{'fraction_non_alphanumeric': 0.04566467017727218, 'fraction_numerical': 0.016400691401696345, 'mean_word_length': 4.654090909090909, 'pattern_counts': {'":': 0, '<': 0, '<?xml version=': 0, '>': 0, 'https://': 0, 'lorem ipsum': 0, 'www.': 0, 'xml': 0}, 'pii_count': 0, 'substrings_counts': 0, 'word_list_counts': {'cursed_substrings.json': 0, 'profanity_word_list.json': 0, 'sexual_word_list.json': 0, 'zh_pornsignals.json': 0}}
Laboulbeniales on semiaquatic Heteroptera. VIII. Monandromyces, a new genus based on Autophagomeces microveliae(Laboulbeniales)

A new genus of Laboulbeniales (Laboulbeniaceae: Stigmatomycetinae), Monandromyces, was described. Its type species, M. hemipteralis, was based on Autophagomyces hemipteralis. The latter, which parasitizes a riparian bug, a species of Microvelia (Heteroptera: Veliidae), was characterized by Roland Thaxter in 1931. Ten new species of Monandromycestaken from members of three genera of Veliidae-were described as follows: M. australis, M. falcatus, M. polhemorum, M. protuberans, M. tenuistipitis, and M. umbonatus (on Microvelia spp.); M. neoalardi (on Neoalardus sp.); and M. elongates, M. longispinae, and M. pseudoveliae (on Pseudovelia spp.). Keys to the species were given and all were illustrated with line drawings.

During my studies of Laboulbeniales, I have accumulated specimens and data pertaining to Autophagomyces sensu lato and, on the basis of new information, have revised the taxonomic status of several of the species cited above.
During visits to Dr. John T. Polhemus's laboratory at the Colorado Entomological Museum, Englewood, in April 1991 and May 1993, I was permitted to examine, for Laboulbeniales, accumulations of semiaquatic Heteroptera collected by John and/or his son Dr. Dan A. Polhemus in Africa, Australia, Indonesia, and Malaysia. On species of three genera of Veliidae, Microvelia Westwood, Neoalardus Distant, and Pseudovelia Hoberlandt, I found specimens of 10 distinctive Laboulbeniales having a combination of characteristics matching those of Thaxter's Autophagomyces microveliae, characters that clearly distinguish them generically from the type species of Autophagomyces, A. platensis, and its allies on Anthicidae, as well as those species parasitizing Cryptophagidae, Phalacridae, and Pselaphidae.
The purpose of this paper is to describe, illustrate, and comment briefly on the morphology and development of a new genus to accommodate Autophagomyces microveliae and ten new species.

MATERIALS AND MET HODS
Except for the host of Autophagomyces microveliae, Microvelia albolineolata Bueno, all other hosts of fungi described in this study were collected in the field by John and/or Dan Polhemus and preserved in 70-80% ethyl alcohol. Some specimens later had been pinned (affixed to paper points), dried, and placed in museum boxes, and any of these insects found to bear fungi were relaxed before removal of the parasites. Labels and insects were carefully separated from the pin. Labels were set aside in vials for eventual reunion with the insects. Infected insects were placed directly into 0.2 % saline or a fluid consisting of benzene (5%), ethyl alcohol (45%), water (35%), and ethyl acetate (15%) for a few hours or overnight after which they were removed to 70 % ethyl alcohol for storage along with a duplicate set of labels. Unsorted lots of insects still in fluid were carefully examined with a dissecting microscope and those bearing parasites were segregated and placed, along with appropriate labels, in vials containing fresh 70 % ethyl alcohol.
Using methods described previously (Benjamin 1971: 101-102 [up to step e];1986: 247;1993: 560), I carefully removed the fungi from hosts and mounted them in glycerine on slides. All observations related to the descriptions and illustrations were made using a Leitz Dialux microscope having differential interference contrast optics. In the citations of specimens taken from insects collected by John and/or Dan Polhemus, CL numbers, i.e., collection localities as described in their field notebooks, are given.
Loan of the slides bearing Thaxter's type and isotypes of Autophagomyces m icroveliae was kindly ar-ranged by Dr. Donald H. Pfister, curator of the Farlow Herbarium.
Terminology and abbreviations used in describing the ascoma (used here for the entire perithecium-bearing thallus) are defined in the text or in a separate key to labeling preceding the first set of legends for figures, and mostly are those outlined by Tavares (1985: 43 1-434 ). Ascomata of all of the taxa discussed in this paper are bilaterally symmetrical and usually are oriented on slides so that they can be viewed only laterally. Thus, with few exceptions, descriptions and measurements of thalli were based on study of specimens as seen from one side or the other. When thalli are mounted on slides, the true relationship of the perithecium relative to the receptacle and appendage may, at times, be ± altered. This occurs when the perithecial stalk cell twists one way or the other when the cover glass is applied. An attempt was made to interpret correctly such distortions in preparing the descriptions. With reference to the perithecium and receptacle, anterior (or forward) is in the direction away from the appendage whereas posterior (or backward) is in the direction toward the appendage. In referring to cells of the receptacle, stalk and basal cells of the perithecium, and cells of the appendage, inner (or inward) is in the direction of the longitudinal axis of the ascoma, outer (or outward) is in the direction away from the axis. In all measurements, length (height) precedes width (breadth). Total length of the thallus, where in some instances the receptacle or perithecium may be ± curved, bent, or reflexed, was measured along the median axis. Occasionally, length is the only dimension given for some cells where orientation made meaningful determination of width uncertain, e.g., cell VII, cells n, m, n', and outer wall cells. The length of cells I and II of the receptacle, which are separated by a strongly diagonal cross wall, was, for cell I, measured upward from the tip of the foot and, for cell II, downward from the base of cell VI to the tips of the region of overlap of each cell with the other. Because perithecial basal cell n gives rise to two vertical rows of outer wall cells instead of only one row each as do basal cells m and n', the relationship of a cell (cells) of one of the n-basal-cell-derived rows of outer wall cells to those derived from basal cells morn' is given as ex n adj. m or ex n adj. n', i.e., adjacent to the mor n'-basal-cell-derived row of outer wall cells. TAXONOMY Monandromyces R. K. Benj., gen. nov.
Receptacle conststmg of three in-line superposed cells (1, II, ID) bearing distally on one side a stalked perithecium and on the other side a simple, free appendage; basal (I) and suprabasal (II) cells strongly obliquely superposed ; cell II subtending the perithecium; tenninal cell (ID) subtending the appendage. Appendage consisting of three superposed cells and a single, tenninal, simple, flask-shaped antheridium; basal cell small, broader than long to nearly isodiametric; median cell elongate, several times longer than broad; tenninal cell small, nearly isodiametric or ± elongate, always less than length of median cell; antheridium flask shaped with an elongate, slender efferent tube. Perithecium with two stalk cells (VI , VII), three persistent basal cells (m, n, n ' ), and four vertical rows of outer wall cells of five cells each. Trichogyne fonning slender, divaricate branchlets tenninally; body consisting of a single slender cell or of two superposed cells separated by a median transverse cross wall, the pair of cells abruptly constricted above and below. Ascogenic cell single. Ascospores 1-septate. ) on which the above description is based. Monandromyces microveliae (Fig. 1) differs from the other species parasitic on Microvelia by its erect, nearly straight habit; its relatively long, slender stalk (cells I and II); the pronounced external convexity of the cells comprising the lower three of the five tiers of outer wall cells, especially the median tier; and the ± distally rounded lip cells, which are nearly equal in length.
Etymology.-From umbonatus (L.), in reference to the umbonate terminal outer wall cell. Notes.-Fifteen mature individuals of Monandromyces umbonatus were recovered from a single female host. Features that most readily distinguish this species from others in the genus are: 1) the prominent umbo arising from the posterior cell of the terminal tier of outer wall cells (Fig. 11 , 13); 2) the slightly sigmoid appendage with its asymmetrical antheridial venter (Fig. 12); and 3) the ± strongly convex cells of the lower three tiers, especially the median tier, of outer wall cells (Fig. 13).
Etymology.-Named for John T. and Dan A. Polhemus, father and son respectively, students of Heteroptera, who collected the host.  Notes.-Monandromyces polhemorum was found on the same host as M. umbonatus, which grew on the mid, left, lateral surface of the abdomen. Twelve mature, three nearly mature, and six immature individuals (two with trichogynes [ Fig. 17]) of M. polhemorum were recovered. As the thallus matures, it appears to undergo a slight natural twist of ca. 90°, and in slide mounts of mature specimens the relationship of the appendage to the base of the perithecium could only be observed in far or near view, as depicted in Fig.  15; the distortion appears not to be an artifact of preparation. Except for a slight external convexity of the median cell of its stalk, the appendage is slightly but uniformly tapered upward from the base (Fig. 16). Another distinguishing feature of M polhemorum is the ± pronounced distal inflation of cell ll of the receptacle (Fig. 15).
Ascoma: Erect, nearly straight, except basal cell of receptacle :t strongly curved; receptacle and appendage pale yellowish brown; perithecium and basal cell of appendage dark orange-brown. Length from tip of foot to tip of perithecium 245-260 J.Lm. Receptacle: Elongate, slender, comprising ca. ¥.! of length of thallus, 96-98 J.Lm long from tip of foot to base of appendage; basal cell (I) nearly uniform in width above blackened foot, except slightly externally rounded distally, 30-33 X 8-9 J.Lm; suprabasal cell (II) elongate, slender, nearly uniform in width above juncture with cell I, 65-75 X 7-8 IJ.m; cross wall separating cells I and II angled upward from rear to front; terminal cell small, slightly longer than wide, 10-11 X 8 J.Lm, adnate on inside to base of perithecial stalk cell (VI). Appendage: Free, 52-56 IJ.m long; basal cell slightly wider than high, 5 X 8 IJ.m; median cell as wide as basal cell, externally :t convex, 16-17 X 8 J.Lm; terminal cell slightly higher than wide, 6-7 X 5-{) IJ.m, narrower than basal and median cells; antheridium relatively narrow, 24-25 IJ.m long, venter ca. 5 IJ.m wide, efferent tube 14-15 X 2 J.Lm. Perithecium: Primary stalk cell (VI) relatively long, broadest above, slightly constricted near base, 45 IJ.m long, 14-15 IJ.m wide distally; secondary stalk cell (VII) ca. 8-10 IJ.m high, perithecial basal cells (m, n, n') comprising ca. 25% of total length of body above stalk cells; body straight, uniformly inflated upward from base, broadest near middle, then evenly tapered to apex, ca. 121 IJ.m long above stalk cells, ca. 45 IJ.m wide, basal outer wall cell (ex n adj. m) bearing near its base a very short, broadbased, dome-shaped protuberance, estimated at ca. 2 IJ.m high; perithecial basal cells and cells of basal and suprabasal tiers of outer wall cells with narrow transverse striae of varying lengths arranged randomly or in closely spaced vertical rows adjacent to lateral margins of cells; one cell of terminal tier of outer wall cells (ex n adj. m) slightly longer than others. Ascospores 40-46 X 5 IJ.m. Notes.-Only one very immature and two mature specimens of this distinctive species were found (Fig.  22, 23); they parasitized the same host as M. falcatus, which grew on the left lateral surface of the pronotum. The slender stipe (i.e., cells I, II, and VI) subtending the perithecial body is a prime distinguishing feature of M. tenuistipitis (Fig. 23). Unfortunately, the orientation of both mature individuals in the slide mount precluded accurate study of the small dome-shaped prominence arising from basal outer wall cell ex n adj . m. Its height of 2 IJ.m given in the description is only an estimate; precise dimensions of the prominence must await discovery of additional specimens. 7. Monandromyces protuberans R. K. Benj., sp. nov.
Ascoma: More or less curved or bent forward abruptly near base of appendage; pale yellowish ALISO brown, except perithecium and basal cell of appendage, which are dark yellow brown. Length from tip of foot to tip of perithecium 205-230 J.LID. Receptacle: Nearly straight, ca 40% of length of thallus; 85-110 ~ long from tip of foot to base of appendage; basal cell (I) including blackened foot 36-46 X 8-11 J.LID; suprabasal cell (II) about one third longer than cell I, 59-76 J.Lm long, 13-15 J.Lm wide distally; cross wall separating cells I and II angled upward from rear to front ; terminal cell (Ill) adnate on inside to base of perithecial stalk cell (VI), nearly isodiarnetric, 9-16 X 9 -12 J.LID. Appendage: Free, 65-75 J.Lm long; basal cell nearly isodiarnetric, 7-9 X 9-11 ~;median cell 19-30 X 9-10 J.LID; terminal cell isodiarnetric, 7-9 X 7-  Notes.-Material available for my study of M. protuberans included 12 mature, two nearly mature, and 11 immature thalli, one bearing a trichogyne (Fig. 26). The broad-based protuberance arising from basal outer wall cell ex n adj. m (Fig. 27) readily distinguishes M. protuberans from other species of the genus including M. tenuistipitis, the only other species parasitizing Microvelia that forms a projection, albeit very small, on the same basal outer wall cell. 8. Monandromyces neoalardi R. K. Benj., n. sp. Fig. 30-35.
Etymology.-From elongatus (L.) Notes.-Like several species described in this work, specimens of M. elongatus available for study were limited, only four mature individuals being foun d; however, the species should be readily distinguished from all others in the genus. It differs in several ways from the other two taxa on Pseudovelia in its greater size; its small, median, domelike perithecial outgrowth (Fig. 41); and the precise arrangement of its perithecial sculpturing (Fig. 41). However, like M. pseudoveliae the septum between cells I and II of the receptacle of M. elongatus angles upward from front to rear (Fig.  4 1). Also, in M. elongatus the wall of the elongate median cell of the stalk of the appendage is ± uniform in thickness (Fig. 42), lacking the alternating thick and thin areas characterizing the appendages of M. pseudoveliae (Fig. 39) and M. longispinae (Fig. 48).
Etymology.-From the combination of longi-(in L. comp.), long, + spina (L.), spine,. in reference to the long spine formed by the suprabasal wall cell of the thallus. mature individuals. Those from Africa were fewer in number, including but one mature and three immature individuals from Malawi and two mature and three immature individuals from Tanzania.
The elongate, acuminate perithecial projection (Fig.  47) is the primary characteristic distinguishing M. longispinae from all species in the genus.
The three mature individuals of M. longispinae from Malawi and Tanzania-neighboring countries in mainland SE Africa located slightly NW of Madagascar across the Mozambique Channel-are identical morphologically to one another but deviate in several ways from those collected in Madagascar. They have relatively short perithecial projections compared to those from Madagascar, 63-73 1-Lm vs. 80-125 1-Lm long. The stalk of the appendage is gradually tapered rather than being ± cylindrical, and its median cell has a thicker wall. Also, the perithecial basal cells and outer wall cells are much less conspicuously striate. Finally, the trichogyne of a juvenile from Tanzania ( Fig. 46, but see M. neoalardi [Fig 30, 31]) lacks the basal, septate enlargement of a juvenile from Madagascar (Fig. 45). Because such differences may reflect only minor changes that have occurred following isolation of the precursor of the populations now found in Madagascar and mainland Africa, I have chosen not to separate the latter from the former taxonomically.

Ascospores
Ascospores of Monandromyces spp. were hyaline, ± aciculate, and consisted of two somewhat unequal cells (Fig. 8,14,18,21,25,29,35,40,43,49). The shorter cell, depending on the species, comprised from about two fifths to one third of the length of the body of the spore and was oriented downward in the perithecium prior to spore discharge (e.g., Fig. 9, 11, 32, 39, 41). The sheath enveloping each spore, as it appeared in nonliving, mounted specimens, was in ± close contact with the shorter cell of the body until gradually expanding towards the apex of the longer cell where it extended slightly beyond the tip. (e.g., Fig. 7, 13, 34). Following spore discharge, the distal part of the longer cell differentiated the foot, which served to affix the developing thallus to the host integument (Fig. 22).

Ascoma
Immature individuals with a range of growth stages sufficient for studying thalloid development of one or more species of Monandromyces were unavailable. The juvenile of M. tenuistipitis shown in Fig. 22 was the youngest of any found in the taxa studied. The suprabasal cell (II) had given rise to the perithecial initial (d) and the terminal cell (ID) subtended the fourcelled appendage in which the cell terminating the three-celled stalk had not yet been transformed into an antheridium (an). However, juveniles of several species at the one-to two-outer-wall-cell stage of perithecial development and with well-developed trichogynes were recovered (Fig. 3, 17, 26, 30 , 45, 46). Also, a few young thalli at still later stages of growth aided in interpreting the relationship to one another of perithecial basal and outer wall cells as well as other features of perithecial development (Fig. 44).
Receptacle.-The three cells (1, II, ID) comprising the receptacle in Monandromyces were superposed in line (e.g., Fig. 1, 19, 32, 41). However, early in development the terminal cell (ill) became offset laterally and was united below with cell II by a ± diagonal cross wall and on the inside to the base of the perithecial primary stalk cell (VI), which was subtended by cell II (Fig. 3, 22, 30, 44). From the beginning, apparently, the cross wall separating cells I and II was strongly diagonal and was oriented upward from rear to front in all but two of the species, i.e., M. pseudoveliae and M. elongatus, where the reverse was true (Fig. 39, 41).
There was no evidence of a spine, i.e., the indurated remnant of the tip of the original ascospore, on either the terminal antheridium or another cell of the appendage.
Perithecium.-Although only a few immature individuals were available among the several species of Monandromyces, those that were found provided information on several stages of early perithecial development. The perithecial primordium (d) arose distally from cell II of the receptacle (Fig. 22). Products of division of this cell led to formation of the two perithecial stalk cells (VI and VII) and the three perithecial basal cells (m, n, and n'). The latter gave rise to the four vertical rows of outer (w) and inner (p) wall cells, and the female organ-three superposed organelles consisting of. a basal carpogenic cell (cp), a median trichophoric cell (tc) , and a terminal trichogyne (tr) (Fig. 3). Figures 3, 26, and 30 depict immature individuals at nearly the same stage of development showing the first permanent tier of outer wall cells (w 1 ) , a tier of outer-wall-cell primordia (o), the first tier of inner wall cells (p ), and the female organ, the lower part of which was surrounded by the upwardly growing wall cells.
Trichogynes were present on juveniles of five of the 11 species of Monandromyces studied. In all instances these were terminated by a small cluster of short, ± digitate, coenocytic branchlets, which sometimes emanated directly from a small, ± ovoid or globoid enlargement (Fig. 31 , 45). In three species, M. australis (Fig. 3), M. polhemorum (Fig. 17), and M. protrudens (Fig. 26), the complex of branchlets was subtended by a relatively large, medianly septate, ovoid body that was abruptly constricted basally and distally. In one species, M. neoalardi, only a short, slender nonseptate stalk separated the group of trichogynic branchlets from the tip of the perithecium (Fig. 30, 3 1). However, both types of trichogynes were found in M. longispinae (Fig. 45 [from Madagascar],46 [from Malawi and Tanzania]). In a juvenile of M. longispinae where the perithecium had reached the four-outer-wall stage and the centrum had begun to develop, the trichogyne had degenerated and only a small remnant persisted about midway from tip to base on the surface of primordial outer wall cell o derived from basal cell m (Fig. 44). Such remnants, always small and inconspicuous and often difficult to detect, were seen on mature perithecia of several species. These were located distally on the surface of the tier-four outer wall cell of the vertical row derived from basal cell m near the juncture of this cell with the adjacent tier-four outer wall cell of the row derived from basal cell n (Fig. 13).

DiSCUSSION
Information on the three slides of Autophagomyces microveliae  is written in ink or pencil on a pair of labels affixed left and right on each slide and is inscribed in Thaxter's own hand. At the top of each left-hand label Thaxter wrote, in ink, " No. 3302," i.e., his accession number. Near the top of each right-hand label he wrote, in ink, " Monandromyces." Immediately above and to the right he added " n. g." At the midpoint of each he wrote " on post. leg riparian bug." At the bottom of each he wrote, in two lines, " Fort de Kock Sumatra/Jacobson. Jan. 1925." Thus, in addition to giving data regarding where on the host the fungus grew and where, when, and by whom the host had been collected, Thaxter indicated that his first impulse was to place the fungus in a new genus. However, he subsequently drew a line through "Monandromyces" and immediately below wrote on one slide, in pencil, " Bordea hemipteralis"; on the other two slides he wrote, also in pencil, only " Bordea." Thus, he later considered assigning the fungus to the genus proposed earlier by Maire (1916). Finally, he overwrote the penciled notations, in ink, with " Autophagomyces hemipteralis." However, when he described the species in the last part of his monograph (Thaxter 1931: 96), he did not retain the epithet hemipteralis, using instead microveliae after the host genus, which by then had been identified. Because of Thaxter's practice not to prepare new slide labels after formally describing the Laboulbeniales he studied, his original, hand-written labels often reflect his changing views regarding their taxonomic disposition (see Benjamin 1995: 53).
Because it is available and suitable, I chose Monandromyces-tentatively coined by Thaxter when he first studied his A. microveliae-as the name for the genus described in this study.
My reasons for earlier removing several taxa from Autophagomyces, i.e., A. sarawakensis, A. p eyerimhoffii, and A. sericoderi (to Corylophomyces), and A. p oissonii (to Triceromyces) have been presented elsewhere (Benjamin 1986(Benjamin , 1995 and will not be paraphrased here.
Discovery of ten additional species-from localities ranging from eastern Africa to southwestern Australia-all having the same basic characteristics and parasitizing hosts belonging to the same family of true bugs as those of Thaxter's Autophagomyces microveliae firmed up my long-held opinion that the latter should be distinguished generically, an idea that, as mentioned above, crossed Thaxter's mind when he first studied the species.
Two thalloid characteristics common to all species of Monandromyces distinguish the genus from other possibly closely related genera of Stigmatomycetinae (sensu Tavares 1985) that have simple appendages lacking sterile branchlets: 1) the strongly diagonally superposed basal and suprabasal cells of the receptacle and 2) the considerably greater length of the median cell of the three-celled stalk of the appendage relative to the length of the basal and terminal cell. Also, it may be significant in that all species of Monandromyces known at present parasitize only riparian bugs of the family Veliidae (Heteroptera).
In a paper now in preparation, I will present a detailed study, based on known taxa and several to be described. that will restrict the generic limits of Autophagomyces to those taxa currently known on Anthicidae, Phalacridae, and perhaps Cryptophagidae (Coleoptera: Cucujiforrnia) and reinstate the genus Bordea to include taxa parasitizing Pselaphidae (Coleoptera: Staphyliniforrnia). Structural characteristics of the receptacle, appendage, and especially the trichogyne ser ve readily to separate these genera from one another and from Monandromyces.