Pseudothecia globosa vel obovoidea vel e latere compressa, erecta, ostreiformia vel dolabrata, margine substrato affixa, parietibus lateralibus plus minusve convexis, carinam longitudinalem et apicem cristatum formantes. Peridium scleroparenchymatosum, tenuitunicatum; hamathecium pseudoparaphysium trabeculatarum matrice gelatinosa circumdatarum  (maturitate saepe invisibilium). Asci bitunicati e basi (raro e latere) oriundi, 8 ascosporas quasi uniseriatas vel biseriatas vel in duobus fasciculis aggregatas continentes. Ascosporae forma diversa, scolecosporae vel didymosporae vel phragmosporae vel dictyosporae, hyalinae, cito brunnescentes, plerumque bipolariter symmetricae.

Pseudothecia globoid to obovoid to strongly laterally compressed, erect, oyster-shaped (conchate) or hatchet-shaped (dolabrate), standing on edge, with lateral walls more or less connivent and extended vertically to a prominent longitudinal keel or cristate apex. Peridium scleroparenchymatous and thin-walled, enclosing a hamathecium of narrow trabeculate pseudoparaphyses, borne in a gel matrix, which are often sparse to lacking at maturity. Bitunicate asci borne in a basal, rarely lateral, orientation within the centrum, containing eight ascospores, overlapping uniseriate, biseriate or in one or two fascicles. Ascospores diverse, ranging from scolecospores to didymospores to phragmospores to dictyospores, hyaline, yellow to dark brown in pigmentation, and generally showing bipolar symmetry.

Ascomata a modified hysterothecium, progressively dichotomously branched, laterally anastomosed along their length to form radiating flabelliform or pseudo-stellate composites, seated upon a conspicuous brown felt-like subiculum, sometimes borne in a stroma. In vertical section, hysterothecia globose to obovoid, typically with a thick three-layered peridium, but fragile, unlike the robust peridium of the Hysteriaceae, composed of small pseudoparenchymatous cells, the outer layer heavily encrusted with pigment and often longitudinally striate on the surface, the middle layer lighter in pigmentation and the inner layer distinctly thin-walled, pallid and compressed. The hamathecium is composed of persistent narrow cellular pseudoparaphyses, often borne in a gel matrix, with tips darkened or branched at maturity. Bitunicate asci are borne in a basal layer and at maturity are typically clavate to cylindric, bearing eight ascospores, overlapping biseriate, ranging from hyaline to light yellow, uniseptate, conspicuously constricted at the septum, fusoid in outline, with at least one end, often both, acuminate, and showing bipolar asymmetry.

Remarks: Corda (1842) originally proposed the Gloniaceae Corda as a sub-familial taxonomic rank under the family Hysteriaceae, in which he placed both Hysterographium and Glonium. Here, we emend and restrict this sub-familial rank and elevate it to family rank. We include the genus Glonium as circumscribed first by von Höhnel (1918) and then by Petrak (1923a); Hysterographium we retain under the Hysteriaceae (Zogg 1962). We feel justified in retaining the Gloniaceae and, more importantly, raising it to family rank for a single genus because of the high statistical support the group receives in our four-gene analysis (Figs. 2 & 3 in Boehm et al. 2009a). Also, the isolates used to define the group are geographically diverse, from Tasmania and the United States, and, most importantly, include the type species as well (Table 1). However, at present we do not include the Gloniaceae within the newly created order Mytilinidiales. This is because of the highly divergent morphology associated with the genus Glonium, as compared to the current genera within the Mytilinidiaceae. These include character states associated with pseudothecial shape, peridial wall thickness, hamathecial type and ascospore symmetry (Barr 1987). Additionally, not enough taxa have been sequenced for the Gloniaceae to establish whether in fact they deserve ordinal status. Thus, for the present, we propose that the family Gloniaceae be considered Pleosporomycetidae incertae sedis. Nevertheless, our four-gene phylogeny does indicate that, despite highly divergent morphologies, especially those related to the hamathecium (i.e., persistent cellular pseudoparaphyses in the Gloniaceae versus narrow trabeculate pseudoparaphyses in the Mytilinidiaceae), the two families are in fact closely related (Boehm et al. 2009a). 

 

NEW GENERA: 

Oedohysterium E.W.A. Boehm & C.L. Schoch, gen. nov. 

Etymology: Greek, Oedo- meaning swollen, referring to the swollen supra-median cell of the ascospores and Hys- from Hysterium.

Hysterothecia solitaria vel gregaria, iuvenia erumpentia, deinde superficialia, navicularia, nonnumquam linearia, plus minusve parallela, neque confluentia, nonnumquam angulo inserta, raro flexuosa vel furcata,  plerumque utrinque obtuse, et fissura longitudinali prominente praedita. Latitudo altitudine minor vel major. Peridium crassum, carbonaceum, maturum fragile, per longitudinem striatum, basim versus incrassatum, sursum attenuatum, bistratosum. Pseudoparaphyses cellulares, 1-2.5 µm latae, hyalinae, septatae, sursum ramosae, vulgo epithecium pigmentatum ascos obtegens formantes. Asci cylindrici vel clavati, bitunicati. Ascosporae irregulariter biseriatae, phragmoseptatae (dictyoseptatae), fusiformes, curvatae, utrinque angustatae, ad septum medium constrictae, (4-)6-8(-11) septis divisae, primum pallide luteae, deinde brunnescentes. Cellula (raro duo cellulae) ascosporarum supramediana conspicue inflata. Anamorphe ad Septonema pertinens.

Hysterothecia isolated to gregarious, erumpent when young, superficial when mature, navicular, sometimes linear in more or less parallel rows, but non confluent laterally, or sometimes situated at angles, rarely flexuous or bifurcating, usually with obtuse ends, and with a prominent longitudinal slit. Sometimes, taller than wide, other times wider than tall. Peridium thick, carbonaceous, brittle with age, longitudinally striated on the margins, thickened towards base, less thick apically, composed of two to three distinct layers, the inner compressed and pallid, the outer thickened and pigmented. Pseudoparaphyses cellular, 1-2.5 μm wide, hyaline, septate, branched above, forming a usually pigmented epithecium above the asci. Asci cylindrical to clavate, usually short stipitate, and bitunicate. Ascospores irregularly biseriate in ascus, typically phragmospores, in one case dictyospores, curved, fusiform, with tapering apices, constricted at the median septum, with (4-)6-8(-11) septa, at first hyaline-yellow, then pigmented sepia to brown at maturity. Genus characterised by a swollen or tumid supra-median cell, rarely with two cells swollen. Anamorph: Septonema Corda. 

Type: Hysterobrevium mori (Schwein.) E.W.A. Boehm & C.L. Schoch, comb. nov.

Etymology: Hystero- from Hysterographium, Latin brevis, short, referring to the spores of the type, Hb. mori.

Hysterothecia navicularia, fissura longitudinali prominente praedita, utrinque acuminata vel obtusa, linearia vel flexuosa, solitaria vel gregaria, vulgo per longitudinem striata, nonnumquam erecta, quasi stipitata, superficialia vel partim in substrato immersa. Asci bitunicati, cyindrici vel clavati. Dictyosporae pigmentatae vel hyalinae, plerumque breviores quam 25 µm, ad septum medium constrictae; ascosporae hyalinae vel luteae iuvenes vulgo strato mucido circumdatae; pigmentatae pallide brunneae, pariete levi; ascosporae ovoideae vel obovoideae, apice obtuso vel acuminato, 3-4(-6) septis transversalibus et 1-2 longitudinalibus divisae.

Hysterothecia navicular, with a prominent longitudinal slit, variable with acuminate to obtuse ends, linear to flexuous, solitary to densely gregarious, surface usually longitudinally striate, sometimes erect, superficial, almost stipitate, to erumpent and partially embedded in substrate, the latter especially when gregarious. Asci bitunicate, cylindrical to clavate. Ascospores pigmented or hyaline dictyospores, usually less than 25 μm long, constricted at least at the median septum. If hyaline to pale-yellow, then typically associated with a gelatinous sheath when young, dissipating with age. If pigmented then lightly so, transparent clear brown, walls smooth; ascospores generally ovoid to obovoid, with either obtuse or acuminate ends, 3-4(-6) transverse septa, and 1-2 longitudinal septa, these mostly associated with the two central cells, but highly variable and sometimes at oblique angles in the end cells.  

Psiloglonium Höhn., Annales Mycologici 16(1,2): 147. 1918.

Boehm EWA, Schoch CL, & Spatafora JW (2009). On the evolution of the Hysteriaceae and Mytilinidiaceae (Pleosporomycetidae, Dothideomycetes, Ascomycota) using four nuclear genes. Mycological Research 113: 468.  

? Glonium subgen. Psiloglonium Höhn., Annales Mycologici 16 (1,2): 145. 1918.

Typus: Psiloglonium lineare (Fr.) Petr., Annales Mycologici 21 (3,4): 227. 1923.

Hysterothecia isolated or gregarious, erumpent to entirely superficial, navicular to linear to highly flexuous, even triradiate, often striated on the outer margin, sometimes arranged in parallel orientation and confluent linearly to some degree, but never dichotomously branched and never associated with a subiculum or stroma. In vertical section, hysterothecia are globose to obovoid, typically with a thick three-layered peridium, composed of small pseudoparenchymatous cells, the outer layer carbonaceous, the middle layer lighter in pigmentation and the inner layer distinctly thin-walled, pallid and compressed. Hysterothecium stout, and stronger than that found in the Gloniaceae. Cellular pseudoparaphyses are persistent, narrow, often borne in a gel matrix, and with tips darkened or branched at maturity above the asci. Asci cylindric, clavate, and octosporous. Ascospores uniseptate, hyaline to pale yellow in mass, obliquely uniseriate or sub-biseriate, oblong or ovoid, the lower cell often narrower and constricted at the septum, usually with at least the apex of the upper larger cell broadly rounded, although there are instances where both ends of the ascospore may be acuminate, as in the genus Glonium.

Etymology: Named after the late Dr Margaret E. Barr, preeminent American mycologist.

Ascomata inconspicue hysterothecioidea, modice compressa e latere in parte superiore, paulo conniventia, sulco inconspicuo angusto, latera paucis striis profundis praedita; ascomata recta vel flexuosa, sessilia, raro furcata, matura altiora quam lata, 1-2.5 mm longa, 250-450 µm alta, 200-300 µm lata. Pseudoparaphyses hyalinae, cellulares, 1-2 µm latae, supra ascos ramosae epithecium formantes. Asci bitunicati, cylindrici, breviter stipitati, (110-)125-135 x 15-20 µm. Phragmosporae fusiformes, angustae, rectae vel paulo curvatae, primum hyalinae, maturae pallide luteae, quaque cellula guttulis magnis refringentibus repleta, (7-)9(-11)-septatae, (35-)40-45(-55) x (7-)9-10(-12)  μm.

Ascomata atypically hysterithecioid, somewhat laterally compressed in the upper region, slightly connivent, sulcus very shallow, existing as a narrow rim, sides laterally striate, striae few and deep, straight to flexuous, sessile on the substrate, rarely bifurcating, taller than wide at maturity: 1-2.5 mm long x 250-450 μm high, 200-300 μm wide. Pseudoparaphyses hyaline, cellular, 1-2 μm wide, branched above the ascal layer to form an epithecium. Asci bitunicate, cylindrical, short-stipitate, (110-)125-135 x 15-20 μm (n = 9). Phragmospores fusiform, narrow, hyaline and straight when young, becoming pale-yellow to lightly clear-brown, and curved when mature, highly guttulate, with guttulae large, highly refractive, present in every cell, with (7-)9(-11) septa, measuring (35-)40-45(-55) x (7-)9-10(-12) μm when mature (n = 27).

Specimens examined: U.S.A., Tennessee, Sevier Co., Great Smoky Mountains National Park, Elkmont, Little River Trail, 35˚ 39' 13.4'' N, 83˚ 34' 44.7'' W, 686 m elev., 5 Nov. 2007, A.N. Miller, S.M. Huhndorf, J.L. Crane, T.J. Atkinson, I. Promputtha, M. Grief, G.K. Mugambi, & P. Chaudhary, deposited as ILLS 59908 (ANM 1495) = holotype; BPI 879783 = paratype; Tennessee, Sevier Co., Great Smoky Mountains National Park, Chimney Tops Picnic Area, Cove Hardwood Loop Trail, 35˚ 38' 10.7'' N, 83˚ 29' 32.1'' W, 4 Nov. 2007, A.N. Miller, S.M. Huhndorf, J.L. Crane, T.J. Atkinson, I. Promputtha, M. Grief, G.K. Mugambi & P. Chaudhary, deposited as ILLS 59907 (ANM 1442), and BPI 879784.

Notes: A superficial resemblance exists between Hysterium barrianum in Clade C, with H. sinense in Clade D. The phragmospores of H. barrianum (Fig. 2N-Q) have a similar number of septa, (7-)9(-11), as those of H. sinense (Fig. 3H), the latter with (3-)5-9(-11) septa. The two species also have spores of similar length. However, the width measurements of H. barrianum, (35-)40-45(-55) x (7-)9-10(-12) μm, serve to separate it from H. sinense, (34-)38-50 x 11-15 μm. Most importantly, H. barrianum does not possess a swollen or tumid supra-median cell, as does H. sinense and the closely related H. insidens. Furthermore, H. barrianum is highly guttulate, and lightly pigmented at maturity, whereas H. sinense and H. insidens possess few if any guttulae, and are much darker in pigmentation at maturity. Lastly, molecular data place the species in different groups within the Hysteriaceae. 

 
Gloniopsis arciformis E.W.A. Boehm, G.K. Mugambi, S.M. Huhndorf & C.L. Schoch, sp. nov.

MycoBank MB515331

Boehm E.W.A., Mugambi G., Miller AN, Huhndorf SM, Marincowitz S, Spatafora JW & Schoch CL. 2009b. A phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae & Gloniaceae, with keys to world species. Studies in Mycology 64: 65.  

Etymology: Latin arcus, a bow or arch, referring to the arcuate or arciform dictyospores.

Hysterothecia solitaria vel pauca aggregata, recta vel flexuosa, carbonacea, plerumque erecta, conspicue applanata et altiora quam lata, (0.5-)1-2.5 mm longa, 250-350 μm lata, 400-600 μm alta, per longitudinem striata, sulco inconspicuo maturitate clauso. Peridium 40-75 µm crassum in medio, basim versus crassius, sursum tenuius, bistratosum. Pseudoparaphyses cellulares 1-1.5 µm latae, ramosae, sursum magis crassitunicatae, epithecium pigmentatum ascos obtegens formantes. Asci cylindrici vel clavati, stipite sinuoso, bitunicati, 50-75 × 14-18 µm; ascosporae irregulariter biseriatae, dictyosporae, pigmentatae, tenuitunicatae, fragiles, facile dilabentes, conspicue arcuatae, 3-5(-7)-septatae, 1-2(-3) septis verticalibus divisae; cellulis centralibus multo maioribus quam distales, ad septa haud constrictae, (10-)12-18(-22) x 6-10 μm.

Hysterothecia solitary to sparsely aggregated, straight to flexuous, carbonaceous, mainly erect, distinctly flattened and taller than wide, (0.5-)1-2.5 mm long, 250-350 μm wide, by 400-600 μm high, longitudinally striated, with an inconspicuous sulcus remaining closed at maturity. Peridium 40-75 µm thick medially, thicker towards the base, thinner towards the sulcus, composed of two layers, the inner thin, compressed and hyaline, the outer denser, and darkly pigmented. Pseudoparaphyses cellular 1-1.5 μm wide, branched and thicker-walled distally towards the top, forming a pigmented epithecium above the asci. Asci cylindrical to clavate, with a sinuous stalk, bitunicate, 50-75 x 14-18 µm (n = 7), ascospores irregularly biseriate. Ascospores pigmented, thin-walled, dictyospores, fragile, easily breaking under the slightest pressure, pronouncedly arcuate or bent (arciform), and thus highly asymmetric, 3-5(-7)-septate, with 1-2(-3) vertical septa, these mostly associated with the mid cells, which are much larger and swollen than the end-cells, no septal constrictions, measuring (10-)12-18(-22) x 6-10 μm (n = 17).

Specimen examined: Kenya, Coast Province, Malindi District, Arabuko-Sokoke National Park, 6 Nov. 2006, G.K. Mugambi. Deposited as BPI 879774, holotype [formerly, GKM L166A (EA)]. 

Notes: Gloniopsis arciformis is represented by a single specimen (BPI 879774) of only ~30 fruitbodies in the protected crevice of a small piece of decorticated hardwood, collected in Arabuko-Sokoke National Park, Malindi District, Kenya. Although the material is sparse, it does permit the description of a new species on account of the highly unusual arcuate dictyospores. Gloniopsis arciformis resides in Clade D, and is phylogenetically closely associated with two other species of Gloniopsis (Gp. praelonga and Gp. subrugosa), as well as with an additional new species described below.  

 

Hysterothecia navicularia, carbonacea, recta vel flexuosa, utrinque obtusa, dense aggregata, erumpentia, ad latera inconspicue striata vel levia, (0.5-)1-3 mm longa, 250-350 µm lata, 250-350 µm alta. Peridium prope basim ad 100 µm crassum, bi- vel tristratosum, stratum internum compressum, hyalinum, strata exteriora densiora et fusca. Pseudoparaphyses cellulares, septatae, 1-1.5 µm latae, sursum ramosae et anastomosantes, epithecium pigmentatum ascos obtegens formantes. Asci cylindrici vel clavati, bitunicati, 60-80 x 12-16 µm, ascosporas irregulariter biseriatas continentes. Ascosporae dictyoseptatae, pigmentatae, obovoideae, tenuitunicatae, fragiles, polis asymmetricis: apice obtuso, ad basim acuminatae vel nonnumquam protrudentes, 3(-4)-septatae, 1-2 septis verticalibus, utrinque saepe septis obliquis divisae, ad septa vix constrictae, iuvenes guttulis repletae, (12-)15-18(-19) x 5-7(-8) μm.

Hysterothecia navicular, carbonaceous, straight to flexuous, with obtuse ends, densely aggregated, erumpent, slightly striated laterally to smooth, (0.5-)1-3 mm long, 250-350 μm wide, by 250-350 μm high. Peridium up to 100 µm thick at base, composed of composed of two to three layers, the inner thin, compressed and hyaline, the outer two progressively denser, and darkly pigmented. Pseudoparaphyses cellular, septate, 1-1.5 μm wide, branched, anastomosed distally, forming a pigmented epithecium above the asci. Asci cylindrical to clavate, bitunicate, 60-80 x 12-16 μm (n = 5), ascospores irregularly biseriate. Ascospores pigmented dictyospores, in outline obovoid, thin-walled, very fragile, spore apices asymmetric, the upper obtuse, the lower acuminate and sometimes drawn out, 3(-4[rarely])-septate, with 1-2 vertical septa, often with oblique septa in end cell, hardly constricted at the septa, highly gutulate when young, (12-)15-18(-19) x 5-7(-8) μm (n = 14). Known from only one collection, from Kenya, East Africa.

Specimen examined: Kenya, Coast Province, Malindi District, Arabuko-Sokoke National Park, 6 Apr. 2005, G.K. Mugambi. Deposited as BPI 879775 = holotype; GKM 1010 (EA) = paratype.

Notes: Molecular data indicate that both Gp. kenyensis and Gp. arciformis are closely associated, adjacent to Gp. praelonga and Gp. subrugosa in Clade D. The spores of all four taxa, however, are different, and thus their association would not have been predicted based on traditional morphology. The spores of Gp. kenyensis do bear a close resemblance, however, to those of Hb. mori. Both have predominantly 3-septate, thin-walled, pigmented dictyospores, with 1-2 vertical septa, often with oblique septa in the end cell. They can be differentiated on spore size: (12-)14-22(-26) x (5-)7-10(-11) μm for Hb. mori, versus (12-)15-18(-19) x 5-7(-8) μm for Gp. kenyensis. The spores of Hb. mori are usually longer and wider, and also show prominent septal constrictions, especially when fresh and hydrated. Additionally, Gp. kenyensis is highly guttulate when young, where this is rarely observed in Hb. mori. Molecular data indicate that they are not related. 

MycoBank MB515422.

Boehm E.W.A., Mugambi G., Miller AN, Huhndorf SM, Marincowitz S, Spatafora JW & Schoch CL. 2009b. A phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae & Gloniaceae, with keys to world species. Studies in Mycology 64: 59.       

Basionym: Hysterium insidens Schwein., Trans. Amer. Philos. Soc., New Series 4(2): 244 (1832).
        ≡ Hysterographium insidens (Schwein.) Sacc., Syll. Fung. 2: 778 (1883).
= Hysterium complanatum Duby, Mém. Soc. Phys. Hist. nat. Genève 16(1): 38 (1862).
= Hysterium depressum Berk. & M.A. Curtis, Grevillea 4(29): 10 (1875).  
= Hysterium fusigerum Berk. & M.A. Curtis, Grevillea 4(29): 11(1875) (as ‘fusiger').
= Hysterium berengeri Sacc., Syll. Fung. 2: 751 (1883). 
= Hysterium janusiae Rehm, Hedwigia 37: 299 (1898). 
= Hysterium apiculatum Starbäck, Bih. K. Svensk. Vet.-Akad. Handl. 25(1): 19 (1899).
= Hysterium batucense Speg., Revista Fac. Agron. Univ. Nac. La Plata 6(1): 116 (1910). 
= Hysterium andicola Speg., Anal. Mus. Nac. Hist. Nat. B. Aires 23: 85 (1912).
= Hysterium atlantis Maire,  Mém. Soc. Sci. Nat. Maroc. 45: 35 (1937). 
= Hysterium lavandulae Urries, Anal. Jard. Bot. Madrof 1: 64 (1941).

Hysterothecia isolated to gregarious, variably erumpent to sessile, 0.5-2.5 mm long, 0.2-0.5 mm high, lying parallel, but not confluent laterally, generally in line with the grain of the wood, and striated laterally with age.  Pseudoparaphyses hyaline, cellular, 1-2.0 μm wide, walls thickened at apices, forming an epithecium, borne in mucilage, above the ascal layer, often encrusted with dark, pigmented crystals. Asci bitunicate, cylindrical, 8-spored, irregularly biseriate, 130-150 x 15-24 μm, short stipitate, and with a prominent apical nasse, especially when young. Ascospores phragmospores transversely (4-)6-8(-11)-septate, constricted at the median septum, inequilateral, slightly curved, at first hyaline-yellow, then brown at maturity, with a prominent swollen supra-median cell. If 5-septate, then swollen cell located at the second position; if 6-septate, then often the third from the top, measuring (20-)23-28(-38) x (5-)7-10(-13) μm. Principally North- and South-America, and Europe (Italy), from bark and old wood of Pinus, Larix, Castanea, Quercus, Eucalyptus, Fraxinus, Aspidosperma, and Lavandula (Zogg 1962). Also reported from South Africa (van der Linde, 1992). Anamorph: Septonema spilomeum Berk. 

 

Oedohysterium sinense (Teng) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515423.

Boehm E.W.A., Mugambi G., Miller AN, Huhndorf SM, Marincowitz S, Spatafora JW & Schoch CL. 2009b. A phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae & Gloniaceae, with keys to world species. Studies in Mycology 64: 59.      

Basionym: Hysterium sinense Teng, Sinensia 4: 134 (1933).

= Hysterium macrosporum Teng, Sinensia 4: 134 (1933), non Peck, Rep. N.Y. St. Mus. nat. Hist. 26: 83 (1874) (1873).

Hysterothecia scattered to subgregarious, linear, sometimes parallel but non-confluent laterally, more often lying at irregular angles, depending on the grain of the substrate, striated in age, usually of a similar size (2-3.5 mm in length), that is, maturing synchronously in a given colony. Pseudoparaphyses hyaline to pale-yellow, cellular, 2-2.5 μm wide, apically branched, walls of even thickness along length, forming a darkened gelatinous epithecium above the ascal layer, +/- encrusted with pigmented crystals. Asci bitunicate, cylindrical, 8-spored, irregularly biseriate, 140-170 x 26-30 μm, short-stipitate, ascospores biseriate to sub-seriate in ascus, with a prominent apical nasse, especially when young, but sometimes persisting through maturity. Ascospores large, fusiform, asymmetric, curved phragmospores, at first hyaline, then pale-yellow to -brown, finally deep brown at maturity, with (3-)5-9(-11) septa, with a medial septal constriction, measuring (34-)38-50 x 11-15 μm, and, like Od. insidens, with a prominent swollen or tumid supra-median cell, usually located just above the median septum. From North America (Boehm, unpubl. data), Europe (Zogg 1962), China (Teng 1933), and South Africa (van der Linde 1992), on decorticated hardwood trees and structures (e.g., aged fence posts).

Notes: Species of Oedohysterium belonging to Clade D (Boehm et al. 2009b) are characterised by elongate asymmetric spores with more than 3 septa, typically showing a swollen or tumid supra-median cell. In this study, two single-ascospore isolates of Od. sinense, one from South Africa (CBS 123345 / BPI 878730), and one from the United States, New Jersey (EB 0339 / BPI 879800), cluster with two isolates of Od. insidens, both from the United States, Massachusetts (CBS 238.34) and Tennessee (ANM 1443 / BPI 879799). Both species have remarkably similar phragmospores (e.g., Fig. 3D versus Fig. 3H). As these two taxa belong to Clade D and are far removed from the type species, H. pulicare, in Clade C, we propose that they be accommodated in the new genus Oedohysterium. An additional new combination is proposed below.

 

Oedohysterium pulchrum (Checa, Shoemaker & Umaña) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515424.

Boehm E.W.A., Mugambi G., Miller AN, Huhndorf SM, Marincowitz S, Spatafora JW & Schoch CL. 2009b. A phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae & Gloniaceae, with keys to world species. Studies in Mycology 64: 59.   

Basionym: Hysterographium pulchrum Checa, Shoemaker & Umaña, Mycologia 99: 289 (2007). 

Notes: The newly described Hg. pulchrum from Costa Rica (Checa et al. 2007) also falls within Clade D (Boehm et al. 2009b) and is here transferred to Oedohysterium, as Od. pulchrum (DQ 402184 / DAOM 234345). This is because molecular data indicate a close association with the two species of Oedohysterium, Od. insidens and Od. sinense. At first surprising, on further consideration, this sub-clade forms a natural assemblage premised on morphological features. The spores of all three taxa show a remarkable degree of similarity in morphology, which includes being similarly pigmented, slightly curved and fusiform, with a common number of transverse septa. The sole difference is the presence of one or two vertical septa in Od. pulchrum, a feature noted by the authors to be absent in some spores (Checa et al. 2007). Most importantly, like Od. insidens and Od. sinense, Od. pulchrum also possesses a swollen supra-median cell. Interestingly, a striking resemblance to the phragmospores of Od. insidens can be seen for those spores of Od. pulchrum that do not possess vertical septa (Checa et al. 2007). This is based on similarities in shape (e.g., curved and fusiform), size [(20-)23-28(-38) x (5-)7-10(-13) μm versus 22-25(-27) x 5-6 μm], and in the number of transverse septa (4-)6-8(-11) versus (5-)6, for Od. insidens and Od. pulchrum, respectively. As molecular data indicate that the presence or absence of vertical septa should be considered a sympleisiomorphic character state within the Hysteriaceae (Boehm et al. 2009), we feel justified in including both phragmospores and dictyospores within the genus Oedohysterium.

 

Basionym: Hysterium mori Schwein., Trans. Amer. Philosoph. Soc. 4(2): 244 (1832).
        ≡ Hysterographium mori (Schwein.) Rehm, Ascomyceten no. 363 (1876).
= Hysterium grammodes De Not., Giorn. Bot. Ital. 2 (7-8): 55 (1847).
        ≡ Hysterographium grammodes (De Not.) Sacc., Syll. Fung. 2: 782 (1883).
= Hysterium rousselii De Not., Piren. Ister. 2(7-8): 19 (1847).
≡ Hysterographium rousselii (De Not.) Sacc., Syll. Fung. 2: 779 (1883).
= Hysterium vulgare De Not., Piren. Ister. 2(7-8): 18 (1847).
= Hysterium australe Duby, Mém. Soc. Phys. Hist. nat. Genève 16(1): 44 (1862).
= Hysterium lesquereuxii Duby, Mém. Soc. Phys. Hist. nat. Genève 16(1): 41 (1862).
≡ Hysterographium lesquereuxii (Duby) Sacc., Syll. Fung. 2: 779 (1883).
= Hysterium gerardi Cooke & Peck, Bull. Buffalo Soc. Nat. Sci. 3: 33 (1875).
        ≡ Hysterographium gerardi (Cooke & Peck) Sacc., Syll. Fung. 2: 783 (1883).
= Hysterium viticolum Cooke & Peck, Bull. Buffalo Soc. Nat. Sci. 3: 33 (1875).
        ≡ Hysterographium viticola (Cooke & Peck) Rehm, Ascomyc. No. 316, in Sacc., Syll. Fung. 2: 782 (1883).
= Hysterium variabile Cooke & Peck, Bull. Buffalo Soc. Nat. Sci. 3: 33 (1875).
        ≡ Hysterographium variabile (Cooke & Peck) Sacc., Syll. Fung. 2: 780 (1883).
= Hysterium formosum Cooke, in Harkness & Cooke Grevillea 7: 3 (1878).
≡ Hysterographium formosum (Cooke) Sacc., Syll. Fung. 2: 783          (1883).
= Hysterium putaminum Cooke Grevillea 7: 48 (1878).
≡ Hysterographium putaminum (Cooke) Sacc., Syll. Fung. 2: 783 (1883).
= Hysterographium portenum Speg., Anal. Soc. Cient. Argent. 9(4): 185 (1880).
= Hysterographium grammodes var. minus Sacc., Syll. Fung. 2: 783 (1883).
= Hysterographium pumilionis Rehm, Discom. 1(3): 21 (1887).
= Hysterographium guaraniticum Speg., Anal. Soc. cient. Argent. 26(1): 56 (1888).
= Hysterographium punctiforme Pat., Bull. Soc. Mycoll. France 4: 120 (1888).
= Hysterographium ruborum Cooke, in Rehm, Ascom., No. 918 (1888).
= Hysterium insulare P. Karst. & Har., Rev. Mycol. Toulouse No. 47: (1890).
= Hysterographium incisum Ellis & Everh., Bull. Torrey Bot. Club  24:  462 (1897).
= Hysterographium ziziphi Pat., Cat. Rais. Pl. Cell. Tunisie: 112 (1897) (as ´zizyphi´).
= Hysterographium rousselii var. piri Feltg., Vorst. Pilz. Luxemb. Nachtr. 3: 111 (1903).

Hysterothecia erumpent-superficial, ellipsoidal, oblong, linear or cylindrical, 1-2(-3.5) mm long, 220-275(-440) μm wide, by 190-330 μm high, mostly straight and lying parallel, but not confluent laterally, often gregarious and crowded so as to cover the substrate, longitudinally striate in age, navicular with tapering ends. Two types of hysterothecial aggregations regularly observed, depending on substrate: (1) Colonies on weathered, whitened decorticated hardwood often forming large oval colonies, with acuminate ends, measuring 5-15 cm in length, with hysterothecia gregarious in the center, densely packed in longitudinal formations, showing multiple stages of development, and darkening the adjacent substrate; when young, prior to emergence of hysterothecia, smaller colonies are seen, but still presenting darkened oval patches, often with coelomycetous anamorph present. (2) Colonies on bark (i.e., corticolous) less gregarious, not darkening the substrate, hysterothecia often situated at angles, rather than in parallel orientation. Peridium 30-60 μm thick medially, to 100+ μm at the base, distinctly three-layered in cross-section, the outer layer darkly pigmented, the middle less so, and the inner layer, thin-walled, pallid and compressed. Pseudoparaphyses cellular, septate, persistent, 1-2 μm wide, hyaline, thickened apically, branched and forming an epithecium in a gelatinous matrix above the ascal layer. Asci cylindrical to clavate, bitunicate, short-stipitate, (50-)80-110 x 10-18 μm. Ascospores pigmented, thin-walled dictyospores, obovoid, ends obtuse, 3-(5-7)-septate, with 1-2(-3) vertical septa usually associated with mid-cells, but on occasion also present obliquely in end cells, constricted at the median septum, sometimes, when fully hydrated, at additional, more distal septa, measuring (12-)14-22(-26) x (5-)7-10(-11) μm. Anamorph coelomycetous, Aposphaeria-like in nature, in culture conidiomata as irregular locules, with conidiogenous cells 8-10 x 1.5-2 μm; conidia (2-)2.5-3.5(-4) x 1-2 μm (Lohman 1932). Cosmopolitan, on aged, usually decorticated, weathered wood or bark of Pinus, Juniperus, Salix, Ostrya, Castanea, Quercus, Ulmus, Morus, Pyrus, Amelanchier, Crataegus, Rubus, Cercocarpus, Prunus, Gleditsia, various Fabaceae, Melia, Pistacia, Cotinus, Rhus, Acer, Ziziphus, Vitis, Fraxinus, Olea, and Aspidosperma (Zogg 1962).

 

Basionym: Gloniopsis constricta N. Amano, Trans. Mycol. Soc. Japan 24: 289 (1983).

Notes: Amano (1983) described a small-spored species of Gloniopsis from Japan, Gp. constricta, noting a prominent median septal constriction. The measurements of the dictyospores were given as 10.4-13.2 x 4.4-5.8 μm, usually with 3-4 transverse and one vertical septum that passes through one to three cells. Although not mentioned (Amano 1983), the illustrations depict a very thick wall and dictyospores highly symmetric in outline and septation. Amano (1983) stated of the spores "...hyaline, later becoming brown...", but did not mention the presence of a gelatinous sheath. He also noted that the closest resemblance is with Hb. smilacis (as Gp. curvata), the latter however with slightly larger spores.  In this study, we were fortunate to obtain a specimen from New Zealand (SMH 5211.1, deposited in F; Fig. 5A-E) that corresponds to the published description given by Amano (1983), but differs on several counts. Like Gp. constricta, the hyaline dictyospores in SMH 5211.1, are highly symmetric and thick-walled, (1-)3(-4)-septate, with 1(-2) vertical septa, but the constriction at the median septum in SMH 5211.1, while present, is not prominent. Also unlike Gp. constricta, the spores in SMH 5211.1 have an obvious gelatinous sheath when young, but this quickly dissipates with age, and may be completely absent in mature specimens. In SMH 5211.1, the spores measure (18-)20(-23) x 10-12 μm, which is considerably larger than those of Gp. constricta. Nevertheless, these differences, in our opinion, are not sufficient to warrant a new species, and we choose here to simply expand the spore measurements to (11-)13-20(-23) x 5-12 μm, rather than describe a new species, proposing instead the new combination Hb. constrictum. 

Basionym: Hysterium subrugosum Cooke & Ellis, Grevillea 5: 54 (1876).
                ≡ Hysterographium subrugosum (Cooke & Ellis) Sacc., Syll. Fung. 2: 780 (1883).
= Hsyterographium hiascens Rehm, Ber. Nat. Hist. Ver. Augsburg 26: 780 (1881).
= Hysterographium kansense Ellis & Everh., Erythea 2: 22 (1894).
= Hysterographium cylindrosporum Rehm, Bih. Kongl. Svenska Vetensk. Akad. Handl. 25(6): 11 (1899).
= Hysterographium minutum M.L. Lohman, Pap. Michigan Acad. 17: 267 (1933).

Hysterothecia erumpent to superficial, scattered to densely crowded, navicular, straight to flexuous, with tapered ends, surface not striated in age, but smooth to sub-rugose in texture, 1-2 mm long, 250-350 μm diam. Peridium composed of small pseudoparenchymatous cells, heavily pigmented at the surface, not showing a distinct number of layers, relatively smooth on outer surface. Pseudoparaphyses narrowly cellular, septate, 1-1.5 μm in diam., hyaline, branched above the asci, borne in a gelatinous matrix. Asci cylindrical to clavate, bitunicate, short-stipitate, 80-150 x 18-25 μm, with a prominent apical nasse, especially when young. Ascospores pigmented thin-walled, dictyospores (22-)25-34(-45) x (6-)8-12(-17) μm, mostly with 7-11 transverse and 1-2 vertical septa, hardly constricted at septa, clear brown, ends paler at times, slightly asymmetric in outline. Anamorph coelomycetous, Aposphaeria-like (Lohman 1933a). Less frequently collected, but reported from North America (Barr 1990b), Europe (Zogg 1962), Argentina (Messuti & Lorenzo 2003) and from South Africa (van der Linde 1992) as well. Old wood and bark of Populus, Quercus, Celtis, Crataegus, Rosa, and Cotinus (Zogg 1962), as well as on weathered fence posts and old planks (Boehm, unpubl. data).

Notes: In Boehm et al. 2009 b, we were able to include three geographically diverse isolates of Gp. praelonga (Table 1), two from South Africa (CBS 112415 and CMW 19983 / PREM 57539), and one from the United States, New Jersey (CBS 123337 / BPI 878725). These isolates cluster together in Clade D and associate with one isolate of Gp. subrugosa from South Africa (CBS 123346 / BPI 878735). Both Gp. praelonga and Gp. subrugosa are somewhat similar in the shape, size and septation of their dictyospores, hyaline in the former (Fig. 6B), pigmented in the latter (Fig. 6G). However, Gp. praelonga tend to have acuminate spore apices, whereas in Gp. subrugosa the apices are obtuse. The spores of Gp. praelonga are (16-)20-32(-34) x (6-)9-12(-15) μm, and those of Gp. subrugosa are (22-)25-34(-45) x (6-)8-12(-17) μm. Septation is also similar in both species, with 5-11 transverse and 1-3 vertical septa in Gp. praelonga and 7-11 transverse and 1-2 vertical septa in Gp. subrugosa. They differ in pigmentation and the presence of a gelatinous sheath in the type. Molecular data indicate that they are closely related.

Basionym: Hysterium andinense Messuti & Lorenzo, Mycological Research 101(3): 303. 1997.

Remarks: Messuti & Lorenzo (1997) described a new species of Hysterium from Patagonia, Argentina, collected from the bark of Austrocedrus chilensis (D. Don.) Florin & Boutlelje, designated as H. andinense Messuti & Lorenzo. However, molecular data presented here (Fig. 3) indicate that this taxon actually belongs to the genus Mytilinidion, in the Mytilinidiaceae, (Mytilinidiales) (Table 1). Although sections were not made, a re-examination of the original material, on which the single ascospore culture was based, showed fragile, thin-walled, globoid pseudothecia, with no prominent slit, enclosing a hamathecium of narrow trabeculate pseudoparaphyses, and three to seven (-nine) septate pigmented phragmospores, features congruent with the genus Mytilinidion. 

MycoBank No.: MB 512022 

Boehm EWA, Schoch CL, & Spatafora JW (2009). On the evolution of the Hysteriaceae and Mytilinidiaceae (Pleosporomycetidae, Dothideomycetes, Ascomycota) using four nuclear genes. Mycological Research 113: 469.  

Basionym: Glonium clavisporum Seaver, Mycologia 17(1): 4. 1925.

Remarks: Zogg (1962) synonymised Glonium simulans W.R. Gerard under G. lineare (Fr.) De Not. However, Lohman (1937) presented evidence as to why G. simulans (= P. simulans) should be considered distinct from G. lineare (= P. lineare). Although ascospores in both species are of similar length, the upper cell of the ascospore in P. lineare is broadly rounded, whereas the upper cell in P. simulans is more acuminate and approaches the appearance observed in the genus Glonium. The ascospores of both species differ markedly in outline as illustrated by Lohman (1932a). P. lineare possesses a pycnidial Plenodomus anamorph, but no anamorph has ever been recorded for G. simulans (Lohman 1937). Lastly, hysterothecia of P. lineare are confluent laterally to some degree, forming parallel, linear flat arrays, and are often partly submerged into the substrate. In contrast, hysterothecia of P. simulans are entirely superficial, never forming lines, but ofter assume flexuous configurations on the substrate, as in P. clavisporum. Here we remove the synonymy proposed by Zogg (1962) and reinstate G. simulans in the genus Psiloglonium. 

 

Notes: Zogg (1962) described this species as G. pusillum from Juniperus phoenicea and Pinus sylvestris from Southern France, noting that it was quite rare. Zogg (1962) stated that this species may or may not be associated with a subiculum, and hence was one of the factors behind his transfer of Petrak's (1923a, b) Psiloglonium species to Glonium. Psiloglonium pusillum has ascospores only slightly larger than those of P. abbreviatum, measuring (9-)10-12(-13) x 4-5(-6) μm. Lee & Crous (2003) also identified this fungus from Proteaceae and Restionaceae in South Africa, and Sivanesan & Hsieh (1989) reported it from Taiwan. It has also been found in North America (Boehm, unpubl. data).

 

Psiloglonium chambianum (Guyot) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515320.

Basionym: Glonium chambianum Guyot, Ann. Serv. Bot. Agric. Tunisie 28: 90 (1955).

Notes: Originally from North Africa, on Lonicera implexa (Caprifoliaceae), the fungus has since been reported from the Proteaceae in South Africa (Lee & Crous 2003). Zogg (1962) gave the spore measurements for G. chambianum as (14-)16-18(-21) x (6-)8-9(-10) μm, whereas Lee & Crous (2003) gave slightly larger measurements, (18-)20-21(-23) x (4-)5-6(-7) μm. Spores ellipsoid to oblong, with upper cell broader than the lower, and with an obovoid, obtuse apex. Psiloglonium chambianum possesses larger spores than P. lineare, P. simulans, and P. clavisporum, but smaller than P. uspallatense.

 

Psiloglonium uspallatense (Speg.) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515321.

Basionym: Glonium uspallatense Speg., Anal. Mus. Nac. Buenos Aires, Ser 3, v. 12, 19: 436 (1909).

Notes: Zogg (1962) listed the species a "doubtful", but Messuti & Lorenzo (2007) reinstated G. uspallatense after locating the original holotype material. They gave the spore measurements as 18-24 x 10-12 μm, placing it intermediate between P. chambianum and P. sasicola.

 

Psiloglonium sasicola (N. Amano) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515322.

Basionym: Glonium sasicola N. Amano, Trans. Mycol. Soc. Japan 24: 287 (1983).

Notes: Amano (1983) described this species from dead culms of Sasa sp. (Bambusaceae) in Japan. The ascospore measurements were given as 25-32 x 5-8 μm, with a rounded apical cell, placing it between P. uspallatense and P. ephedrae. Amano (1983) further reported that ascospores of this species are associated with a gelatinous sheath, previously not known among these didymospored fungi. 

 

Psiloglonium ephedrae (Henn.) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515323.

Basionym: Glonium ephedrae Henn., Öfvers. K. Vet. Akad. Förhandl. 2: 328 (1900).
                = Glonium costesi Speg., Bol., Acad. Nci. Ci., Córdoba 25: 78-79 (1921).

Notes: Messuti & Lorenzo (2007) reinstated G. ephedrae with the synonym G. costesi, after locating and comparing original type materials. Psiloglonium ephedrae possesses very large didymospores, measuring 26-35 x 8-15 μm, the upper cells broadly ovate. It has been collected from Ephedra andicola, and, as G. costesi, from Proustia pyrifolia in Chile.

 

Psiloglonium hysterinum (Rehm) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515324.

Basionym: Glonium hysterinum Rehm, Hedwigia 37: 298 (1898).

Notes: Rehm (1898) originally described a species of Glonium from Southern Brazil with large fusiform didymospores, prominently constricted at the septum, and with acuminate spore apices ("Enden zugespitzt"). The spore measurements were given as 45 x 9 μm.

 

Psiloglonium colihuae (Lorenzo & Messuti) E.W.A. Boehm & C.L. Schoch, comb. nov.

MycoBank MB515325.

Basionym: Glonium colihuae Lorenzo & Messuti, Mycol. Res. 102: 1104 (1998).

Notes: Lorenzo & Messuti (1998) described a new species on culms of Chusquea culeou from the Argentine Nothofagus rainforests. The spore measurements were given as 30-43 x 4-9.8 μm, and, although the spores are fusiform in outline, they possess moderately acuminate apices. In comparing this species to other acuminate-spored species of Glonium, the authors noted that the greatest degree of similarity was with the slightly smaller-spored G. caucasicum.

 

Psiloglonium araucanum (Speg.) E.W.A. Boehm, S. Marincowitz & C.L. Schoch, comb. nov.

MycoBank MB515326.

Basionym: Glonium araucanum Speg., Rev. Fac. Agron. Veter. La Plata 6: 110 (1910).
= Gloniella caucasica Rehm, Vestn. Tiflissk. Bot. Sada 25:12 (1912).
        ≡ Glonium caucasicum (Rehm) H. Zogg, Beitr. Kryptfl. Schweiz 11(3): 67 (1962).

Notes: Messuti & Lorenzo (2007) transferred Glonium caucasicum to G. araucanum, after examining the types for both species. Previously, Zogg (1962) had transferred Gloniella caucasica to Glonium. Here we transfer G. araucanum to Psiloglonium. This taxon possesses fusiform spores with highly acuminate apices. Messuti & Lorenzo gave the spore measurements as 22-28 x 8-10 μm, whereas Zogg (1962) gives them as (19-)22-25(-27) x (6-)7-9(-10) μm. Although originally European in distribution (Zogg 1962), the taxon has subsequently been collected from South (Messuti & Lorenzo 2007) and North America (Boehm unpubl. data), and from South Africa (Lee & Crous 2003).

Lee & Crous (2003) identified a series of isolates from South Africa on the Restionaceae as Glonium compactum (CBS 112412, CMW 18760, CMW 17941). However, in their study they did not note the presence of subicula, nor a stromal crust. These features were stressed for this taxon by Zogg (1962). These same isolates were used in Boehm et al. (2009), and were shown to associate, with high branch support, with two species of Psiloglonium, P. clavisporum and P. simulans, distant from the other species of Glonium surveyed (e.g., G. stellatum and G. circumserpens). Thus, a new combination was proposed, Psiloglonium compactum. However, it is now realised that this new combination was made in error and is hereby retracted. It must be concluded that the South African isolates (Lee & Crous 2003) were not G. compactum, due to the absence of subicula and stroma, but rather, we suspect, the cosmopolitan P. araucanum, which has similar, but slightly smaller, fusiform acuminate didymospores. Lee & Crous (2003) give the ascospore measurements for the South African "G. compactum" as (24-)26-27(-30) x (4-)5-6(-7) μm, which matches closely those given above for P. araucanum. Furthermore, the illustrations in Lee & Crous (2003) closely match P. araucanum, and not those of G. compactum, as given by Zogg (1962). If we are correct in assuming that the South African isolates used in Boehm et al. (2009) are in fact P. araucanum (Type II) and not G. compactum (Type III), then this would provide a high degree of support for the inclusion of species with acuminate spore apices, belonging to Type II, in the genus Psiloglonium, along with species with obtuse spore apices, belonging to Type I (e.g., P. simulans and P. clavisporum). A reanalysis of the original South African herbarium specimens from which the sequences were derived (PREM 57570, PREM 57569, PREM 57566), by S. Marincowitz, has confirmed that they do indeed correspond to P. araucanum and not to G. compactum. Molecular data thus supports the association of Types I and II within the genus Psiloglonium.