The genus Psiloglonium Höhn.
von Höhnel, Annales Mycologici 16: 145 (1918)
by Eric W.A. Boehm
The genus Psiloglonium (Clade B).
A-D. Psiloglonium simulans (ANM 1557 [BPI 879803], USA).
E-H. Psiloglonium clavisporum (GKM 344A [BPI 879801], Kenya).
I-M. Psiloglonium lineare (ANM 117 [ILLS], USA).
N-Q. Psiloglonium araucanum (ANM 42 [ILLS], USA).
Scale bar (habitat) = 500 μm; Scale bar (spores and asci) = 10 μm.
Fig. 8 from Boehm et al. 2009b.
A discussion of the genus Psiloglonium (von Höhnel 1918; Petrak 1923 a, b) by necessity must begin with the genus Glonium. This is because Zogg (1962) synonymised a number of species under the genus Glonium that were originally classified in Psiloglonium by von Höhnel (1918) and Petrak (1923a, b). Both Psiloglonium and Glonium possess hyaline to yellow didymospores, somewhat constricted at the septum, with obtuse or acuminate ends, typically with cells unequal in size, borne in hysterothecia. Von Höhnel (1918) was the first to view the genus Glonium as comprised of two distinct morphological types, and stressed the importance of subicula, using it to divide the genus, at first, into two subgenera, Glonium and Psiloglonium, and, further in the same article, into two separate genera, with or without subicula, respectively. Petrak (1923a) recognised that von Höhnel (1918) had established the genus Psiloglonium, both at sub-generic and generic rank, but it was Petrak (1923a) who explicitly designated the type species for Psiloglonium as P. lineare (Fr.) Petr. (Fig. 8I-M), retaining G. stellatum Muhl. (Fig. 12A-E) as the type species for the genus Glonium sensu von Höhnel (1918). Petrak (1923a, b) eventually placed a number of species in Psiloglonium, all subsequently transferred to Glonium by Zogg (1962). Müller & von Arx (1950) originally accepted the genus Psiloglonium, but later reduced it to a synonym of Glonium (von Arx & Müller 1975). Lohman (1933a, 1937) also did not support Psiloglonium, based on the observation that similar anamorphs were shared between species of the two subgenera. Barr (1987), was the only modern author to retain the genus Psiloglonium, as distinct from the subiculate Glonium.
Although von Höhnel (1918) and Petrak (1923a, b) both stressed the importance of subicula as a major morphological distinction between Psiloglonium and Glonium, Zogg (1962) noted that some species previously classified as Psiloglonium by Petrak (1923a) do in fact possess subicula on occasion (e.g., P. lineare). Zogg (1962) further noted an additional two species that were occasionally associated with subicula, namely G. pusillum H. Zogg and G. graphicum (Fr.) Duby, stating: “…ohne Subiculum oder auf ziemlich deutlichem Subiculum sitzend…” Hence, Zogg (1962) considered subicula not to be a synapomorphic character state, and transferred those species previously classified by Petrak (1923 a, b) in Psiloglonium [e.g., P. lineare, P. microspermum Höhn., P. ruthenicum (Petr.) Petr., and P. finkii Petr.] to the genus Glonium. Although Zogg (1962) did not support Psiloglonium, he did in fact recognise three distinct morphological forms within his concept of Glonium, two of which (Types I and II) we incorporate in Psiloglonium, the third (Type III) forming the basis for the Gloniaceae (Boehm et al. 2009). Zogg (1962) arranged the species of Glonium based on (1) didymospore shape: spore apices obovoid to rounded (Type I) versus spores fusiform with acuminate apices (Type II and III); and (2) the degree of complexity surrounding the architecture of the hysterothecia, simple, linear, solitary to gregarious (Types I, II) versus complex bifurcating, laterally anastomosing to form flabelliform pseudostellate composites, sometimes associated with a thin stromal crust (Type III). Thus, the genus Glonium sensu Zogg (1962) was comprised of two groups of species, one with obovoid to rounded spores apices borne in regular hysterothecia (Type I) versus those with acuminate spore apices borne in complex bifurcating or modified hysterothecia (Type III). Species belonging to Type II possess fruitbodies of Type I, but spores of Type III; the assumption was that they constituted an intermediate, perhaps transitional, morphological group. This, then, de-emphasised the presence or absence of subicula per se, as stressed by von Höhnel (1918) and Petrak (1923 a, b). Nevertheless, Zogg (1962) maintained all three types within the genus Glonium. Molecular data presented here (see below), indicate that Types I & II are closely related, with Type III forming a distant clade in the Gloniaceae (Boehm et al. 2009).
Type I: This type is characterised by hysterothecia that may be solitary to gregarious, erumpent to entirely superficial, navicular to linear to highly flexuous, even triradiate, sometimes arranged in parallel orientation and confluent linearly to some degree, but never dichotomously branched, or associated with a stromal crust, as found in the Gloniaceae (Type III). These species correspond to Psiloglonium sensu von Höhnel (1918). Here, the didymospores are relatively small, hyaline, and have at least one, if not both ends, obovoid to obtuse (Type I), rather than acuminate (Types II and III). Zogg (1962) recognised five species, listed here by increasing ascospore length: Glonium abbreviatum (Schwein.) M.L. Lohman, G. pusillum H. Zogg, G. lineare, G. chambianum A.L. Guyot, and G. curtisii (Duby) M.L. Lohman. Barr (1975) transferred the last species to Ostreichnion, as O. curtisii (Duby) M.E. Barr in the Mytilinidiaceae, since transferred to the Hysteriaceae (Boehm et al. 2009). A sixth species, G. finkii (Petr.) M.L. Lohman, was included by Zogg (1962), based on ascospore shape, but placed apart in the key due to the unusual arrangement of the ascospores within the upper part of the ascus (Lohman 1937).
Psiloglonium lineare was previously reinstated within the Hysteriaceae, listing G. lineare as a synonym (Boehm et al. 2009). Here we also reinstate Psiloglonium finkii Petr. An additional two species are included in Type I, namely G. simulans W.R. Gerard and G. clavisporum Seaver, synonymised by Zogg (1962) under G. lineare, but earlier recognised by Lohman (1932a, 1937) to be distinct from G. lineare. Boehm et al.(2009) proposed new combinations for these taxa, based on morphological as well as molecular data, as P. simulans (W.R. Gerard) E.W.A. Boehm, C.L. Schoch & J.W. Spatafora (Fig. 8A-D) and P. clavisporum(Seaver) E.W.A. Boehm, C.L. Schoch & J.W. Spatafora (Fig. 8E-H). To these species can also be added G. sasicola N. Amano from Japan, the first report of a gelatinous sheath in the genus (Amano 1983). In this same publication Amano (1983) proposed an additional new species, G. macrosporum N. Amano, also from Japan. The spore measurments were given as 13.1-16.8 x 4.0-5.6 μm, nearly identical to those of P. simulans at (10-)14-16(-18) x (4.5-)5-6 μm (Lohman 1937). Moreover, the illustrations given by Amano (1983) match closely those given by Lohman (1932a) for P. simulans. We therefore synonymise G. macrosporum under P. simulans.
More recently, Lorenzo & Messuti (1998), in a reappraisal of the type specimens collected by Spegazzini and Hennings from Argentina and Chile, have reinstated Glonium costesii Speg. In a later publication, Messuti & Lorenzo (2007) synonymised G. costesii under the earlier epithet G. ephedrae Henn. With spore measurements of 26-35 x 8-15 μm, G. ephedrae possesses the largest spores in Type I. In the same publication, Messuti & Lorenzo (2007) also accepted two additional species, G. chilense Speg. and G. uspallatense Speg., previously considered by Zogg (1962) to be doubtful species. The spores of G. chilense measure 15-16 x (5-)7-8 μm, which places it very close to P. lineare, the latter with slightly smaller spores, (10-)12-14(-18) x (4-)5-7(-8) μm (Zogg 1962). However, G. chilense has almost identical ascomatal and spore measurements as P. simulans, given above. We therefore synonymise G. chilense with the earlier name G. simulans, as P. simulans. For G. uspallatense, Messuti & Lorenzo (2007) gave spore measurements of 18-24 x 10-12 μm, intermediate between G. chambianum, (14-)16-18(-21) x (6-)8-9(-10) μm (Zogg 1962), and G. sasicola, 25-32 x 5-8 μm (Amano 1983).
Recently, Mugambi & Huhndorf (2009) proposed a new genus, Anteaglonium, outside of the Hysteriales but within the Pleosporales, to accommodate A. abbreviatum (Schwein.) Mugambi & Huhndorf (Fig. 9A-E), A. globosum Mugambi & Huhndorf (Fig. 9F-I), A. parvulum (W.R. Gerard) Mugambi & Huhndorf (Fig. 9J-M), and A. latirostrum Mugambi & Huhndorf (Fig. 9N-R). The first three species are characterised by hyaline didymospores that belong to Type I, as defined by Zogg (1962), and are less than 8 μm in length. The fourth species, A. latirostrum, belongs to Type II (see below), with longer spores. Although phylogenetically unrelated to Psiloglonium, these species share a similar morphology and thus are included in the key below.
Type II: This type is characterised by relatively large didymospores, distinctly fusoid in outline, prominently constricted at the septum, and with acuminate apices. Zogg (1962) recognised two species, namely Glonium caucasicum (Rehm) H. Zogg and the much larger-spored, neotropical G. hysterinum Rehm, to which can be added the newly described G. colihuae Lorenzo & Messuti, on Chusquea culeou from Argentina (Lorenzo & Messuti 1998). Glonium caucasicum has recently been synonymised under the earlier name G. araucanum Speg. by Messuti & Lorenzo (2007), based on a comparison of the type specimen of G. caucasicum to Spegazini’s earlier type of G. araucanum from Chile.
Type III: This type corresponds to von Höhnel’s (1918) and Petrak’s (1923a, b) circumscription of the genus Glonium, and includes species with fusiform spores, with acuminate apices, typically producing complex laterally anastomosing hysterothecia, forming stellate composites, usually with prominent subicula, with or without stroma. Zogg (1962) included the type, G. stellatum Muhl. (Fig. 12A-E), G. compactum Kern, and G. graphicum (Fr.) Duby, the later sometimes variably associated with subicula. Zogg (1962) also stated that G. compactum possesses a subiculum, much like G. stellatum, and with similar spore size, but whereas hysterothecia in G. stellatum are merely seated on the subiculum, in G. compactum the hysterothecia are embedded in and arise from a thin stromal crust, which is itself seated on subicula. Recently, a fourth species was added, based on molecular evidence (Boehm et al. 2009), namely G. circumserpens (Nyl.) Kantvilas & Coppins (Fig. 12F-H), from Tasmania (Kantvilas & Coppins 1997).
Recent sequence data (Boehm et al. 2009a, b; Mugambi & Huhndorf 2009), clearly indicate that the genus Glonium sensu Zogg (1962) actually comprises three entirely unrelated lineages within the Pleosporomycetidae, one within the Hysteriaceae and two forming clades outside of the family. The first lineage corresponds to Psiloglonium sensu von Höhnel (1918), and forms a highly supported monophyletic clade in this study (Clade B in Fig. 1). This clade includes: Psiloglonium clavisporum, with four single-ascospore isolates from New Jersey, the United States, (CBS 123338 / BPI 878726, CBS 123339 / BPI 878727, CBS 123340 / BPI 878728 and CBS 123341 / BPI 878729), and two from Kenya (GKM 344A / BPI 879801, GKM L172A in EA), P. simulans, with two isolates from the United States, one from Michigan (CBS 206.34), deposited in 1934 by M.L. Lohman, and a more recent collection from Tennessee (ANM 1557 / BPI 879803), and, lastly, P. araucanum, with three isolates from South Africa, two from Kirstenbosch (CBS 112412 / PREM 57570, CMW 18760/ PREM 57569) and one from Jonkershoek (CMW 17941 / PREM 575566). Psiloglonium clavisporum and P. simulans belong to Type I, whereas P. araucanum belongs to Type II. Both are phylogenetically related and reside in Clade B (Fig. 1). Recently, a second lineage has been shown to be associated with the Pleosporales, now accommodated in the new genus Anteaglonium(Mugambi & Huhndorf 2009), for which we include six accessions representing four species (Table 1). The third lineage corresponds to Glonium (Type III), in the Gloniaceae (Boehm et al. 2009), for which we have included four isolates, representing two species (Table 1). We treat here all species of Glonium sensu Zogg (1962), belonging to Types I and II, outside of Anteaglonium, as belonging to Psiloglonium. Since the generic name Glonium is reserved for species in the Gloniaceae (Boehm et al. 2009), we propose eight new combinations for the genus Psiloglonium. In addition to the 12 currently recognised species in Psiloglonium, the following key also includes entries for the unrelated Gloniaceae, Anteaglonium and Ostreichnion curtisii.
Key to the species of Psiloglonium and Anteaglonium
- Asci ovoid, +/- cylindrical; ascospores borne in the upper portion of the ascus, not evenly distributed; ascospores (12-)13-15 x 6-7 μm; Puerto Rico → Psiloglonium finkii Petr.
- Asci typically cylindrical to club-shaped; ascospores in one row or distichous in the asci, but always regularly arranged for its full length → 2
- Ascospores obovoid, with at least one, often both, ends obtuse, typically with upper cell larger, +/- constricted at the septum (Type I) → 3
- Ascospores fusiform (i.e., spindle-shaped), with both ends acuminate, usually constricted at the septum (Types II and III) → 14
- Ascospores small, 8 μm or less in length (Anteaglonium, in part) → 4
- Ascospores longer than 8 μm (PsilogloniumType I) → 6
- Ascospores 6-8 x 2.5-3 μm; hysterothecia with apices acuminate, but not associated with a darkened crust; no KOH-soluble pigments; New Zealand, East Africa, North America → Anteaglonium parvulum (W.R. Gerard) Mugambi & Huhndorf
Note: A. parvulum lies within the Pleosporales (Mugambi & Huhndorf 2009).
- Not with the above combination of characters → 5
- Ascospores (5-)6-7(-8) x 2-3(-3.5) μm (as in A. parvulum); but hysterothecia with apices truncated, and associated with a darkened crust (tending to darken the substratum); minute amounts of soluble pigment in KOH (easily missed); Europe, East Africa, North America → Anteaglonium abbreviatum (Schwein.) Mugambi & Huhndorf
Note: A. abbreviatum lies within the Pleosporales (Mugambi & Huhndorf 2009).
- Ascospores 6-7 x 2-3 μm (as in A. parvulumand A. abbreviatum); but hysterothecia globose with roughened walls, an indistinct slit, and associated with sparse, short subicula, and also with short tomentum on the walls of the ascomata; like A. abbreviatumalso associated with a darkened crust on substrate; producing a strong green soluble pigment in KOH; eastern and mid-western North America → Anteaglonium globosum Mugambi & Huhndorf
Note: A. globosum lies within the Pleosporales (Mugambi & Huhndorf 2009).
- Ascospores (9-)10-12(-13) x 4-5(-6) μm; Europe, Africa, North America → Psiloglonium pusillum (H. Zogg) E.W.A. Boehm & C.L. Schoch
- Ascospores slightly larger → 7
- Ascospores (10-)12-14(-18) x (4-)5-7(-8) μm; ascomata +/- confluent laterally, in parallel rows, semi-immersed to erumpent; cosmopolitan → Psiloglonium lineare (Fr.) Petrak
- Ascospores similar in length; ascomata not confluent laterally, usually entirely superficial → 8
- Ascospores (10-)14-16(-18) x (4.5-)5-6 μm; cosmopolitan → Psiloglonium simulans (W.R. Gerard) E.W.A. Boehm, C.L. Schoch & J.W. Spatafora
- Ascospores slightly larger → 9
- Ascospores (15-)16-18(-20) x 5-6(-7) um; Sporidesmium stygiumanamorph usually present; North and South America, Africa → Psiloglonium clavisporum (Seaver) E.W.A. Boehm, C.L. Schoch & J.W. Spatafora
- Ascospores slightly larger in length and breadth → 10
- Ascospores (14-)16-18(-21) x (6-)8-9(-10) μm; North America, Europe and North Africa → Psiloglonium chambianum (Guyot) E.W.A. Boehm & C.L. Schoch
- Ascospores slightly larger → 10
- Ascospores 18-24 x 10-12 μm; Argentina → Psiloglonium uspallatense (Speg.) E.W.A. Boehm & C.L. Schoch
- Ascospores slightly larger → 12
- Ascospores 25-32 x 5-8 μm, with a gelatinous sheath; Japan → Psiloglonium sasicola(N. Amano) E.W.A. Boehm & C.L. Schoch
- Ascospores slightly larger → 13
- Ascospores 30-35 x 8-15 μm; Chile → Psiloglonium ephedrae(Henn.) E.W.A. Boehm & C.L. Schoch
- Ascospores (59-)62-68(-76) x 13-15 μm; North & South America → Ostreichnion curtisii(Duby) M.L. Lohman
Note: The genus Ostreichnion, previously placed in the Mytilinidiaceae, has been transferred to the Hysteriaceae (Boehm et al. 2009).
- Hysterothecia usually borne in/on subicula, typically bifurcated, forming radiating flabelliform or pseudo-stellate composites, with or without a stroma (Type III) → Gloniaceae
Note: A key to the species of the Gloniaceae is provided under that family.
- Hysterothecia not bifurcated, forming radiating flabelliform or pseudo-stellate composites, nor with a stroma → 15
- Ascospores less than 30 μm long → 16
- Ascospores more than 30 μm long → 17
- Ascospores (19-)22-25(-27) x (6-)7-9(-10) μm, both ends acuminate, with a prominent septal constriction; cosmopolitan (Type II) → Psiloglonium araucanum (Speg.) E.W.A. Boehm, S. Marincowitz & C.L. Schoch
- Ascospores 22-28 x 4-6 μm, acuminate, 1-septate, hyaline and with a mucilaginous sheath when young, but acquiring additional septa and pigmentation with age, to become 3-5-septate and pale brown at maturity; Kenya → Anteaglonium latirostrum Mugambi & Huhndorf
Note: A. latirostrum lies within the Pleosporales (Mugambi & Huhndorf 2009).
- Ascospores 30-43 x 4-9.8 μm; Argentina (Type II) → Psiloglonium colihuae (Lorenzo & Messuti) E.W.A. Boehm & C.L. Schoch
- Ascospores about 45 x 9 μm; Brazil (Type II) → Psiloglonium hysterinum (Rehm) E.W.A. Boehm & C.L. Schoch
Psiloglonium pusillum (H. Zogg) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515327.
Basionym: Glonium pusillum H. Zogg, Beitr. Kryptfl. Schweiz 11(3): 62 (1962).
Notes: Zogg (1962) described this species as G. pusillum from Juniperus phoenicea and Pinus sylvestris from Southern France, noting that it was quite rare. Zogg (1962) stated that this species may or may not be associated with a subiculum, and hence was one of the factors behind his transfer of Petrak’s (1923a, b) Psiloglonium species to Glonium. Psiloglonium pusillum has ascospores only slightly larger than those of P. abbreviatum, measuring (9-)10-12(-13) x 4-5(-6) μm. Lee & Crous (2003) also identified this fungus from Proteaceae and Restionaceae in South Africa, and Sivanesan & Hsieh (1989) reported it from Taiwan. It has also been found in North America (Boehm, unpubl. data).
Rare. Southern French Maritime Alps. Collected from old needle scales protuberances of living Juniperus phoenicea & from the bark of living Pinus silvestris (Zogg 1962). But also collected from South Africa by Lee & Crous (2003).
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).”
Psiloglonium chambianum (Guyot) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515320.
Basionym: Glonium chambianum Guyot, Ann. Serv. Bot. Agric. Tunisie 28: 90 (1955).
Notes: Originally from North Africa, on Lonicera implexa (Caprifoliaceae), the fungus has since been reported from the Proteaceae in South Africa (Lee & Crous 2003). Zogg (1962) gave the spore measurements for G. chambianum as (14-)16-18(-21) x (6-)8-9(-10) μm, whereas Lee & Crous (2003) gave slightly larger measurements, (18-)20-21(-23) x (4-)5-6(-7) μm. Spores ellipsoid to oblong, with upper cell broader than the lower, and with an obovoid, obtuse apex. Psiloglonium chambianum possesses larger spores than P. lineare, P. simulans, and P. clavisporum, but smaller than P. uspallatense.
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 ofLeucospermum 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.”
Psiloglonium uspallatense (Speg.) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515321.
Basionym: Glonium uspallatense Speg., Anal. Mus. Nac. Buenos Aires, Ser 3, v. 12, 19: 436 (1909).
Notes: Zogg (1962) listed the species a “doubtful”, but Messuti & Lorenzo (2007) reinstated G. uspallatense after locating the original holotype material. They gave the spore measurements as 18-24 x 10-12 μm, placing it intermediate between P. chambianum and P. sasicola.
Psiloglonium sasicola (N. Amano) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515322.
Basionym: Glonium sasicola N. Amano, Trans. Mycol. Soc. Japan 24: 287 (1983).
Notes: Amano (1983) described this species from dead culms of Sasa sp. (Bambusaceae) in Japan. The ascospore measurements were given as 25-32 x 5-8 μm, with a rounded apical cell, placing it between P. uspallatense and P. ephedrae. Amano (1983) further reported that ascospores of this species are associated with a gelatinous sheath, previously not known among these didymospored fungi.
Japan. Collected from dead culm of Sasasp.
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.”
Psiloglonium ephedrae (Henn.) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515323.
Basionym: Glonium ephedrae Henn., Öfvers. K. Vet. Akad. Förhandl. 2: 328 (1900).
= Glonium costesi Speg., Bol., Acad. Nci. Ci., Córdoba 25: 78-79 (1921).
Notes: Messuti & Lorenzo (2007) reinstated G. ephedrae with the synonym G. costesi, after locating and comparing original type materials. Psiloglonium ephedrae possesses very large didymospores, measuring 26-35 x 8-15 μm, the upper cells broadly ovate. It has been collected from Ephedra andicola, and, as G. costesi, from Proustia pyrifolia in Chile.
Psiloglonium hysterinum (Rehm) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515324.
Basionym: Glonium hysterinum Rehm, Hedwigia 37: 298 (1898).
Notes: Rehm (1898) originally described a species of Glonium from Southern Brazil with large fusiform didymospores, prominently constricted at the septum, and with acuminate spore apices (“Enden zugespitzt“). The spore measurements were given as 45 x 9 μm.
Psiloglonium colihuae (Lorenzo & Messuti) E.W.A. Boehm & C.L. Schoch, comb. nov. MycoBank MB515325.
Basionym: Glonium colihuae Lorenzo & Messuti, Mycol. Res. 102: 1104 (1998).
Notes: Lorenzo & Messuti (1998) described a new species on culms of Chusquea culeou from the Argentine Nothofagus rainforests. The spore measurements were given as 30-43 x 4-9.8 μm, and, although the spores are fusiform in outline, they possess moderately acuminate apices. In comparing this species to other acuminate-spored species of Glonium, the authors noted that the greatest degree of similarity was with the slightly smaller-spored G. caucasicum.
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. Indeed it falls into this part of the key, belonging to the second clade (see above). Also see table in Lorenzo & Messuti (1998).
Psiloglonium araucanum (Speg.) E.W.A. Boehm, S. Marincowitz & C.L. Schoch, comb. nov. MycoBank MB515326, Fig. 8N-Q.
Basionym: Glonium araucanum Speg., Rev. Fac. Agron. Veter. La Plata 6: 110 (1910).
= Gloniella caucasica Rehm, Vestn. Tiflissk. Bot. Sada 25:12 (1912).
≡ Glonium caucasicum (Rehm) H. Zogg, Beitr. Kryptfl. Schweiz 11(3): 67 (1962).
Boehm et al. 2009b: “Messuti & Lorenzo (2007) transferred Glonium caucasicum to G. araucanum, after examining the types for both species. Previously, Zogg (1962) had transferred Gloniella caucasica to Glonium. Here we transfer G. araucanum to Psiloglonium. This taxon possesses fusiform spores with highly acuminate apices. Messuti & Lorenzo gave the spore measurements as 22-28 x 8-10 μm, whereas Zogg (1962) gives them as (19-)22-25(-27) x (6-)7-9(-10) μm. Although originally European in distribution (Zogg 1962), the taxon has subsequently been collected from South (Messuti & Lorenzo 2007) and North America (Boehm unpubl. data), and from South Africa (Lee & Crous 2003). Lee & Crous (2003) identified a series of isolates from South Africa on the Restionaceae as Glonium compactum (CBS 112412, CMW 18760, CMW 17941). However, in their study they did not note the presence of subicula, nor a stromal crust. These features were stressed for this taxon by Zogg (1962). These same isolates were used in Boehm et al. (2009), and were shown to associate, with high branch support, with two species of Psiloglonium, P. clavisporum and P. simulans, distant from the other species of Glonium surveyed (e.g., G. stellatum and G. circumserpens). Thus, a new combination was proposed, Psiloglonium compactum. However, it is now realised that this new combination was made in error and is hereby retracted. It must be concluded that the South African isolates (Lee & Crous 2003) were not G. compactum, due to the absence of subicula and stroma, but rather, we suspect, the cosmopolitan P. araucanum, which has similar, but slightly smaller, fusiform acuminate didymospores. Lee & Crous (2003) give the ascospore measurements for the South African “G. compactum” as (24-)26-27(-30) x (4-)5-6(-7) μm, which matches closely those given above for P. araucanum. Furthermore, the illustrations in Lee & Crous (2003) closely match P. araucanum, and not those of G. compactum, as given by Zogg (1962). If we are correct in assuming that the South African isolates used in Boehm et al. (2009) are in fact P. araucanum (Type II) and not G. compactum (Type III), then this would provide a high degree of support for the inclusion of species with acuminate spore apices, belonging to Type II, in the genus Psiloglonium, along with species with obtuse spore apices, belonging to Type I (e.g., P. simulans and P. clavisporum). A reanalysis of the original South African herbarium specimens from which the sequences were derived (PREM 57570, PREM 57569, PREM 57566), by S. Marincowitz, has confirmed that they do indeed correspond to P. araucanum and not to G. compactum. Molecular data thus supports the association of Types I and II within the genus Psiloglonium”.
Psiloglonium finkii Petrak
= Glonium finkii (Petrak) Lohman, M.L. 1937. Bull. Torr. Bot. Club 64: 66.
Collected from old wood in Puerto Rico.
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.”
Psiloglonium lineare (Fr.) Petrak
Hysterium lineare Fries, 1823
Psiloglonium lineare (Fr.) Petrak, 1923
Glonium clavisporum Seaver, 1925a [Note: Reinstated as P. clavisproum (Seaver) Boehm, Schoch & Spatafora]
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 W.R. Gerard, 1876 [Note:Reinstated as P. simulans (W.R. Gerard) Boehm, Schoch & Spatafora]
Glonium lineare (Fr.) De Not. 1847
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.
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, and 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.”
Psiloglonium simulans (W.R. Gerard) E.W.A. Boehm, C.L. Schoch & J.W. Spatafora
Mycological Research 113: 469. 2009. MycoBank No.: MB 512018
Basionym: Glonium simulans W.R. Gerard, Bull. Torrey Club 6: 78. 1876.
Boehm et al. (2009a): “Zogg (1962) synonymised Glonium simulans W.R. Gerard under G. lineare (Fr.) De Not. However, Lohman (1937) presented evidence as to why G. simulans (= P. simulans) should be considered distinct from G. lineare (= P. lineare). Although ascospores in both species are of similar length, the upper cell of the ascospore in P. lineare is broadly rounded, whereas the upper cell in P. simulans is more acuminate and approaches the appearance observed in the genus Glonium sensu von Höhnel (1918). The ascospores of both species differ markedly in outline as illustrated by Lohman (1932a). P. lineare possesses a pycnidial Plenodomus anamorph, but no anamorph has ever been recorded for G. simulans (Lohman 1937). Lastly, hysterothecia of P. lineare are confluent laterally to some degree, forming parallel, linear flat arrays, and are often partly submerged into the substrate. In contrast, hysterothecia of P. simulans are entirely superficial, never forming lines, but ofter assume flexuous configurations on the substrate, as in P. clavisporum. Here we remove the synonymy proposed by Zogg (1962) and reinstate G. simulans in the genus Psiloglonium“.
Lohman (1937) for Glonium simulans Gerard: “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.”
Psiloglonium calvisporum (Seaver) E.W.A. Boehm, C.L. Schoch & J.W. Spatafora
Mycological Research 113: 469. 2009. MycoBank No.: MB 512022
Basionym: Glonium clavisporum Seaver, Mycologia 17(1): 4. 1925.
Boehm et al. (2009a): “Zogg (1962) synonymised Glonium clavisporum Seaver under G. lineare (Fr.) De Not., despite convincing evidence presented by Lohman (1932a, 1937) as to why G. clavisporum (= P. clavisporum) should be considered distinct from G. lineare (= P. lineare). Foremost among these, are differences in the associated anamorph. P. clavisporum possesses a dinstinct dematiaceous anamorph, Sporidesmium stygium Berk. & M.A. Curtis, present among hysterothecia, as well as in culture. Whereas, P. lineare possesses a pycnidial Plenodomus anamorph, in nature and in culture (Lohman 1933a, 1937). Although both P. clavisporum and P. lineare possess similar shaped ascospores, they differ in length: 16 – 20 μm vs 12 – 15 μm, respectively (Lohman 1937). Finally, hysterothecia of P. lineare are confluent laterally to some degree, forming parallel, linear flat arrays, typically not completely erumpent, being sunken to some degree in the substrate. Whereas, hysterothecia in P. clavisporum are superficial, and never anastomose laterally to form linear arrays. Thus, morphological features serve to differentiate P. clavisporum from P. lineare. P. clavisoporum is exceedingly abundant in New Jersey. It can be found growing on the trunks of the Planetree Plantanus occidentalis (e.g., Nomahegan Park, Union Co., Grover Cleaveland Park, Essex Co., Watchung Reservation, Union Co.) The fungus forms irregular black patches on the bark, measuring several centimeters in diameter, and typically displays abundant dematiaceous conidia of the Sporidesmium stygium state. As these colonies develop, in the center emerges the ascigerous stage with irregular hysterothecia, that are gregarious, but rarely fused, often flexuous along their length. Specimens deposited with the CBS and with BPI.
Lohman (1937) for Glonium clavisporum Seaver: “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.”