Glonium

4. The genus Glonium Muhl. : Fr.
syst. Myc. II, 1823, 594-595

Solenarium Sprengel, 1827
Psiloglonium (von Hohnel) Petrak, 1923b

Unidentified species of Glonium from NJ.

The genus Glonium Muhl. : Fr.
by E.W.A. Boehm

The genus Glonium Muhl. : Fr. is characterized by hyaline to yellowish, uniseptate, slighlty asymmetric ascospores, constricted at the septum, with obtuse or acuminate ends, and typically with cells unequal in size. The type species, Glonium stellatum Muhl. : Fr., is characterized by ascospores with acuminate ends and hysterothecia that are progressively dichotomously-branched, laterally anastomosed to form radiating labelliform or stellate composites, seated upon a conspicuous brown felt-like subiculum – features not found in most other members of the genus (Lohman 1937; Zogg 1962). De Notaris (1847) thus extended the original generic concept, based on G. stellatum, to include species with and without subicula, stressing ascospore morphology instead, an idea supported by Zogg (1962). Other workers, however, have historically viewed the genus Glonium as comprised of two distinct morphological series. von Höhnel (1918) stressed the importance of subicula, using it to divide the genus into two subgenera, Glonium and Psiloglonium, with or without subicula, respectively. Petrak (1923) raised Psiloglonium (von Höhnel) Petr. to generic rank, while Müller and von Arx (1962) originally accepted the genus Psiloglonium, but later reduced it to a synonym of Glonium (Müller and von Arx 1975). Lohman (1933) also did not support von Höhnel’s Psiloglonium, based on the observation that similar anamorphic states were shared between species of the two subgenera. However, Barr (1987) was one of the only modern authors to retain the genus Psiloglonium, as distinct from the subiculate Glonium. In addition to lack of subicula, species within Psiloglonium typically possess ascospores with at least one end obtuse, whereas subiculate species within Glonium, have ascospores typically with both ends acuminate, and are also usually characterized by laterally fused, dichotomously branched hysterothecia, in some cases associated with a stroma (e.g., G. compactum Kern; Zogg 1962). The North American species were reviewed by Lohman (1937). Zogg (1962) accepted eleven species in the genus. The Psiloglonium series, listed here by increasing ascospore length, include: G. abbreviatum (Schw.) Lohman, G. pussilum Zogg, G. lineare (Fr.) de Not., G. chambianum Guyot, G. finkii (Petrak) Lohman and G. curtisii (Duby) Lohman, the latter transferred to Ostreichnion in the Mytilnidiaceae (Barr 1975). The second series, corresponding to von Hohnel’s circumscription of the genus Glonium inlcude: G. caucasicum (Rehm) Zogg, G. hysterinum Rehm, G. graphicum (Fr.) Duby, G. stellatum, and G. compactum. An additional four species have been recently described or reinstated for the genus, namely: G. sasicola Amano from Japan, the first report of a gelatinous sheath in the genus (Amano 1983), the reinstated G. costesi Speg., and the newly described G. colihuae Lorenzo & Messuti from Argentina (Lorenzo and Messuti 1998) – all belonging to the Psiloglonium series. The recently described subiculate, acuminate-spored G. circumserpens Kantvilas & Coppins from Tasmania, belongs to the Glonium series (Kantvilas and Coppins 1997). This is a highly unusual member of the Hysteriaceae, because it occurs not only on wood, but is also terricolous and saxicolous and apparently facultatively lichenized as well (Kantvilas and Coppins 1997). Clearly, molecular data is needed to substantiate whether indeed the genus Glonium is comprised of two separate lineages, Psiloglonium and Glonium. This relates to the phylogenetic importance placed on subicula and on the characteristic manner by which hysterothecia anastomose and bifurcate in the type (G. stellatum) and related species, belonging to von Höhnel's circumscription of the genus Glonium. Also, the question of Actidiographium, in the Hysteriaceae, and Actidium, in the Mytilinidiaceae, and their affinity or lack thereof with the genus Glonium needs to be addressed with molecular data.

Amano (1983): “Glonium Muhlenberg: Fries emend. de Notaris. Muhlenberg, Cat. Am., p. 101, 1813; Syst. mycol. 2: 594, 1823, emend., de Notaris, Giorn. bot. ital. II, 2 (7-8): 27, 1847.Type species: Glonium stellatum Muhlenb.: Fr., Syst. mycol. 2: 595, 1823. Although ascomata of Glonium stellatum, the type species of the genus Glonium, are embedded in subicula, de Notaris (1847) extended the original generic concept to include species with ascomata not embedded in subicula; he placed more emphasis on characteristics of the ascospores. Von Hohnel (1918) considered the presence or absence of subiculum to be taxonomically important. He subdivided the genus Glonium sensu de Notaris (1847) into two subgenera Glonium & Psiloglonium, with or without subiculum, respectively. Petrak (1923) raised the subgenus Psiloglonium to generic rank. Muller & von Arx (1962) accepted the genus Psiloglonium but later reduced it to a synonym of Glonium (von Arx & Mueller, 1975). Zogg (1962) regarded Psiloglonium as a synonym of Glonium, because both subiculate & non-subiculate ascomata were observed in the same species. von Arx & Muller (1975) state 10 species for Glonium.”

Wehmeyer (1975): “Glonium has ellipsoid to elongate or branched ascostromata, often clustered upon a subiculum. The species without a subiculum have been placed in a genus Psiloglonium by some authors. The ascospores are two celled & hyaline. Several species have pycnidial states of the form genera Sphaeronaema or Plenodomus. Glonium clavisporum has a Sporidesmium state with irregular, brown, muriform conidia borne singly on slender conidiophores. Lohman (1931, 1937) has reported upon the life histories of the genus. (The genus Lophiosphaera, in the Lophiostomataceae, differs only in the ascostromata being sunken in the substrate)…[re: centrum development (Luttrell (1953)]…This is typical of the bitunicate & ascolocular type of development found in the Dothideales. The presence of true pseudoparaphyses is somewhat anomalous but would probably indicate a relationship with the Pleosporales near the Lophiostomataceae. It indicates also the sort of variation which confused the original attempt to separate the types of centrum development.”

Dichotomous Key to species of Glonium Muhl. : Fr.
(modified from Zogg 1962)

A. Psiloglonium series: 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: corresponds to the Psiloglonium series sensu von Hohnel (1918) & Petrak (1923), designated as: Psiloglonium Höhn., Annales Mycologici 16: 145 (1918). Typus: Psiloglonium lineare (Fr.) Petr., Annales Mycologici 21: 227 (1923).

I. Asci cylindrical to club-shaped. Ascospores in one row or distichous in asci, but regularly arranged. Alternative (II) tropical.

(a) Ascospores oval to oblong; one end of spore rounded or obtuse, the other weakly (+/-) acuminate; upper rounded cell larger; +/- constricted at septum.

1. Ascospores (5) 6 – 7 (8) x 2 – 3 (3.5) μm:

= Glonium abbreviatum (Schw.) Lohman

2. Ascospores (9) 10 – 12 (13) x 4 – 5 (6) μm:

= Glonium pusillum Zogg

3. Ascospores (10) 12 -14 (18) x (4) 5 – 7 (8) μm:

= Glonium lineare (Fr.) De Not.

4. Ascospores (14) 16-18 (21) x (6) 8 – 9 (10) μm:

= Glonium chambianum Guyot

5. Ascospores + gelatinous sheath, 25 – 32 x 5 – 8 μm (Japan):

= Glonium sasicola N. Amano

6. Ascospores 30 – 35 x 8 – 15 μm (Chile):

= Glonium costesi Speg.

7. Ascospores (59) 62 – 68 (76) x 13 – 15 μm (SE USA):

= Glonium curtisii (Duby) Lohman

(b) Ascospores spindle-shaped, with both ends acuminate; constricted at septum.

1. Ascospores (19) 22 – 25 (27) x (6) 7 – 9 (10) μm:

= Glonium caucasicum (Rehm) Zogg

2. Ascospores 30 – 43 x 4 – 9.8 μm (Argentina):

= Glonium colihuae Lorenzo & Messuti

3. Ascospores about 45 x 9 μm:

= Glonium hysterinum Rehm

II. Asci ovoid, +/- cylindrical; ascospores borne in upper portion of ascus (not evenly distributed), unevenly regimented; ascospores (12) 13 – 15 x 6 – 7 μm (Tropical):

= Glonium finkii (Petrack) Lohman

B. Glonium series: 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 are globose to obovoid, typically with a thick three-layered peridium (although more fragile than those of Psilogolonium): the following correspond to the series Glonium sensu von Hohnel (1918) & Petrak (1923): Typus: Glonium stellatum Muhl. : Fr., Cat. Pl. Amer. Sept.: 101 (1813); Fries, Syst. Mycol. 2: 594-5 (1823).

1. Hysterothecium branched, irregular, “graphoid”. Without subiculum or sitting upon a good development of subiculum on the substrate. Ascospores oblong to spindle-shaped; upper cell pear-shaped, constricted at septum; both ends acuminate. Ascospores (13) 15 – 18 (21) x (3) 5 – 6 μm:

= Glonium graphicum (Fries) Duby

2. Hysterothecium mostly star-shaped or stellate in overall appearance, with branched & laterally anastomosing hysterothecia. Distinct creeping appearance. Mostly associated with a well-developed subiculum. Ascospores oblong to spindle-shaped; upper cell pear-shaped, constricted at septum; both ends acuminate. Ascospores (18) 21 – 26 (28) x (4) 5 – 6 (7) μm:

= Glonium stellatum Muhl. : Fr.

3. Hysterothecium borne within a crust-like stroma; subiculum embedded in stroma. Ascospores spindle-shaped with upper cell slightly swollen & larger than lower cell. Ascospores 24 – 28 x 5 – 6 μm:

= Glonium compactum Kern

4. Hysterothecium lirelliform, laterally-anastomosed, branched-elongate, on soil, saxicolous or lignicolous (Tasmania); Ascospores hyaline, constricted at the septum, apices pointed, (15-) 16 – 17 x 6 – 7 μm:

= Glonium circumserpens (Nyl.) Kantvilas & Coppins

Detailed Dichotomous Key to species of Glonium Muhl. : Fr.
(modified from Zogg 1962)

I. Asci cylindrical to club-shaped. Ascospores in one row or distichous in asci, but regularly arranged. Alternative (II) tropical.

(a) Ascospores oval to oblong; one end of spore rounded or obtuse, the other weakly (+/-) acuminate; +/- constricted at septum.

1. Ascospores (5) 6 – 7 (8) x 2 – 3 (3.5) μm:

= Glonium abbreviatum (Schw.) Lohman

Lohman. 1937. Bull. Torr. Bot. Cl.: 64, 64. Cosmopolitan (Europe, Americas, Java). On old wood & bark of: Araucaria, Palmae, Alnus, Ulmus, Magnolia, Liriodendron, Eriobotrya, Citrus, Acer, Berchemia, Tilia, Eucalyptus, Rhododendron.

Hysterium abbreviatum Schw., 1834
Hysterium medium Cooke, 1879
Glonium medium (Cooke) Sacc., 1883
Psiloglonium microspermum v. Hohnel, 1924
Glonium macrosporum Sacc., 1875
Hysterium parvulum Gerard, 1874b
Glonium parvulum (Ger.) Sacc., 1883
Psiloglonium parvulum (Ger.) E. Muller, 1962
Glonium ruthenicum Petrak, 1921
Psiloglonium ruthenicum Petrak, 1925

Lohman (1937): “Hysterothecia short oblong, up to 0.5mm in length (usually less than 1mm in length; rarely to 1.5mm), smooth with obtuse ends, erumpent from between the fibers of the wood in more or less parallel series, +/- sunken into substrate, usually densely crowded & blackening the substratum; on firm weathered wood. Paraphyses filiform, simple Asci short-cylindric, 40-50 x 4-5 μm. Ascospores +/- ovoid or oblong (at least with the upper end rounded), the lower cell usually narrower & often tapering; ascospores oblong or ovoid with the lower cell somewhat the narrower, obliquely uniseriate, constricted at the septum, 6.5-8 x 2.5-3.5 μm. Pycnidia Sphaeronaema parvulum Lohman (1933) superficial, single or clustered, 150-200 μm in diameter, 225-300 μm in height including the beak; conidia hyaline, ovate to inequilateral or elliptic-oblong, 1 x 1.5 μm, acrogenetic on simple conidiophores, 6-9 x 1.5-2 μm. Pycnidia rarely found accompanying hysterothecia, but have been obtained in ascospore derived cultures from Acer, Liriodendron, Tilia, & Ulmus. Common throughout the Eastern USA, occurring on firm weathered wood, esp. Quercus, Magnolia & Liriodendron in the S Appalachians. The short hysterothecia & small spores are distinctive features. Pycnidial stage relates it to G. stellatum.”

Lorenzo & Messuti (1998) “Hysterothecia solitary to gregarious, erumpent to superficial, smooth, ellipsoid with rounded extremes, straight, unbranched, 0.26 – 1.0 x 0.1 – 0.4mm. Pseudoparaphyses hyaline, septate, branched, filiform, ca. 1 – 1.5 μm, longer than the asci. Asci cylindrical, octospored, 40 – 75 x 3.29 – 7.0 μm. Ascospores hyaline to yellowish-brown, irregularly uniseriate, transversely uni-septate, slightly constricted at the septum, upper cell broader than lower cell, oval to oblong, guttulate, (5-) 6 – 11 x 2 – 4 μm.”

van der Linde (1992): “Fruitbodies arranged in groups, closely associated, erumpent or superficial, long-ellipsoid with rounded ends, straight, not branched, up to 1.2 x 0.25mm. Peudoparaphyses hyaline, filiform, very thin & slender, septate, not markedly thickened at the tips. Asci cylindrical, irregularly uniseriate, 8-spored, 55 – 60 x 7um. The asci are small in comparison with those of other species in this genus. Ascospores hyaline, 1-septate, upper cell broader than lower cell, constricted at the septum, measuring 7 – 8 x 3 um.”

2. Ascospores (9) 10 – 12 (13) x 4 – 5 (6) μm:

= Glonium pusillum Zogg

Zogg H. 1962. Die Hysteriaceae s. str. und Lophiaceae. Beitr Kryptogamenfl Schweiz 11: 1-190. Rare. Southern French Maritime Alps. Collected from old needle scales protuberances of living Juniperus phoenicea & from the bark of living Pinus silvestris.

Lee & Crous (2003): “Ascomata hysterothecioid, subglobose to globose in vertical section, solitary to gregarious, erumpent to superficial with base immersed, straight to flexuous, ellipsoidal with pointed extremes, opening by a depressed longitudinal slit, 0.2 – 0.3 mm long, 0.1 – 0.2 mm wide. Peridium consisting of three layers, up to 17.5 μm in diameter, pseudoparenchymatous, carbonaceous, outer layer dark, inner layer hyaline, compressed, middle layer consisting of 1 – 2 rows of thick walled, brown cells. Pseudoparaphyses cellular, hyaline, septate, flexuous,, branched, 1.5 – 2 μm wide. Asci bitunicate, cylindric to claviform, stipitate, octosporous, 33 – 48 x 8 – 11 μm. Ascospores (9-) 11 – 12 (-13) x 4 – 5 μm, ellipsoid with upper cell often larger, hyaline, with one traverse septum. Zogg (1962) first described this species from Juniper & Pine collected in Europe. An additional record was made from Taiwan by Sivanesan & Hsieh (1989. Mycol. Res. 93: 340-351). The asci in the original description (26 – 35 x 10 – 12 μm) are smaller than in our collections, but the ascospore dimensions (9 – 13 x 4 – 6 μm) are similar. This species is similar to G. lineare in ascospore shape & its small ascomata, but differs in ascospore dimensions (10 -1 8 x 4 – 8 μm).”

3. Ascospores (10) 12 -14 (18) x (4) 5 – 7 (8) μm:

= Glonium lineare (Fr.) De Not.

de Notaris. 1847. Giorn. Bot. Ital. II: 27-28. Cosmopolitan. Old bark & wood of: Taxus, Salix, Carpinus, Corylus, Alnus, Fagus, Castanea, Quercus, Ficus, Pyrus, Crataegus, Rosa, Prunus, Robinia, Buxus, Pistacia, Cotinus, Ilex, Acer, Aesculus, Kalmia, Fraxinus.

Hysterium lineare Fries, 1823
Psiloglonium lineare (Fr.) Petrak, 1923
Glonium clavisporum Seaver, 1925a
Hysterium confluens Wallroth, 1833
Glonium confluens (Wallr.) Duby, 1862
Hysterium fraetum Dunfour (in Herb. Fries)
Glonium interruptum Saccardo, 1873
Hysterium Kalmiae Schweinitz, 1834
Glonium simulans Gerard, 1876
Glonium macrosporum N. Amano? (proposed syn. E. Boehm)

Ellis & Everhart (1892) for Glonium lineare: “Hysterothecia more or less crowded, sometimes longitudinally confluent, mostly lying parallel, subimersed in the wood which is often blackened, linear, rather flat, straight or flexuous, ends obtuse, black, smooth, lips slightly swollen 1-1.5mm long x 0.5 mm wide. Asci clavate, cylindrical, 75-90 x 12-14 μm. Paraphyses conglutinate, their tips united above & forming a dark colored, coarsely granular stratum (epithecium). Ascopores uniseriate, ovate, uniseptate, ad slightly constricted at the septum, hyaline, 12-15 x 6-8 μm. Common on old decorticated wood of various deciduous trees.”

Lee & Crous (2003): “Ascomata hysterothecioid, solitary to gregarious, almost rectangular in vertical section, erumpent to superficial, straight to flexuous, ellipsoidal with blunt to pointed extremes, opening by a depressed longitudinal slit, 0.2 – 0.4mm long, 0.1 – 0.2mm wide, up to 0.1mm high. Peridium consisting of three layers, up to 12.5μm in dia, pseudoparenchymatous, carbonaceous, outer layer dark, present only in sides & top, inner layer hyaline, compressed, middle layer consisting of 1 – 2 rows of thick-walled, brown cells between outer & inner layers at sides & 3 – 4 rows at the bottom. Pseudoparaphyses cellular, hyaline, septate, flexuous branched, 1.5 – 2.5μm wide. Asci bitunicate, cylindric to clavate, stipitate, octosporous, 43 – 55 x 9 - 13μm. Ascospores (12-) 13 – 14 (-15) x (4-) 5μm (Mean = 13.5 x 4.7μm), ellipsoid with upper cell often broader, hyaline with one transverse septum: hyaline didymospores.” This S. African G. lineare is atypical, with very small hysterothecia.

Lohman (1937): “Hysterothecia usually in parallel series on tangential surfaces of firm wood & accompanied by a thin black crust, variably elongated, averaging about 1.5mm, but often confluent linearly, at first narrow with a fine longitudinal fissure, later up to 0.4mm broad with the black epithecium more or less exposed, developed within the wood & not easily freed, with the flat stromatic top scarcely above the surface at maturity & the lateral walls poorly developed & light colored below. Paraphyses branched & united above into a bluish brown granular mass forming the epithecium. Asci cylindric, clavate, when not extended measuring (55) 60-75 (90) x (10) 12-14 μm. Ascospores obliquely uniseriate or sub-biseriate, oblong or ovoid, the lower cell often narrower, constricted at the septum, measuring (10) 12-15 x (5) 6-8 μm. The long, flat fructifications, dark epithecium & broad ascospores are distinctive features. Pycnidia preceding & bordering hysterothecial aggregates, arising within the wood & then erumpent by their short papillae, (150) 200-300 μm in diameter. Conidia hyaline, 1-celled, elliptic oblong, 2.5-3 x 1.5-2 μm, acrogenetic on slender conidiophores. This is the Plenodomus type (Lohman, 1933), described previously (Hilitzer, 1929) as Hysteropycnis globularis.”

Lohman (1937) for Glonium simulans Gerard, synonimized by Zogg (1962) into Glonium lineare, Lohman writes: “Hysterothecia superficial, scattered to closely aggregated, 0.5-2.0 x 0.3-0.4mm, obtuse at the ends when mature, rugose, punctuate or finely longitudinally striate, usually seated upon a think black crust. Asci cylindric (60) 65-80 (85) x (8) 10-12 (14) μm. Paraphyses branched above forming a pale, yellowish granular epithecium. Ascospores elliptic-biconical, slightly curved, some becoming yellow-brown in age, measuring (10) 14-16 (18) x (4.5) 5-6 μm; the upper cell usually broader than the lower at the septum. No secondary stage observed. Cultures produce only sterile mycelia. Species appears more closely related to Glonium clavisporum than to Glonium abbreviatum & Glonium lineare, with which it has often been confused.”

Lohman (1937) for Glonium clavisporum Seaver, synonimized by Zogg (1962) into Glonium lineare, Lohman writes: “Hysterothecia superficial, prominent, straight or flexuous, rounded above & with obtuse ends, smooth or longitudinally striate, up to 2.2 mm in length x 0.4mm in width; apparently highly variable. Asci cylindric, 90-110 x 8-10 (12) μm. Paraphyses branched above, oily granular, pale to greenish yellow in mass. Ascospores elliptic to clavate-fusoid, slightly curved, constricted at the septum, uniseriate but overlapping, measuring (15) 16-18 (20) x 5-6 μm. Hysterothecia associated with scattered clusters of large dark conidia of Sporidesmium stygium B. & C. These measure (65-90) 120-220 (300) x (30) 50-90 μm , averaging 140-190 x 60-85 μm.” Lohman further states: “While hysterothecia variable, ascospore morphology & size quite uniform. Hysterothecia when on tangential surfaces of wood are more elongate, often confluent laterally & straight; whereas when seated on bark, they are variable in length, irregularly scattered, the longer ones flexuous, smooth or striate, solitary or densely crowded, with or without associated black crust or with or without the associated Sporidesmium stygium stage!” Habitat: “Bark of logs & living trees & on firm weathered wood of Vitis, Liquidambar, Sassafras, Platanus, Quercus, Acer.”

Glonium clavisporum Seaver. Note the Sporidesmium stygium B. & C. anamorphic state surrounding the hysterothecia. Glonium lineare & Glonium simulans, to which this most closely resembles, possess a different anamorphic state or do not possess one altogether. Zogg synonimized both under G. lineare.

Note: Again, Zogg synonimized G. simulans & G. clavisporum into G. lineare, however different anamorphic states (Plenodomus/Hysteropycnis for G. lineare & Sporidesmium for G. clavisporum, suggest that at least these two species are distinct.

4. Ascospores (14) 16-18 (21) x (6) 8 – 9 (10) μm:

= Glonium chambianum Guyot

Guyot. 1955. Ann. Serv. Bot. et Agr. Tunisie 28: 90-91. Rare. Collected from decayed branch of Lonicera implexa from North Africa.

Lee & Crous (2003): “Ascomata hyeterothecioid, subglobose to globose in vertical section, solitary to gregarious, erumpent to superficial with base immersed, straight to flexuous, simple to arely branched, ellipsoid with blunt extremes, opening a depressed longitudinal slit, 0.4 – 0..6mm long, 0.1 – 0.2mm wide, 0.1 – 0.2mm high. Peridium consisting of two layers, up to 40 μm in diameter, carbonaceous, outer layer dark, inner layer hyaline, compressed. Pseudoparaphyses cellular, hyaline, septate, flexuous, branched, 0.5 – 1.5 μm wide. Asci bitunicate, cylindric to clavate, stipitate, octosporous, biseriate, 73 – 88 x 13 – 18 μm. Ascospores (18-) 20 – 21 (-23) x (4-) 5 – 6 (-7) μm, ellipsoid to oblong, with upper cell often broader, hyaline, with one transverse septum, slightly constricted at the septum. From: dead flower heads of Protea susanne, twigs of Leucospermum praecox. The fist record of this species was from northern Africa on dead twigs of Lonicera implexa (Caprifoliacaeae). Species is similar to G. lineare but with larger ascospores.”

5. Ascospores + gelatinous sheath, 25 – 32 x 5 – 8 μm (Japan):

= Glonium sasicola N. Amano

Amano, N. 1983. Trans. mycol. Soc. Japan 24: 283-297. Japan. Collected from dead culm of Sasa sp.

Amano (1983): “Ascoma oval or elongated with somewhat acute ends, dispersed, straight or slightly curved, not branched, sometimes faintly longitudinally striate, superficial or immersed at base, opening by a longitudinal slit, black, carbonaceous, 0.15 – 1.5mm long, 180 – 240um wide, 130 – 150um high. Tissues of ascomatal walls of textura epidermoidea, brown or dark brown. Pseudopharaphyses filiform, septate, branched, anastomosed, forming brownish, epithecium above. Asci bitunicate, clavate or cylindrical, short-stipitate, eight-spored, 85 – 105 x 15 – 18um. Ascospores irregularly biseriate, fusiform with hyaline gelatinous sheath, one-septate, constricted at the septum, hyaline, with rounded ends, 25 – 32 x 5 – 8um. On dead culm of Sasa sp. Japan. Sterile in culture. G. sasicola is mainly characterized by one-septate hyaline ascospores with a hyaline gelatinous sheath. No species of Glonium has been reported to have ascospores with gelatinous sheath.”

6. Ascospores 30 – 35 x 8 – 15 μm (Chile):

= Glonium costesi Speg.

Spegazzini. 1921. Bol. Acad. nac. Cs. Cordoba 25: 78. On live twigs of Proustia pirifolia, in the surroundings of Los Perales, Chile

Lorenzo & Messuti (1998): “Hysterothecia without subiculum, solitary, superficial, slightly striated, ellipsoid (=cymbiform), straight, 0.5 – 0.75 x 0.15 – 0.25 mm. Pseudoparaphyses hyaline, scarce, septate, branched, anastomosed, irregularly thickened, ca. 1.5 μm in dia. Ascospores hyaline, biseriate, transversely uni-septate, constricted at the mid-septum, fusiform, with rounded ends, the upper cell larger than the lower, broadly ovate, 30 – 35 x 8 – 15 μm. Spegazzini describes this as growing on live twigs of Proustia pirifolia, in the surroundings of Los Perales, Chile.” Note: species validated & removed from Zogg’s (1962) “doubtful species” list by Lorenzo & Messuti.

7. Ascospores (59) 62 – 68 (76) x 13 – 15 μm:

= Glonium curtisii (Duby) Lohman

Lohman, M.L. 1937. Bull. Torr. Bot. Club 64: 66. Rare. South & North America (Southern States). Old wood of: Sabal, Smilax, Castanea, Quercus, Persea, Rubus, Prunus, Cyrilla, Berchemia, Vitis.

Hysterium Curtisii Duby, 1862
Gloniella Curtisii (Duby) Sacc., 1883
Hysteroglonium Curtisii (Duby) Earl, 1901
Hysterium chlorinum Berk et Curt. (nach Sacc, 1883)
Glonium chloriunm (B. et C.) Sacc, 1883
Hysterium cyrillae Berk. et Curt. (nach Sacc., 1883)
Glonium cyrillae (B. et C.) Sacc. 1883
Psiloglonium curillae (B. et C.) Muller (Muller & von Arx, 1962)
Glonium pigasporum Ell. et Ev. (in Herb. Ellis)
Glonium macrosporium Tracy et Earle, 1896

Note: Barr has transferred Glonium Curtisii (Duby) Lohman to Ostreichnion Curtisii (Duby) M.E. Barr [1975. Mycotaxon 3(1): 86] in the Mytilinidiaceae.

Lohman (1937): “Hysterothecia scattered, breaking through the bark or periderm but appearing superficial at maturity, narrow to broad elliptic, sub-conchiform, 0.5-0.7 (1) x 0.5 mm, black, smooth or finely striate. Asci with walls thickened apically, broad cylindric, 155-180 x 30-35 μm with short or inconspicuous stalk & fine filiform granular paraphyses branched above in an oily-granular matrix, the mass sordid white to pale citron. Ascospores cylindric with rounded ends, slightly curved, hyaline to pale greenish or yellow-brown, biseriate, measuring (59) 62-68 (76) x 13-15 μm, constricted at the septum with the lower cell about one third the length of the spore & the end walls, especially the upper, thickened with age in such a manner tat the endoplasts appear cup-shaped at the ends & the cells pseudoseptate. From North Carolina to Florida & Louisiana on small twigs & stems of various broadleafed species: Berchemia, Castanea prunella, Cyrilla, Persea, Prunus, Quercus aquatica & Q. nigra, Rubus, Sabal, Smilax & Vitis”. A highly unusual member of the Hysteriaceae.

(b) Ascospores spindle-shaped, with both ends acuminate; constricted at septum.

1. Ascospores (19) 22 – 25 (27) x (6) 7 – 9 (10) μm:

= Glonium caucasicum (Rehm) Zogg

Zogg H. 1962. Beitr Kryptogamenfl Schweiz 11: 1-190. Rare. Europe. Old climber of Rubus in the Caucuses.

Gloniella caucasica Rehm, 1912c

2. Ascospores 30 – 43 x 4 – 9.8 μm (Argentina):

= Glonium colihuae Lorenzo & Messuti

Lorenzo & Messuti. 1998. Mycol. Res. 102: 1101-1107. Collected from Chusquea culeou (Colihuetium) from Argentina.

Lorenzo & Messuti (1998): “Hysterothecia without subiculum, predominantly solitary to gregarious, erumpent to superficial at maturity, slightly striate, ellipsoidal with rounded extremes (= cymbiform) to sub-globose, usually straight, unbranched, 0.3 – 1.21 x 0.13 – 0.44 mm. Pseudoparaphyses hyaline, abundant, septate, flexuous, branched, narrow up to 1 μm in dia. Asci cylindric-claviform, octospored, 84 – 151 x 13 – 20 μm. Ascospores hyaline, biseriate to triseriate, transversely uniseptate, constricted at the mid-septum, fusiform, with rounded extremes, both cells approximately of the same length, the upper cell slightly broader than the lower cell, narrowly ovate, with two droplets in each cell, at 30 – 43 x 4 – 9.8 μm.” Collected from Chusquea culeou (Colihuetium) from Argentina. Related to species of: Glonium caucasicum (Rehm) Zogg, G. costesi Speg. & G. hysterinum Rehm. See table in Lorenzo & Messuti (1998).

3. Ascospores about 45 x 9 μm:

= Glonium hysterinum Rehm

Rehm. 1898. Hedwigia 37: 298. Rare. Neotropics. Collected on old wood in Southern Brazil.

II. Asci ovoid, +/- cylindrical; ascospores borne in upper portion of ascus (not evenly distributed), unevenly regimented; ascospores (12) 13 – 15 x 6 – 7 μm:

= Glonium finkii (Petrack) Lohman

Lohman, M.L. 1937. Bull. Torr. Bot. Club 64: 66. Collected from old wood in Puerto Rico.

Psiloglonium Finkii Petrak, 1923

From Lohman (1937): “Hysterothecia deep seated on tangential surface of the wood & not easily separable, 1 – 2.5mm long, flattened , stromatic, with poorly developed lateral walls hyaline below; Paraphyses delicate, branched above, forming a dark granular epithecium. Asci broad saccate, tapering to a short stalk, thick walled at the apex. Ascospores irregularly grouped in upper part of the ascus, elliptic oblong, the upper cell broad with rounded end & the lower narrower, measuring (12) 13-15 μm long, 6-7 μm wide in the upper cell, (3.5) 4-5 in the lower.”

1. Hysterothecium branched, irregular, “graphoid”. Without subiculum or sitting upon a good development of subiculum on the substrate. Ascospores oblong to spindle-shaped; upper cell pear-shaped, constricted at septum; both ends acuminate. Ascospores (13) 15 – 18 (21) x (3) 5 – 6 μm:

= Glonium graphicum (Fries) Duby

Duby. 1862. Mem. Soc. Phys. et Hist. Nat. 16: 47. Rare, Old wood & bark of Pinus & Juniperus. Europe (Germany).

Hysterium graphicum Fries, 1823
Hysterium contortum Ditm. (in Sturms Fl.)

2. Hysterothecium mostly star-shaped or stellate in overall appearance, with branched & laterally anastomosing hysterothecia. Distinct creeping appearance. Mostly associated with a well-developed subiculum. Ascospores oblong to spindle-shaped; upper cell pear-shaped, constricted at septum; both ends acuminate. Ascospores (18) 21 – 26 (28) x (4) 5 – 6 (7) μm:

= Glonium stellatum Muhl. : Fr.

Fries, E. 1823. Syst. myc. II: 595. Quite rare. Collected from old wood of Quercus. North America.

Glonium accumulatum Schw., 1834
Solenarium byssoideum Sprengel, 1827
Solenarium Muhlengergii Kunze, 1817

Lohman (1937): “Hysterothecia superficial, variable in length, following progressively on dichotomously branched & often radiating hyphal str&s & forming irregular, labelliform or stellate composites seated upon a conspicuous brown felt-like subiculum or a black crust, single, merged or overlapping. Asci cylindric, 70-90 (100) x 8-10 (12) μm. Ascospores biseriate, elliptic fusiform, slightly curved, constricted at the septum (occasionally as much as four-celled when arrested in germination), measuring (18) 20-24 (28) x (4) 5-6 (7) μm. Pycnidia Sphaeronaema byssoideum Lohman (1933), common on black crust. More common northward, especially in Appalachians on oak & chestnut. In it’s ascospores, G. stellatum must be grouped with G. simulans & G. clavisporum; in it’s pycnidia with G. abbreviatum.”

Ellis & Everhart (1892): “Subiculum effused, brownish black , indeterminate, two or more centimeters broad, composed of slender, branching, interwoven fibers. Hysterothecia adnate to the subiculum, anastomosing, radiating & forming patches 1 -2 inches across, the branches crowded so as to entirely over the subiculum, narrowly cleft, lips mostly closed. Asci cylindrical, 75-80 x 10 μm, short stipitate, paraphysate. Ascospores eight in ascus, overlapping sub-biseriate, fusoid, hyaline, uniseptate & constricted at the septum, 20-22 x 5-6 μm. On rotten wood, northern USA & Canada, south to the Carolinas; probably to be met with in other parts of the country. The large patches of radiate hysterothecia are generally made up of severl smaller, orbicular patches, 3-4mm dia., confluent at their adjacent margins & presenting a very neat appearance.”

3. Hysterothecium borne within a crust-like stroma; subiculum embedded in stroma. Ascospores spindle-shaped with upper cell slightly swollen & larger than lower cell. Ascospores 24 – 28 x 5 – 6 μm:

= Glonium compactum Kern

Kern. 1959. Ber. Schweiz. Bot. Ges. 69: 277. Rare. Tropical. Collected from Ivory Coast Africa on Strychnos aculeate. Also from South Africa.

Lee & Crous (2003): “Ascomata hysterothecioid, subglobose to globose in vertical section, solitary to gregarious, immersed, straight to flexuous, rarely branched (!), ellipsoidal withy pointed extremes, opening by a depressed longitudinal slit, 0.3 – 0.6mm long, 0.2 – 0.3mm wide. Peridium consisting of two layers, up to 50 μm in dia., pseudoparenchymatous, carbonaceous, outer layer dark, inner layer hyaline, compressed. Pseudoparaphyses cellular, hyaline, septate, flexuous, branched, 0.5 – 1.5 μm wide. Asci bitunicate, clavate, stipitate, octosporous, 65 – 69 x 12 - 15 μm. Ascospores (24-) 26 – 27 (-30) x (4-) 5 – 6 (-7) μm. fusiform, often curved, hyaline, with one transverse septum, slightly constricted at the septum. The first record of the species from Africa was on Strychnos aculeate (Loganiaceae), a host on which it was rarely encountered (Zogg, 1962). This species differs from most Glonium species, which have erumpent to almost superficial ascomata with immersed bases, by having ascomata that are deeply imbedded in the host substratum. The present species was the most common Glonium species encountered during the survey of Fynbos, & was collected from Restios only.” Note: Zogg (1962) figures a very different hysterothecial arrangement from that described here (Lee & Crous, 2003). Zogg illustrates hysterothecia embedded in a stroma, with subiculum subtending & intertwined within stroma. Closest affinities with G. stellatum. Whereas, Lee & Crous state that hysterothecia are immersed in the substrate & not borne in a stroma. Note: in Zogg (1962) Figs. 8 & 9 reversed. G. compactum = Fig. 8.

4. Hysterothecium lirelliform, laterally-anastomosed, branched-elongate, on soil, saxicolous or lignicolous (Tasmania); Ascospores hyaline, constricted at the septum, apices pointed, (15-) 16 – 17 x 6 – 7 μm:

= Glonium circumserpens (Nyl.) Kantvilas & Coppins

Kantvilas & Coppins. 1997. Lichenologist 29: 525-531.

Kantvilas & Coppins (1997): “Ascomata lirelliform, mostly 1 – 3 (-4 – 5) mm long, 0.15 – 0.3mm wide, 0,35 – 0.5mm tall, black, often glossy, very brittle, superficial over the substratum (soil, rocks, lignum), simple, occasionally branched, straight or slightly curved, dispersed, crowded & sme3times overlapping or, most commonly, laterally contiguous in radiating, +/- stellat clusters up to 10 – 20mm wide; lips convergent to a very narrow slit or fused. Exciple in section black, completely carbonized & very brittle, 20 – 80um thick at the sides, 50-100um thick at the base, unchanged or very weakly olive in KOH, unchanged or weakly reddish brown in HNO3, consisting of irregular, connate cells, 4.5 – 11.5um wide, becoming colourless or pale brown in the upper part & merging into a hypothecium up to c. 55um thick. Hymenium colorless, 95 – 150um thick, I minus with or without pretreatment in KOH. Asci eight-spored, narrow cylindrical, 85 – 120um long, 11 – 15 um wide, non-amyloid. Paraphyses (paraphysoids) separating easily, serpentine, sparsely branched and anastomosing, 1 – 2 (-2 – 5)um thick; apices blunt, neither pigmented nor swollen. Ascospores hyaline, lacking a perispores, thin walled, one-septate, noticeably constricted at the septum, sometimes with apices rather pointed: (15-) 16 – 17 x 6 – 7um.” Potentially facultatively licheninzed; hyphae in associated with green algal cells on substrate. Highly unusual member of the Hysteriaceae, it is both terricolous and saxicolous, as well as lignicolous. Collected from Tasmania.

Glonium circumserpens (Nyl.) Kantvilas & Coppins, courtesy of Dr. Gintaras Kantvilas, of the Tasmanian Herbarium,
Hobart, Tasmania. Note the laterally anastomosed hysterothecia and the accuminate ascospores; also the central
sulcus that runs along the long axis of the fruitbody. Subiculum present but not visible in these figures.