyces R. K. Benj.(Benjamin 1992a), Filariomyces Shanor (Shanor 1952), ldiomyces Thaxt.(Benjamin 1983), Laboulbenia Thaxt.(Thaxter 1896;Tavares 1985), Microsomyces Thaxt.(Benjamin 1985), Phalacrichomyces R.K. Benj.(Benjamin 1992b), Prolixandromyces R. K. Benj.(Benjamin 1981), Rhizopodomyces Thaxt.(Benjamin 1979), Stigmatomyces Karsten (Thaxter 1896), Synandromyces Thaxt.(Benjamin 1984), Tavaresiella Majewski (Benjamin 1993 ), and Triceromyces Majewski (Benjamin 1986), in that two stalk cells form three basal cells (n, n', and m) that give rise to four vertical rows of outer wall cells: two rows from cell n, and one row each from cells n' and m.Thaxter considered this a feature of what he termed the "normal" type of perithecial development in Laboulbeniales, for it is the one most often encountered in the order.

Within the Laboulbenieae, Tavares (1985) recognizes a subtribe, Amorphomycetinae, that includes four clearly dioecious genera, morphomyces, Dioicomyces Thaxt., Rhizopodomyces, and Tetrandromyces Thaxt (1912), and one genus, Nanomyces Thaxt.(1931) that may include both monoecious and dioecious taxa.In the context of Tavares's system of classification, Corylophomyces can be placed in Amorphomycetinae.

The receptacle of the female of Amorphomyces and Rhizopodomyces consists of but two cells, whereas that of the other four genera consists of three cells.Species of Rhizopodomyces parasitize Hebridae (Heteroptera) (Benjamin 1979;Thaxter 1931); those of Amorphomyces parasitize Staphylinidae (Coleoptera), where they are known only on species of subfamilies Aleocharinae and Oxytelinae (Tavares 1985;Thaxter 1931).

Eleven species of Amorphomyces have been characterized (Tavares 1985;Thaxter 1931).Tavares describes in detail the development of the receptacle and appendage of the female of Amorphomyces falagriae Thaxt.(Tavares 1970, Fig. 8-12), which undoubtedly is representative of the genus as a whole.The basal cell of the receptacle proper comprises the apparent receptacle.The suprabasal cell, termed cell II-III by Tavares, represents a combination of cells II and III of the three- elled receptacle as found in Dioicomyces and Tetrandromyces.Cell II-III gives rise to the perithecium on one side and a very small, rudimentary, bicellular primary appendage on the other side, and it is closely associated, at the same level, with the true stalk cells and basal cells of the perithecium.Early in the development of the perithecium, the primary appendage becomes embedded in the perithecial wall and is virtually undetectible.The lower segment of the male spore divides and forms a two-celled receptacle; the small, inconspicuous terminal segment, which is separated from the lower segment by a diagonal cross wall and was not detected by Thaxter (1908Thaxter ( , 1931)), gives rise to a single, simple antheridium in line with the receptacle (e. g., Tavares 1970, Fig. 1-3).

Rhizopodomyces includes seven described species (Benjamin 1979;Tavares 1985).In the female, the two-celled receptacle is well separated from the body of the perithecium by the usually greatly elongate primary stalk cell (VI) (e. g., Benjamin 1979, Fig. l A, 2 K).The suprabasal cell, II (actually a II-III cell), typically forms much of the receptacle proper and bears, in addition to the perithecium, a free, well-developed, unicellular appendage, which may or may not persist as the ascoma ages (e. g., Benjamin 1979, Fig. 2 E-K).The basal cell, I, usually forms one or more laterally directed, rhizoidlike outgrowths (e. g., Benjamin 1979, Fig. 1 A, 2 E--K, 4 H).The male consists of three or sometimes four superposed cells in which the basal cell, derived from the lower segment of the spore, constitutes the receptacle.The terminal cell forms a simple antheridium directly (e. g, Benjamin 1979, Fig. 21, 41) or a sterile appendage, which is subtended by one or two divergent antheridia (e. g., Benjamin 1979, Fig. 4D F, 6C).

Nanomyces is represented by three relatively small species that parasitize Labiidae (Dermaptera).Thaxter (1931) characterizes the presumed females as resembling depauperate forms of Dimeromyces Thaxter (1896).However, as Tavares (1985) points out, unlike the female of Dimeromyces, the perithecium has a greater number of wall cells and well-defined basal perithecial stipe, whereas the secondary stalk cell typically is small, relatively inconspicuous, and closely associa ed with the basal cells.Ascospores are more or less obliquely septate and often strongly dimorphic (e. g., Benjamin 1970, Fig. 3 D;Thaxter 1908, Pl. XLII, Fig. 24, 25, 29).The lower cell of the male spore remains undivided and forms the receptacle; the upper cell forms three superposed cells, with the distal cell being converted into a simple antheridium having a terminal or subterminal discharge tube ( . g., Rossi 1993, Fig. 5;Thaxter 1908, Pl. XLII, Fig. 23, 28, 33, 36;1931, Pl. XII, Fig. 6, 7).Thaxter (1912: 168) based Tetrandromyces on a single species, T. brachidae Thaxter found on one of the Staphylinidae, Brachida reyi Sharp, from Argentina.The female of this species (Thaxter 1931, Pl. XIII, Fig. 21, 22) is nearly identical in structure to that of species of Dioicomyces having symmetrical appen ages.The male (Thaxter 1931, Pl. XIII, Fig. 23, 24 ), however, resembles that of Dioicomyces only in having an undivided receptacle.The upper spore segment forms three superposed cells followed above by a pair of cells, united on the inside, bearing four adnate antheridia with only the discharge tubes free.I have in my collection single male and female individuals of an unnamed Tetrandromyces (RKB 396), which is nearly identical to T. brachidae, taken from an unidentified staphylinid from Mindanao, Philippine Islands.

In addition to Dioicomyces, Thaxter named two other dioecious genera from Anthicidae, Dicrandromyces (Thaxter 1931: 72), with one species from the Philippine Islands (Thaxter 1931, Pl. XIII, Fig 16-20), and Triandromyces (Thaxter 1931: 69), with three species from Africa (Thaxter 1931, Pl. XIII, Fig 1-15), which, also like Tetrandromyces, have females closely resembling those of Dioicomyces with symmetrical appendages.The male spore of Triandromyces resembles that of Dioicomyces and Tetrandromyces in having a more or less oblique septum (Thaxter 1931: 61, 6~), whereas in Dicrandromyces the septum is nearly transverse (Thaxter 1931: 72).The mature male of Dicrandromyces and Triandromyces differs from that of Dioicomyces and Tetrandromyces n having a two-celled rather than a one-celled receptacle.Unlike Dioicomyces and like Tetrandromyces, the male in these genera characteristically forms more than one antheridium.In Dicrandromyces, the upper spore segment produces a single cell that subtends a pair of free cells each of which bears a pair of free antheridia.Thus, the male of this genus superficially resembles that of Tetrandromyces, but differs in its two-celled receptacle and free antheridia borne on free supporting cells.In Triandromyces, the upper spore segment forms two or three, but usually two, superposed cells; the distal cell bearing a pair of free antheridia, the subtending cell or cells forming single free antheridia.Tavares (1985), in her treatise on Laboulbeniales, relegated Dicrandromyces and Triandromyces to synonymy with Tetrandromyces, acting, in part, on an earlier query by Thaxter (1931) as to whether these genera and Dioicomyces should be united under a single name.At this time I agree, on the basis of its distinctive mal , that Tetrandromyces sensu Thaxter should be retained as a genus separate from Dioicomyces; however, also on the basis of the male, I regard as uncertain the synonymy of Dicrandromyces and Triandromyces with Tetrandromyces.The males and females of Dioicomyces italic us Speg.(=D.formicomi Thaxt., fide Rossi [Rossi 1993]) (Thaxter 1931, Pl. XII, Fig. 1, 2, 6, 7) and D. indentatus Thaxt.(Thaxter 1931, Pl. XIII, Fig. 26-29) resemble those of Dicrandromyces and Triandromyces except for the male having only one antheridium and a single-celled receptacle.The discovery and study of additional taxa fitting Thaxter's concepts of Dicrandromyces and Triandromyces could provide a better understanding of the taxonomic significance of the male in these genera.It may be that Dicrandromyces and Triandromyces should be united with Dioicomyces or placed in a single genus distinct from either Dioicomyces or Tetrandromyces.

In consideration of the characteristics, briefly summarized above, of the genera Tavares included in her Amorphomycetinae, it i my opinion that Nanomyces and Rhizopodomyces are not closely related to the other three genera or to one another.Dioicomyces and Tetrandromyces (including Dicrandromyces and Triandromyces) probably are closely related in view of the marked similarity of their males and especially their females.Amorphomyces, with its diagonally septate spores, which is a feature common to many speices of Dioicomyces and Tetrandromyces, suggests its relationship to the latter genera, especially Dioicomy- ces (Tavares 1970;Thaxter 1931).However, the modified receptacle and two-celled, albeit vestigial, appendage of the female of Amorphomyces, clearly distinguish the genus from other Amorphomycetinae.

Corylophomyces is here induced in Amorphomycetinae because of features it has in common with Dioicomyces and its presumed allies, e. g., a simple male, a female having a three-celled receptacle, and a small, determinate appendage.Its ascoma has a similar perithecium with four or five cells in each row of outer wall cells, a relatively elongate primary stalk cell, and a smallish secondary stalk cell closely associated with the basal cells.The trichogyne of C. sarawakensis and C. sericoderi resembles that of D. anthici Thaxter (Thaxter 1908, Pl. XLII, Fig. 23) and other speci s of Dioicomyces (unpublished observations) in consisting of two relatively large cells separated by a transverse cross wall and ending in a receptive branch.Downward growth of the trichogyne toward the male, as seen in these taxa, probably should not in itself be ALISO viewed as an indication of close relationship, for this phenomenon appears to be characteristic of other dioecious Laboulbeniales as well, e. g., Aporomyces un- iflagellatus Thaxt.(Benjamin 1989, Fig. 1 ); Rhizopodomyces californicus Benjamin (Benjamin 1979, Fig. 2 H, F g. 3 C-E); and the female of the dioecious morph of Triceromyces biformis Benjamin (Benjamin 1986, Fig. 50, 51).

TAXONOMY

Corylophomyces R. K. Benj., gen nov.Dioecius.Mas.-Receptaculum cellu!arum duarum superpositarum constans; antheridio uno simplici terminali.Femina.-Receptaculumcellularum trium constans; cellulo basali (I) et cellulo suprabasali (II) oblique separatis; cellulo terminali (HI) et cellulo I unitis; basi cellulae VI perithecii et cellua III plus minusve unitis.Appendix primaria cellularum duarum vel trium constans; apice rotundato vel acuto.Perithecium cellulis basilaribus tribus persistentibus et ccllulis parietis externis in quatuor ordinibus longitudinalibus numusquique quatuor vel quinque cellularum.Trichogyna valde deflexa cellularum duarum constans; basi constricta; mmulo tenninali.Ascosporae 1septatae.


2.

Corylophomyces sericoderi (Santam.)R. K. Benj., comb.nov.Fig. 10-19 ==Autophagomyces sericoderi Santam.Nova Hedwigia 56: 412, 1993.


Fig. l- 9 .Fig
9
Fig. l-9.Corylophomyces peyerimhoffii (Fig. 1, 6--9: BCB-SS-1623a; Fig. 2-5: BCB-SS-1713).-I-2.Two males, the one on the right with the foot missing.-3--4.Two juvenile females showing early stages of development of the perithecium.-5-7.Three mature females.Cell VI of the perithecium of the ascoma shown in Fig. 7 was twisted and the perithecial body rotated 180° when the specimen was mounted; thus.the positions of cells VII and m in this individual are reversed in comparison to the same cells in the individual shown in Fig. 6.-8-9.Two ascospores.(Bars 10 J..Lm: A, Fig. 5-7; B. Fig. 1--4, 8-9.)




3. Corylophomyces sarawakensis (Thaxt.)R. K. Benj., comb.nov.Fig. 20-35 =Autophagomyces sarawakensis Thaxt., Mem.Amer.Acad.Arts.16: 94, 1931.


Fig. 20 -
20
Fig. 20-35.Corylophomyces sarawakensis (Fig. 23, 28, 30-35: Thaxter 2374; Fig. 20-2 , 24-27, 29: RKB 1438B).-20-22,29.Four  males.-23-28.Six juvenile females showing several stages of development of the pcrithecium and trichogyne.Details are given in the text.-30-31.Two females in intermediate and late stages of development, respectively.Note position of the trichogynic remnant, and, in Fig. 31, the development of the two distal prominences, characteristic of the species, from primordial outer wall cell ( o) of the row of outer wall cells derived from basal cell n adjacent to the row of cells derived from basal cell m.-32.Mature female.Division into two cells of the distal cell of the row of outer wall cells derived from basal cell n. which gave rise to the terminal prominences, could not be determined and the cell is here designated w"-5 .-33-35.Three ascospores.(Bars= 10 j.Lm: A, 20-31, 33-35; B, 32.)




Fig. 43-45.Corylophomyces weirii (Weir 491).-43-44.Two mature females (holotype: Fig. 43).When mounted, the individual shown in Fig. 44 was compressed slightly, causing a slight longitudinal separation of some of the cells of the perithecial body, which was pushed downward against the upper end of the appendage.-45.Sketch, slightly enlarged, of the tip of the perithecium depicted in Fig. 44.The relationship of three of the four terminal outer wall cells to the vertical rows of outer wall cells derived from basal cells m, n', and n is indicated.The fourth terminal outer wall derived from basal cell n is out of view on the opposite side of the perithecial tip.The apex of the terminal basal-cell-n-derived outer wall cell (ex n) adjacent to the terminal cell derived from basal cell m (ex m) forms a pair of terminal prominences, one much more pronounced than the other.(Bars 10 )Lm: A, 43, 44; B, 45.)




Type species.-Corylophomycessericoderi (Santam.)R.K. Benj..-From the name of the host family, Corylophidae, + myces, fungus.KEY TO THE SPECIES OF CORYLOPHOMYCES A. Appendage of female consisting of two cells . . . . . . . . .B Appendage of female consisting of three cells . . . . . . . .C B. Perithecial apex broadly rounded; terminal outer wall cells hardly protrudent, one with a barely perceptible upgrowth on one side . . . . . . . . . . . . . . . . . . . . . .( 1) C. peye rimhoffii -Perithecial apex attenuate; terminal outer wall cells ± protrudent, one with a distally rounded, curved prolongation . . . . . . . . . . .• . . . . . . . . . . . . . . . . . . . .(2) C. sericoderi C. Female ascoma nearly straight; perithecial apex asymmetric, bearing distally two pairs of shon, rounded prominences . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(3) C. sarawakensis -Female ascoma sharply inclined or recurved; perithecial apex
Etymology emale ascoma strongly inclined; perithecium straight, sym-metric; apex broadly rounded . . . . . . . . . . . . . (5) C. weirii1. Corylophomyces peyerimhoffi.i (Maire) R. K.Benj., comb. nov.Fig. l-9=Cryptandromyces peyerimhoffi Maire. Bull. Soc. Hist. Nat. Af-rique N. 11: 38, 1920.=Autophagomyces peyerimhoffi (Maire) Maire in Thaxt. Mem.Amer. Acad. Ans 16: 366, 1931.
± symmetric . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .D D. Female ascoma strongly refiexed; perithecium curved, asymmetric, apex abruptly nanowed, bearing a single blunt subterminal prominence on one side . . . . . . . . .(4) C. rejlexus




Specimens examined.-MALAYSIA.Sarawak: Kuching, 1912, J. C. Moulton coli., on the base of the left elytron of a minute unidentified species of Corylophidae ("Phalacrid allied to Olibrus" fi e Thaxter[1931]), Thaxter 2374 (HOLOTYPE: FH, Ace.No.4037; !SOTYPES: FH, Ace.Nos.4038,4039); on the base of the left elytron of duplicates of the host that bore the holotype, i.e., Thaxter 2374, RKB 1331B (RSA); "Sarawak": the only data other than Thaxter's accession number, i.e., Thaxter 1820, accompanying the hosts (undoubtedly received from J. C. Moulton to whom Thaxter attributed insects collected in Sarawak[Thaxter 1931: 6)); on the base of the left elytron of an unidentified species of Corylophidae, RKB 1438B (RSA).




Ascoma valde intro recurvatum, auriantiaco-brunneum praeter corpus perithecii fuscatum infra apicem.Thallus 104-136 fLm pede ad marginem antica perithecii.Receptaculum: Plus minusve triangulare, 17-35 ~J.m altum, latitudine distali 12-20 ~J.m; cellula basalis (I) 23-28 X 9-12 fLm, externe convexa, interne concava; c