Lophium Fries (pro parte)

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!

The genus Glyphium Nitschke ex Lehmann
by Eric W.A. Boehm

The genus Glyphium, with erect dolabrate ascomata, was originally included by Zogg (1962), Barr (1987, 1990) and others (e.g., Goree 1974; Lorenzo & Messuti 2005; Sutton 1970) in the Mytilinidiaceae. However, recent molecular evidence, based only on a single isolate of G. elatum (Grev.) Zogg (CBS 268.34), 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 (Geiser et al. 2006; Lucking et al. 2004; Schmitt et al. 2005). The Chaetothyriales represent an extreme case of convergent evolution, where the bitunicate ascus and pseudothecium have evolved independently of the Dothideomycetes and constitute an entirely unrelated group within the Eurotiomycetes. Thus, it may not be that improbable that Glyphium indeed represents an extreme case of homoplasy in the mytilinidiaceous pseudothecium. But this all rests on the analysis of one single isolate (see Lindemuth et al. 2001)! Clearly, more isolates are needed to confirm the phylogeny of this very interesting and beautiful fungus. From previous reports (Lohman 1933a; Sutton 1970), apparently the ascospores do not germinate on media, necessitating "tissue cultures" to be made from subicular hyphae, from which conidia can be derived. However, Sutton (1970) states that "although conidia germinated, growth ceased almost immediately". Contrary to these earlier reports, recently a number of cultures have been initiated from discharged ascospores of G. elatum, originating from Salix and Malus. The filiform ascospores germinated readily on PDA. Interestingly, the colonies formed bear little resemblance to those of CBS 268.34.

Dichotomous Key to species of Glyphium Nitschke ex Lehmann
(modified from Sutton 1970, Barr 1990, Goree 1974 & Zogg 1962)

A. Ascospores not disintegrating into part-spores in the ascus:

1. Fruitbody 1 – 3 mm in height, 0.2 – 0.3 mm in width.

= Glyphium elatum (Grev.) Zogg

2. Fruitbody smaller, 0.3 – 0.5 mm in height, 0.2 – 0.3 mm in width. Florida on Tillandsia.

= Glyphium tillandsiae (Cash) Zogg

B. Ascospores disintegrating into part-spores in the ascus:

1. Ascospore part-spores or "fragments" consistently 3-septate or less, slightly narrower than G. schizosporum; 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.

= Glyphium corrugatum (Ell.) Goree

2. Ascospore part-spores or "fragments" usually with more than 3 septa per fragment, 1 - 2 um wider than G. corrugatum; 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.

= Glyphium schizosporum (Maire) Zogg

Detailed Dichotomous Key to species of Glyphium Nitschke ex Lehmann
(modified from Sutton 1970, Barr 1990, Goree 1974 & Zogg 1962)

A. Ascospores not disintegrating into part-spores in the ascus:

1. Fruitbody 1 – 3 mm in height, 0.2 – 0.3 mm in width.

= Glyphium elatum (Grev.) Zogg

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

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

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.”
 

2. Fruitbody smaller, 0.3 – 0.5 mm in height, 0.2 – 0.3 mm in width. Florida on Tillandsia.

= Glyphium corrugatum (Ell.) Goree

Goree, H. (1974. Can. J. Bot. 52: 1265).

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

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. Asci 125 - 300 x 10 – 11 um, 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) um. 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 um long x 10 – 17 um wide. Conidiophores are single or fasciculate, simple, rarely branched except at the base, 2 - 5 septate, brown, thick-walled, irregularly verrucate, up to 20 um long x 3 – 5 um 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).”

2. Ascospore part-spores or "fragments" usually with more than 3 septa per fragment, 1 - 2 um wider than G. corrugatum; 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.

= Glyphium schizosporum (Maire) Zogg

Zogg, 1962. Rarely collected. Switzerl&, Southern France, North Africa. Collected from old wood of Betula, Rhamnus.

Lophium schizosporum Maire, 1917

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) um. 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 um. 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 um long x 8 - 11 um 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."