Glyphium

The genus Glyphium Nitschke ex Lehmann
Nova Acta Caes. Leop. -Carol. German. Nat. Cur. 50: 139 (1886)

by Eric W.A. Boehm

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Glyphium elatum (Grev.) Zogg. Many thanks to Guy Marson of Luxembourg who sent me these stereophotos and granted me permission to use them here. These wonderful “bouquets” of erect ligulate to dolabrate pseudothecia were collected from Malus (apple) [top] & from Salix caprea [center & bottom] near Luxembourg City and Hesperange. Apparently common. Thanks Guy!

Lophium Fries (pro parte)

Fungi classified in the genus Glyphium Nitschke ex Lehmann posses erect, strongly laterally compressed, transversely striate, carbonaceous pseudothecia, that are ligulate (strap- like) to dolabrate (hatchet-shaped) in outline (Zogg 1962). Ascomal dehiscence is characterized by a longitudinal slit borne upon a crest-like apex, parallel to the compressed axis of the fruitbody. Although ascoma are superficial, at times the base is embedded in the substrate and may be associated with subtending black interwoven hyphal strands that anchor the base to a subicular matrix. Hamathecial elements consist of trabeculate pseudoparaphyses, borne in a gel-matrix, that are thin to lacking at maturity (Barr 1990). The filiform ascospores are multiseptate and light brown at maturity, occupying the greater length of the ascus, and are often spirally arranged, at least for part of their length. In some species, the ascospores disarticulate into part-spores, fragmenting before maturity while still within the ascus (Goree 1974; Sutton 1970). A dematiaceous hyphomycetous anamorph has been described for these species, belonging to the form genus Peyronelia, and is characterized by verruculose conidia borne in chains, transversely septate with few longitudinal septa (Lohman 1933a; Sutton 1970). Species of Glyphium are lignicolous or corticolous, and have been recovered from a wide range of angiosperms, including, most commonly, Salix, Populus, Rhus, Alnus, Rhamnus, Fagus, Pyrus and Malus. One species from Florida has been recovered from Tillandsia.

Currently the genus consists of five species, separated primarily by whether the ascospores disassociate into part-spores. Ascospores not disintegrating into part-spores include the cosmopolitan Glyphium elatum(Grev.) Zogg, the type species, which occurs in both North (Lohman 1933a) and South (Lorenzo & Messuti 2005) America, Europe (Bisby & Ellis 1952; Zogg 1962), the Russian Far East (Vasilyeva 2001), China (Teng 1936) and Taiwan (Chen & Hsieh 1996),  and the diminutive G. tillandsiae (Cash) Zogg, known only from the type locality of Highlands Hammock, near Sebring, Florida, which occurs on Tillandsia fasciculata(Cash 1943). The three species with ascospores disintegrating into part-spores, with a Peyronelia anamorph, include: the newly described Glyphium grisonense G. Mathiassen from Norway (Mathiassen 1993), G. corrugatum (Ellis) Goree, which has been collected primarily in the Western United States and Canada (Goree 1974; Sutton 1970), and G. schizosporum (Maire) Zogg, which has been reported from Algeria, France, Switzerland and Norway (Mathiassen 1989; Sutton 1970; Zogg 1962).

The genus Glyphium was originally placed by Zogg (1962) in the Lophiaceae Zogg ex von Arx & Muller (von Arx & Muller 1975), later consigned to the earlier name Mytilinidiaceae (Barr 1990). However, current classification of the Ascomycota (Eriksson 2006; Lumbsch & Huhndorf 2007) excludes the genus Glyphium from the Mytilinidiaceae. This is because recent molecular evidence, using the nuclear small (nuSSU) and large (nuLSU) ribosomal subunits, as well as the mitochondrial small (mtSSU) and large (mtLSU) ribosomal subunits (Lindemuth et al. 2001; Lumbsch et al. 2005) has removed the genus Glyphium to the Chaetothyriales in the Eurotiomycetes. The unusual placement of the genus Glyphium in the Chaetothyriales of the Eurotiomycetes (Lindemuth et al. 2001; Lumbsch et al. 2005) has been confirmed and restated in a number of subsequent publications (Geiser et al. 2006; Kodsueb et al. 2006; Lucking et al. 2004; Schmitt et al. 2005), including the Assembling the Fungal Tree of Life (AFTOL) Project (Lutzoni et al. 2004).

All of these studies, however, were based on sequences derived from a single isolate (CBS 268.34; AFTOL #1145) labeled as Glyphium elatum, recovered from Salix in Colorado by M.L. Lohman and deposited in 1934. Recently, environmental studies of DNA isolated from the biodeterioration of cathedral marble in Italy have detected G. elatum, based on BLAST hits to the GenBank accessions of the CBS 268.34 isolate (Cappitelli et al. 2007). Similarly, molecular detection of fungal signature sequences in atherosclerotic plaques has also identified G. elatum, again based on the CBS 268.34 isolate (Ott et al. 2007). These environmental studies seem at odds with the lignicolous or corticolous substrate preferences known to exist within the genus Glyphium, and further underscore the need to verify the provenance of CBS 268.34.

A study currently underway has secured fresh European material, from which a number of genes have been sequenced. The intent is to determine whether indeed the genus Glyphium belongs to the Chaetothyriales in the Eurotiomycetes, as attested to by CBS 268.34, or whether the genus actually lies within the Dothideomycetes. Essentially, we were interested in whether morphological features historically used to classify the genus Glyphium within the Mytilinidiaceae were phylogenetically informative in the context of sequence-based phylogenies.

Barr (1990): “Ascomata superficial with bases at times embedded in substrate, separate or gregarious, erect ligulate or dolabrate, usually laterally compressed, medium to large sized; apex compressed, opening by long slit; surface black, often transversely striate, with dependent hyphae near base anchoring to substrate & mingling with subiculum; peridium carbonaceous, brittle, narrow, composed of small cells. Asci basal, cylindric. Trabeculae sparse in gel matrix. Ascospores hyaline to light brown, filiform, end obtuse, multiseptate, at times separating into several celled part-spores; wall thin, smooth; granular; in a fascicle & often spirally wound in the ascus. Anamorphs hyphomycetous (Peyronelia chains of brown, transversely & sometimes longitudinally septate, verruculose conidia borne from short brown conidiophores, associated with ascomata in nature, developing from conidia in culture. On branches & wood of angiosperms. Type species: Glyphium dolabriformis (Wallroth) Lehmann = Glyphium elatum (Greville) Zogg. Species of Glyphium had been included under Lophium by most authors until Zogg (1962) pointed out the reasons for separating two genera. Goree (1974) recognized two species in western North America: Glyphium elatum & Glyphium corrugatum. In Europe, Glyphium schizosporum (Maire) Zogg has ascospores that separate into part-spores within the ascus, as do those of Glyphium corrugatum. Sutton (1970) separated the two species on the basis of part-spore size & septation as well as differences in conidia. In addition, Glyphium tillandsiae (Cash) Zogg was described from Tillandsia in Florida (Cash, 1943). It is a smaller fungus than Glyphium elatum, but like that species has ascospores that do not separate into part spores in the ascus. Eriksson (1981) described & illustrated the habit & peridium of Glyphium elatum. The outermost layer of peridium breaks vertically along the narrow sides into two transversely furrowed plates & the incompletely covered inner peridium near the base gives rise to tufts of hyphae that grow downward to the substrate. Lohman (1933) obtained only sterile cultures bearing sclerotial bodies on the surface of the medium; his cultures were tissues cultures because ascospores of his material did not germinate. Sutton (1970) also found that ascospores would not germinate. He cultured conidia that were borne on hyphae of the subiculum, morphologically connected to the ascomata. These in pure culture grew & produced similar conidia. Sutton described the conidia of Glyphium corrugatum (as G. leptotrhecium) with up to 55 transverse septa & usually one or two longitudinal septa, up to 230 x 10 – 17 μm, & of Glyphium schizosporum with up to 50 transverse septa & an occasional longitudinal septum, up to 220 x 8 – 11 μm. These details aid in separating the two species.”

 

 

 

Key to species of Glyphium Nitschke ex Lehman

  1. Ascospores not disintegrating into part-spores in the ascus → 2

1′. Ascospores disintegrating into part-spores within the ascus  → 3

  1. Ascocarps scattered to sometimes aggregated, erect, ligulate to sometimes dolabrate, sessile, unbranched, superficial or sometimes with the base immersed, 0.25 – 0.60 (length) x 0.1 – 0.25 (width) x 0.65 – 2.0 (height) mm. Ascocarp wall black (inner layers only slightly colored), carbonaceous, prosenchymatous, the surface distinctly transversely striate (striations parallel to the ostiole). Ostiole a narrow fissure with the edges connivent. Asci bitunicate, filiform, 250 – 718 x 7 – 10 μm, mostly long-stipitate, eight-spored. Pseudoparaphyses filiform, hyaline, septate, branched, anastomosed, 0.5 – 1.5 μm wide. Ascospores filiform, tapering toward the obruse ends, pale yellowish or olive-tan to light brown, many-septate, septa (3-) 5 – 8 (-12) μm apart, constricted or unconstricted at the septa, 232 – 416 (& probably up to 650) x (1-) 1.5 – 2 (-3) μm, fasiculate, usually with regular or irregular spiraling for part of their length, & remaining intact within the ascus. Imperfect states unknown. Cosmopolitan (Goree 1974; Zogg 1962) → Glyphium elatum(Grev.) Zogg

2′. Fruitbody smaller. Ascomata superficial, sparse, flabelliform, laterally compressed & narrowed toward the base, fuscous to fuscous black, slightly paler above, 0.3 – 0.5 mm high, 0.2 – 0.3 mm wide, 0.1 mm thick, opening by a slit along the top, minutely roughened & sometimes faintly transversely striate; asci terete, the wall thickened at the apex, short pedicillate, 8-spored, 250 – 300 x 10 – 12 um; ascospores nearly the length of the ascus, 2.5 – 3 um thick, hyaline or subhyaline, parallel, multiseptate, spore sections oblong, 1-guttulate, 4 – 5 um long; paraphyses filiform, irregularly branched, pale brownish. On Tillandsia fasciculata from Southern Florida. No known anamorph (Cash 1943) → Glyphium tillandsiae (Cash) Zogg

 

  1. Ascocarps scattered to aggregated, erect, ligulate to sometimes dolabrate, sessile, unbranched, superficial or sometimes with the base immersed, 0.25 – 0.75 (length) x 0.1 – 0.25 (width) x 0.85 – 1.75 (height) mm. Ascocarp wall black (inner layers only slightly colored), carbonaceous, prosenchymatous, the surface distinctly transversely striate (striations parallel to the ostiole). The outer layers of the two lateral halves of the ascocarp wall sometimes bowed out & split away from the rest of the ascocarp except at the apex & base. Ostiole a narrow fissure with the edges connivent. Ascospore part-spores or “fragments” consistently 3-septate or less, slightly narrower than the next two species. Ascospores filiform, tapering toward the obtuse ends, many septate, fasciculate & usually with regular or irregular spiraling for part of their length, fragmenting in the ascus; spore fragments cylindrical with the ends truncate to more commonly obtuse, (8-) 14 – 20 (-37) x (1.5-) 2 (-3) μm, pale yellowish or olive tan to light brown, (one-) to three- (occasionally up to seven-) septate; septa (3-) 4 – 5 (-8) μm apart, usually constricted at the septa of wider fragments, but usually not constricted at the septa of narrower fragments. Peyronelia conidial state with up to 55 transverse septa & usually one or two longitudinal septa, up to 230 x 10 – 17 μm, wider and prone producing lateral buds or stubs; North American: Western Canada & the United States (Goree 1974) → Glyphium corrugatum(Ell.) Goree

3′. Ascospore part-spores or “fragments” usually with more than 3 septa per fragment. European in distribution → 4

 

  1. Subiculum dense, or very sparsely developed, composed of branched, thick-walled, septate brown hyphae ca. 3.5 um diam., formed from the base or the lower half of pseudothecium. Pseudothecia 0.7 – 1.5(2) mm high, seated upright in subiculum, scattered or clustered, laterally flattened, dolabriform, but often +/- obpyriform in outline with age, black,, with a 200 – 750 um long slit along the sharp upper edge. Peridium 80 – 115 um thick, tow layered, outer layer 30 – 55 8m thick, dense, carbonized and brittle, early bursting vertically along the narrow sides of pseudothecium into two laterally transversely striate plates. Ascospores 270 – 420 um long, filiform, multiseptate, slightly spiralling, fragmenting before maturity into cylindrical, pale brown (2-) 3 – 7(-11) -septate units, measuring (13.8-) 15.1 – 42.8 (-48) x 2.9 – 5.5 (-6) um, mean 24.2 x 4.3 um.  Anamorph seen as effuse, black colonies on naked wood, in close connection with pseudothecia and subiculum. Conidial chains unbranched thick-walled, multiseptate, irregularly constricted, verrucose and dark brown. Maturing acropetally. The ascomata of G. schizosporumare narrow, and never develop to become obpyriform in outline, the asci are very long, but only 7 – 12 um wide, the spores are very narrow, and the conidiophores slightly different from those of G. grisonense. Only known from North Norway and from one high altitude locality in Switzerland (Mathiassen 1993) → Glyphium grisonense G. Mathiassen

4′. Pseudothecia 0.1 – 0.15 x 0.025 – 0.05 cm, superficial, scattered, gregarious or crowded, arising amongst the subicle, conidiophores, or from bare wood, erect or slanting, dolabriform and flattened laterally, narrower towards the base, black, with several transverse parallel ridges and subicular hyphae extending down towards the substrate from the vertical edges, lenticular in longitudinal sections taken at 90 degrees to the long axis, unilocular, lateral walls composed of outer sclerotioid black prosenchyma becoming thin-walled, pale brown pseudoprenchyma. Ascospores parallel in the ascus, fragmenting before maturity into cylindrical, pale olive to pale brown, 1 – 9 septate units (mean 5-septate) measuring 14 – 26 (9 – 36) x 3 – 4 (mean 18 x 3.5) μm. Peyronelia conidial state with up to 50 transverse septa & an occasional longitudinal septum, up to 220 x 8 – 11 μm, not prone to producing buds, but simply straight chains; European: known from a few high altitude localities in Switzerland, France and Algeria (Goree 1974; Sutton 1970; Zogg 1962) → Glyphium schizosporum (Maire) Zogg

 

 

Glyphium elatum (Grev.) Zogg

Zogg, 1962. Beiträge zur Kryptogamenflora der Schweiz, Band 11(3): 99.

 

Lophium elatum Greville, 1825

Lophium dolabriforme Wallroth, 1833

Glyphium dolabriforme (Wallr.) Nitschke ex Lehmann, 1886

Lophium leptotrhecium Tracy et Earle, 1901

Acrospermum fultum Harkness, 1884

Zogg, 1962. Rarely collected. European alpine belt & Denmark. On old weathered wood of Salix, Alnus, Fagus & Pyrus.

The following is unpublished amaterial nd under copyright protection by Eric W.A. Boehm, as it presently forms the basis for a manuscript under preparation. It is included here, “pre-press” for general release, but its use is highly restricted.

Eric Boehm (2009 unpublished): “Glyphium elatum (Grev.) Zogg – In his Scottish Cryptogamic Flora, Vol. III (1825), R.K. Greville published an account of an unusual lignicolous fungus collected from Appin, Scotland, stating: “Perithecia, in my specimen, not placed upon a crust; black, erect, about a line high, stipitate; the stalk nearly cylindrical at the base, very gradually dilating and passing into the compressed, transversely striated, black, wedge-shaped perithecium: the whole very similar to the head of a long-shaped hatchet”. Greville (1825) established Lophium elatum Grev., on account of the erect, dolabrate ascomata, to which he gave the common name of “Elongated Lophium“. Greville (1825) was the first to illustrate the filiform nature of the multiseptate ascospores. Greville also sent material to Fries, who validated the binomial in his Elenchus Fungorum (1828), and thus the type has subsequently been designated as Lophium elatum Grev. : Fr. Early authors also reported specimens from Sedan and Briançon, France (Desmazières 1842; Montagne 1834). Meanwhile, Wallroth (1833) described Lophium dolabriforme Wallr. from the bark of Pyrus and Alnus in Germany, but noted that his specimens may have been conspecific with Greville’s, stating: “Lophium elatum Grev. haud multum differre videtur“. Duby (1862) examined and illustrated Wallroth’s material, but gave no spore measurements. It remained for Rehm, in his Revision der Hysterineen im herb. Duby (1886) to reexamine Wallroths’ material and give the spore measurements as 300 x 2 μm. Later, in L. Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz, Rehm (1896) revised the measurements to 150 – 320 x 2 – 3 μm. Rehm (1896) considered L. elatum Grev. a doubtful species, maintaining instead L. dolabriforme for the dolabrate forms of Lophium. This situation was maintained by subsequent authors (e.g., Fuckel 1873; Lindau 1922), while others (e.g., Massee 1895) maintained L. elatum, and did not list L. dolabriforme. Massee (1895) found L. elatum at Herb. Berk., Kew, in a package simply labeled “Scotland” in Greville’s handwriting, and, presuming that this material represented the type, gave the spore measurements as 180 – 200 x 1.5 μm. Lehman (1886) studied Fuckel’s German material of L. dolabriforme, collected from Pyrus communis and Prunus spinosa (Fuckel 1873), and concluded that the ascomata were sufficiently different from those of Lophium mytilinum Pers. : Fr., that he proposed the new genus Glyphium Nitschke, ascribed as such because Nitschke had first used it as a herbarium name (Sutton 1970). Thus, Glyphium Nitschke ex Lehmann was establihed to accommodate L. dolabriforme, as G. dolabriforme (Wallr.) Lehmann. The species L. elatum Grev. : Fr. was retained within the genus Lophium (Lehman 1886). Bisby & Ellis (1952) collected material in Guernsey, the Channel Islands, from Lonicera periclymenum, and determined that the spores measured 150 – 330 x 2 – 3 μm, agreeing well with Rehm’s (1896) earlier measurements for L. dolabriforme. Most remarkably, however, Bisby & Ellis (1952) were also able to relocate Greville’s original type material at Kew, and determined that, unlike the measurements given by Massee (1895), many of the spores were actually greater than 300 μm in length. Therefore, Bisby & Ellis (1952) concluded that the two names were conspecific, with L. elatum Grev. : Fr. 1828 anteceding L. dolabriforme Wallr. 1833. Although they preferred to retain L. elatum within a broad interpretation of the genus Lophium, they did note that the name Glyphium, provided by Lehmann (1886), was available. Following Bisby and Ellis (1952), Zogg (1962) transferred three species from Lophium to the genus Glyphium, noting that the dolabriform nature of the ascocarp differed sufficiently from the conchate ascocarp found in Lophium. The first of these transfers involved the type species. Although Greville’s specimen at Kew was not examined, Zogg (1962) did examine Fuckel’s (1873) material, upon which Lehman (1886) erected the genus Glyphium, and thereby transferred L. elatum Grev. : Fr., to G. elatum (Grev.) Zogg, listing also L. dolabriforme Wallr. 1833, and G. dolabriforme (Wallr.) Nitschke ex Lehmann 1886 as synonyms”. © Eric W.A. Boehm 2009

Goree, H. (1974. Can. J. Bot. 52: 1265): “Ascocarps scattered to sometimes aggregated, erect, ligulate to sometimes dolabrate, sessile, unbranched, superficial or sometimes with the base immersed, 0.25 – 0.60 (length) x 0.1 – 0.25 (width) x 0.65 – 2.0 (height) mm. Ascocarp wall black (inner layers only slightly colored), carbonaceous, prosenchymatous, the surface distinctly transversely striate (striations parallel to the ostiole). Ostiole a narrow fissure with the edges connivent. Asci bitunicate, filiform, 250 – 718 x 7 – 10 μm, mostly long-stipitate, eight-spored. Pseudoparaphyses filiform, hyaline, septate, branched, anastomosed, 0.5 – 1.5 μm wide. Ascospores filiform, tapering toward the obruse ends, pale yellowish or olive-tan to light brown, many-septate, septa (3-) 5 – 8 (-12) μm apart, constricted or unconstricted at the septa, 232 – 416 (& probably up to 650) x (1-) 1.5 – 2 (-3) μm, fasiculate, usually with regular or irregular spiraling for part of their length, & remaining intact within the ascus. Imperfect states unknown. On bark & decorticated wood (& occasionally leaves) of Angiospermae. Glyphium elatum is known from Denmark, France, Germany, Switzerland, & the United Kingdom (Bisby & Ellis 1952; Zogg 1962) & in North America from western Canada & the United States. It has been collected rather infrequently; nevertheless in the USA & Canada it appears to be primarily a coastal species. It is found also in WA in the lowlands surrounding the Olympic Mnts., & extends eastward in to the Cascade Mnts. in the valleys & passes. It extends eastward along the USA-Canadian boarder into N. Idaho, perhaps because of the oceanic influence of the westerlies. Asci & ascospores of American materials on average longer than those of European collections. Still greater in length of asci is Lophium caulicola Teng (Sinensia 7: 492, 1936) from China containing dolabrate hysterothecia with non-fragmenting, filiform, multiseptate ascospores nearly as long as the asci (700 – 800 μm) which may be a species of Glyphium distinct from G. elatum.”

Barr (1990): “Ascomata 650 μm to 2 mm in height, 250 – 600 μm wide, compressed & 100 – 250 μm in side view. Asci 200 – 700 x 6 – 10 μm. Ascospores 200 – 416 (-650) x (1-) 1.5 – 2 (-3) μm, light yellowish brown, multiseptate, not separating into part-spores in the ascus. Anamorph not known. Note: Goree (1974) examined & cited collections from British Columbia, Idaho, Washington & California. He concluded that this species was primarily coastal in western North America. In Europe it is know from Britain, Denmark, France, Germany & Switzerland. On wood, periderm & occasionally leaves of angiosperms, north temperate zone. Collected by Barr in CA from Umbellularia californica.”

 

Glyphium tillandsiae (Cash) Zogg

Zogg, 1962. Beiträge zur Kryptogamenflora der Schweiz, Band 11(3): 103.

Lophium tillandsiae Cash, 1943

Rare. Collected in Florida by E.K. Cash (1943. Mycologia 35: 595-96) on Tillandsia.

The following is unpublished material and under strict copyright protection by Eric W.A. Boehm, as it presently forms the basis for a manuscript under preparation. It is included here, “pre-press” for general release, but its use is highly restricted.

  1. tillandsiae(Cash) Zogg– “Zogg (1962) also transferred Lophium tillandsiae Cash to the genus Glyphium, as G. tillandsiae (Cash) Zogg. This diminutive species of Glyphium was collected by C.L. Shear from Tillandsia fasciculata in Highlands Hammock State Park, west of Sebring, Florida and described by Cash (1943). Although ascoma measure only 0.3 – 0.5 mm in height, they contain ascospores of somewhat comparable size to other species of Glyphium, measuring nearly the length of the ascus, given as 250 – 300 μm in length (Cash 1943)”. © Eric W.A. Boehm 2009

Cash (1943): “Ascomata superficial, sparse, flabelliform, laterally compressed & narrowed toward the base, fuscous to fuscous black, slightly paler above, 0.3 – 0.5 mm high, 0.2 – 0.3 mm wide, 0.1 mm thick, opening by a slit along the top, minutely roughened & sometimes faintly transversely striate; asci terete, the wall thickened at the apex, short pedicillate, 8-spored, 250 – 300 x 10 – 12 um; ascospores nearly the length of the ascus, 2.5 – 3 um thick, hyaline or subhyaline, parallel, multiseptate, spore sections oblong, 1-guttulate, 4 – 5 um long; paraphyses filiform, irregularly branched, pale brownish. On Tillandsia fasciculata, Mar. 3-4, 1941, C.L. Shear 1386 & 1387 & on Tillandsia sp., Mar. 1, 1937, 725 (type), all from Highlands Hammock, near Sebring. Lophium tillandsiae differs in shape from other species of Lophiumhaving apothecia less than 1 mm in height, also from L. schizosporum Maire in the absence of a black subiculum, & in shorter asci. Few fungi have been found reported on Tillandsia.”

 

Glyphium corrugatum (Ell.) Goree
Goree, 1974. Can. J. Bot. 52: 1265.

Acrospermum corrugatum Ellis, 1881
Lophium leptothecium Earle, 1901
Glyphium leptothecium (Earle in Greene) Sutton
Lophium apoclastosporum Solh., 1949

The following is unpublished material and under strict copyright protection by Eric W.A. Boehm, as it presently forms the basis for a manuscript under preparation. It is included here, “pre-press” for general release, but its use is highly restricted.

  1. corrugatum(Ellis) Goree – “Zogg (1962) also placed as a synonym under G. elatum, the binomial Lophium leptotheciumEarle, with a question mark, based on a description by Saccardo (1902). L. leptothecium was originally described by Earle (1901) from Colorado. Sutton (1970) examined Earle’s material, designating one a lectotype, and compared this to specimens recently collected in Manitoba and Saskatchewan, from Populus tremuloides. Sutton (1970) concluded that they matched Earle’s lectotype and noted that the spores differed from those of G. elatum, in that they disarticulated within the ascus, prior to discharge, forming 0 – 3 septate spore-fragments, measuring 12 – 24 (8 – 43) x 2 – 3 μm. Sutton (1970) therefore removed L. leptothecium from synonymy with G. elatum, and made the transfer to G. leptothecium (Earle) Sutton. Also associated with this fungus was an anamorphic state assigned to the form genus Peyronelia; whereas no anamorph is known to be associated with G. elatum. The Peyroneliaanamorph of G. leptothecium possesses verruculose chains of reddish-brown conidia, which mature acropetally, with up to 55 transverse septa, usually one or two longitudinal septa, and are prone to producing lateral buds or side branches (Sutton 1970). Earlier, Lohman (1933a) had collected a similar fungus from Salix in Colorado which he described as L. dolabriforme Wallr., but noted that the spores disarticulated within the ascus into one- to three-septate part-spores, measuring 12 x 2 μm. Sutton (1970) examined Lohman’s specimens of L. dolabriforme at MICH (Lohman No. 196), and concluded that they in fact corresponded to G. leptothecium, and not to G. elatum. Both Lohman (1933a) and Sutton (1970) were not able to induce ascospore germination for G. leptothecium. Lohman (1933a) however did manage to establish a culture from No. 196 based on inoculum derived from rhizoidal strands subtending the base of the fruitbody. Sutton (1970) noted that although isolated conidia of G. leptothecium germinated, growth ceased almost immediately, but he was able to conclusively demonstrate a physical linkage between the Peyronelia anamorph and the subtending rhizoidal strands. It should be noted at this point, that the culture labeled Glyphium elatum CBS 268.34 (AFTOL #1145), upon which the transfer of the genus Glyphium to the Chaetothyriales in the Eurotiomycetes was based (Lindemuth et al. 2001; Lumbsch et al. 2005), and restated in a number of subsequent publications (Geiser et al. 2006; Kodsueb et al. 2006; Lucking et al. 2004; Schmitt et al. 2005), including the AFTOL project (Lutzoni et al. 2004), does not actually represent G. elatum. Rather, CBS 268.34, although labeled as No. 196, L. dolabriforme, (Lohman 1933a) was in point of fact later determined by Sutton (1970) to be a species of G. leptothecium. Sutton (1970) based his identification on the fact that spores within the ascus disarticulated into 9 – 21 (mean 14) x 2 μm fragments with 1 – 3 septa, and a Peyronelia anamorph was present. The misidentification of No. 196 by Lohman (1933a) as L. dolabriforme, is surprising, given that spores were noted to disarticulate in the ascus, and given the presence of an anamorph, features known not to be associated with the type, G. elatum (Sutton 1970). Furthermore, the culture was not initiated from a single ascospore, but rather, represents a vegetative culture derived from rhizoidal hyphae subtending the base of the fruitbody (Lohman 1933a). This confusion as to the provenance of CBS 268.34 thus, forms a central component to our paper, and will be revisited later. The story of G. leptothecium however does not end here. Shortly after Sutton (1970), Goree (1974) transferred G. leptothecium (Earle) Sutton to G. corrugatum (Ellis) Goree. This was based on the precedence of the name Acrospermum corrugatum Ellis 1881, over the name L. leptothecium Earle 1901. Ellis (1881) originally collected A. corrugatum from unidentified lignum in Utah, and was the first to note the disarticulating ascospores in the ascus. Meanwhile, Harkness (1884) collected a similar fungus from Eucalyptus in San Francisco, California, naming it A. fultum Harkness, noting that his material differed from Ellis’ in that the spores did not disarticulate in the ascus. Later, Ellis and Everhardt (1887), as well as Harkness (1885), considered A. fultum as conspecific with A. corrugatum. It was Harkness’ California material that was distributed by Ellis in his North American Fungi, Series II, No. 2055, as A. corrugatum (Goree 1974). Goree examined this material and determined it to possess non-fragmenting ascospores, thus corresponding to G. elatum. Thus, what was distributed was actually G. elatum, at that time understood as either L. elatum or L. dolabriforme, but labeled as A. corrugatum. Upon examination of the holotype for A. corrugatum in the Ellis Collection at NY (listed as SJ Harkness, No. 73), but representing material originally collected by Ellis in Utah, Goree (1974) noted fragmenting ascospores, corresponding to G. corrugatum, and agreeing with an examination of more than 80 specimens from the Western United States and Canada. Goree (1974) also included under G. corrugatum, as a synonym, Lophionema apoclastosporum Solh., a fungus collected by Solheim (1949) from Wyoming. As an aside, much earlier, Riddle (1920) had also examined the same material of A. corrugatum as examined by Goree, that is, that issued in Ellis and Everhart’s North American Fungi, Series II, No. 2055, and concluded that the fungus did not belong to the genus Acrospermum, based on a number of features associated with the fruitbody. Riddle (1920), however, designated the material as L. dolabriforme, and was not cited by Goree (1974). Apparently G. corrugatum is a very common species, distributed throughout the western United States and Canada”. © Eric W.A. Boehm 2009

Goree (1974): “Ascocarps scattered to aggregated, erect, ligulate to sometimes dolabrate, sessile, unbranched, superficial or sometimes with the base immersed, 0.25 – 0.75 (length) x 0.1 – 0.25 (width) x 0.85 – 1.75 (height) mm. Ascocarp wall black (inner layers only slightly colored), carbonaceous, prosenchymatous, the surface distinctly transversely striate (striations parallel to the ostiole). The outer layers of the two lateral halves of the ascocarp wall sometimes bowed out & split away from the rest of the ascocarp except at the apex & base. Ostiole a narrow fissure with the edges connivent. Asci bitunicate, filiform, (310-) 458 – 553 (-701) x (7-) 8 – 10 (-12) μm, short- to long-stipitate, eight-spored. Pseudoparaphyses filiform, hyaline, septate, branched, anastomosed, 0.5 – 1.5 (-2) μm wide. Ascospores filiform, tapering toward the obtuse ends, many septate, fasciculate & usually with regular or irregular spiraling for part of their length, fragmenting in the ascus; spore fragments cylindrical with the ends truncate to more commonly obtuse, (8-) 14 – 20 (-37) x (1.5-) 2 (-3) μm, pale yellowish or olive tan to light brown, (one-) to three- (occasionally up to seven-) septate; septa (3-) 4 – 5 (-8) μm apart, usually constricted at the septa of wider framents, but usually not constricted at the septa of narrower framents. Imperfact state a Peyronelia (Sutton, 1970). Subiculum sometimes present. On bark & decorticated wood of Angiospermae. More than 80 specimens were examined. Glyphium corrugatum is known only from western Canada & the United States. It is a wide-spread inland species coming nearest to the Pacific Coast in the chaparral of southern CA. It is perhaps the commonest hysteriaceous fungus in the West, & should be sought south of the United States.”

Sutton (1970), as Glyphium leptothecium (Earle in Greene) Sutton: “Pseudothecia 0.1-0.15 x 0.025-0.05 cm, superficial, scattered, gregarious or crowded, arising amongst the subicle, conidiophores, or from the bare wood, erect or slanting, dolabriform and flattened laterally, narrower towards the base, black, with several transverse parallel ridges, lenticular in longitudinal sections taken at 90 degrees to the long axis, unilocular, lateral walls composed of outer sclerotioid black prosenchyma becoming thin-walled and hyaline towards the locule, basal region of isodiametric, thin-walled, pale brown pseudoparenchyma. Asci125 – 300 x 10 – 11 μm, bitunicate, clavate with an obtuse apex and tapered gradually to a narrow base. Ascospores parallel in the ascus, fragmenting before maturity into cylindrical, pale olive to pale brown, 0 – 3 septate units (occasionally 4-septate units present) measuring 12 – 24 (8 – 43) x 2 – 3 (mean 17 x 2.2) μm. Pseudoparaphyses numerous, filiform, hyaline, irregularly branched, 1 um wide. Colonies forming conidia are effuse and black. Conidia are produced both from the conidiophores and the subicular hyphae, although the modification of the subiculum in this manner is very rare and only results in single short conidia. Conidia from definite conidiophores are poorly differentiated in long chains which mature acropetally. Rarely do the chains break up into separate conidia; in the majority of cases the distinction between individual conidia in the chains is difficult to make, only periodic minor constrictions along their length indicate the probable circumscription of single conidia. Chains are frequently branched irregularly with up to 55 transverse septa and the majority of cells further divided by 1 – 2 longitudinal or oblique septa, thick-walled, irregularly verrucate, reddish brown, up to 230 μm long x 10 – 17 μm wide. Conidiophores are single or fasciculate, simple, rarely branched except at the base, 2 – 5 septate, brown, thick-walled, irregularly verrucate, up to 20 μm long x 3 – 5 μm wide. On decorticated wood of deciduous trees, including Populus tremuloides from Saskatchewan and Alberta, Canada; Populus sp. from Alberta, Canada; Salix sp. from Wyoming and Colorado, USA; Amelanchier sp., Rhus sp., and Quercus sp. from Colorado, USA; and undetermined wood from Saskatchewan and British Columbia, Canada.”

Barr (1990): “Ascomata 850 μm to 1.75 mm in height, 250 – 750 μm wide, compressed, 100 – 250 μm in side view. Asci measure (310-) 450 – 553 (-700) x (7-) 8 – 10 (-12) μm. Ascospores are about the length of the ascus, separating into (1-) 3- (7-) septate part-spores in the ascus, measuring (8-) 14 – 20 (-37) x (1.5-) 2 – 3 μm, in ascus, light yellowish brown. Distribution on wood & periderm of angiosperms, mostly western North America: Populus tremuloides Colorado, Idaho, Maine, Montana. Note: Goree (1974) studied many collections from western North America & determined that this species is widespread in inland regions of Canada & USA. The specimen cited from Maine is sparse in numbers of ascomata, but they are characteristic of the species (i.e., indicating that it is also on the East Coast).”

 

Glyphium grisonense G. Mathiassen 
Mathiassen. 1993. Sommerfeltia 20.

Mathiassen (1993): Description: “Subiculum dense, or very sparsely developed, composed of branched, thick-walled, septate brown hyphae ca. 3.5 um diam., formed from the base or the lower half of pseudothecium. Pseudothecia 0.7 – 1.5(2) mm high, seated upright in subiculum, scattered or clustered, laterally flattened, dolabriform, but often +/- obpyriform in outline with age, black,, with a 200 – 750 um long slit along the sharp upper edge. Peridium 80 – 115 um thick, tow layered, outer layer 30 – 55 8m thick, dense, carbonized and brittle, early bursting vertically along the narrow sides of pseudothecium into two laterally transversely striate plates. (This out layer is unable to grow as fast as the rest of the peridium, thus uncovering larger parts of the inner layer as the pseudothecium develops). Inner layer 35 – 60 um thick, consisting of +/- isodiametric, thin-walled, dark brown cells, the innermost usually becoming narrower and somewhat lighter. Asci 340 – 459 x 12.4 – 18.7 um, mean 396.3 x 15.9 um (n=50), cylindrical-clavate, gradually tapering towards base, bitunicate, thick-walled, at first 8 spored, later polysporous. Paraphysoids ca. 1.5 um in diam., numerous, long, branched, anastomosing, septate, usually with many small guttulae. Ascospores 270 – 420 um (n=118) long, filiform, multiseptate, slightly spiralling, fragmenting before maturity into cylindrical, pale brown (2-) 3 – 7(-11) -septate units, measuring (13.8-) 15.1 – 42.8 (-48) x 2.9 – 5.5 (-6) um, mean 24.2 x 4.3 um, Q=5.8 (n=132). Anamorph seen as effuse, black colonies on naked wood, in close connection with pseudothecia and subiculum. Conidial chains unbranched thick-walled, multiseptate, irregularly constricted, verrucose and dark brown. Maturing acropetally. Nomenclatural and taxonomic notes: Glyphium grisonense was previously treated as G. cf. schizosporum (Mathiassen 1989). The type material of G. schizosporum was not examined, and the determination was based on descriptions given by other authors (Maire 1917; Zogg 1962; Sutton 1970). The type collection of G. schizosporum is kept in Montpellier (MPU), but I have not seen it, as MPU does not lend out types. Slides of the type collection are kept in Edmonton (CFB cf. Sutton 1970, Homgren et al. 1990), but unfortunately I have not received these slides yet. However, I have examined all the known collections of G. schizosporum in Europe. With one exception, they were all different from my material, and matched the description of G. schizosporum perfectly. Therefore, these collections are undoubtedly identical with the type collection. Glyphium grisonense is closely related to G. schizosporum, but the differences are sufficiently pronounced to justify the rank of different species. The ascomata of G. schizosporum are narrow, and never develop to become obpyriform in outline, the asci are very long, but only 7 – 12 um wide, the spores are very narrow, and the conidiophores slightly different from those of G. grisonense. Only known from a few high altitude localities in Switzerland, France and Algeria (Zogg 1962). These species are thus separated on several morphological characters, and in different geographical distribution. One of the two known samples from Switzerland (Kt. Graubunden, Arosa, on Betula), turned out to be Glyphium grisonense. Although both Zogg (1962) and Sutton (1970) refer to this sample in their publications, they have hardly examined it microscopically. Their descriptions of e.g., the asci fits only G. schizosporum: 7 – 9 um wide (Zogg 1962: 102), 9.5 um wide (Sutton 1970: 260). The mature ascomata of G. schizosporum are very similar to young and immature ascomata of G. grisonense, but even young, immature and empty asci are considerable wider than mature asci of G. schizosporumHosts: In this investigation found on Salix caprea ssp. sericea (9 col.), S. myrsinifolia ssp. myrsinifolia (11 coll.), S. pentandra (18 coll.). So far only known from Betula and Salix(see also Mathiassen 1989: 49). Ecology: A predominantly lignicolous species, but in some samples also found on bark. Most frequent on dead twigs, probably a primary saprophyte, but also able to sporulate on decayed substrate. Usually accompanied with the anamorph (Peyronelia sp., see Sutton 1970: 263). Distribution: Only known from North Norway and from one high altitude locality in Switzerland. Fits within the disjunct distribution pattern which I earlier discussed for Glyphium cf. schizosporum and Hypoxylon macrosporum (cf. Mathiassen 1989: 21). More samples were found in central Scandinavia than in Troms, but as the majority of these samples were restricted to only one, rather small geographical area, it seems to be more common in the north. In Troms (Mathiassen 1989) it was distributed in al the vegetation regions, except in pure LA, and I therefore expect it to occur in northern Sweden too. This large region has so far been very little investigated, except for a few small areas. This probably explains its absence in the Swedish pyrenomyete lists and herbaria”.

 

Glyphium schizosporum (Maire) Zogg
Zogg, 1962. Beiträge zur Kryptogamenflora der Schweiz, Band 11(3): 101.

Lophium schizosporum Maire, 1917

Rarely collected. Switzerland, Southern France, North Africa. Collected from old wood of Betula, Rhamnus.

The following is unpublished material and under strict copyright protection by Eric W.A. Boehm, as it presently forms the basis for a manuscript under preparation. It is included here, “pre-press” for general release, but its use is highly restricted.

  1. schizosporum(Maire) Zogg– “Glyphium corrugatum is morphologically very similar to the only other species with disarticulating spores, namely Lophium schizosporum Maire, which was the second species Zogg (1962) transferred to the genus Glyphium, as G. schizosporum (Maire) Zogg. G. schizosporum has been reported from Algeria, France, Switzerland and Norway (Mathiassen 1989; Sutton 1970; Zogg 1962). Aside from geographical differences, the two species may be distinguished as follows: the part-spores of G. schizosporum consist of fragments that are slightly larger in diameter, with more septa than those found in G. corrugatum (1- to 9-septate, mean 5-septate, versus 0- to 3- (4-)septate, respectively). Additionally, although close in measurement, the verruculose conidia of the Peyronelia state in G. schizosporum typically have up to 50 transverse septa with only an occasional longitudinal septum, and produce only straight chains that do not branch; whereas those of G. corrugatum have up to 55 transverse septa & usually one or two longitudinal septa, and are prone to producing lateral buds or side branches (Sutton 1970). It is of unknown significance why dematiaceous hyphomycetous anamorphs are found only in the two species of Glyphium with disarticulating ascospores. However, cultural work with other members of the Mytilinidiaceae by Lohman (1933a, b) has demonstrated similar anamorphic states for the genus Lophium Fr. and Mytilinidion Duby. Although not congeneric, the anamorphs of Glyphium, Lophium and Mytilinidion tend belong to the same developmental series of the Hughes classification scheme (Hughes 1953). That is, they belong to Section I, characterized by catenate conidia with acropetal development, where an apical meristem results in the oldest conidia at the base of the chain (Sutton 1970). Lohman (1933b) described a Septonema toruloideum Cooke & Ellis anamorphic state for M. scolecosporum M.L. Lohman, a fungus belonging to subgenus Lophiopsis Lohman that morphologically trends into the genus Lophium (1932). Bisby and Hughes (1952) also found Septonema secedens Corda (Hughes 1951) as the anamorph for M. karstenii Sacc. (= M. rhenanum Fuckel; Zogg 1962). Lohman (1933a) found a Papulospora mytilina (Pers.) Lohman state for L. mytilinum Pers. : Fr., although this was later not recognized by Tubaki (1958). All of these dematiaceous anamorphs possess catenate conidia that arise in acropetal succession (Hughes 1953)”. © Eric W.A. Boehm 2009

Sutton (1970): “Pseudothecia 0.1 – 0.15 x 0.025 – 0.05 cm, superficial, scattered, gregarious or crowded, arising amongst the subicle, conidiophores, or from bare wood, erect or slanting, dolabriform and flattened laterally, narrower towards the base, black, with several transverse parallel ridges and subicular hyphae extending down towards the substrate from the vertical edges, lenticular in longitudinal sections taken at 90 degrees to the long axis, unilocular, lateral walls composed of outer sclerotioid black prosenchyma becoming thin-walled, pale brown pseudoprenchyma. Asci up to 250 um long x 9.5 um wide, bitunicate, clavate, with an obtuse apex and tapered gradually to a narrow base. Ascospores parallel in the ascus, fragmenting before maturity into cylindrical, pale olive to pale brown, 1 – 9 septate units (mean 5-septate) measuring 14 – 26 (9 – 36) x 3 – 4 (mean 18 x 3.5) μm. Pseudoparaphyses numerous, filiform, hyaline, irregularly branched, 1 um wide. Colonies producing conidia are effuse and black. Conidiophores single, more rarely fasciculate, simple, unbranched, up to 3-septate, brown, thick-walled, irregularly verrucate, 10 – 15 x 4 – 6 μm. Conidia from definite conidiophores mature acropetally and are poorly differentiated. Rarely do the chains break up into separate conidia and in most cases it is difficult to distinguish individual conidia. Chains are not regularly constricted and remain unbranched, with up to fifty transverse septa and the occasional longitudinal septum, thick-walled, irregularly verrucate, dark brown, up to 220 μm long x 8 – 11 μm wide. On decorticated wood of deciduous trees, including Betula, Rhamnus alpina and Salix appendiculata and indt. wood from Switzerland; Ilex aquifolium from North Africa; and indet. wood from France.”