Pleosporales

Molecular phylogeny

In total, 278 pleosporalean taxa are included in the phylogenetic analysis. These form 25 familial clades in the dendrogram, i.e. Aigialaceae, Amniculicolaceae, Arthopyreniaceae, Cucurbitariaceae/Didymosphaeriaceae, Delitschiaceae, Didymellaceae, Dothidotthiaceae, Hypsostromataceae, Lentitheciaceae, Leptosphaeriaceae, Lindgomycetaceae, Lophiostomataceae, Massariaceae, Massarinaceae, Melanommataceae, Montagnulaceae, Morosphaeriaceae, Phaeosphaeriaceae, Pleomassariaceae, Pleosporaceae, Sporormiaceae, Testudinaceae/Platystomaceae, Tetraplosphaeriaceae, Trematosphaeriaceae and Zopfiaceae (Plate 1). Of these, Lentitheciaceae, Massarinaceae, Montagnulaceae, Morosphaeriaceae and Trematosphaeriaceae form a robust clade in the present study and in previous studies (Schoch et al. 2009; Zhang et al. 2009a, b). We thus emended the suborder, Massarineae, to accommodate them.

Pleosporales suborder Massarineae Barr, Mycologia 71: 948. (1979a). emend.

Habitat freshwater, marine or terrestrial environment, saprobic. Ascomata solitary, scattered or gregarious, globose, subglobose, conical to lenticular, immersed, erumpent to superficial, papillate, ostiolate. Hamathecium of dense or rarely few, filliform pseudoparaphyses. Asci bitunicate, fissitunicate, cylindrical, clavate or broadly clavate, pedicellate. Ascospores hyaline, pale brown or brown, 1 to 3 or more transverse septa, rarely muriform, narrowly fusoid, fusoid, broadly fusoid, symmetrical or asymmetrical, with or without sheath.

Accepted genera of Pleosporales

Acrocordiopsis Borse & K.D. Hyde, Mycotaxon 34: 535 (1989). (Pleosporales, genera incertae sedis)

Generic description

Habitat marine, saprobic. Ascomata seated in blackish stroma, scattered or gregarious, superficial, conical to semiglobose, ostiolate, carbonaceous. Hamathecium of dense, long trabeculate pseudoparaphyses. Asci 8-spored, cylindrical with pedicels and conspicuous ocular chambers. Ascospores hyaline, 1-septate, obovoid to broadly fusoid.

Anamorphs reported for genus: none.

Literature: Alias et al. 1999; Barr 1987a; Borse and Hyde 1989.

Type species

Acrocordiopsis patilii Borse & K.D. Hyde, Mycotaxon 34: 536 (1989). (Fig. 1)

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Ascomata 1–2 mm high × 1.8–3 mm diam., scattered or gregarious, superficial, conical or semiglobose, with a flattened base not easily removed from the substrate, ostiolate, black, very brittle and carbonaceous and extremely difficult to cut (Fig. 1a and b). Peridium 250–310 μm thick, to 600 μm thick near the apex, thinner at the base, comprising three types of cells; outer cells pseudoparenchymatous, small heavily pigmented thick-walled cells of textura epidermoidea, cells 0.6–1 × 6–10 μm diam., cell wall 5–9 μm thick; cells near the substrate less pigmented, composed of cells of textura prismatica, cell walls 1–3(−5) μm thick; inner cells less pigmented, comprised of hyaline to pale brown thin-walled cells, merging with pseudoparaphyses (Fig. 1c, d and e). Hamathecium of dense, long trabeculate pseudoparaphyses, ca. 1 μm broad, embedded in mucilage, hyaline, anastomosing and sparsely septate. Asci 140–220 × 13–17 μm (\( \bar{x} = { 165}.{3 } \times { 15}.{6 }\mu {\text{m}} \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with short pedicels, 15–25(−40) μm long, with a large and conspicuous ocular chamber (Fig. 1f and g). Ascospores 17.5–25 × 12.5–15(−20) μm (\( \bar{x} = { 21}.{5 } \times { 13}.{6 }\mu {\text{m}} \), n = 10), uniseriate to partially overlapping, ovoid or ellipsoidal, hyaline, 1-septate, not constricted at the septum, smooth-walled (Fig. 1h and i).

Anamorph: none reported.

Material examined: INDIA, Indian Ocean, Malvan (Maharashtra), on intertidal wood of Avicennia alba Bl., 30 Oct. 1981 (IMI 297769, holotype).

Notes

Morphology Acrocordiopsis was formally established by Borse and Hyde (1989) as a monotypic genus represented by A. patilii based on its “conical or semiglobose superficial carbonaceous ascomata, trabeculate pseudoparaphyses, cylindrical, bitunicate, 8-spored asci, and hyaline, 1-septate, obovoid or ellipsoid ascospores”. Acrocordiopsis patilii was first collected from mangrove wood (Indian Ocean) as a marine fungus, and a second marine Acrocordiopsis species was reported subsequently from Philippines (Alias et al. 1999). Acrocordiopsis is assigned to Melanommataceae (Melanommatales sensu Barr 1983) based on its ostiolate ascomata and trabeculate pseudoparaphyses (Borse and Hyde 1989). Morphologically, Acrocordiopsis is similar to Astrosphaeriella sensu stricto based on the conical ascomata and the brittle, carbonaceous peridium composed of thick-walled black cells with rows of palisade-like parallel cells at the rim area. Ascospores of Astrosphaeriella are, however, elongate-fusoid, usually brown or reddish brown and surrounded by a gelatinous sheath when young; as such they are readily distinguishable from those of Acrocordiopsis. A new family (Acrocordiaceae) was introduced by Barr (1987a) to accommodate Acrocordiopsis. This proposal, however, has been rarely followed and Jones et al. (2009) assigned Acrocordiopsis to Melanommataceae.

Phylogenetic study

Acrocordiopsis patilii nested within an unresolved clade within Pleosporales (Suetrong et al. 2009). Thus its familial placement is unresolved, but use of the Acrocordiaceae could be reconsidered with more data.

Concluding remarks

Acrocordiopsis, Astrosphaeriella sensu stricto, Mamillisphaeria, Caryospora and Caryosporella are morphologically similar as all have very thick-walled carbonaceous ascomata, narrow pseudoparaphyses in a gelatinous matrix (trabeculae) and bitunicate, fissitunicate asci. Despite their similarities, the shape of asci and ascospores differs (e.g. Mamillisphaeria has sac-like asci and two types of ascospores, brown or hyaline, Astrosphaeriella has cylindro-clavate asci and narrowly fusoid ascospores, both Acrocordiopsis and Caryosporella has cylindrical asci, but ascospores of Caryosporella are reddish brown). Therefore, the current familial placement of Acrocordiopsis cannot be determined. All generic types of Astrosphaeriella sensu stricto, Mamillisphaeria and Caryospora should be recollected and isolated for phylogenetic study.

Aigialus Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85: 699 (1985). (Aigialaceae)

Generic description

Habitat marine, saprobic. Ascomata mostly subglobose in front view, fusoid in sagittal section, rarely subglobose, scattered, immersed to erumpent, papillate, ostiolate, ostiole rounded or slit-like, periphysate. Peridium 2-layered. Hamathecium of trabeculate pseudoparaphyses. Asci 8-spored, cylindrical, pedicellate, with an ocular chamber and conspicuous apical ring. Ascospores ellipsoidal to fusoid, muriform, yellow brown to brown, with terminal appendages.

Anamorphs reported for genus: none.

Literature: Eriksson 2006; Jones et al. 2009; Kohlmeyer and Schatz 1985; Lumbsch and Huhndorf 2007.

Type species

Aigialus grandis Kohlm. & S. Schatz, Trans. Br. Mycol. Soc. 85: 699 (1985). (Fig. 2)

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Ascomata 1–1.25 mm high × 1–1.3 mm diam. in front view, 250–400 μm broad in sagittal section, vertically flattened subglobose, laterally compressed, scattered, immersed to semi-immersed, papillate, with an elongated furrow at the top of the papilla, wall black, carbonaceous, ostiolate, ostiole filled with branched or forked septate periphyses (Fig. 2a). Peridium 70–100 μm thick laterally, up to 150 μm thick at the apex, thinner at the base, comprising two cell types, outer layer composed of small heavily pigmented thick-walled pseudoparenchymatous cells, cells 1–2 μm diam., cell wall 2–5 μm thick, inner layer thin, composed of small hyaline cells (Fig. 2b). Hamathecium of dense, very long trabeculate pseudoparaphyses, 0.8–1.2 μm broad, embedded in mucilage, anastomosing and branching above the asci. Asci 450–640 × 22–35 μm (\( \bar{x} = 505 \times 30\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a long furcate pedicel, 90–180 μm long, with a low truncate ocular chamber and a refractive apical apparatus (to 12 μm wide × 4 μm high) (Fig. 2e and f). Ascospores 75–95 × 15–26 μm (\( \bar{x} = 84.3 \times 17.5\mu m \), n = 10), obliquely uniseriate and partially overlapping, broadly fusoid to fusoid with narrowly rounded ends in front view, flat on one side from side view (14–20 μm thick), yellowish brown, apical cells usually hyaline, muriform, with 14–17(−18) transversal septa, 1–3 longitudinal septa in most cells, slightly constricted at the septa, with a gelatinous cap at each end (Fig. 2c and d).

Anamorph: none reported.

Material examined: BELIZE, Wee-Wee Cay, on submerged wood of roots and branches of Rhizophora mangle L., Mar. 1983, leg. J. Kohlmeyer (NY, J.K. 4332b, isotype).

Notes

Morphology Aigialus was formally established by Kohlmeyer and Schatz (1985) based on its immersed or semi-immersed ascomata with periphysate ostiole, trabeculate pseudoparaphyses, cylindrical and fissitunicate asci, and distinctive muriform ascospores with gelatinous sheath or caps. There are five accepted species in the genus, namely A. grandis, A. mangrovei Borse, A. parvus S. Schatz & Kohlm., A. rhizophorae Borse and A. striatispora K.D. Hyde (Jones et al. 2009). Aigialus was first assigned to the Melanommatales, but its familial status was uncertain (Kohlmeyer and Schatz 1985). Barr (1990b) included Aigialus in Massariaceae based on its conspicuous apical ring in the asci and ascospore characters, and this has subsequently been widely followed (Eriksson 2006; Hawksworth et al. 1995; Kirk et al. 2001; Lumbsch and Huhndorf 2007).

Phylogenetic study The generic type of Aigialus (A. grandis) together with other three marine species, i.e. A. mangrovei, A. parvus as well as A. rhizophorae form a robust clade on the phylogenetic tree. Thus a new family, Aigialaceae, was introduced to accommodate Aigialus together with Ascocratera and Rimora (Suetrong et al. 2009).

Concluding remarks The pleosporalean status of Aigialus has been phylogenetically verified, and the single branch containing Aigialus, Ascocratera and Rimora represents a familial rank of Aigialaceae (Suetrong et al. 2009).

Amniculicola Yin. Zhang & K.D. Hyde, Mycol. Res. 112: 1189 (2008). (Amniculicolaceae)

Generic description

Habitat freshwater, saprobic. Ascomata solitary, scattered, or in small groups, initially immersed, becoming erumpent, to nearly superficial, globose, subglobose to conical, wall black, roughened; apex well differentiated into two tuberculate flared lips surrounding a slit-like ostiole. Peridium thin, 2-layered, outer layer composed of small heavily pigmented thick-walled cells of textura angularis, inner layer composed of hyaline thin-walled cells of textura angularis. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage, anastomosing between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical to narrowly fusoid, short pedicellate, with an ocular chamber and a small apical apparatus. Ascospores fusoid, hyaline, 1-septate, constricted at the septum, surrounded by an irregular hyaline gelatinous sheath.

Anamorphs reported for genus: Anguillospora longissima, Spirosphaera cupreorufescens and Repetophragma ontariense (Zhang et al. 2008c, 2009c).

Literature: Zhang et al. 2008c, 2009a, c.

Type species

Amniculicola lignicola Ying Zhang & K.D. Hyde, Mycol. Res. 112: 1189 (2008). (Fig. 3)

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Ascomata 350–450 μm high × 300–500 μm diam., solitary, scattered, or in small groups of 2–3, initially immersed, becoming erumpent, to nearly superficial, with basal wall remaining immersed in host tissue, globose, subglobose, broadly or narrowly conical, often laterally flattened, with a flattened base not easily removed from the substrate, wall black, roughened, often bearing remnants of wood fibers; apex well differentiated into two tuberculate flared lips surrounding a slit-like ostiole, 150–250 μm long, filled with a purplish amorphous matter, oriented in the axis of the wood fibers; underlying wood stained pale purple (Fig. 3a and b). Peridium 40–55 μm thick laterally, up to 120 μm thick at the apex, thinner at the base, coriaceous, 2-layered, outer layer composed of small heavily pigmented thick-walled cells of textura angularis, cells 4–9 μm diam., cell wall 2–3 μm thick, apex cells smaller and walls thicker, inner layer composed of hyaline thin-walled cells of textura angularis, 8–16 μm diam., in places with columns of textura prismatica, oriented perpendicular to the ascomatal surface, and larger, paler cells of textura prismatica towards the interior and at the base, 10–25 μm (Fig. 3c, d and e). Hamathecium of dense, long trabeculate pseudoparaphyses <1 μm broad, embedded in mucilage (Indian ink), anastomosing between and above the asci. Asci 140–184 × 9–10 μm, 8-spored, bitunicate, fissitunicate, cylindrical to narrowly fusoid, with a short, narrowed, twisted, furcate pedicel which is 15–25 μm long, with a low truncate ocular chamber and a small inconspicuous apical apparatus barely seen in water (Fig. 3f). Ascospores (20.5-)28–32 × (6-)8(−9) μm, obliquely uniseriate and partially overlapping, broadly fusoid to fusoid with broadly to narrowly rounded ends, hyaline, 1-septate, deeply constricted at the median septum, the upper cell often shorter and broader than the lower one, smooth, containing four refractive globules, surrounded by an irregular hyaline gelatinous sheath 4–8.5 μm thick, best seen in India ink, released senescent ascospores are greyish and 3-septate, strongly constricted at all septa (Fig. 3g).

Anamorph: none reported.

Colonies slow growing, reaching 4 cm diam. after 70 d growth on Malt Extract Agar (MEA) at 25°C, flat, with irregular to rhizoidal margin, off-white to grey, reverse reddish purple to deep reddish purple, the medium is stained pale yellow.

Material examined: FRANCE, Ariège, Prat Communal, Ruisseau de Loumet, 1000 m, on partly submerged wood of Fraxinus excelsior, 8 Aug. 2006, leg. Jacques Fournier (PC 0092661, holotype); 3 Sept. 2004 (BPI 877774; CBS: H-17932); Rimont, Ruisseau de Peyrau, 400 m, on driftwood of Alnus glutinosa (L.) Gaertn., 23 Jul. 2006 (HKU(M) 17515, isotype).

Notes

Morphology

Amniculicola is a freshwater genus which stains the woody substrate purple (Zhang et al. 2008c, 2009a, c). This genus appears only to be reported from Europe. A detailed description of the generic type was provided by Zhang et al. (2008c).

Phylogenetic study

Three species of Amniculicola cluster together with Anguillospora longissima, Spirosphaera cupreorufescens and Repetophragma ontariense as well as Pleospora rubicunda Niessl (current name Murispora rubicunda (Niessl) Y. Zhang ter, J. Fourn. & K.D. Hyde) and Massariosphaeria typhicola (P. Karst.) Leuchtm. (current name Neomassariosphaeria typhicola (P. Karst.) Yin. Zhang, J. Fourn. & K.D. Hyde). A new family, i.e. Amniculicolaceae, was introduced to accommodate these taxa (Zhang et al. 2008c, 2009a, c).

Concluding remarks

All of the five teleomorphic taxa within Amniculicolaceae are from freshwater in Europe and their ascomata stain the woody substrate purple. Purple staining makes taxa of this family easily recognized in the field.

Anomalemma Sivan., Trans. Br. Mycol. Soc. 81: 328 (1983). (?Melanommataceae)

Generic description

Habitat terrestrial, fungicolous. Ascomata gregarious, superficial, papillate, ostiolate. Peridium composed cells of pseudoparenchymatous. Asci clavate, 8-spored. Hamathecium of dense, filliform pseudoparaphyses. Ascospores 1- (rarely 2- to 3-) septate, fusoid, reddish brown, constricted at the main septum.

Anamorphs reported for genus: Exosporiella (= Phanerocorynella) (Sivanesan 1983).

Literature: Berkeley and Broome 1866; Keissler 1922; Massee 1887; Saccardo 1878a; Sivanesan 1983.

Type species

Anomalemma epochnii (Berk. & Broome) Sivan., Trans. Br. Mycol. Soc. 81: 328 (1983). (Fig. 4)

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≡ Sphaeria epochnii Berk. & Broome, Ann. Mag. nat. Hist., Ser. 3 18: 128 (1866).

Ascomata 340–500 μm high × 170–286 μm diam., gregarious on the intertwined hyphae, superficial, papillate, wall black, coriaceous, roughened (Fig. 4a). Peridium composed of two types of cells, outer layer 17–22 μm wide, composed of heavily pigmented thick-walled cells of textura angularis, cells up to 8 × 13 μm diam., cell wall 1–1.5 μm thick, inner layer 30–34 μm thick, composed of hyaline thin-walled cells (Fig. 4d). Hamathecium of dense, long cellular pseudoparaphyses, 2–4 μm broad, septate. Asci 75–108 × 9.5–12.5 μm (\( \bar{x} = 92.8 \times 11.1\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, dehiscence not observed, cylindro-clavate to clavate, with a furcate pedicel up to 6–25 μm long, with a small ocular chamber best seen in immature asci (ca. 2 μm wide × 1 μm high) (Fig. 4b and c). Ascospores 20–25(−30) × 5–7.5 μm (\( \bar{x} = 23.1 \times 6.3\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, fusoid to narrowly fusoid with narrowly rounded ends, brown, 1-septate, rarely 2- to 3-septate, deeply constricted at the median septum, smooth (Fig. 4e, f, g and h).

Anamorph: Exosporiella fungorum (Fr.) P. Karst. (Sivanesan 1983).

= Epochnium fungorum Fr., Syst. mycol. 3: 449 (1832).

Mycelium composed of branched, septate, pale brown hyphae. Stroma none. Conidiophores macronematous or semi-macronematous, mononematous, hyaline, smooth, branched towards the apex. Conidiogenous cells monoblastic, cylindrical or doliform. Conidia cylindrical or ellipsoidal, dry, 3-4-septate, smooth, hyaline or pale brown.

Material examined: UK, England, Warleigh near Bath, on fungus on bark (Epochnium sp.), Mar. 1866, leg. Warbright? (K(M):143936, syntype, ex herb. C.E. Broome).

Notes

Morphology

Sphaeria epochnii was first described and illustrated by Berkeley and Broome (1866) from Britain and the anamorphic stage is the hyphomycetous Epochniella fungorum. Sphaeria epochnii has subsequently been transferred to Melanomma (as M. epochnii (Berk. & Broome) Sacc.; Saccardo 1878a), Byssosphaeria (as B. epochnii (Berk. & Broome) Cooke; Massee 1887) and Chaetosphaeria (as C. epochnii (Berk. & Broome) Keissl.; Keissler 1922). The deposition of Sphaeria epochnii in Chaetosphaeria is obviously unacceptable, as Chaetosphaeria has unitunicate asci. Melanomma has been reported having Aposphaeria or Pseudospiropes anamorphs, which differs from Exosporiella (Sivanesan 1983). In addition, the presence of well developed prosenchymatous stroma in Sphaeria epochnii can also readily distinguish it from Melanomma (Sivanesan 1983).

The gregarious ascomata and formation of prosenchymatous stroma of Anomalemma resembles those of Cucurbitaria, but the pleosporaceous dictyosporous ascospores of Cucurbitaria readily distinguish it from Anomalemma epochnii. In addition, the pseudoparenchymatous peridium, fungicolous habitat and brown 1-septate ascospores, which later becoming 3-septate differ from any other pleosporalean genus. Thus a new genus, Anomalemma, was introduced to accommodate it (Sivanesan 1983). Anomalemma is presently monotypic.

Phylogenetic study

None.

Concluding remarks

Anomalemma epochnii certainly resembles Byssosphaeria in its ascomata clustering together in groups on closely intertwined hyphae and brown ascospores, and may well be included in this genus. Its fungicolous habitat, however, distinguishes it from Byssosphaeria.

Appendispora K.D. Hyde, Sydowia 46: 29 (1994a). (?Didymellaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata small, clustered, immersed, subglobose or irregularly pyriform. Peridium thin. Hamathecium of dense, long trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical, apical rounded with ocular chamber and faint ring, with short pedicels. Ascospores uniseriate to partially overlapping, fusoid, brown, 1-septate, slightly constricted at the septum.

Anamorphs reported for genus: none.

Literature: Hyde 1994a.

Type species

Appendispora frondicola K.D. Hyde, Sydowia 46: 30 (1994a). (Fig. 5)

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Ascomata 120–280 μm high × 180–280 μm diam., clustered, immersed with minute ostioles visible through cracks or blackened dots on the host surface, subglobose or irregularly pyriform (Fig. 5a and b). Peridium 40 μm thick, comprising two types of cells; outer cells, small heavily pigmented thick-walled cells of textura angularis, inner cells compressed, hyaline. Hamathecium of dense, very long trabeculate pseudoparaphyses, ca. 1 μm broad, embedded in mucilage, hyaline, anastomosing (Fig. 5e). Asci 130–144 × 11–13 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with an ocular chamber and faint ring, with short pedicels (Fig. 5c and d). Ascospores 21–30 × 7–9 μm, uniseriate to partially overlapping, fusoid, brown, 1-septate, slightly constricted at the septum, with an irregular ridged ornamentation and 3–5 narrow appendages at each end (Fig. 5f and g).

Anamorph: none reported.

Material examined: BRUNEL, Jalan, Muara, Simpang 835, on dead rachis of Oncosperma horridum on forest floor, Nov. 1992, K.D. Hyde 1652 (BRIP 21354, holotype).

Notes

Morphology

Appendispora was described as a saprobe of palm, and is characterized by small, immersed ascomata, bitunicate, fissitunicate asci, trabeculate pseudoparaphyses, brown, 1-septate, appendaged ascospores with irregular wall striations (Hyde 1994a). Based on its trabeculate pseudoparaphyses embedded within gel matrix and its brown ascospores, Appendispora was assigned to Didymosphaeriaceae (Barr 1987b; Hyde 1994a).

Phylogenetic study

None.

Concluding remarks

The saprobic habitat and association with monocots, cylindrical asci, trabeculate pseudoparaphyses as well as its brown, 1-septate ascospores make it difficult to determine a better phylogenetic position than Didymellaceae.

Ascorhombispora L. Cai & K.D. Hyde, Cryptog. Mycol. 28: 294 (2007). (Pleosporales, genera incertae sedis)

Generic description

Habitat freshwater, saprobic. Ascomata solitary or gregarious, superficial, globose to subglobose, dark brown to black, short papillate, ostiolate, coriaceous. Peridium relatively thin, textura angularis in longitudinal section, 2-layered. Hamathecium not observed. Asci 8-spored, obpyriform, broadly clavate to saccate, pedicellate, bitunicate, apex rounded, persistent. Ascospores overlapping 2-3-seriate, broadly fusoid to rhomboid, thick-walled, surrounded by mucilaginous sheath, 3-euseptate, not constricted at septa, median septum wide, forming a darker band, central cells large, trapezoid, dark brown to black, verruculose, polar end cells small and paler.

Anamorphs reported for genus: none.

Literature: Cai and Hyde 2007.

Type species

Ascorhombispora aquatica L. Cai & K.D. Hyde, Cryptog. Mycol. 28: 295 (2007). (Fig. 6)

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Ascomata 140–170 μm high × 150–185 μm diam., solitary or gregarious, superficial, globose to subglobose, dark brown to black, short papillate, ostiolate, ostioles rounded, small, coriaceous. Peridium relatively thin, 10–18 μm wide, textura angularis in longitudinal section, composed of two layers of angular cells, outer later dark brown to black, relatively thick-walled, inner layer hyaline, relatively thin-walled (Fig. 6a and b). Hamathecium not observed. Asci 100–198 × 72–102 μm (\( \bar{x} = 186 \times 88\mu m \), n = 15), 8-spored, obpyriform, broadly clavate to saccate, pedicellate, bitunicate, apex rounded, deliquescent (Fig. 6c, d and e). Ascospores 30.5–45 × 16–26.5 μm (\( \bar{x} = 38.5 \times 21\mu m \), n = 25), overlapping 2-3-seriate, broadly fusoid to rhomboid, thick-walled, surrounded by mucilaginous sheath, 3-euseptate, not constricted at septa, median septum wide, forming a darker band, central cells large, trapezoid, 11–18 μm long, dark brown to black, verruculose, polar end cells small, hemispherical, 3.5–4 μm long, subhyaline to pale brown, smooth (Fig. 6f).

Anamorph: none reported.

Material examined: CHINA, Yunnan, Jinghong, on submerged bamboo in a small forest stream, 26 Jan. 2003, leg. det. L. Cai, CAI-1H31 (HKU(M) 10859, holotype).

Notes

Morphology

Ascorhombispora was introduced as a monotypic genus from freshwater by Cai and Hyde (2007), and is characterized by superficial, coriaceous, non-stromatic ascomata, large, saccate asci; lack of interascal filaments and trapezoid (rhombic), 3-septate, dark brown to black ascospores with smaller end cells which are subhyaline to pale brown. Ascorhombispora is most comparable with Caryospora and Zopfia. But the globose to subglobose ascomata and thin peridium, saccate asci lacking interascal pseudoparaphyses, and the 3-septate, rhomboid ascospores with the paler end cells of Ascorhombispora differs from those of Caryospora (Cai and Hyde 2007).

Phylogenetic study

Phylogenetic analysis based on either SSU or LSU rDNA sequences indicated that Ascorhombispora aquatica belongs to Pleosporales, but its familial placement was left undetermined (Cai and Hyde 2007).

Concluding remarks

The sac-shaped asci and absence of pseudoparaphyses are uncommon in Pleosporales, especially among those from freshwater.

Asteromassaria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368 (1917). (?Morosphaeriaceae)

Generic description

Habitat terrestrial, saprobic. Ascomata medium-sized, clustered, at first immersed and then breaking through the host surface and becoming superficial, globose, subglobose, coriaceous. Peridium 2-layered, thicker near the base. Hamathecium of dense, septate, cellular pseudoparaphyses which branch and anastomosing frequently between and above asci. Asci (4-)8-spored, bitunicate, cylindro-clavate to clavate, with a short truncated pedicel and a small ocular chamber. Ascospores obliquely uniseriate and partially overlapping to biseriate, fusoid to fusoid-ellipsoidal, pale brown when mature, 1-septate, some becoming 3-septate when old, constricted at the median septum.

Anamorphs reported for genus: Scolicosporium (Sivanesan 1984).

Literature: Barr 1982a; b; 1993a; Boise 1985; Shoemaker and LeClair 1975; Sivanesan 1987; Tanaka et al. 2005.

Type species

Asteromassaria macrospora (Desm.) Höhn., F. von, Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. I 126: 368 (1917). (Fig. 7)

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≡ Sphaeria macrospora Desm., Ann. Sci. Nat. Bot. 10: 351 (1849).

Ascomata 400–600 μm high × 450–650 μm diam., 4–20 clustered together, at first immersed and then breaking through the host surface and becoming superficial, globose, subglobose, not easily removed from the substrate, wall black, coriaceous, roughened, apex usually widely porate, with or without papilla (Fig. 7a). Peridium 70–90 μm wide, thicker near the base where it is up to 180 μm wide, comprising two cell types, outer cells composed of heavily pigmented small cells, cells 3–5 μm diam., inner layer composed of less pigmented cells of textura angularis, 10–20 μm diam. (Fig. 7b and c). Hamathecium of dense, septate, 2–3 μm broad, pseudoparaphyses which branch and anastomosing frequently between and above asci (Fig. 7d). Asci (180-)200–280 × 28–43 μm (\( \bar{x} = 230 \times 35\mu m \), n = 10), 8-spored (sometimes 4-spored), bitunicate, fissitunicate dehiscence not observed, cylindro-clavate to clavate, with a short truncated pedicel up to 30 μm, with a small ocular chamber (ca. 3 μm wide × 3 μm high) (Fig. 7e and f). Ascospores 50–58 × (14-)18–21 μm (\( \bar{x} = 55.3 \times 18.2\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, fusoid to fusoid-ellipsoidal, with narrowly rounded ends, lightly brown when mature, 1-septate, some becoming 3-septate when old, constricted at the median septum, the upper cell often broader and longer than the lower one, minutely verrucose (Fig. 7g, h, i and j).

Anamorph: Scolicosporium macrosporium (Berk.) B. Sutton.

Acervuli immersed in bark, brown, discrete, up to 250 μm diam., opening by irregular rupture of the overlaying tissues. Peridium of thin-walled angular cells. Conidiophores cylindrical, 1-2-septate, up to 30 μm long and 3–5 μm wide. Conidiogenous cells holoblastic, 1-2-annellate, cylindrical, hyaline. Conidia 100–190 × 12–15 μm, fusoid, pale brown with paler or hyaline ends, 7–17 transverse septate, smooth-walled, with a tapered apex and truncate base (adapted from Sivanesan 1984).

Material examined: CZECH REPUBLIC, Mährisch-Welвkirchen (Hranice), Wsetin (Vsetin), Berg Čap., on Fagus sylvatica L., Aug. 1938, F. Petrak (L, 1004).

Notes

Morphology

In this study we were unable to obtain the holotype, so we used a collection of Petrak’s. The main morphological characters of Asteromassaria are the medium- to large-sized, globose to depressed ascomata opening with a pore, clavate to oblong asci, narrowly cellular pseudoparaphyses, pale to dark brown, bipolar symmetric, mostly fusoid, distoseptate or euseptate ascospores (Barr 1993a). The bipolar symmetric ascospores of Asteromassaria can readily be distinguished from other genera of this family (Barr 1993a; Tanaka et al. 2005). Currently, it comprises 12 species (Tanaka et al. 2005; http://www.mycobank.org, 28-02-2009).

Phylogenetic study

Asteromassaria pulchra (Harkn.) Shoemaker & P.M. LeClair is basal to Morosphaeriaceae in the phylogenetic tree based on four genes, but its placement is influenced by taxon sampling that was different in several analyses.

Concluding remarks Asteromassaria can be distinguished from other comparable genera, i.e. Pleomassaria and Splanchnonema by 1-septate and pale brown ascospores, thick-walled textura angularis peridium and Scolicosporium anamorphic stage (see under Pleomassaria).

Astrosphaeriella Syd. & P. Syd., Annls mycol. 11: 260 (1913). (?Melanommataceae)

Generic description

Habitat terrestrial, saprobic. Ascomata densely scattered or in small groups, erumpent through the outer layers of the host tissues to nearly superficial, reflexed pieces of the ruptured host tissue usually persisting around the base of the ascomata, often star-like, conical to semiglobose, with a central papilla. Peridium upper wall usually comprising a thick dark brittle pseudoparenchymatous layer, base usually flattened and thin-walled. Hamathecium of dense, filliform, trabeculate pseudoparaphyses, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid. Ascospores narrowly fusoid with acute ends, hyaline, pale brown or brown, 1-3-septate.

Anamorphs reported for genus: Pleurophomopsis (Hyde et al. 2011).

Literature: von Arx and Müller 1975; Barr 1990a; Chen and Hsieh 2004; Hawksworth 1981; Hawksworth and Boise 1985; Hyde and Fröhlich 1998; Hyde et al. 2000; Kirk et al. 2001; Sydow and Sydow 1913; Tanaka and Harada 2005a; b; Tanaka et al. 2009.

Type species

Astrosphaeriella stellata Syd. & P. Syd., Annls mycol. 11: 260 (1913). (Fig. 8)

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Ascomata 360–570 μm high × 860–1150 μm diam., densely scattered or in small groups, erumpent through the outer layers of the host tissues to nearly superficial, reflexed pieces of the ruptured host tissue usually persisting around the base of the ascomata, forming star-like flanges around the ascomata from the surface view; ascomata broadly conical, with a flattened base not easily removed from the substrate, wall black; apex with a central papilla which is black and shiny at maturity, scarcely projecting (Fig. 8a). Peridium 40–70 μm thick, carbonaceous and crisp, 1-layered, composed of very small dark brown thick-walled pseudoparenchymatous cells, cells 2–5 μm diam., cell wall 2–6 μm thick, in places at the base composed of hyaline cells of textura prismatica, cells 5 × 8 μm diam. (Fig. 8b). Hamathecium of dense, very long trabeculate pseudoparaphyses, <1 μm broad, embedded in mucilage (Indian ink), anastomosing between and above the asci. Asci 130–190 × 11.5–15 μm (\( \bar{x} = 161.5 \times 12.8\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindro-clavate to narrowly fusoid, with a short, narrowed pedicel which is 10–35 μm long, with a large ocular chamber (Fig. 8c). Ascospores 35–50 × 5–7.5 μm (\( \bar{x} = 43.4 \times 6\mu m \), n = 10), biseriate, elongate- fusoid, gradually tapering towards the ends, hyaline, turning pale brown when mature, 1(−3)-septate, constricted at the median septum (Fig. 8d,e and f).

Anamorph: none reported.

Material examined: USA, Hawaii, Kapano Gulch, in bamboo culms, 5 Jun. 1947, leg. Kopf & Rogers, det. Miller (BISH 145726, as Astrosphaeriella fusispora Syd. & P. Syd.).

Notes

Morphology

Astrosphaeriella has been treated as a synonym of Microthelia (von Arx and Müller 1975), but the large conical ascomata, numerous trabeculate pseudoparaphyses and 1-septate and elongated ascospores of Astrosphaeriella all disagree with those of Microthelia (Hawksworth 1981). It was assigned to Platystomaceae by Barr (1990a) in Pleosporales or Melanommataceae by Kirk et al. (2001). Following a systematic study of Astrosphaeriella, only four species were accepted, i.e. A. aosimensis I. Hino & Katum., A. stellata, A. trochus (Penz. & Sacc.) D. Hawksw. and A. venezuelensis M.E. Barr & D. Hawksw. (Hawksworth 1981), and it was defined as a tropical genus, occurring exclusively on palms or bamboo. Astrosphaeriella stellata was selected as the type of Astrosphaeriella, and A. fusispora was regarded as a synonym of A. stellata (Hawksworth 1981). More taxa were subsequently added (Barr 1990a; Hawksworth and Boise 1985; Hyde and Fröhlich 1998), and the generic concept extended to include three elements: 1. typical semi-immersed to superficial ascomata with flattened base, cylindro-clavate asci with fusoid ascospores and trabeculate pseudoparaphyses, i.e. Astrosphaeriella sensu stricto (e.g. A. fusispora and A. vesuvius (Berk. & Broome) D. Hawksw. & Boise); 2. Trematosphaeria-like with rounded ascomata (e.g. A. africana D. Hawksw.); and 3. Massarina-like species with immersed ascomata (e.g. A. bakeriana (Sacc.) K.D. Hyde & J. Fröhl.) (Chen and Hsieh 2004; Tanaka and Harada 2005a; b). Currently, a broad generic concept of Astrosphaeriella is accepted, and 47 taxa are included in Astrosphaeriella.

Phylogenetic study

Phylogenetic analysis based on LSU and SSU nurDNA sequence data indicates that Astrosphaeriella is polyphyletic, and located in the basal region of the Pleosporales between Testudinaceae and Zopfiaceae/Delitschiaceae (Tanaka et al. 2009), or basal to Aigialaceae (Schoch et al. 2009). The genus is, however, clearly not related to Trematosphaeria as previously understood (Boise 1985).

Concluding remarks Astrosphaeriella is currently polyphyletic and new collections of the different elements listed above are needed in order to understand the placement of various species. We suggest that some immersed bambusicolous species may belong in Tetraplospheariaceae.

Asymmetricospora J. Fröhl. & K.D. Hyde, Sydowia 50: 183 (1998). (?Melanommataceae)

Generic description

Habitat terrestrial, saprobic. Ascomata solitary or in small groups, immersed, black, lenticular in section, uni- or often multi-locular, with a central ostiole without tissue differentiation. Upper peridium carbonaceous, thicker at sides and apex. Lower peridium composed of irregular-shaped, hyaline cells. Hamathecium of trabeculate pseudoparaphyses, branching and anastomosing between and above asci, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate unknown, clavate, short pedicellate. Ascospores 1-septate, hyaline, constricted at the septum, with a broad, spreading mucilaginous sheath.

Anamorphs reported for genus: none.

Literature: Fröhlich and Hyde 1998.

Type species

Asymmetricospora calamicola J. Fröhl. & K.D. Hyde, Sydowia 50: 184 (1998). (Fig. 9)

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Ascomata 675–950 μm high × 875–1500 μm diam., solitary or in small groups of 2–10, immersed and forming slightly protruding domes on the substrate surface, with near-white rim around the central ostiole; in vertical view lenticular, multi- or rarely unilocular, individual locules 175–270 μm high × 320–400 μm diam., with a flattened base, ostiole a central opening without tissue differentiation (Fig. 9a). Upper peridium 32–70 μm wide, carbonaceous, composed of a few layers of black walled cells of textura angularis. Lower peridium thinner, composed of hyaline cells of textura globulosa or textura prismatica (Fig. 9b). Hamathecium of long trabeculate pseudoparaphyses, 1.2–1.6(−2) μm wide, branching and anastomosing between and above asci, embedded in mucilage. Asci 137.5–207.5 × 26–35 μm (\( \bar{x} = 172.8 \times 31.5\mu m \), n = 20), 8-spored, bitunicate, fissitunicate dehiscence not observed, clavate, with short pedicel (to 25 μm), with ocular chambers (ca. 3 μm wide × 4 μm high) (Fig. 9c and d). Ascospores 35–55 × 10.5–15 μm (\( \bar{x} = 44.7 \times 12.4\mu m \), n = 50), biseriate, navicular to obovoid, hyaline, becoming pale brown when senescent, straight or usually curved, smooth, asymmetric, 1-septate, the upper cell larger with a rounded end, basal cell with a tapering end, constricted at the septum, with spreading mucilaginous sheath (Fig. 9e, f and g) (data from Fröhlich and Hyde 1998).

Anamorph: none reported.

Material examined: AUSTRALIA, North Queensland, Palmerston, Palmerston National Park, on dead rattan of Calamus caryotoides A.Cunn. ex Mart., Mar. 1994, J. Fröhlich (HKU(M) 7794, holotype).

Notes

Morphology

Asymmetricospora was introduced as a monotypic genus represented by A. calamicola based on its “absence of a subiculum, the absence of short dark setae around the papilla and its asymmetric ascospores” (Fröhlich and Hyde 1998). Because of the immersed ascomata, ostiole and peridium morphology, fissitunicate asci and trabeculate pseudoparaphyses, Asymmetricospora was assigned to Melanommataceae (sensu Barr 1990a; Fröhlich and Hyde 1998).

Morphologically Asymmetricospora can be distinguished from its most comparable genus, Astrosphaeriella, by its ostiole, which is a simple opening without tissue differentiation, asymmetric ascospores, and the usually multi-loculate fruiting body (Fröhlich and Hyde 1998).

Phylogenetic study

None.

Concluding remarks

The placement of Asymmetricospora under Melanommataceae remains to be confirmed.

Barria Z.Q. Yuan, Mycotaxon 51: 313 (1994). (Phaeosphaeriaceae)

Generic description

Habitat terrestrial, parasitic. Ascomata small- to medium-sized, solitary or scattered, immersed, globose, subglobose, ostiolate, coriaceous. Apex with or without papilla and with a pore-like ostiole. Peridium 2-layered. Hamathecium of dense, long cellular pseudoparaphyses, septate, embedded in mucilage. Asci bitunicate, fissitunicate, cylindrical to clavate, with a short, furcate pedicel. Ascospores ellipsoid, hyaline at first, turning brown at maturity, 1-septate, strongly constricted at the septum.

Anamorphs reported for genus: none.

Literature: Yuan 1994.

Type species

Barria piceae Z.Q. Yuan, Mycotaxon 51: 314 (1994). (Fig. 10)

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Ascomata 240–370 μm high × 200–320 μm diam., solitary, scattered, immersed, globose, subglobose, coriaceous, apex with or without papilla and with a pore-like ostiole (Fig. 10a). Peridium 20–35 μm thick, comprising two cell types, the outer cells comprising 3–4 layers of brown pseudoparenchymatous cells, cells 4–5 μm diam., cell wall 2–3 μm thick, inner cells comprising 3–4 layers of pale brown compressed cells, cells 2 × 16 μm diam., cell wall 0.5–1.5 μm thick (Fig. 10b). Hamathecium of dense, long cellular pseudoparaphyses, 2–3 μm broad, septate. Asci 135–200(−220) × 14–20 μm (\( \bar{x} = 156 \times 16.6\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to clavate, with a short, furcate pedicel, up to 22 μm long, with a large ocular chamber (ca. 4 μm wide × 3 μm high) (Fig. 10c and d). Ascospores 19–21.5 × 10–12 μm (\( \bar{x} = 20.4 \times 11\mu m \), n = 10), uniseriate to partially overlapping, ellipsoid, hyaline or greenish with numerous small guttules at first and olive green to smoky brown at maturity, 1-septate, strongly constricted at the septum, foveolate, surrounded with sheath (Fig. 10e and f).

Anamorph: none reported.

Material examined: CHINA, Xinjiang Province, Uygur, Urumqi, Tianshan Mountain, on needles of Picea schrenkiana, 1 Jul. 1992, Z.Q. Yuan (NY 92003, isotype).

Notes

Morphology

Barria was established by Yuan (1994) as a monotypic genus represented by B. piceae according to its “two-celled, pigmented ascospores, pseudoparenchymatous peridium and narrowly cellular pseudoparaphyses” thus differing in its combination of characters from all of the morphologically related dothideomycetous genera, such as Didymosphaeria, Didymopleella or Stegasphaeria. The taxon was considered to belong in Phaeosphaeriaceae. Ascomata and colour or shape of ascospores, however, readily distinguish it from other 1-septate Phaeosphaeriaceae genera, i.e. Didymella, Lautitia and Metameris (Yuan 1994). Barria piceae causes blight of spruce needles.

Phylogenetic study

None.

Concluding remarks

The status of Barria with its unusual verrucose ascospores and thick gel coating is uncertain. In many ways it resembles Belizeana, with its cylindrical asci, 1-septate, ellipsoid ascospores with sheath and verruculose surface (Kohlmeyer and Volkmann-Kohlmeyer 1987). However, the latter is a marine genus while Barria causes leaf blight of terrestrial Picea (Yuan 1994). The placement in Phaeosphaeriaceae seems logical based on the parasitic life style, thin and simple peridium, wide cellular pseudoparaphyses and brown ascospores. However, molecular data are needed to confirm this.

Belizeana Kohlm. & Volkm.-Kohlm., Bot. Mar. 30: 195 (1987). (Pleosporales, genera incertae sedis)

Generic description

Habitat marine, saprobic. Ascomata solitary, scattered, or in small groups, medium-sized, immersed to semi-immersed, subglobose to broadly ampulliform, black, ostiolate, carbonaceous. Peridium thin, comprising several layers of brown thin-walled cells of textura angularis. Hamathecium of dense, filliform pseudoparaphyses, rarely branched. Asci 8-spored, bitunicate, fissitunicate, broadly cylindrical to clavate, with a short pedicel and an ocular chamber. Ascospores uniseriate, broadly ellipsoidal, hyaline, turn pale brown when senescent, 1-septate, constricted at the septum, thick-walled, 2-layered, mature spores with tuberculate ornamentation between the two layers.

Anamorphs reported for genus: Phoma-like (Kohlmeyer and Volkmann-Kohlmeyer 1987).

Literature: Kohlmeyer and Volkmann-Kohlmeyer 1987.

Type species

Belizeana tuberculata Kohlm. & Volkm.-Kohlm., Bot. Mar. 30: 196 (1987). (Fig. 11)

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Ascomata 170–300 μm high × 160–290 μm diam., solitary, scattered, or in small groups of 2–3, immersed to semi-immersed, subglobose to broadly ampulliform, carbonaceous, black, pale brown on the sides, ostiolate, epapillate or shortly papillate, ostiolar canal filled with a tissue of hyaline cells (Fig. 11a). Peridium 25–35 μm wide, comprising several layers thin-walled cells of textura angularis, which are hyaline inwardly, near the base composed of a hyaline hyphal mass producing asci, up to 20 μm thick (Fig. 11b, c and e). Hamathecium of dense, ca. 2 μm broad, filliform pseudoparaphyses, rarely branched, embedded in mucilage (Fig. 11g). Asci 145–170 × 20–30 μm (\( \bar{x} = 163 \times 25\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, broadly cylindrical to clavate with a short pedicel, thick-walled, with a small ocular chamber (Fig. 11d, f and h). Ascospores 21–26 × 13–18 μm (\( \bar{x} = 22 \times 15\mu m \), n = 10), uniseriate, broadly ellipsoidal, hyaline, turn pale brown when senescent, 1-septate, constricted at the septum, thick-walled, 2-layered, mature spores with tuberculate ornamentation between the two layers (Fig. 11i and j).

Anamorph: Phoma-like (Kohlmeyer and Volkmann-Kohlmeyer 1987).

Material examined: BELIZE, Twin Cays, on Laguncularia sp., 7 Apr. 1983, leg. & det. J. Kohlmeyer (Herb. J. Kohlmeyer No. 4398, holotype); AUSTRALIA, Towra Point, New South Wales, trunk of eroded tree with oysters and shipworms, intertidal zone, Botany Bay, 23 Aug. 1981 (Herb. J. Kohlmeyer No. 4209, paratype).Notes

Morphology

Belizeana was formally established to accommodate B. tuberculata, an obligate marine fungus, which is characterized by verrucose ascospores (Kohlmeyer and Volkmann-Kohlmeyer 1987). Belizeana tuberculata can be assigned to Pleosporaceae (Pleosporales) according to Luttrell’s (1973) treatment and keys of von Arx and Müller (1975), but cannot resolve a proper family based on Barr (1979a, 1983). The unique morphology together with obligate marine habitat makes B. tuberculata readily distinguishable from all other taxa of Pleosporaceae.

Phylogenetic study None.

Concluding remarks

The ascospores of Belizeana tuberculata are most comparable with those of Acrocordiopsis patilii, but the superficial conical ascomata of A. patilii are distinct from B. tuberculata. Thus, the familial placement of Belizeana is still undetermined.

Biatriospora K.D. Hyde & Borse, Mycotaxon 26: 263 (1986). (Pleosporales, genera incertae sedis)

Generic description

Habitat marine, saprobic. Ascomata large, solitary or gregarious, immersed, subglobose to pyriform, ostiolate, papillate, periphysate, black, branching, carbonaceous. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with apical apparatus. Ascospores uniseriate to partially overlapping, fusoid, hyaline when young, becoming brown to dark brown at maturity, multi-septate towards each end, with a hyaline, globose refractive chamber or appendage at each end, not constricted at the septum.

Anamorphs reported for genus: none.

Literature: Hyde and Borse 1986; Suetrong et al. 2009.

Type species

Biatriospora marina K.D. Hyde & Borse, Mycotaxon 26: 264 (1986). (Fig. 12)

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Ascomata 650–860 μm high × 350–510 μm diam., solitary or gregarious, immersed, subglobose to pyriform, ostiolate, papillate, periphysate, black, carbonaceous (Fig. 12.2a). Hamathecium of dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad, branching, embedded in mucilage. Asci 175–400 × 22–40 μm, 8-spored, bitunicate, fissitunicate, cylindrical, with long pedicels and apical apparatus (Fig. 12.1a, b, 2b). Ascospores 55–82 × 16–25 μm, uniseriate to partially overlapping, fusoid, hyaline when young, becoming brown to dark brown at maturity, 2-4-septate towards each end, and with a hyaline, globose refractive chamber or appendage at each end, 6–8 × 4–6 μm diam., not constricted at the septum (Fig. 12.1c, d, 2c).

Anamorph: none reported.

Material examined: SEYCHELLES, 2 Jan. 1984 (Herb. IMI 297768 holotype).

Notes

Morphology

Biatriospora was introduced to accommodate a marine fungus B. marina, which is characterized by horizontal ascomata and ascospores with polar, globose refractive chambers and polar septa (Hyde and Borse 1986). Polar refractive chambers can also occur in other marine fungi, such as Lulworthia and Aigialus. The chambers have been proposed as important for spore attachment to substrates in a liquid environment (Hyde and Borse 1986).

Phylogenetic study

Multigene phylogenetic analysis indicated that Biatriospora marina forms a separate branch, sister to other families of Pleosporales (Suetrong et al. 2009), and maybe related to species in Roussoella (Plate 1).

Concluding remarks The familial status of Biatriospora can not be determined.

Bicrouania Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (?Melanommataceae)

Generic description

Habitat marine, saprobic. Ascomata immersed gregarious, erumpent to superficial, globose to subglobose, black, periphysate, coriaceous, epapillate or papillate, ostiolate. Peridium thin, 2-layered. Hamathecium of dense, long trabeculate pseudoparaphyses, branching and anastomosing between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a thick, furcate pedicel lacking ocular chamber. Ascospores obliquely uniseriate and partially overlapping, ellipsoidal with broadly rounded ends, reddish brown, 1-septate, thick-walled, constricted at the septum.

Anamorphs reported for genus: none.

Literature: Jones et al. 2009; Kohlmeyer and Volkmann-Kohlmeyer 1990.

Type species

Bicrouania maritima (P. Crouan & H. Crouan) Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 685 (1990). (Fig. 13)

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≡ Sphaeria maritima P. Crouan & H. Crouan, Florule du Finistére, Paris: 27 (1867) non Sphaeria maritima Cooke & Plowright, Grevillia 5: 120 (1877).

Ascomata 320–440 μm high × 370–460 μm diam., gregarious, immersed, mostly erumpent to superficial, globose to subglobose, black, coriaceous, with a rough surface, papillate or epapillate, ostiolate, periphysate (Fig. 13a). Peridium 40–50 μm thick laterally, up to 75 μm thick at the apex, thinner at the base, 2-layered, outer layer composed of small heavily pigmented pseudoparenchymatous cells, inner layer very thin, composed of hyaline thin-walled small cells, merging into pseudoparaphyses (Fig. 13a and b). Hamathecium of dense, very long trabeculate pseudoparaphyses, 0.8–1.2 μm broad, branching and anastomosing between and above the asci. Asci 170–225 × 17.5–22.5 μm (\( \bar{x} = 199.6 \times 20\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a thick, furcate pedicel which is up to 70 μm long, lacking ocular chamber (Fig. 13c, d and e). Ascospores 22–26 × 12–15 μm (\( \bar{x} = 24.5 \times 13.3\mu m \), n = 10), obliquely uniseriate and partially overlapping, ellipsoidal with broadly rounded ends, reddish brown, 1-septate, slightly constricted at the septum, thick-walled, with a thick darkened band around the septum, smooth (Fig. 13c, d and e).

Anamorph: none reported.

Material examined: FRANCE, Finistère, on Halimone portulacoides (IMI 330806, isotype, as Sphaeria maritima).

Notes

Morphology

When Kohlmeyer and Volkmann-Kohlmeyer (1990) studied the four marine Didymosphaeria species, the monotypic Bicrouania was established to accommodate B. maritima (as Didymosphaeria maritima (P. Crouan & H. Crouan) Sacc.), which could be distinguished from Didymosphaeria by its superficial ascomata lacking a clypeus, thick-walled asci and its association with algae (Kohlmeyer and Volkmann-Kohlmeyer 1990). Jones et al. (2009) agreed that it cannot be placed in Didymosphaeria based on its superficial ascomata, but that it does have many similarities with Didymosphaeria. Molecular data are required to determine its relationship with Didymosphaeria and to resolve its higher level placement.

Phylogenetic study None.

Concluding remarks

Besides the morphological differences, its marine and substrate habitats also differ from Didymosphaeria.

Bimuria D. Hawksw., Chea & Sheridan, N. Z. J. Bot. 17: 268 (1979). (Montagnulaceae)

Type species

Bimuria novae-zelandiae Hawksworth, Chea & Sheridan, N. Z. J. Bot. 17: 268 (1979). (Fig. 14)

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Ascomata (185-)200 × 310(-330) μm diam., solitary, scattered, semi-immersed or superficial, globose, hyaline when young, turning dark brown to black when mature, ostiolate, the ostiole more or less sessile or raised into a very short neck. Peridium 5–8(-12) μm thick, comprising 2–3 layers of radically compressed pseudoparenchymatous cells, cells 10–15 μm diam. in surface view, cell wall 2–3 μm thick. Hamathecium consisting of few, 2.5–4 μm broad cellular pseudoparaphyses, embedded in mucilage, rarely anastomosing and branching, septate, 7–13 μm long between two septa. Asci (65-)80–95 × 20–32.5 μm (\( \bar{x} = 75.6 \times 29.4\mu m \), n = 10), (1-)2(-3)-spored, bitunicate, fissitunicate, broadly clavate, with a short and small knob-like pedicel which is up to 13 μm long, ocular chamber best seen in immature asci (Fig. 14a, b, c, d and g). Ascospores accumulating in a subglobose black shiny mass adhering together outside the ostiole, 55–68 × 25–28 μm (\( \bar{x} = 59 \times 26\mu m \), n = 10), broadly ellipsoid but becoming narrowed towards the poles, muriform with (5-)7 transverse septa, cells with (0-)l(-2) longitudinal septa in each cell, no constriction at the septa, dark brown, the apical cells paler with no longitudinal septa, verruculose (Fig. 14e and f).

Anamorph: none reported.

Material examined: NEW ZEALAND, North Island, Wairarapa District, Nutty Farm, isolated from soil, 3 Mar. 1978, Chea Chark Yen & J.E. Sheridan (CBS 107.79, isotype).

Type species

Bricookea sepalorum (Vleugel) M.E. Barr, Mycotaxon 15: 346 (1982). (Fig. 15).

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≡ Metasphaeria sepalorum Vleugel, Svensk bot. Tidskr. 2: 369 (1908).

Ascomata 120–250 μm high × 170–440 μm diam., solitary, scattered, or in small groups, or forming locules in massive stromatic tissues, initially immersed, becoming erumpent, to nearly superficial, depressed globose, black, membraneous, roughened; apex rounded, sometimes very short and almost inconspicuous, with a somewhat slit-like or Y-shaped ostiole (Fig. 15a). Peridium 16–30 μm wide, comprising two types of cells, outer cells heavily pigmented thick-walled textura angularis, cells 4.5–8 μm diam., cell wall 1–1.5 μm thick, inner cells of subhyaline thin-walled textura angularis, cells larger than outer cells (Fig. 15b). Hamathecium of long cellular pseudoparaphyses, 1.5–2 μm broad, embedded in mucilage, anastomosing, septate. Asci 63–83 × 9.5–11 μm (\( \bar{x} = 73.8 \times 10.8\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, oblong, cylindro-clavate or slightly obclavate, with a short knob-like pedicel which is 5–13 μm long, with an ocular chamber (Fig. 15c, d and e). Ascospores (14-)15.5–19 × 5–7 μm (\( \bar{x} = 16.9 \times 5.9\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, ellipsoid to narrowly obovoid, hyaline, 3-septate, constricted at each septum, the cells above central septum often broader than the lower ones, smooth (Fig. 15f, g, h, i and j).

Anamorph: none reported.

Material examined: SWEDEN, on Juncus filliformis, Stockholm, J. Vleugel. Jul. 1907 (S, type as Metasphaeria sepalorum Vleugel).

Type species

Byssolophis byssiseda (Flageolet & Chenant.) Clem., Gen. Fung. (Minneapolis): 286 (1931). (Fig. 16)

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≡ Schizostoma byssisedum Flageolet & Chenant., in Chenantaise, Bull. Soc. mycol. Fr. 35: 125 (1919).

Ascomata 300–450 μm high × 600–750 μm long × 350–420 μm broad, gregarious, semi-immersed to erumpent, coriaceous, ovoid with a flattened base and apex with a elongate slit-like ostiole, up to 700 μm long and 200 μm wide (Fig. 16a). Peridium not observed. Hamathecium of dense, long pseudoparaphyses, up to 1.5–2.5 μm broad, anastomosing and branching between and above the asci (Fig. 16b). Asci 80–105 × (5-)7.5–10 μm (\( \bar{x} = 91 \times 8\mu m \); n = 10), 8-spored, bitunicate, fissitunicate, cylindrical or cylindro-clavate, with a furcate pedicel and a small ocular chamber (J-) (Fig. 16d). Ascospores 18–20(−28) × 4.5–6(−7.5) μm (\( \bar{x} = 20.8 \times 5.7\mu m \), n = 10), uniseriate to biseriate, fusoid, hyaline, turning faintly brown when old, 1-septate, with 1–2 distinct oil drops in each cell and usually with a short terminal appendage at each end (Fig. 16c).

Anamorph: none reported.

Material examined: on decaying wood (K(M):164030, isotype).

Type species

Byssosphaeria keitii (Berk. & Broome) Cooke [as ‘Byssosphaeria keithii’], (1879). (Fig. 17)

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≡ Sphaeria keitii Berk. & Broome [as ‘Sphaeria keithii’], Ann. Mag. Nat. Hist., IV 17: 144 (1876).

Ascomata 360–500(−600) μm high × 420–640 μm diam., scattered or in small groups, superficial with basal subiculum anchoring on the substrate, globose, subglobose to turbinate, non-papillate with pore-like ostiole, ostiolar region sometimes with orange and greenish tint, wall black, roughened, coriaceous (Fig. 17a). Peridium 55–85 μm thick, peridium outside of the substrate comprising two cell types, outer layer composed of brown thick-walled cells of textura epidermoidea, cells 1–3 μm diam., inner layer composed of small hyaline cells, cells 3–5 μm diam., merging into pseudoparaphyses; peridium inside the substrate one layer, composed of large pale brown cells of textura angularis, cells 6–13 μm diam. (Fig. 17c). Hamathecium of dense, long trabeculate pseudoparaphyses, 1–2 μm broad, embedded in mucilage, anastomosing between and above the asci. Asci 90–120(−148) × 10–14 μm, 8-spored, bitunicate, fissitunicate, cylindro-clavate to clavate, biseriate above and uniseriate below, pedicel 15–20(−53) μm long, the immature asci usually with longer and furcate pedicel (−68 μm) (Fig. 17d,e and f). Ascospores 29–34(−38) × 5.5–8(−10) μm, fusoid with narrow ends, mostly straight, sometimes slightly curved, smooth, pale brown, 1-septate, becoming 3-septate after discharge, with hyaline appendages at each acute to subacute end; in some mature spores the appendage may be absent (Fig. 17b).

Anamorph: Pyrenochaeta sp. (Barr 1984; Samuels and Müller 1978).

Pycnidia 70–500 μm diam. Conidiogenous cells phialidic, lining cavity, 5–8 × 4–6 μm to 5–10 × 3–6 μm. Conidia 2.5–3.5(−4) × 1.5–2(−3) μm, hyaline, ellipsoid or subglobose (Barr 1984).

Material examined: ERIE, Dublin, Glasnevin Botanic Garden, on old rope, Jun. 1872, W. Keit (K(M):108784, holotype, as Sphaeria keitii Berk. & Broome).

Type species

Calyptronectria platensis Speg., Anal. Mus. nac. Hist. nat. B. Aires 19: 412 (1909). (Fig. 18)

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Ascomata 120–270 μm high × 170–400 μm diam., solitary, scattered, immersed, lenticular to subglobose, papillate, ostiolate (Fig. 18a and b). Apex with a small and slightly protruding papilla. Peridium 18–30 μm wide, comprising two types of cells, outer layer composed of pseudoparenchymatous cells, cells 3–6 μm diam., cell wall 1–2 μm thick, inner layer comprising less pigmented cells, merging with pseudoparaphyses (Fig. 18b and c). Hamathecium of long, filliform pseudoparaphyses, 1–2 μm broad, branching and anastomosing, embedded in mucilage. Asci 98–140 × 12.5–20 μm (\( \bar{x} = 107 \times 15.4\mu m \), n = 10), 8-spored, sometimes 4-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel, 12–20 μm long, with an ocular chamber (to 4 μm wide × 3 μm high) (Fig. 18e and f). Ascospores 17–22.5 μm × (6.3-)7.5–10 μm (\( \bar{x} = 19.8 \times 7.6\mu m \), n = 10), biseriate above and uniseriate below, ellipsoid to broadly fusoid with broadly to narrowly rounded ends, hyaline, usually with (3-)5 transverse septa, with or without 1–3 longitudinal septa in the central cells, constricted at the median septum, the upper cell often broader than the lower one, smooth, surrounded by an irregular hyaline gelatinous sheath up to 3 μm thick (in dry specimen) (Fig. 18d).

Anamorph: none reported.

Material examined: ARGENTINA, La Plata, on decaying branches of Manihot carthaginensis (Jacq.) Müll., Sept. 1906, Spegazzini (LPS 1209, holotype).

Type species

Carinispora nypae K.D. Hyde, J. Linn. Soc., Bot. 110: 99 (1992). (Fig. 19)

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One or two ascomata per stroma. Ascomata up to 0.8 mm diam., scattered or in small groups, developing beneath the host epidermis, crust-like, as circular spots, wall brown, with a small central ostiole, in section 225–285 μm high × 510–750 μm diam., lenticular, ostiolar canal lacking periphyses (Fig. 19a and b). Peridium 35–45 μm wide at sides, pale brown, at sides composed of a thin layer of thin-walled elongate cells, fusing with the stromatic tissue and host cells, at the base composed of thick-walled cells, forming a textura epidermoidea and fusing with host cells. A wedge of pale brown hyphae forming a textura porrecta is present at the rim (Fig. 19c). Hamathecium of dense, long filliform pseudoparaphyses 1–3 μm broad, embedded in mucilage, anastomosing between and above the asci, rarely septate. Asci 142–207 × 14.2–19.8 μm, 8-spored, bitunicate, fissitunicate, clavate to cylindrical, with a furcate pedicel, up to 40 μm long, apex with an ocular chamber and apical ring (to 2 μm wide × 3 μm high, J-), developing from ascogenous tissue at the base of the ascocarp (Fig. 19d, e, f, g and h). Ascospores 42–66 × 7–10.6 μm, biseriate, narrowly fusoid with broadly to narrowly rounded ends, somewhat curved, yellow to pale brown, yellow in mass, 7-8-septate, constricted at the septa, the two central cells being the largest, surrounded by a gelatinous sheath; the sheath has a central “spine” and curved polar extrusions (Fig. 19i and j).

Anamorph: none reported.

Material examined: BRUNEI DARUSSALAM, Tungit Api Api mangrove, from decaying intertidal fronds of Nypa fruticans Wurmb., 14 Apr. 1987, K.D. Hyde (BRIP 17106, holotype).

Type species

Caryosporella rhizophorae Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94: 356 (1985). (Fig. 20)

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Ascomata 0.8–1.1 mm high × 0.9–1.2 mm diam., densely scattered or gregarious, superficial with a flattened base, not easily removed from the host surface, subglobose, black, short papillate, ostiolate, periphysate, carbonaceous (Fig. 20a and b). Peridium 120–150 μm thick at sides, up to 200 μm thick at the apex, thinner at the base, 3-layered, outer layer composed of golden-yellow, very thick-walled cells of textura epidermoidea, mixed with subglobose, large cells near the surface, cells 7–15 μm diam., middle layer composed of deep brown, very thick-walled cells of textura epidermoidea, inner layer composed of hyaline, thin-walled cells of textura prismatica, up to 50 × 5 μm diam., merging with pseudoparaphyses (Fig. 20b, c and d). Hamathecium of dense, long trabeculate pseudoparaphyses, 1.5-2 μm wide, anastomosing and branching above the asci. Asci 225–250(−275) × 14–17 μm (\( \bar{x} = 137 \times 16.3\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a long, narrowed, pedicel which is up to 75 μm long, apical characters not observed (Fig. 20e). Ascospores 25–28(−30) × 9–13 μm (\( \bar{x} = 26.8 \times 11\mu m \), n = 10), uniseriate to partially overlapping, ellipsoidal to broadly fusoid with narrow hyaline rounded ends, deep reddish brown, thick-walled, 1-septate with hyaline germ pore at each end, slightly constricted at the septum, verruculose, sometimes with “net”-like ridged ornamentations (Fig. 20f and g).

Anamorph: suspected spermatia (Kohlmeyer 1985).

Material examined: BELIZE, Twin Cays, tip of prop root of Rhizophora mangle, 18 Mar. 1984, J. Kohlmeyer (NY. Herb. J. Kohlmeyer No. 4532a, holotype).

Type species

Chaetomastia hirtula (P. Karst.) Berl., Icon. fung. (Abellini) 1: 38 (1890). (Fig. 21)

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≡ Sphaeria hirtula P. Karst., Fungi Fenn. Exs. N. 825 (1869).

Ascomata 214–286 μm high × 210–258 μm diam., scattered or in groups, superficial, globose, wall black; apex often opening with a broad pore within slightly raised papilla, up to 30 μm diam., coriaceous (Fig. 21a). Peridium 20–26 μm thick, 1-layered, composed of heavily pigmented cells of textura angularis, cells up to 5 × 15 μm diam., cell wall up to 3.5 μm thick (Fig. 21b). Hamathecium of dense, long cellular pseudoparaphyses, embedded in mucilage. Asci 90–130 × 12.5–17.5(−22.5) μm (\( \bar{x} = 111 \times 16.3\mu m \), n = 10), mostly 4-spored, bitunicate, fissitunicate, broadly cylindrical, with a furcate pedicel, 18–48 μm long, with a large ocular chamber best seen in immature asci (to 3 μm wide × 3 μm high) (Fig. 21c and d). Ascospores 20.5–27 × 7–10 μm (\( \bar{x} = 23.5 \times 8.2\mu m \), n = 10), uniseriate to partially overlapping, ellipsoid to broadly fusoid with broadly to narrowly rounded ends, brown, 3-septate, verruculose, constricted at all septa, constricted at the median septum, the cell above the central septum often broader than the others (Fig. 21e and f).

Anamorph: none reported.

Material examined: FINLAND, ETELÄ-HÄME (EH/Ta), Tammela, Mustiala, På Rub. id., 8 May 1866. P.A. Karsten (H, FFE 825, kleptotype).