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)
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)
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)
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)
≡ 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)
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)
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)
≡ 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)
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)
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)
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)
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)
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)
≡ 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)
Generic description
Habitat terrestrial, saprobic. Ascomata solitary, superficial, globose, dark brown, epapillate, ostiolate. Peridium thin, pseudoparenchymatous. Hamathecium of few, cellular pseudoparaphyses, embedded in mucilage, rarely anastomosing and branching. Asci bitunicate, fissitunicate, broadly clavate with short pedicels, 2-3-spored. Ascospores muriform, broadly ellipsoid, dark brown with subhyaline end cells, verrucose.
Anamorphs reported for genus: none.
Literature: Barr 1987b; Hawksworth et al. 1979; Lumbsch and Huhndorf 2007.
Type species
Bimuria novae-zelandiae Hawksworth, Chea & Sheridan, N. Z. J. Bot. 17: 268 (1979). (Fig. 14)
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).
Notes
Morphology
Bimuria novae-zelandiae was first isolated from soil of a barley field in New Zealand (Hawksworth et al. 1979). Based on B. novae-zelandiae, the genus is characterized by a very thin peridium, mostly 2-spored and fissitunicate asci as well as the muriform, dark brown, verrucose ascospores (Hawksworth et al. 1979). Because of its unique morphological characters, the familial placement of this genus has been debatable and it has been placed in Pleosporaceae (Hawksworth et al. 1979), in Phaeosphaeriaceae (Barr 1987b) and in Melanommataceae (Lumbsch and Huhndorf 2007).
Morphologically, Bimuria is most comparable with some superficially similar or allied genera, in particular Montagnula (Hawksworth et al. 1979). However, the thick carbonaceous peridium distinguishes Montagnula from that of Bimuria (Hawksworth et al. 1979). In addition, the ascospores of Montagnula are discharged forcibly through the ostiole instead of forming a mass outside of the ostiole as in Bimuria (Hawksworth et al. 1979). Ascomauritiana lignicola V.M. Ranghoo & K.D. Hyde has somewhat similar ascospores in 4-spored asci, but this taxon has unitunicate asci (Ranghoo and Hyde 1999). The morphological characters of Bimuria, such as ascospore release and large, thick-walled ascospores may be an adaptation to its soil-borne habitat (Hawksworth et al. 1979).
Phylogenetic study
Bimuria novae-zelandiae was found to be closely related to Phaeodothis winteri (Niessl) Aptroot (syn. Didymosphaerella opulenta (De Not.) Checa & M.E. Barr) and Montagnula opulenta (De Not.) Aptroot in analysis of combined sequences, i.e. SSU rDNA, LSU rDNA, RPB2 and TEF1 sequences (Schoch et al. 2006, 2009). These two species had been included by Barr (2001) in her new family Montagnulaceae.
Concluding remarks
We agree with Barr (2001) and include the genus in Montagnulaceae based on both morphological and phylogenetic characters.
Bricookea M.E. Barr, Mycotaxon 15: 346 (1982). (?Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic (or parasitic?). Ascomata small- to medium-sized, solitary, scattered, or in small groups, immersed, erumpent to superficial, depressed globose, papillate, ostiolate. Peridium thin. Hamathecium filliform, cellular pseudoparaphyses, embedded in mucilage, anastomosing, septate. Asci bitunicate, fissitunicate, cylindrical, cylindro-clavate or slightly obclavate, with a short knob-like pedicel, with an ocular chamber. Ascospores hyaline, ellipsoid to narrowly obovoid, 3-septate, constricted at each septum.
Anamorphs reported for genus: none.
Literature: Barr 1982a; Berlese 1896; Holm 1957; Shoemaker and Babcock 1989a.
Type species
Bricookea sepalorum (Vleugel) M.E. Barr, Mycotaxon 15: 346 (1982). (Fig. 15).
≡ 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).
Notes
Morphology
Bricookea was formally established by Barr (1982a) as a monotypic genus represented by B. sepalorum based on its “globose to depressed ascomata, slit-like ostiole with labial cells, bitunicate asci, cellular pseudoparaphyses, and hyaline septate ascospores”. Bricookea was morphologically assigned to Phaeosphaeriaceae. Holm (1957) checked the authentic collections from North America and type material from Europe, and observed that the ascospores of collections from North America were significantly larger than those from the type material from Sweden. Thus, Shoemaker and Babcock (1989a) considered that the collections from North America represented a new species, which they introduced as B. barrae Shoemaker & C.E. Babc. Although the short slit-like ostiole has previously been reported (Shoemaker and Babcock 1989a), it is inconspicuous in the type specimen from Sweden. Currently, only two species are accommodated in this genus.
Phylogenetic study
None.
Concluding remarks
The knob-shaped pedicel, slit-like ostiole, hyaline ascospores as well as the herbaceous substrate all disagree with any current pleosporalean family. Thus, we temporarily retain this genus under Phaeosphaeriaceae until DNA sequence comparisons can be carried out.
Byssolophis Clem., in Clements & Shear, Gen. fung., Edn 2 (Minneapolis): 286 (1931). (Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, gregarious, semi-immersed to erumpent, coriaceous, ovoid, with a conspicuous elongate slit-like ostiole on the top. Peridium not observed. Hamathecium of dense, long pseudoparaphyses, anastomosing and branching between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical or cylindro-clavate, with a furcate pedicel. Ascospores fusoid, hyaline, turning faintly brown when old, 1-septate, with a short terminal appendage at each end.
Anamorphs reported for genus: none.
Literature: Clements and Shear 1931; Holm 1986; Müller and von Arx 1962.
Type species
Byssolophis byssiseda (Flageolet & Chenant.) Clem., Gen. Fung. (Minneapolis): 286 (1931). (Fig. 16)
≡ 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).
Notes
Morphology
Byssolophis was introduced as a monotypic genus based on B. byssiseda, which is characterized by its semi-immersed, gregarious, ovoid ascomata, with a conspicuous central apical ostiolar slit (Holm 1986). Subsequently, two more species were introduced, viz. B. ampla (Berk. & Broome) L. Holm and B. sphaerioides (P. Karst.) E. Müll. (Holm 1986; Müller and von Arx 1962).
Phylogenetic study
The current phylogeny places Byssolophis sphaerioides in proximity of Hypsostromataceae without resolving any sister taxa (Plate 1).
Concluding remarks
The slit-like ostiole, cylindrical asci, hyaline and 1-septate ascospores as well as the form of pseudoparaphyses are similar to species in Lophiostoma. Thus, Byssolophis may be a synonym of Lophiostoma.
Byssosphaeria Cooke, Grevillea 7: 84 (1879). (Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, scattered to gregarious, superficial, globose, subglobose to turbinate, non papillate with white, orange, red or green ostiolar region, wall black. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage, anastomosing between and above the asci. Asci bitunicate, fissitunicate, clavate to nearly cylindrical, with a furcate pedicel. Ascospores fusoid with narrow ends, straight or slightly curved, brown, 1-septate when young.
Anamorphs reported for genus: Pyrenochaeta or Chaetophoma-like (Barr 1984; Hawksworth et al. 1995; Samuels and Müller 1978).
Literature: von Arx and Müller 1975; Barr 1984; Boise 1984; Bose 1961; Chen and Hsieh 2004; Cooke and Plowright 1879; Hyde et al. 2000; Luttrell 1973; Mugambi and Huhndorf 2009b; Müller and von Arx 1962; Samuels and Müller 1978.
Type species
Byssosphaeria keitii (Berk. & Broome) Cooke [as ‘Byssosphaeria keithii’], (1879). (Fig. 17)
≡ 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).
Notes
Morphology
Byssosphaeria was introduced by Cooke and Plowright (1879) based on its superficial ascomata seated on a “tomentose subiculum of interwoven threads”, which includes various species in Sphaeria and Byssisedae, and was validly typified by B. keitii (Cooke 1878). Byssosphaeria keitii was treated as a synonym of B. schiedermayeriana (Fuckel) M.E. Barr by Sivanesan (1971), and B. schiedermayeriana exclusively occurs in tropical regions or greenhouse environments in temperate regions (Barr 1984). Morphologically, B. keitii is characterized by its large ascomata with orange to reddish plain apices, and is closely related to B. rhodomphala (Berk.) Cooke (Barr 1984).
For a long time, Byssosphaeria was assigned to Herpotrichia sensu lato, and Byssosphaeria schiedermayeriana was renamed as H. schiedermayeriana Fuckel (von Arx and Müller 1975; Bose 1961; Luttrell 1973; Müller and von Arx 1962; Sivanesan 1971). After studying Herpotrichia in North America, Barr (1984) accepted a relatively narrow generic concept, Herpotrichia sensu stricto, and revived Byssosphaeria; this proposal is supported by phylogenetic study (Mugambi and Huhndorf 2009b). Currently Byssosphaeria comprises 32 species (http://www.mycobank.org, 08-01-2009).
Phylogenetic study
The monophyletic nature of Byssosphaeria is well demonstrated, as well as its familial status in Melanommataceae (Mugambi and Huhndorf 2009b).
Concluding remarks
Orange and greenish plain apices exist in the specimen we examined, which is different from records as “orange, bright or dull reddish plain apices” by Barr (1984). This might be because different specimens have different colours, or there may be a variation of apical colour within a single species, as both orange and green can coexist on the same ascoma (see Fig. 17a). The coloured apical rim, together with the trabeculate pseudoparaphyses as well as the presence of subiculum make Byssosphaeria readily distinguishable from other morphologically comparable genera, e.g. Herpotrichia and Keissleriella (Hyde et al. 2000).
Calyptronectria Speg., Anal. Mus. nac. Hist. nat. B. Aires 19: 412 (1909). (Melanommataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to medium-sized, solitary, scattered, or in small groups, immersed, lenticular to subglobose, papillate, ostiolate. Hamathecium of long, filliform pseudoparaphyses, branching and anastomosing, embedded in mucilage. Asci 4- to 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel. Ascospores muriform, broadly fusoid to fusoid with broadly to narrowly rounded ends, hyaline.
Anamorphs reported for genus: none.
Literature: Barr 1983; Rossman et al. 1999; Spegazzini 1909.
Type species
Calyptronectria platensis Speg., Anal. Mus. nac. Hist. nat. B. Aires 19: 412 (1909). (Fig. 18)
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).
Notes
Morphology
Calyptronectria is a relatively poorly studied genus, which was formally established based on C. argentinensis Speg. and C. platensis, with C. platensis being chosen as the generic type (Spegazzini 1909). Morphologically, Calyptronectria is characterized by its immersed ascomata, trabeculate pseudoparaphyses and hyaline, muriform ascospores as well as its peridium that turns reddish brown in KOH (Rossman et al. 1999) (not shown here). Subsequently, C. indica Dhaware was introduced from India, and Barr (1983) transferred Teichospora ohiensis Ellis & Everh. to Calyptronectria as C. ohiensis (Ellis & Everh.) M.E. Barr. However, this proposal is inappropriate as the type specimen of T. ohiensis is “unitunicate” (Barr 1983; Rossman et al. 1999). Subsequently, Rossman et al. (1999) transferred Calyptronectria ohiensis to Thyridium (as T. ohiense (Ellis & Everh.) Rossman & Samuels).
Phylogenetic study
None.
Concluding remarks
The immersed ascomata, trabeculate pseudoparaphyses, bitunicate asci, hyaline and muriform ascospores as well as the reaction of peridium to KOH (turns reddish brown) make it distinguishable from all other reported genera (Rossman et al. 1999). Thus Calyptronectria is a morphologically well defined genus.
Carinispora K.D. Hyde, J. Linn. Soc., Bot. 110: 97 (1992). (Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic. One or two ascomata per stroma. Ascomata scattered or in small groups, developing beneath the host epidermis, erumpent, lenticular, ostiolate, lacking periphyses. Peridium pale brown, composed of thin-walled elongated cells at the sides and thick-walled cells of textura epidermoidea at the base. Hamathecium of dense, long filliform pseudoparaphyses, embedded in mucilage, anastomosing between and above the asci, rarely septate. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindrical, with a short furcate pedicel, apex with an ocular chamber and apical ring. Ascospores biseriate, narrowly fusoid, yellow to pale brown, multi-septate, constricted at the septa, the two central cells being the largest, surrounded by a gelatinous sheath.
Anamorphs reported for genus: none.
Literature: Hyde 1992a, 1994b.
Type species
Carinispora nypae K.D. Hyde, J. Linn. Soc., Bot. 110: 99 (1992). (Fig. 19)
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).
Notes
Morphology
Carinispora is distinguished from Phaeosphaeria by its saprobic life style and lenticular ascomata formed under the host epidermis, peridium structure and sheath surrounding the ascospores (Hyde 1992a, 1994b). Two species were reported, i.e. C. nypae and C. velatispora K.D. Hyde.
Phylogenetic study
Suetrong et al. (2009) could not resolve Carinispora nypae in a phylogeny based on four genes.
Concluding remarks
Both Carinispora nypae and C. velatispora are reported as marine fungi, which should be taken into consideration for their familial placement.
Caryosporella Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94: 355 (1985). (?Melanommataceae)
Generic description
Habitat marine, saprobic. Ascomata densely scattered or gregarious, superficial, subglobose, black, papillate, ostiolate, periphysate, carbonaceous. Peridium carbonaceous. Hamathecium of dense, long trabeculate pseudoparaphyses, anastomosing and branching above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical. Ascospores ellipsoidal to broadly fusoid with narrowly hyaline rounded ends, deep reddish brown, thick-walled, 1-septate with hyaline germ pore at each end.
Anamorphs reported for genus: suspected spermatia (Kohlmeyer 1985).
Literature: Eriksson 2006; Kohlmeyer 1985; Lumbsch and Huhndorf 2007.
Type species
Caryosporella rhizophorae Kohlm., Proc. Indian Acad. Sci., Pl. Sci. 94: 356 (1985). (Fig. 20)
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).
Notes
Morphology
Caryosporella was formally established by Kohlmeyer (1985) based on the obligate marine fungus, C. rhizophorae, which is characterized by its superficial ascomata, 3-layered peridium, filliform trabeculate pseudoparaphyses, and brown, 1-septate ascospores. Caryosporella was originally assigned to Massariaceae despite several major differences, such as the superficial ascomata, reddish brown ascospores (Kohlmeyer 1985). Subsequently, Caryosporella was assigned to Melanommataceae (Eriksson 2006; Lumbsch and Huhndorf 2007).
Phylogenetic study
Suetrong et al. (2009) showed that a single isolate of Caryosporella rhizophorae does not reside in Pleosporales, but is related to Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. & Volkm.-Kohlm. and placed in Dothideomycetidae incertae sedis.
Concluding remarks
As an obligate marine fungus, the familial placement of Caryosporella rhizophorae is uncertain but it may not belong to Pleosporales.
Chaetomastia (Sacc.) Berl., Icon. fung. (Abellini) 1: 38 (1890). (Teichosporaceae)
≡ Melanomma subgen. Chaetomastia Sacc., Syll. fung. (Abellini) 2: 113 (1883).
Generic description
Habitat terrestrial, saprobic. Ascomata relatively small, scattered, or in small groups, superficial, globose or subglobose, black, papillate, ostiolate, coriaceous. Peridium relatively thin, 1-layered, composed of heavily pigmented cells of textura angularis. Hamathecium of dense, long cellular pseudoparaphyses, embedded in mucilage. Asci mostly 4-spored, bitunicate, fissitunicate, broadly cylindrical with a furcate pedicel, with a large ocular chamber, especially apparent in immature asci. Ascospores ellipsoid to broadly fusoid with broadly to narrowly rounded ends, brown, 3-septate, constricted at all septa.
Anamorphs reported for genus: coelomycetous where known: conidia hyaline or brown, aseptate or 1-septate (Aposphaeria- or Coniothyrium-like) (Barr 1989c).
Literature: Barr 1987b, 1989c; 1993a; b; 2002; Berlese 1890; Clements and Shear 1931; Eriksson 1999; Eriksson and Hawksworth 1987, 1998; Holm 1957; Leuchtmann 1985; Saccardo 1883.
Type species
Chaetomastia hirtula (P. Karst.) Berl., Icon. fung. (Abellini) 1: 38 (1890). (Fig. 21)
≡ 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).
Notes
Morphology
Chaetomastia was introduced by Saccardo (1883) as a subgenus of Melanomma, and five species were included, i.e. M. canescens Speg., M. cucurbitarioides Speg., M. hirtulum (P. Karst.) Sacc., M. hispidulum Sacc. and M. pilosellum P. Karst. Berlese (1890) promoted it to genus rank. Subsequently, Chaetomastia hirtula (P. Karst.) Berl. was selected as the lectotype species of the genus (Clements and Shear 1931). Chaetomastia has been regarded as having unitunicate asci (Eriksson and Hawksworth 1986, 1998; Eriksson 1999). However its bitunicate status was confirmed by Holm (1957). Holm (1957) treated C. hirtula as Melanomma hirtulum (P. Karst.) Sacc., and Leuchtmann (1985) transferred this species to Montagnula sensu lato based on the ascospore morphology and the hyphae surrounding the ascomata. Barr (1987b) suggested that ascoma, peridium structure and ascospore characters pointed Montagnula sensu stricto to Phaeosphaeriaceae, while the characters of ascomata and peridium structure of Chaetomastia were thought to fit the definition of Dacampiaceae (Barr 1987b). In particular, the peridium and ascospore characters of C. hirtula are comparable with those of the generic type of Massariosphaeria (M. phaeospora). Thus, Barr (1989c) accepted Massariosphaeria sensu stricto and assigned the phragmosporous species of Massariosphaeria sensu lato to Chaetomastia.
Barr (2002) later assigned Chaetomastia to Teichosporaceae based on its saprobic or hypersaprobic lifestyle, occurring on woody stems and peridium structure, and this is widely followed (Eriksson 2006; Lumbsch and Huhndorf 2007). Currently, 11 species are accepted in this genus (http://www.indexfungorum.org/).
Phylogenetic study
None.
Concluding remarks
Familial placement of Chaetomastia is undetermined currently but has been included in the Teichosporaceae by authoritative sources (Eriksson 2006; Lumbsch and Huhndorf 2007) or the Dacampiaceae (http://www.indexfungorum.org/).
Chaetoplea (Sacc.) Clem., Gen. Fung. (Minneapolis): 275 (1931). (?Phaeosphaeriaceae)
≡ Pyrenophora subgen. Chaetoplea Sacc., Syll. fung. (Abellini) 2: 279 (1883).
Generic description
Habitat terrestrial, saprobic. Ascomata small to medium, immersed, erumpent to superficial, globose to subglobose, papillate, ostiolate. Peridium not examined. Hamathecium of dense, long, narrowly cellular pseudoparaphyses. Asci 8-spored or 4-spored, bitunicate, fissitunicate, cylindro-clavate, with a thick, furcate pedicel. Ascospores ellipsoid or fusoid, pale brown to brown, phragmosporous or muriform.
Anamorphs reported for genus: Microdiplodia-like (Barr 1990b).
Literature: Barr 1981; 1987a; b; 1990b; Clements and Shear 1931; Ramaley and Barr 1995; Yuan and Barr 1994.
Type species
Chaetoplea calvescens (Fr.) Clem., Gen. Fung. (Minneapolis): 275 (1931). (Fig. 22)
≡ Sphaeria calvescens Fr. Scleromyc. Sueciae 401.
Ascomata not examined. Peridium not examined. Hamathecium of dense, long, narrow cellular pseudoparaphyses, 2–3 μm broad, septate, branching and anastomosing. Asci 90–110 × 10–12 μm, 8-spored, rarely 4-spored, bitunicate, fissitunicate, cylindro-clavate, with a thick, furcate pedicel which is up to 30 μm long (Fig. 22a and b). Ascospores 13–18 × 5.5–7 μm, obliquely uniseriate and partially overlapping, broadly fusoid to oblong with broadly rounded ends, pale brown, 2-3-septate, constricted at the septa, containing four refractive globules (Fig. 22c).
Note: The specimen is only a slide, and no peridium or ascomata information could be obtained.
Anamorph: coelomycetous, conidia yellowish, 1-septate, 9–13 × 4–5(−8) μm (Webster and Lucas 1959); Microdiplodia henningsii Staritz=Chaetodiplodia caudina Karst. (Sutton 1980) (referred to Barr 1990b (p50)).
Material examined: SWEDEN, sub-collection: Curtis Herbarium, verified by R.A. Shoemaker, leg. E.M. Fries 401 (FH-81113, isotype, microscope slide).
Notes
Morphology
Chaetoplea was introduced based on C. calvescens, which has been regarded as similar to Pleospora or Leptosphaeria (Eriksson and Hawksworth 1987; Wehmeyer 1961; von Arx and Müller 1975). Based on the differences in ascomata, peridium structure, pseudoparaphyses as well as its anamorphic stage, Chaetoplea was maintained as a separate genus (Barr 1990b; Yuan and Barr 1994). Chaetoplea sensu lato was accepted by Barr (1990b), which included some species of Teichospora as well as the subgenus Pleospora subg. Cylindrosporeae.
The following is from the label of specimen.
“Sphaeria calvescens, Scler. Suecicae (Ed. 2) 401. No specimen of Scler. Suecicae 401 is now at Uppsala according to R. Santesson 1966. This Curtis Herbarium specimen in the Farlow Herbarium is isotype. Wehmeyer (1961) in his Pleospora monograph did not study any portion of the Scler. Suecicae exsiccatus 401, nor did Webster & Lucas in the taxonomic and life-history study (Trans. Brit. Myc. Soc. 42, 332–342. 1959) of this species.
The specimen has most of the features described by Webster & Lucas including the presence of the conidial state Microdiplodia henningsii Staritz. I did not see vertical septa in the ascospores. Webster & Lucas note that vertical septa may be occasionally be lacking. The fungus is otherwise as they describe it although some perithecia collapse and appear cupulate.”—by R.A. Shoemaker.
Phylogenetic study
None.
Concluding remarks
The substrate of Chaetoplea sensu Barr (1990b) can be herbaceous stalks, decorticated wood or periderm, or old cotton cloth and string, which may indicate its heterogeneous nature. The ascospores seem very much like Phaeosphaeria which may be an earlier name; more details concerning the ascomatal, peridial and hamathecial structures are needed to make any conclusion.
Cilioplea Munk, Dansk botanisk Arkiv 15: 113 (1953). (Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to medium-sized, solitary, scattered or in small groups, immersed, globose or subglobose, papilla covered with short and blackish setae, coriaceous. Peridium thin, comprising small heavily pigmented thick-walled cells of textura angularis. Hamathecium of cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, furcate pedicel, and small ocular chamber. Ascospores fusoid to narrowly fusoid with narrowly rounded ends, pale brown to reddish brown, multi-transverse septa, usually with one longitudinal septum in some central cells, constricted at the primary septum.
Anamorphs reported for genus: none.
Literature: Barr 1990b, 1992b; Crivelli 1983; Lumbsch and Huhndorf 2007; Müller 1951; Munk 1953, 1957.
Type species
Cilioplea coronata (Niessl) Munk, Dansk botanisk Arkiv 15: 113 (1953). (Fig. 23)
≡ Pleospora coronata Niessl, Notiz. Pyr.: 16 (1876).
Ascomata 170–290 μm high × 200–410 μm diam., solitary, scattered, or in small groups, immersed, globose or subglobose, wall black, papilla raised, 50–80 μm high, with short and blackish setae, coriaceous (Fig. 23a). Peridium 9–15 μm thick laterally, up to 28 μm thick at the apex, thinner at the base, 1-layered, composed of small heavily pigmented thick-walled cells of textura angularis, cells up to 4 × 2.5 μm diam., cell wall 2–3 μm thick, apex cells smaller and walls thicker (Fig. 23b). Hamathecium of long cellular pseudoparaphyses, 2–3 μm broad. Asci (60-)80–108 × 10–15 μm (\( \bar{x} = 85.3 \times 12.1\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, thick, furcate pedicel, 5–15 μm long, and a small ocular chamber (to 3 μm wide × 2 μm high) (Fig. 23c and d). Ascospores 21–27.5 × 5.5–7.5 μm (\( \bar{x} = 24 \times 6.7\mu m \), n = 10), biseriate to uniseriate at base, fusoid to narrowly fusoid with narrowly rounded ends, pale reddish brown, 5–7 transverse septa (mostly 5), usually with one longitudinal septum in some central cells, deeply constricted at the median septum, the part above the primary septum shorter and broader, smooth-walled.
Anamorph: none reported.
Material examined: GERMANY, Hadiberg. on Reseda lutea Hadiberg, 20 Sept. 1875, Niessl (M 175-89-290, lectotype; M 175-89-291, type).
Notes
Morphology
Cilioplea was introduced by Müller (1951) as a subgenus of Pleospora, and this was followed by Munk (1957), who had earlier proposed it as a separate genus typified by C. coronata based on its hairy papilla, clavate asci as well as its “perfectly paraphysoid” (see Munk 1953). A relatively narrow concept of Pleospora was accepted by Crivelli (1983), and four species was assigned under the separate genus Cilioplea, viz. C. coronata, C. genisticola (Fautrey & Lambotte) Crivelli, C. kansensis (Ellis & Everh.) Crivelli and C. nivalis (Niessl) Crivelli. Subsequently, another six species were added (Barr 1990b, 1992b). Currently, ten species are included under Cilioplea.
Phylogenetic study
None.
Concluding remarks
The most striking character of Cilioplea is its setose papilla, which has been shown to have no phylogenetic significance in Lentitheciaceae (Zhang et al. 2009a). Cilioplea was assigned under Lophiostomataceae (Lumbsch and Huhndorf 2007), but there is little morphological similarity with the Lophiostomataceae sensu stricto (Zhang et al. 2009a). Thus its familial placement needs further study.
Crivellia Shoemaker & Inderb., in Inderbitzin, Shoemaker, O’Neill, Turgeon & Berbee, Can. J. Bot. 84: 1308 (2006). (Pleosporaceae)
Generic description
Habitat terrestrial, hemibiotrophic or parasitic. Ascomata small- to medium-sized, scattered, immersed, erumpent to nearly superficial, papillate, ostiolate. Peridium thin, composed of two cells types, outer cells of thick walled and textura angularis, inner cells thin-walled, yellow. Hamathecium of dense, long and thin pseudoparaphyses. Asci
(4-)8-spored, bitunicate, fissitunicate dehiscence not observed, broadly cylindrical to cylindrical, with a short, furcate pedicel and an ocular chamber. Ascospores fusoid to broadly fusoid, pale brown, septate, sometimes with one or two vertical septa in the middle cells, constricted at the septa.
Anamorphs reported for genus: Brachycladium (Inderbitzin et al. 2006).
Literature: Inderbitzin et al. 2006.
Type species
Crivellia papaveracea (De Not.) Shoemaker & Inderb., Can. J. Bot. 84: 1308 (2006). (Fig. 24)
≡ Cucurbitaria papaveracea De Not., Sfer. Ital.: 62 (1863).
Ascomata 210–260 μm high × 300–380 μm diam., densely scattered, immersed, erumpent to nearly superficial, flattened globose, dark brown, papillate, ostiolate (Fig. 24a). Peridium 25–30 μm thick, thicker near the apex and thinner at the base, composed of two cell types, outer cells of thick-walled and textura angularis, cells up to 10 × 5 μm diam., cell wall 2–4 μm thick, inner cells thin-walled, yellow (Fig. 24b). Hamathecium of dense, long, 1–2 μm broad, rarely septate pseudoparaphyses. Asci 85–125 × 10–13 μm (\( \bar{x} = {1}0{6} \times {11}\mu {\text{m}} \), n = 10), (4-)8-spored, bitunicate, fissitunicate dehiscence not observed, broadly cylindrical to cylindrical, with a short, furcate pedicel, with a relatively large ocular chamber (Fig. 24c and d). Ascospores (16-)19–24 × 5–7.5 μm (\( \bar{x} = 20.4 \times 6.3\mu m \), n = 10), overlapping uniseriate to rarely biseriate, fusoid to broadly fusoid, pale brown, 3-septate, sometimes with one or two vertical septa in the middle cells, constricted at the septa, the upper cell often broader than the lower one, smooth-walled.
Anamorph: Brachycladium penicillatum (Corda) Fr. (Inderbitzin et al. 2006).
Material examined: AUSTRIA, Vienna, on decaying stems of Papaver rhoeas L., 28 Oct. 2001, W. Jaklitsch (UBC F14995, epitype).
Notes
Morphology
Crivellia was separated from Pleospora and introduced as a new genus by Inderbitzin et al. (2006) based on their differences in ascospore morphology and anamorphic stages. Crivellia is characterized by having small- to medium-sized ascomata, and yellow, 3-septate ascospores with one or two vertical septa in central cells. Its Brachycladium anamorphic stage with phragmosporous conidia also differs from that of Stemphylium, which is the anamorphic stage of Pleospora (Inderbitzin et al. 2006). Currently, two species are included within Crivellia, i.e. C. homothallica Inderb. & Shoemaker and C. papaveracea.
Phylogenetic study
Crivellia papaveracea was shown to be closely related to some species of Alternaria, and its pleosporaceous status was confirmed following molecular studies (Inderbitzin et al. 2006).
Concluding remarks
Crivellia seems to belong to Pleosporaceae, and may be closely related to Pleospora.
Decaisnella Fabre, Annls Sci. Nat., Bot., sér. 6 9:112 (1878). (Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata medium to large, immersed to erumpent, clypeate, papillate, ostiolate. Hamathecium of dense, long, cellular pseudoparaphyses, rarely septate, embedded in mucilage. Asci mostly 4- or 8-spored, rarely 2-spored, cylindrical to cylindro-clavate, with a furcate pedicel. Ascospores muriform, dark brown, oblong with broadly rounded ends.
Anamorphs reported for genus: none.
Literature: Barr 1986; 1990a; b; Fabre 1878; Saccardo 1883.
Type species
Decaisnella spectabilis Fabre, Annls Sci. Nat., Bot., sér. 6 9: 112 (1879). (Fig. 25)
Ascomata 520–680 μm high × 430–600 μm diam., solitary, scattered, or in small groups of 2–3, immersed to erumpent, clypeate, globose or subglobose, black, roughened, with a blunt papilla up to 170 μm high, apex with a round ostiole, coriaceous (Fig. 25a). Peridium 70–90 μm thick at sides, thicker near the apex, comprising two types of cells; part immersed in host tissue, outer layer pseudoparenchymatous, 55–65 μm thick, pigmented, inner layer composed of lightly pigmented to hyaline thin-walled compressed cells, 15–23 μm thick, cells 3.5–7 μm diam., part above host tissue heavily pigmented covered by clypeus tissues (Fig. 25b). Hamathecium of dense, long, cellular pseudoparaphyses, 1.5–3 μm broad, rarely septate, embedded in mucilage. Asci 150–200 × 15–25(−33) μm (\( \bar{x} = 181 \times 20.6\mu m \), n = 10), (2-)4-spored, bitunicate, fissitunicate, broadly cylindrical, with a short, thick, furcate pedicel which is 20–40 μm long, no apical apparatus observed (Fig. 25e). Ascospores 37–45 × 12–17 μm (\( \bar{x} = 43 \times 15\mu m \), n = 10), uniseriate and sometimes slightly overlapping, oblong with broadly rounded ends, dark brown, verrucose or smooth, 7–9 transverse septa and 1–3 longitudinal septa in some of the cells, no constriction at the septa (Fig. 25c and d).
Anamorph: none reported.
Material examined: GERMANY, Valsalpe in der Ramsau, Bayer, Alpen, on Rhamnus pumila Turra., Jul. 1913, Karl Arnold (NY2082, syntype as Teichospora megalocarpa Rehm).
Notes
Morphology
Decaisnella was formally established by Fabre (1879), but was treated as a synonym of Teichospora by Saccardo (1883). This was followed by several mycologists over a long time. The main morphological differences between Decaisnella and Teichospora include the size and septation of ascospores, shape of ascomata, structure of peridium and type of pseudoparaphyses (Barr 1986). Thus Barr (1986) revived Decaisnella and assigned it to Massariaceae based on the shape of ascomata and large, distoseptate ascospores. Currently, 15 species are accepted under Decaisnella (http://www.mycobank.org/MycoTaxo.aspx). Neither the size of ascomata nor the ascospore characters have proven sufficient to place taxa at the family level in Pleosporales (Zhang et al. 2009a), and therefore familial placement of Decaisnella remains uncertain.
Phylogenetic study
Decaisnella formosa resided in the clade of Lophiostomataceae and in proximity to Lophiostoma macrostomoides De Not. (Plate 1).
Concluding remarks
The muriform ascospores, saprobic life style and 4-spored asci point Decaisnella spectabilis to Montagnulaceae, but this can only be confirmed following a molecular phylogenetic study.
Delitschia Auersw., Hedwigia 5: 49 (1866). (Delitschiaceae)
Generic description
Habitat terrestrial, saprobic (coprophilous). Ascomata medium- to large-sized, solitary or scattered, immersed to erumpent, globose or subglobose, apex with or without papilla, ostiolate. Peridium thin, composed of compressed cells. Hamathecium of dense, long pseudoparaphyses, anastomosing and branching. Asci 8-spored, cylindrical to cylindro-clavate, with short pedicel. Ascospores uni- to triseriate, pale to dark brown, ellipsoid, 1-septate, usually constricted at the septum, smooth, with a full length germ slit in each cell.
Anamorphs reported for genus: none.
Literature: Auerswald 1866; Barr 2000; Cain 1934; Dennis 1968; Eriksson 2006; Griffiths 1901; Hyde and Steinke 1996; Kirschstein 1911; Kruys et al. 2006; Luck-Allen and Cain 1975; Lumbsch and Huhndorf 2007; Moreau 1953; Munk 1957; Romero and Samuels 1991; Schoch et al. 2006; Winter 1887.
Type species
Delitschia didyma Auersw., Hedwigia 5: 49 (1866). (Fig. 26)
Ascomata 400–800 μm diam., solitary or scattered, immersed, globose or subglobose, black, papilla short, 70–130 μm broad, central, with a wide opening, coriaceous (Fig. 26a). Peridium ca. 15 μm thick laterally, up to 35 μm thick at the apex, up to 30 μm at the base, comprising a single layer of small lightly pigmented thin-walled cells of textura angularis, cells 4–10 μm diam., cell wall <1 μm thick, apex cells smaller and wall thicker (Fig. 26b). Hamathecium of dense, very long pseudoparaphyses, 1.5–2 μm broad, anastomosing and branching. Asci 290–380 × 35–45 μm (\( \bar{x} = 357.5 \times 40.6\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with short, narrowed pedicels which are rounded at the base, 25–60 μm long, apex with a wide ocular chamber (Fig. 26d and e). Ascospores 50–58 × 20–22.5 μm (\( \bar{x} = 54 \times 21.3\mu m \), n = 10), obliquely uniseriate and partially overlapping, ellipsoid with narrowly rounded ends, reddish brown, 1-septate, slightly constricted at the septum, smooth-walled, each cell with a full length germ slit (Fig. 26c).
Anamorph: none reported.
Material examined: GERMANY, Near Königstein, in forest, rare, Oct. 1904, W. Krieger (L, 1950).
Notes
Morphology
Delitschia was established by Auerswald (1866), and assigned to Sphaeriaceae. It was considered to be closely related to Sordariaceae and Amphisphaeriaceae. Winter (1887) assigned Delitschia under Sordariaceae, and this placement is followed in several subsequent studies (Griffiths 1901; Kirschstein 1911). Cain (1934) suggested that Delitschia might belong in Pleosporaceae, and this proposal was supported by Moreau (1953) and Dennis (1968). Finally, Munk (1957) established Sporormiaceae (Pseudosphaeriales), and Delitschia was assigned therein. Luck-Allen and Cain (1975) reviewed and redefined the genus as having bitunicate asci, pigmented and 1-septate ascospores with an elongated germ slit in each cell and surrounded by a gelatinous sheath, and in particular, the coprophilous habitat. Luck-Allen and Cain (1975) accepted 46 species. Subsequently, some wood-inhabiting species were also described (Hyde and Steinke 1996; Romero and Samuels 1991). Three genera, i.e. Delitschia, Ohleriella and Semidelitschia were separated from Sporormiaceae, and a new family, Delitschiaceae, was introduced by Barr (2000) to accommodate them. Delitschiaceae is characterized by a periphysate ostiole, wide endotunicate asci with a wide ocular chamber and ascospores having cells with germ slits. Delitschiaceae has been subsequently accepted (Eriksson 2006; Lumbsch and Huhndorf 2007).
The genus comprises 83 names (Index Fungorum) and is estimated to comprise 51 species (Kirk et al. 2008). Keys to Delitschia can be found in Luck-Allen and Cain (1975) and Hyde and Steinke (1996).
Phylogenetic study
Delitschia didyma and D. winteri (W. Phillips & Plowr.) Sacc. form a robust phylogenetic clade within Delitschiaceae, which is basal to other members of Pleosporales (Kruys et al. 2006; Schoch et al. 2006) except for Massariaceae (Voglmayr and Jaklitsch 2011). This might indicate its early derivation (Zhang et al. 2009a).
Concluding remarks
Morphologically, Delitschia is a well defined genus, and each cell of the ascospore has a full length germ slit. Currently, most species of this genus are coprophilous, although a few species are reported from wood (Hyde and Steinke 1996; Luck-Allen and Cain 1975). Whether the lignicolous habitat is an important character that might separate these taxa from the main coprophilous group, needs to be addressed, however, the morphological characters are similar.
Didymosphaeria Fuckel, Jb. nassau. Ver. Naturk. 22–23: 140 (1870). (Didymosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or parasitic. Ascomata solitary, scattered, or in small groups, immersed to erumpent, globose to ovoid, papillate, ostiolate, periphysate. Ostiole with a pore-like opening. Peridium 1-layered, thin, composed of brown pseudoparenchymatous cells of textura angularis. Hamathecium of dense, trabeculate, anastomosing mostly above the asci. Asci (2-)4-spored or 8-spored, bitunicate, cylindrical, with a furcate pedicel. Ascospores uniseriate, ellipsoid, brown, 1-distoseptate.
Anamorphs reported for genus: Dendrophoma, Fusicladiella and Phoma (Aptroot 1995).
Literature: Aptroot 1995; Barr 1989a, b, 1990a, 1992a, b; 1993a; b; Fuckel 1870; Hawksworth 1985a, b; Hawksworth and Boise 1985; Hawksworth and Diederich 1988; Hyde et al. 2000; Lumbsch and Huhndorf 2007; Saccardo 1882; Scheinpflug 1958; Sivanesan 1984.
Type species
Didymosphaeria futilis (Berk. & Broome) Rehm, Hedwigia 18: 167 (1879). (Fig. 27)
≡ Sphaeria futilis Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 9: 326 (1852).
Ascomata 190–230 μm high × 240–340 μm diam., scattered, or in small groups, immersed to slightly erumpent, subglobose to ovoid, membraneous, near-hyaline, under clypeus, papillate, periphysate (Fig. 27a and c). Papilla central, up to 100 μm high, black, with a pore-like ostiole (Fig. 27a and c). Peridium 30–40 μm wide upper part, 6–23 μm wide near the base, 1-layered, composed of brown pseudoparenchymatous cells of textura angularis, cell wall 2–3 μm thick (Fig. 27b). Hamathecium of dense, long trabeculate pseudoparaphyses, 0.8–1.5 μm broad, anastomosing mostly above the asci, embedded in mucilage (Fig. 27d). Asci 90–110 × 7.5–10 μm (\( \bar{x} = 97 \times 9\mu m \), n = 10), 2–4-spored, rarely 8-spored, bitunicate, fissitunicate, cylindrical, with a furcate pedicel, 17.5–27.5 μm long, with a large ocular (to 2.5 μm wide × 4 μm high) (Fig. 27d, e and f). Ascospores 14–15.5 × (5.5-) 6–7.5 μm (\( \bar{x} = 14.8 \times 6.9\mu m \), n = 10), uniseriate, ellipsoid with obtuse ends, brown, 1-septate, distoseptate, slightly to not constricted, capitate (Fig. 27g).
Anamorph: Dendrophoma sp., Fusicladiella sp. vel aff. (Sivanesan 1984).
Material examined: UK, England, Norfolk, King’s Cliffe; on dead stem (in ramis emortuis) Rosa sp., Mar. 1850, M.J. Berkeley (K(M): 147683, holotype).
Notes
Morphology
Didymosphaeria is a widely distributed genus with wide host range (Aptroot 1995). Didymosphaeria was formally established by Fuckel (1870) based on six ascomycetous species, and D. epidermidis (Fries) Fuckel (or D. peltigerae Fuckel) has been chosen as the lectotype species (see comments by Aptroot 1995). Hawksworth and David (1989: 494) proposed to conserve the genus with a lectotype specimen, Fungi Rhenani 1770. The genus had been considered as a depository to accommodate all types of didymosporous pyrenocarpous ascomycetes. Many workers have tried to redefine the genus and excluded some species. Saccardo (1882) restricted the genus to brown-spored species, and about 100 species have been excluded subsequently (Barr 1989a, b, 1990a, 1992a, b, 1993b; Hawksworth 1985a, b; Hawksworth and Boise 1985; Hawksworth and Diederich 1988; Scheinpflug 1958). Over 400 epithets of Didymosphaeria were included until the monograph of Aptroot (1995).
Aptroot (1995) examined more than 3000 specimens under the name Didymosphaeria. The type specimen of Didymosphaeria (Fungi Rhenani 1770) represents the widespread and common D. futilis (Aptroot 1995). In this study, we did not get the lectotype specimen, but described the type of D. futilis (Sphaeria futilis). Using a narrow concept (ignoring differences of host or country of origin), Aptroot (1995) accepted only seven species, which were closely related with the generic type of Didymosphaeria with over 100 synonyms distributed among them. Many taxa were found to belong to other groups, i.e. Aaosphaeria, Amphisphaeria, Astrosphaeriella, Dothidotthia, Flagellosphaeria, Kirschsteiniothelia, Megalotremis, Montagnula, Munkovalsaria, Mycomicrothelia, Parapyrenis or Phaeodothis. Didymosphaeria is mainly characterized by a peridium consisting of flattened or irregular cells or completely hyphae; a hamathecium consisting of narrow, trabeculate paraphysoids or paraphyses, richly anastomosing above the asci; and brown thinly distoseptate ascospores. Didymosphaeriaceae was maintained as a separated family within Pleosporales by Aptroot (1995) because of the distoseptate ascospores and trabeculate pseudoparaphyses mainly anastomosing above the asci. This proposal, however, has not received much support (Lumbsch and Huhndorf 2007).
Phylogenetic study
There have been few molecular investigations of Didymosphaeria when compared to the morphological studies. Didymosphaeria futilis resided in the clade of Cucurbitariaceae (or Didymosphaeriaceae) (Plate 1). The correct identification of the Didymosphaeria strain used for sequencing, however, has not been verified.
Concluding remarks
Didymosphaeria is a well established genus represented by D. futilis. Of particular significance are the narrow pseudoparaphyses which anastomose above the asci and brown 1-septate ascospores with indistinct distosepta. Familial placement of Didymosphaeria is unclear yet because of insufficient molecular data.
Dothidotthia Höhn., Ber. Deutsch. Bot. Ges. 36: 312 (1918). (Didymellaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary, clustered or somewhat gregarious, erumpent, subglobose, apex somewhat papillate to depressed, coriaceous. Peridium composed of a few layers of dark brown cells of textura angularis, and giving rise dark brown, thick-walled hyphae in the basal region, 2-layered. Hamathecium septate pseudoparaphyses branched in upper part above asci. Asci 8-spored, bitunicate, clavate, straight to curved. Ascospores biseriate to obliquely uniseriate, ellipsoid, pale brown, 1-septate.
Anamorphs reported for genus: Dothiorella and Thyrostroma (Hyde et al. 2011; Phillips et al. 2008).
Literature: Barr 1989b; Phillips et al. 2008.
Type species
Dothidotthia symphoricarpi (Rehm) Höhn., Ber. Deutsch. Bot. Ges. 36: 312 (1918). (Fig. 28)
≡ Pseudotthia symphoricarpi Rehm, Ann. Mycol. 11: 169 (1913).
Ascomata up to 500 μm high × 550 μm diam., gregarious clustered, rarely solitary, erumpent, subglobose, apex somewhat papillate to depressed, coriaceous (Fig. 28a). Peridium 20–80 μm thick, composed of 3–6 layers of dark brown cells of textura angularis, giving rise dark brown, thick-walled hyphae in the basal region, 2-layered, outer layer wall thicker and inner layer wall thinner (Fig. 28b). Hamathecium hyaline, septate pseudoparaphyses, 2–3 μm wide, branched in upper part above asci. Asci 70–120 × 15–22 μm, 8-spored, bitunicate, clavate, straight to curved (Fig. 28c, d and e). Ascospores (20-)22–23(−26) × (8-)9–10(−11) μm, biseriate to obliquely uniseriate and partially overlapping, ellipsoid tapering towards subacutely rounded ends, pale brown, 1-septate, constricted at the septum, smooth (Fig. 28f) (description referred to Phillips et al. 2008).
Anamorph: Thyrostroma negundinis (Phillips et al. 2008).
Material examined: USA, North Dakota, on branches of Symphoricarpos occidentalis Hook. (NY, holotype); Colorado, San Juan Co, c. 0.5 mile up Engineer Mountain Trail from turnoff at mile 52.5, Hwy 550, dead twigs of Symphoricarpos rotundifolius A. Gray, 24 Jun. 2004, A.W. Ramaley 0410 (BPI 871823, epitype).
Notes
Morphology
Dothidotthia was formally established to accommodate Pseudotthia symphoricarpi (Montagnellaceae, Dothideales) (von Höhnel 1918a). Many mycologists considered Dothidotthia closely related to a genus of Venturiaceae such as Dibotryon by Petrak (1927), or Gibbera by von Arx and Müller (1954) and Müller and von Arx (1962). Dothidotthia had been treated as a synonym of Gibbera (von Arx 1954; Müller and von Arx 1962), which was followed by Shoemaker (1963) and Eriksson and Hawksworth (1987). Based on the coelomycetous anamorphic stage and peridium structure, shape of asci, as well as morphology of pseudoparaphyses, Barr (1987b, 1989b) retrieved Dothidotthia, and considered it closely related to Botryosphaeria (Botryosphaeriaceae). Currently, 11 species are included within Dothidotthia (http://www.mycobank.org, 01–2011).
Phylogenetic study
Based on a multi-gene phylogenetic analysis, Dothidotthia formed a separate familial clade (Phillips et al. 2008). Thus Dothidotthiaceae was introduced to accommodate it (Phillips et al. 2008).
Concluding remarks
By comparing the morphological characters and phylogenetic dendrograms by Phillips et al. (2008) and de Gruyter et al. (2009), Dothidotthia seems closely related to Didymellaceae, but Dothidotthiaceae should still be treated as a separate family.
Dubitatio Speg., Anal. Soc. cient. argent. 12: 212 (1881). (Arthopyreniaceae (or Massariaceae))
Generic description
Habitat terrestrial, saprobic. Ascomata medium-sized, solitary, densely scattered, or in small groups of 2–4, immersed, covered with white crystaline rim, papillate, ostiolate. Hamathecium of dense pseudoparaphyses, long, 2–3 μm broad, branching and anastomosing. Asci cylindrical, pedicellate, with furcate pedicel. Ascospores 1-septate, asymmetrical, reddish to dark brown.
Anamorphs reported for genus: Aplosporella-like (Rossman et al. 1999).
Literature: Barr 1979b, 1987b; Müller and von Arx 1962; Rossman et al. 1999; Spegazzini 1881.
Type species
Dubitatio dubitationum Speg., Anal. Soc. cient. argent. 12: 212 (1881). (Fig. 29)
Ascomata 350–530 μm high × 550–700 μm diam., solitary, densely scattered, or in small groups of 2–4, immersed, with a protruding papilla, 110–160 μm high, 160–250 μm diam., globose or subglobose, black, covered with white crystalline material which becomes hyaline and gel-like in water, ostiolate (Fig. 29a and b). Peridium 18–25 μm thick laterally (excluding the rim), up to 35 μm thick at the apex, thinner at the base, 1-layered, composed of small pale brown thin-walled cells of textura prismatica, cells 5–12 × 3–5 μm diam., cell wall up to 1 μm thick, apex cells smaller and walls thicker (Fig. 29b). Hamathecium of dense, long pseudoparaphyses, 2–3 μm broad, branching and anastomosing between and above the asci. Asci 150–190(−230) × 12.5–15 μm (\( \bar{x} = 172.5 \times 13.4\mu m \), n = 10), (6-)8-spored, rarely 4-spored, bitunicate, fissitunicate, cylindrical, with a furcate pedicel which is up to 40 μm long, ocular chamber not observed (Fig. 29c, d and e). Ascospores 19–22.5 × 10–12 μm (\( \bar{x} = 20.2 \times 11.4\mu m \), n = 10), uniseriate to obliquely uniseriate and partially overlapping, broadly ellipsoid with broadly to narrowly rounded ends, reddish brown, 1-septate, constricted at septum, asymmetric with a larger upper cell, thick-walled, possibly distoseptate (Fig. 29f, g and h).
Anamorph: Aplosporella-like (for detailed description see Rossman et al. 1999).
Conidiomata globose, ca. 300 μm diam. Conidia holoblastic, broadly fusoid, 13–15 × 7–10 μm, dark brown, finely spinulose (Rossman et al. 1999).
Material examined: ARGENTINA, Buenos Aires, Tuyu, on Celtis tala Gill., Jan. 1881, leg. det. C. Spegazzini (NY, isotype; LPS, holotype).
Notes
Morphology
When established Dubitatio, Spegazzini (1881) considered it as intermediate between Sphaeriaceae and Nectriaceae as has been mentioned by Rossman et al. (1999). Müller and von Arx (1962) treated Dubitatio as a synonym of Passerinula, while the differences of ascomata and ascospores could easily distinguish these two genera (Rossman et al. 1999). After checking the type specimen, Dubitatio was assigned to Dothideomycetes, and considered closely related to Dothivalsaria in the Massariaceae (Barr 1979b, 1987b). Dubitatio chondrospora was assigned to Pseudomassaria (as P. chondrospora (Ces.) Jacz.) (Barr 1964; Müller and von Arx 1962).
Phylogenetic study
None.
Concluding remarks
The black ascomata with white crystalline covering and central white ostiolar region as well as the asymmetrical reddish brown ascospores are striking characters of Dubitatio dubitationum. The genus cannot be assigned to any family with certainty based on morphological characters and fresh collections are needed for sequencing.
Entodesmium Reiss, Hedwigia 1: 28 (1854). (Phaeosphaeriaceae)
Generic description
Habitat terrestrial, saprobic (or parasitic?). Ascomata scattered or in small groups, immersed, papillate, ostiolate, periphysate. Peridium thin, comprising one cell type of pigmented pseudoparenchymatous cells. Hamathecium of dense, long pseudoparaphyses, septate, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with furcate pedicel. Ascospores ellipsoid to filliform, multi-septate, deeply constricted at the primary septum (usually near apex), breaking into partspores.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Barr 1992b; Eriksson 1967a; b; Holm 1957; Liew et al. 2000; Shoemaker 1984a, b.
Type species
Entodesmium rude Reiss, Hedwigia 1: 28 (1854). (Fig. 30)
Ascomata 160–250 μm high × 150–300 μm diam., in groups, immersed with long and protruding cylindrical papilla, globose to subglobose, black, coriaceous (Fig. 30a). Papilla 100–220 μm long, 70–120 μm broad, cylindrical, with periphysate ostiole. Peridium 25–33 μm wide, comprising pseudoparenchymatous cells, cells up to 10 × 7.5 μm diam., cell wall up to 2 μm thick, beak cells smaller and wall thicker (Fig. 30b and c). Hamathecium of dense, long pseudoparaphyses, septate, 2–3 μm wide, embedded in mucilage. Asci 100–175 × 8–13 μm (\( \bar{x} = 147.5 \times 11.3\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a furcate pedicel which is 18–50 μm long, and with a low ocular chamber (ca. 1 μm wide × 1 μm high) (Fig. 30e,f, g and h). Ascospores 108–138 × 3–3.5 μm (\( \bar{x} = 123 \times 3.2\mu m \), n = 10), filliform, brown, multi-septate, breaking into 22–28 partspores, 5–7 × 3–3.5 μm diam. (Fig. 30d).
Anamorph: none reported.
Material examined: GERMANY, Königstein, on stems of Coronilla varia L., 20 May 1895, W. Krieger (H, Krieger 1070).
Notes
Morphology
Entodesmium is characterized by having immersed ascomata dark cylindrical, periphysate papillae, numerous clavate to cylindrical asci surrounded by narrowly cellular pseudoparaphyses, and ellipsoidal to filliform multi-septate ascospores (Barr 1992b; Shoemaker 1984b). Currently, five species, viz. Entodesmium eliassonii L. Holm, E. lapponicum (L. Holm) L. Holm, E. mayorii (E. Müll.) L. Holm, E. niessleanum (Rabenh. ex Niessl) L. Holm and E. rude are accepted in this genus (Holm 1957; Shoemaker 1984b). Von Arx and Müller (1975) assigned Entodesmium to the Pleosporaceae sensu lato, and Shoemaker (1976) assigned E. rude (as Ophiobolus rudis) to Ophiobolus sensu lato based on the fragmenting filliform ascospores. According to the short, blackish beak and periphysate ostiole, Barr (1992b) assigned Entodesmium to Lophiostomataceae. The hosts of Entodesmium are restricted to stems of legumes (Barr 1992b; Shoemaker 1984b).
Phylogenetic study
Limited phylogenetic studies indicate that Entodesmium rude may have affinities to Phaeosphaeriaceae (Liew et al. 2000; Plate 1).
Concluding remarks
Species of Entodesmium share several morphological characters, such as immersed, papillate ascomata, periphysate ostioles, pale yellow to light yellowish brown, multi-septate (≥ 3), narrowly fusoid to filliform ascospores, and are specific to legumes. All of the above similarities indicate a close relationship among members of Entodesmium. We do not agree with Barr (1992b) who assigned Entodesmium to Lophiostomataceae because the ascomata are immersed, the papilla are not laterally compressed and the peridium comprises a single type of cells of textura angularis. These characters plus multi-septate, lightly pigmented ascospores, which break up into partspores and host specificity to legumes support inclusion in Phaeosphaeriaceae. Entodesmium multiseptatum (G. Winter) L. Holm and E. niessleanum were originally described as Leptosphaeria species (Shoemaker 1984b) indicating their similarity to Phaeosphaeria with which Leptosphaeria is commonly confused (Shoemaker 1984a; Shoemaker and Babcock 1989b). Phylogenetic study has also shown that Entodesmium rude is related to members of Phaeosphaeriaceae (Liew et al. 2000). Thus we assign Entodesmium to Phaeosphaeriaceae as a separate genus until further phylogenetic analysis is carried out on verified specimens.
Eudarluca Speg., Revta Mus. La Plata 15: 22 (1908). (?Phaeosphaeriaceae)
Generic description
Habitat terrestrial, parasitic. Ascomata small, solitary, scattered, immersed to erumpent, subglobose, ostiolate, papillate. Peridium thin, composed of a few layers cells of textura prismatica. Hamathecium of dense, cellular pseudoparaphyses, septate. Asci 8-spored, bitunicate, fissitunicate, cylindrical to fusoid, with a furcate pedicel. Ascospores broadly fusoid to fusoid, hyaline to pale yellow, rarely 1- or 3- septate, mostly 2-septate, constricted at the primary septum.
Anamorphs reported for genus: Sphaerellopsis (Sivanesan 1984).
Literature: Bayon et al. 2006; Eriksson 1966; Katumoto 1986; Ramakrishnan 1951; Spegazzini 1908.
Type species
Eudarluca australis Speg., Revta Mus. La Plata 15: 22 (1908). (Fig. 31)
Ascomata 160–190 μm high × 180–290 μm diam., solitary, scattered, or in small groups, semi-immersed to erumpent, subglobose to broadly ellipsoid, wall black, ostiolate, apex with a short papilla, 40–70 μm broad (Fig. 31a and b). Peridium < 10 μm wide laterally, up to 25 μm thick at the apex, thinner at the base, composed of lightly pigmented thin-walled cells of textura prismatica, cells up to 12 × 4 μm diam., cell wall <1 μm thick, apex cells heavily pigmented, smaller and walls thicker (Fig. 31b and c). Hamathecium of dense, long cellular pseudoparaphyses, 1.5–2.5 μm broad, septate. Asci 50–70 × 7.5–10 μm (\( \bar{x} = 61.4 \times 8.4\mu m \), n = 10), 8-spored, with a short, thick, furcate pedicel, up to 12.5 μm long, bitunicate, fissitunicate, cylindrical to fusoid, no obvious ocular chamber (Fig. 31d, e, f and g). Ascospores 16–20 × 4–6 μm (\( \bar{x} = 17.3 \times 5\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, broadly fusoid to fusoid, hyaline to pale yellow, 2-septate, sometimes 1- or 3-septate, constricted at the two main septa, the medium cell often broader than the others, smooth (Fig. 31h).
Anamorph: Sphaerellopsis filum (Biv.) B. Sutton (Sivanesan 1984).
Material examined: BRAZIL, Sao Paulo, on leaves of Canna sp., 1905, leg. Usteri, nro; det. Ove Eriksson (LPS 5.415, type).
Notes
Morphology
Eudarluca was introduced based on E. australis (Spegazzini 1908), and E. australis was subsequently treated as a synonym of E. caricis (Biv.) O.E. Erikss. (Eriksson 1966). The most striking character of E. australis is its 2-septate ascospores, which is quite rare in Pleosporales. Sphaerellopsis filum, anamorph of E. caricis, is a cosmopolitan hyperparasite associated with a large number of rust species (Płachecka 2005).
Phylogenetic study
A detailed phylogenetic study was conducted on Sphaerellopsis filum, the anamorphic stage of Eudarluca australis based on both AFLP and ITS sequences, and only limited variation between different isolates was detected (Bayon et al. 2006).
Concluding remarks
By blasting within GenBank, ITS sequences of E. caricis (= E. australis, strain MullMK, GB, access AY836374) are most comparable with species in Leptosphaeria and Phoma. Thus Eudarluca appears to be related to Leptosphaeriaceae pending further study.
Falciformispora K.D. Hyde, Mycol. Res. 96: 26 (1992). (Trematosphaeriaceae)
Generic description
Habitat freshwater, saprobic. Ascomata small, scattered to gregarious, erumpent to nearly superficial, depressed globose to ovoid, black, ostiolate, epapillate, coriaceous. Peridium thin, comprising two cells types, outer layer composed of thick-walled cells of textura angularis, inner layer composed of hyaline compressed cells. Hamathecium long and cellular pseudoparaphyses, septate, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, broadly clavate to fusoid, with a short, thick pedicel. Ascospores fusoid to somewhat clavate, hyaline, usually slightly curved, multi-septate.
Anamorphs reported for genus: none.
Literature: Hyde 1992b; Raja and Shearer 2008.
Type species
Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96: 27 (1992). (Fig. 32)
Ascomata 180–270 μm high × 250–340 μm diam., scattered to gregarious, erumpent and eventually superficial, depressed globose to ovoid, black, ostiolate, epapillate, coriaceous (Fig. 32a). Peridium up to 35 μm wide, comprising two cell types, outer layer composed of thick-walled cells of textura angularis, up to 8 μm diam., cell wall up to 5 μm thick, inner layer composed of hyaline compressed cells, cells 12 × 3 μm diam., cell wall 1–1.5 μm thick (Fig. 32a). Hamathecium long and cellular pseudoparaphyses, 2–3 μm broad, septate, embedded in mucilage. Asci 115–130 × 23–31 μm, 8-spored, bitunicate, fissitunicate, broadly clavate to fusoid, with a short, thick pedicel, 8–15 μm long, with an ocular chamber (to 5 μm wide × 3 μm high) (Fig. 32b and c). Ascospores 42–50 × 8–10 μm, 2–3 seriate, fusoid to somewhat clavate, hyaline, usually slightly curved, 6–8-septate, mostly 7-septate, slightly constricted at all septa, smooth-walled, surrounded by a thin mucilaginous sheath which is longer at the base (up to 20–30 μm) (Fig. 32d, e and f).
Anamorph: none reported.
Material examined: MEXICO, Nova Hispania, mangrove near Boca de Pascuales, saprobic on immersed intertidal mangrove wood, Mar. 1988, K.D. Hyde (BRIP 16972, holotype).
Notes
Morphology
Falciformispora was formally established by Hyde (1992b) as a monotypic genus and was assigned to Pleosporaceae by comparing with Setosphaeria, but Setosphaeria has the anamorphic stage of Exserohilum and is exclusively parasitic on Gramineae unlike Falciformispora. The setae on the ascomata of Setosphaeria could also serve as a distinguishing character from Falciformispora. Raja and Shearer (2008) also collected this species from freshwater in Florida. They considered that the species was more closely related to Chaetomastia than Setosphaeria, but that Falciformispora differed in having hyaline ascospores.
Phylogenetic study
Phylogenetic analyses in Schoch et al. (2009) and Suetrong et al. (2009) placed Falciformispora lignatilis in Trematosphaeriaceae in proximity to another marine species associated with mangroves, Halomassarina thalassiae.
Concluding remarks
Phylogenetic work confirmed that the saprobic habitat of Falciformispora is inconsistent with most other members of Pleosporaceae. The hyaline multi-septate ascospores with a mucilaginous sheath indicate affinities to Lophiostomataceae but this is not supported in DNA sequence comparisons. Carinispora is also similar and may be related.
Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989). (?Phaeosphaeriaceae)
Generic description
Habitat terrestrial (or freshwater?), saprobic. Ascomata small- to medium-sized, solitary, scattered, or in groups, immersed to nearly superficial, globose to subglobose, papillate. Peridium thin, comprising pseudoparenchymatous cells. Hamathecium dense, narrowly cellular, embedded in mucilage. Asci bitunicate, fissitunicate, oblong to ovoid, with a short pedicel. Ascospores ellipsoid to broadly fusoid with narrow ends, reddish brown, multi-septate, constricted at the primary septum.
Anamorphs reported for genus: Zalerion (Tanaka and Harada 2003a).
Literature: Boise 1984, 1989; Fisher and Webster 1992; Shearer and Crane 1971; Tanaka and Harada 2003a; Webster 1993.
Type species
Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn 49: 310 (1989). (Fig. 33)
≡ Trematosphaeria fallax Mouton, Bull. Soc. R. Bot. Belg. 25: 155, (1886).
Ascomata 130–240 μm high × 200–330 μm diam., solitary, scattered or in groups, initially immersed, becoming erumpent to nearly superficial, with basal wall remaining immersed in host tissue, not easily removed from the substrate, globose or subglobose, roughened, papillate, coriaceous (Fig. 33a). Peridium 30–45 μm wide, comprising cells of pseudoparenchymatous, up to 12.5 × 9 μm diam. (Fig. 33b and c). Hamathecium of dense, narrowly cellular pseudoparaphyses, 1–2 μm broad, embedded in mucilage. Asci 150–200 × 40–60 μm (\( \bar{x} = 171.5 \times 48\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, oblong to ovoid, with a short pedicel, 10–24 μm long, with a ocular chamber (to 5 μm wide × 6 μm high) (Fig. 33d). Ascospores 55–80 × 16–22 μm (\( \bar{x} = 67.1 \times 18.1\mu m \), n = 10), biseriate to 4-seriate, ellipsoid to broadly fusoid with narrow ends, reddish brown with paler end cells, 8-septate, constricted at the primary septum, smooth-walled (Fig. 33e, f, g, h and i).
Anamorph: Zalerion sp. (Tanaka and Harada 2003a).
Material examined: BELGIUM, Beaufays, on cut off, still hard wood. Oct. 1922, V. Mouton (BR, Capsa: K 7534, holotype). (Note: The specimen is not in good condition, only a few ascomata left).
Notes
Morphology
Boise (1989) formally established Hadrospora to accommodate Trematosphaeria fallax and T. clarkia (Sivan.) Boise, and Hadrospora fallax (syn. T. fallax) was selected as the generic type. Hadrospora is a widely distributed species that has been reported from Belgium, China, Italy, Japan, Switzerland and the United States (Boise 1989; Fisher and Webster 1992; Shearer and Crane 1971; Tanaka and Harada 2003a; Webster 1993). Hadrospora was temporarily assigned to Phaeosphaeriaceae based on its “small, thin-walled ascomata and narrow cellular pseudoparaphyses” (Boise 1989), which is distinguished from other genera of Phaeosphaeriaceae by its “large, stout ascospores that form within oblong-ovoid asci” (Boise 1989). Currently, Hadrospora includes two species, i.e. H. fallax and H. clarkii (Sivan.) Boise differentiated by ascospore size.
Phylogenetic study
None.
Concluding remarks
Hadrospora seems not closely related to Phaeosphaeriaceae.
Halotthia Kohlm., Nova Hedwigia 6: 9 (1963). (?Zopfiaceae)
Generic description
Habitat marine, saprobic. Ascomata large, solitary, gregarious or confluent, broadly conical to subglobose, flattened at the base, carbonaceous, immersed to erumpent, ostiolate, epapillate. Peridium plectenchymatous. Hamathecium of dense, long, cellular pseudoparaphyses, septate, branching. Asci 8-spored, bitunicate, cylindrical, with a short pedicel. Ascospores uniseriate, ellipsoidal, subcylindrical or obtuse-fusoid, dark brown, 1-septate, constricted at the septum.
Anamorphs reported for genus: none.
Literature: Kohlmeyer 1963; Suetrong et al. 2009.
Type species
Halotthia posidoniae (Durieu & Mont.) Kohlm., Nova Hedwigia 6: 9 (1963). (Fig. 34)
≡ Sphaeria posidoniae Durieu & Mont. Exploration scientifique de l’Algérie, pp. 502–503, Taf. 25, Abb. 8a-i, 1849.
Ascomata 0.8–1.1 mm high × 1.5–2.1 mm diam., solitary, gregarious or confluent, broadly conical to subglobose, flattened at the base, carbonaceous, immersed to erumpent, ostiolate, epapillate (Fig. 34a). Peridium 165–275 μm thick at sides, thicker near the apex, plectenchymatous. Hamathecium of dense, long cellular pseudoparaphyses, 1.5–2 μm broad, septate, branching. Asci 275–290 × 25–35 μm, 8-spored, bitunicate, cylindrical, with a short pedicel (Fig. 34b, c and d). Ascospores 37–60.5 × 16.5–26 μm, uniseriate, ellipsoidal, subcylindrical or obtuse-fusoid, dark brown, 1-septate, constricted at the septum (Fig. 34e, f, g and h) (adapted from Kohlmeyer and Kohlmeyer 1979).
Anamorph: none reported.
Material examined: ITALY, in rhizomes of Posidonia oceanica (Posidoniaceae), 1861, Caldesi (S, isotype of Sphaeria posidoniae)
Notes
Morphology
Halotthia was introduced to accommodate the marine fungus, H. posidoniae (as Sphaeria posidoniae), which is characterized by immersed to erumpent, large, carbonaceous ascomata, thick peridium, bitunicate, 8-spored, cylindrical asci, ellipsoidal, 1-septate, and dark brown ascospores (Kohlmeyer 1963). Morphologically, Halotthia is most comparable with Bicrouania maritima, but the conical ascomata with flattened base of H. posidoniae can be readily distinguished from B. maritima.
Phylogenetic study
Phylogenetically, Halotthia posidoniae, Pontoporeia biturbinata and Mauritiana rhizophorae form a robust clade, which may represent a potential family (Suetrong et al. 2009).
Concluding remarks
Currently the familial status of Halotthia is unresolved (Suetrong et al. 2009).
Helicascus Kohlm., Can. J. Bot. 47: 1471 (1969). (Morosphaeriaceae)
Generic description
Habitat marine, saprobic. Ascostromata lenticular, immersed, black, carbonaceous, enclosing several loculi, pseudoclypeus composed of host cells enclosed in black stromatic fungus material. Ascomata depressed ampulliform, horizontally arranged under a black pseudoclypeus, ostiolate, torsellioid ostioles, papillate. Peridium absent, partitions between loculi formed of brown, isodiametric or elongated cells of the stroma. Hamathecium of dense, long pseudoparaphyses. Asci 8-spored, bitunicate, subcylindrical to oblong clavate, with a short pedicel and conspicuous apical ring. Ascospores uniseriate, obovoid, brown, 1-septate, at each end with a germ pore, surrounded with dissolving sheath.
Anamorphs reported for genus: none.
Literature: Kohlmeyer 1969; Kohlmeyer and Kohlmeyer 1979; Suetrong et al. 2009.
Type species
Helicascus kanaloanus Kohlm., Can. J. Bot. 47: 1471 (1969). (Fig. 35)
Ascostromata 0.6–0.78 mm high × 1.25–2.75 mm diam., lenticular, immersed, black, carbonaceous, enclosing 3–4(−5) loculi, pseudoclypeus composed of host cells enclosed in black stromatic fungus material (Fig. 35a). Ascomata 235–370 μm high × 440–800 μm diam., depressed ampulliform, horizontally arranged under a black pseudoclypeus, ostioles 70–170 μm diam., torsellioid ostiole (Adams et al. 2005), papilla slightly rising over the surface of the pseudoclypeus, subconical,canal filled with thick, bright orange to yellowish periphyses, 270–435 μm high, 255–300 μm diam. Peridium absent, partitions between loculi formed of brown, isodiametric or elongated cells of the stroma. Hamathecium of dense, very long pseudoparaphyses. Asci 250–335 × 25–30 μm, 8-spored, subcylindrical, finally oblong-clavate (400–480 μm long), with a short pedicel, bitunicate, thick-walled, physoclastic, apically multi-layered and annulate, ectoascus forming a third, thin permeable outer layer around the base, endoascus swelling in water and becoming coiled at maturity, finally stretching and pushing the ascus into the ostiolar canal (Fig. 35b). Ascospores 36.5–48.5 × 18–22.5 μm, uniseriate, obovoid, brown, 1-septate, at each end with a germ pore, surrounded with dissolving sheath, 2.7–5.4 μm thick, with funnel-shaped, apical indentations (Fig. 35c and d) (adapted from Kohlmeyer and Kohlmeyer 1979).
Anamorph: none reported.
Material examined: USA, Hawaii, Oahu, Kaneohe Bay, Heeia Swamp, on Rhizophora mangle, 4 Jun. 1968 (Herb. J. Kohlmeyer No. 2566, holotype; No. 2565, 2567, paratype).
Notes
Morphology
Helicascus is another marine genus, which is characterized by its thin additional sheath around the base of the asci, the coiling and stretching mechanism of the basal part of the endoascus and its conspicuous apical apparatus which is not that common in bitunicate asci (Kohlmeyer 1969). The immersed stroma comprising several loculi sharing one common ostiole is another striking character of Helicascus.
Phylogenetic study
Multigene phylogenetic analysis indicated that both Helicascus kanaloanus and H. nypae K.D. Hyde nested within Morosphaeriaceae (Suetrong et al. 2009).
Concluding remarks
Helicascus is a well defined marine genus.
Herpotrichia Fuckel, Fungi rhenani exsic.: no. 2171 (1868). (Melanommataceae)
Generic description
Habitat terrestrial, parasitic, hyperparasitic or saprobic. Ascomata medium-sized, immersed, erumpent to nearly superficial, scattered to gregarious, globose to subglobose with a broad pore. Peridium composed of pseudoparenchymatous cells. Hamathecium of dense, long pseudoparaphyses, embedded in mucilage, septate, branching. Asci cylindrical to cylindro-clavate, with a furcated pedicel. Ascospores fusoid, ellipsoid or oblong with broadly to narrowly round ends, 1-septate, constricted at the septum, uni- to biseriate.
Anamorphs reported for genus: Pyrenochaeta or Pyrenochaeta-like (Sivanesan 1984).
Literature: von Arx and Müller 1975; Barr 1984; Cannon 1982; Freyer and van der Aa 1975; Mugambi and Huhndorf 2009b; Samuels 1973; Samuels and Müller 1978; Sivanesan 1971, 1984.
Type species
Herpotrichia rubi Fuckel, Fungi rhenani exsic 2171. (1868). (Fig. 36)
Ascomata 220–430 μm high × 240–390(-530) μm diam., scattered to gregarious, immersed to erumpent, rarely superficial, globose to subglobose, wall black, coriaceous, apex with a small sometimes inconspicuous papilla, usually with a pore, lacking periphyses (Fig. 36a and b). Peridium 32–45 μm wide at the sides, up to 60 μm wide at the apex, basal wall thinner, all walls comprising cells of textura angularis, cells 2.5–4 μm diam., cell wall 2–4(−7) μm thick, exterior cells more thick-walled and pigmented, inner cells thin-walled and less pigmented, comprising thin-walled cells up to 9 μm diam., apex cells smaller and walls thicker (Fig. 36b and c). Hamathecium of dense, long pseudoparaphyses, 2–3 μm broad, embedded in mucilage, septate, branching (Fig. 36e). Asci 105–150 × 12.5–15 μm (\( \bar{x} = 137.5 \times 13.8\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a furcate pedicel that is 20–42.5 μm long, and ocular chamber up to 2.5 μm wide × 2.5 μm high (Fig. 36d and f). Ascospores 17.5–25 × (5.5-)6.3–9 μm (\( \bar{x} = 20.5 \times 7.3\mu m \), n = 10), biseriate to partially overlapping uniseriate near the base, fusoid with narrowly rounded ends, hyaline when immature and becoming pale brown, 1-septate, deeply constricted at the septum, the upper cell often broader than the lower one, verruculose (Fig. 36g and h).
Anamorph: Pyrenochaeta rhenana Sacc. (Sivanesan 1984).
Material examined: AUSTRIA, on Rubus idaeus L., very rarely in the spring, in the Oestreicher meadow forest (G, F. rh. 2171, type).
Notes
Morphology
Herpotrichia was established by Fuckel (1868) comprising two species H. rhenana Fuckel and H. rubi Fuckel, but no generic type was assigned. Bose (1961) designated H. rhenana as the lectotype species with H. rubi as a synonym. This proposal was followed by Müller and von Arx (1962) and Sivanesan (1971). Herpotrichia rubi was later assigned as the generic type (Holm 1979) as it was found to be validly published 2 years earlier than H. rhenana, thus having priority (Cannon 1982). However, Cannon (1982) reported that Sphaeria herpotrichoides Fuckel (1864, cited as a synonym of H. rhenana) was the earliest name. Thus he made a new combination as H. herpotrichoides (Fuckel) P.F. Cannon and cited H. rubi as the synonym. Herpotrichia rubi is maintained as the type of the genus (Holm 1979; Cannon 1982), but the current name is H. herpotrichoides.
Herpotrichia is a morphologically well studied genus (Barr 1984; Bose 1961; Müller and von Arx 1962; Pirozynski 1972; Samuels and Müller 1978; Sivanesan 1971, 1984), and Herpotrichia sensu lato is characterized by having subglobose, pyriform to obpyriform ascomata and a peridium of textura angularis or comprising thick-walled polygonal cells with thin-walled hyaline cells towards the centre. Asci are clavate to cylindrical, 4–8-spored and ascospores are hyaline at first, becoming pale to dark brown, one to many septate, constricted or not at the septa and often surrounded by a mucilaginous sheath. Several morphologically distinct genera were synonymized under Herpotrichia using the above broad circumscription (Barr 1984; Müller and von Arx 1962; Sivanesan 1984). In particular, Barr kept Lojkania as a separate genus after studying its type material (Barr 1984, 1990a). Sivanesan (1984) was also of the opinion that Lojkania and Neopeckia were distinct genera as several of their characters differed. Byssosphaeria and Pseudotrichia have subsequently been assigned to Melanommataceae, Lojkania to Fenestellaceae and Neopeckia to Coccoideaceae (Barr 1984). Herpotrichia sensu stricto is represented by H. rubi and has erumpent to superficial ascomata or immersed in a subiculum, clavate to cylindrical, 4–8-spored, stalked asci with a conspicuous apical “nasse”, hyaline, 1-septate ascospores, usually becoming pale brown and several septate, constricted or not constricted at septa, usually surrounded by sheath (Sivanesan 1984). Currently, about 90 species are included in this genus (http://www.indexfungorum.org/, 12/01/2009).
Phylogenetic study
Herpotrichia diffusa (Schwein.) Ellis & Everh., H. juniperi (Duby) Petr., H. herpotrichoides and H. macrotricha have been shown to have phylogenetic affinity with the generic types of Byssosphaeria schiedermayeriana, Melanomma pulvis-pyrius and Pleomassaria siparia, which had been assigned under Melanommataceae (Kruys et al. 2006; Mugambi and Huhndorf 2009b; Schoch et al. 2006, 2009; Zhang et al. 2009a). In this study, Pleomassaria siparia together with its closely related species of Prosthemium is kept in a separate family, viz Pleomassariaceae.
Concluding remarks
Even species under Herpotrichia sensu stricto (according to Sivanesan 1984) have diverse hosts (such as gymnosperms (H. coulteri (Peck) S.K. Bose and H. parasitica (R. Hartig) Rostr.) and angiosperms (H. diffusa and H. villosa Samuels & E. Müll.)) or substrates (like dead or living leaves, bark or decorticated wood) (Sivanesan 1984). Species of Herpotrichia sensu stricto are also reported from various locations such as Europe, Asia or America, and they have various life styles, e.g. parasitic, hyperparasitic or saprobic (Sivanesan 1984). Additional factors (like hosts or locations) may need to be considered in order to get a natural concept for Herpotrichia.
Immotthia M.E. Barr, Mycotaxon 29: 504 (1987). (Teichosporaceae)
Generic description
Habitat terrestrial, hyperparasitic. Ascomata gregarious, globose, superficial, ostiolate, periphysate. Hamathecium of cellular pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical, with a short pedicel. Ascospores 1-seriate, ellipsoidal, brown to reddish brown, 1-septate, constricted at the septum, smooth.
Anamorphs reported for genus: none.
Literature: Barr 1987a, 2002; Wang et al. 2004.
Type species
Immotthia hypoxylon (Ellis & Everh.) M.E. Barr, Mycotaxon 29: 504 (1987). (Fig. 37)
≡ Amphisphaeria hypoxylon Ellis & Everh., J. Mycol. 2: 41 (1886).
Ascomata gregarious, globose, superficial, ostiolate, periphysate, papillate (Fig. 37a). Hamathecium of cellular pseudoparaphyses, 2–2.5 μm broad, septate. Asci 60–82 × 7–9 μm, 8-spored, bitunicate, cylindrical, with a short pedicel (Fig. 37b, c and d). Ascospores 10–13 × 4.4–5.4 μm, 1-seriate, ellipsoidal, brown to reddish brown, 1-septate, constricted at the septum, smooth (Fig. 37f, g and h) (adapted from Wang et al. 2004).
Anamorph: none reported.
Material examined: USA, Louisiana, Pointe a la Hache, on decaying wood, a branch of Carya oliviformis (Juglandaceae) lying on the ground in grass (parasitic on some effused Hypoxylon), 30 Dec. 1885, A.B. Langlois, No. 138 (NY, holotype of Amphisphaeria hypoxylon Ellis & Everh.).
Notes
Morphology
Immotthia was introduced to accommodate a species of Amphisphaeria (A. hypoxylon), which has bitunicate asci, and is characterized by superficial, ostiolate, periphysate, papillate ascomata, cellular pseudoparaphyses, bitunicate, 8-spored, cylindrical asci, ellipsoid, smooth, brown to reddish brown, 1-septate ascospores (Barr 1987a; Wang et al. 2004).
Phylogenetic study
None.
Concluding remarks
It seems that those Amphisphaeria species with bitunicate asci should be assigned to Pleosporales. Morphologically, Immotthia is somewhat comparable with Herpotrichia.
Isthmosporella Shearer & J.L. Crane, Mycologia 91: 141 (1999). (Pleosporales, genera incertae sedis)
Generic description
Habitat freshwater, saprobic. Ascomata small- to medium-sized, scattered, immersed, erumpent to superficial, globose, papillate, ostiolate, periphysate, membranous. Peridium 2-layered, outer layer composed of brown, pseudoparenchymatic, fusoid-cylindric cells, inner layer composed of fusoid, subhyaline to pale brown, compressed cells. Hamathecium of rare, broad, septate, interascal pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, oblong to clavate, with a short pedicel, ocular chamber not observed. Ascospores 3–4 seriate, cylindrical to fusoid, isthmoid at centre, constricted at septa, isthmus 1-septate, surrounded by a gelatinous sheath.
Anamorphs reported for genus: none.
Literature: Shearer and Crane 1999.
Type species
Isthmosporella pulchra Shearer & J.L. Crane, Mycologia 91: 142 (1999). (Fig. 38)
Ascomata 240–330 μm diam., scattered on decorticated wood, immersed, erumpent to superficial, globose, black, papillate, papilla short, cylindrical, 60 μm long × 55 μm wide, ostiolate, periphysate, membranous (Fig. 38a). Peridium 2-layered, outer 3–4 cell layers composed of brown, pseudoparenchymatic, fusoid-cylindric cells, 2–6.5 μm long; inner layer composed of 5–7 rows of fusoid, subhyaline to pale brown compressed cells, 11–20 × 2–3.5 μm diam. (Fig. 38a and b). Hamathecium of rare, broad, septate, interascal pseudoparaphyses (Fig. 38f). Asci (95-)135–160(−175) × (25-)30–45(−60) μm, 8-spored, bitunicate, fissitunicate, oblong to clavate, with a short pedicel, ocular chamber not observed (Fig. 38c, d and e). Ascospores 80–105(−110) × (7-)8–10 μm, 3–4-seriate, cylindrical to fusoid, isthmoid at centre, sometimes bent at isthmus and becoming u- or v- shaped, end cells tapering, 12–17-phragmoseptate, constricted at septa, isthmus 1-septate, 2–5.5 × 2–4.5 μm diam., hyaline, frequently fragmenting to form partspores; filled with lipid droplets that merge to form large guttules; surrounded by a gelatinous sheath with a dense region near the isthmus, sheath greatly enlarging in water (Fig. 38g, h, i and j).
Anamorph: none reported.
Colonies on yeast soluble starch agar containing balsa wood sticks effuse, white. Hyphae hyaline, septate.
Material examined: USA, New York, Adirondack Park. Piercefield. Tupper Lake at public boat launch from Rt. 30, UTM Zone 18, 539840 mE, 4892100mN; 44°10″59″N, 80°31′6″W, on submerged, decorticated wood, 7 Jul. 1994, J.L. Crane & C.A. Shearer A-254-1 (ILLS 53086, holotype).
Notes
Morphology
Isthmosporella was described as a freshwater genus typified by I. pulchra, and is characterized by globose, pseudoparenchymatous ascomata, sparse, septate pseudoparaphyses, fissitunicate asci and hyaline, cylindrical to fusoid, phragmoseptate, isthmoid ascospores surrounded with a gelatinous sheath (Shearer and Crane 1999). Based on the morphological characters, i.e. small, globose ascomata, peridium with small pseudoparenchymatous cells and sparse pseudoparaphyses, Isthmosporella was assigned to the Phaeosphaeriaceae (Shearer and Crane 1999). The aquatic habitat of Isthmosporella, however, disagree with the Phaeosphaeriaceae. Isthmosporella seems less likely to belong to Pleosporineae.
Phylogenetic study
None.
Concluding remarks
Molecular phylogenetic studies should be conducted to explore its familial placement within Pleosporales.
Kalmusia Niessl, Verh. nat. Ver. Brünn 10: 204 (
1872). (Montagnulaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to medium-sized, solitary, scattered or in small groups, immersed to erumpent, globose or subglobose, often laterally flattened, coriaceous, wall black, with or without papilla. Hamathecium of dense, filliform, delicate, septate pseudoparaphyses, branching and anastomosing between and above asci, embedded in mucilage. Asci bitunicate, fissitunicate unknown, clavate, with a long, furcate pedicel. Ascospores narrowly ovoid to clavate, pale brown, 3-septate, distoseptate.
Anamorphs reported for genus: Cytoplea (Petrak and Sydow 1926).
Literature: Barr 1987b, 1990a, 1992a; Lindau 1897; von Niessl 1872.
Type species
Kalmusia ebuli Niessl, Verh. nat. Ver. Brünn 10: 204 (1872). (Fig. 39)
Ascomata 290–360 μm high × 300–520 μm diam., solitary, scattered, or in small groups, immersed to erumpent, globose or subglobose, coriaceous, wall black, with or without papilla, ostiolate (Fig. 39a). Papilla small, up to 100 μm high, with small ostioles (Fig. 39a). Peridium 15–40 μm wide, comprising one cell type of small, pigmented, thick-walled cells of textura prismatica to textura angularis, cells ca. 5 × 3 μm diam., cell wall 2–3 μm thick (Fig. 39b and c). Hamathecium of dense, delicate pseudoparaphyses, 1–1.5 μm broad, septate, branching and anastomosing between and above asci, embedded in mucilage. Asci 75–125 × 10–15 μm (\( \bar{x} = 90.5 \times 12\mu m \), n = 10), 8-spored, bitunicate, fissitunicate unknown, clavate, with a long, narrowed, furcate pedicel which is up to 45 μm long, and a low ocular chamber (ca. 2 μm wide × 1 μm high) (Fig. 39d, e and f). Ascospores 15–18 × 5.5–6.5 μm (\( \bar{x} = 16.3 \times 5.8\mu m \), n = 10), biseriate, narrowly ovoid to clavate, pale brown, 3-distoseptate, without constriction, smooth-walled (Fig. 39g, h and i).
Anamorph: none reported.
Material examined: BELGIUM, Dolembreux, on branchlets and pieces of stumps of Sarothamnus scoparius from woodland, Oct. 1922, V. Mouton (BR 101525–63, holotype).
Notes
Morphology
Kalmusia was formally established by von Niessl (1872), and is mainly characterized as “immersed, sphaeroid ascoma with central, stout papilla, surrounded by hyphae in the substrate, stipitate asci with septate pseudoparaphyses, and brown, 3-septate, inequilateral ascospores” (Barr 1992a).
The most morphologically comparable genus to Kalmusia is Thyridaria, which had been treated as a subgenus under Kalmusia (Lindau 1897), and was subsequently transferred to Platystomaceae in Melanommatales (Barr 1987b, 1990a). Compared to Thyridaria, Kalmusia has sphaeroid ascomata, a peridium of small pseudoparenchymatous cells, basal asci and very thin pseudoparaphyses, thus it was assigned to Phaeosphaeriaceae of the Pleosporales by Barr (1990a), and the genus is utilized to accommodate both K. ebuli and K. clivensis (Berk. & Broome) M.E. Barr, as well as closely related species, i.e. K. utahensis (Ellis & Everh.) Huhndorf & M.E. Barr and K. coniothyrium (Fuckel) Huhndorf (Barr 1992a). But this proposal is questionable, as the clavate, distoseptate ascospores, as well as the clavate asci with very long pedicels are uncommon in Phaeosphaeriaceae, and most recent phylogenetic study indicated that some species of Kalmusia reside outside of Phaeosphaeriaceae (Zhang et al. 2009a).
Phylogenetic study
Both Kalmusia scabrispora Teng Kaz. Tanaka, Y. Harada & M.E. Barr and K. brevispora (Nagas. & Y. Otani) Yin. Zhang, Kaz. Tanaka & C.L. Schoch reside in the clade of Montagnulaceae (Zhang et al. 2009a). Familial placement of Kalmusia can only be verified after the DNA sequences of the generic type (K. ebuli) are obtained.
Concluding remarks
Kalmusia is distinct amongst the Pleosporales as it has pale brown ascospores with indistinct distosepta and clavate asci with long pedicels. Although both K. scabrispora and K. brevispora reside in the clade of Montagnulaceae, they both lack the distoseptate ascospores that are possessed by the generic type (K. ebuli). Thus, the familial placement of Kalmusia is still undetermined.
Karstenula Speg., Decades Mycologicae Italicae ad no. 94 (in sched.) (1879). (Montagnulaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata rarely small-, usually medium-sized, immersed usually under thin clypeus, scattered to gregarious, with flattened top and rounded pore-like ostiole, coriaceous. Peridium 2-layered, outer layer composed of reddish brown to dark brown small cells, inner layer of pale compressed cells. Hamathecium of dense, cellular pseudoparaphyses. Asci cylindrical to cylindro-clavate with short furcate pedicel. Ascospores muriform, ellipsoid to fusoid, reddish brown to dark brown.
Anamorphs reported for the genus: Microdiplodia (Constantinescu 1993).
Literature: Barr 1990a; Eriksson and Hawksworth 1991; Kodsueb et al. 2006a; Munk 1957; Zhang et al. 2009a.
Type species
Karstenula rhodostoma (Alb. & Schwein.) Speg., Decades Mycologicae Italicae no. 94. (1879). (Fig. 40)
≡ Sphaeria rhodostoma Alb. & Schwein., Consp. fung. (Leipzig): 43 (1805).
Ascomata 250–430 μm high × 450–650 μm diam., scattered or gregarious, immersed in the subiculum which sometimes sloths off, globose or subglobose, black, flattened top often white or reddish and sometimes slightly protruding out of the substrate surface, usually with a wide opening ostiole after removing the cover, coriaceous (Fig. 40a and b). Peridium 30–40 μm wide, comprising two cell types, outer region 1-layered, composed of relatively small heavily pigmented thick-walled compressed cells, cells 2–4 × 5–10 μm diam., cell wall 2–4 μm thick, inner layer cells larger and wall thinner, comprising cells of textura angularis, merging with pseudoparaphyses (Fig. 40c and d). Hamathecium of dense, long cellular pseudoparaphyses 2–3.5 μm broad, septate, branching or anastomosing not observed. Asci 150–210 × 12.5–15 μm (\( \bar{x} = 182 \times 13.1\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a broad, furcate pedicel which is 12–35 μm long, and with an ocular chamber (to 4 μm wide × 3 μm high) (Fig. 40e and f). Ascospores 20–26 × 7.5–10 μm (\( \bar{x} = 22.4 \times 8\mu m \), n = 10), obliquely uniseriate and partially overlapping, ellipsoid, reddish brown, with 3 transverse septa and a vertical septum in one or two central cells, constricted at the septa, verruculose (Fig. 40g, h and i).
Anamorph: Microdiplodia frangulae Allesch. (Constantinescu 1993).
Conidiomata globose to subglobose, 330–495 μm diam., in subiculum. Conidia 9–13 × 4–5 μm, reddish brown, 1-septate (information obtained from Barr 1990a).
Material examined: Fries, Suecia (received by herbarium in 1834) (PH 01048835, type, as Sphaeria rhodostoma Alb. & Schwein.).
Notes
Morphology
Karstenula is an ambiguous genus, which has been synonymized under Pleomassaria (Lindau 1897; Winter 1885). Some of the ascomata characters are even comparable with those of Didymosphaeria, such as ascomata seated in subiculum or beneath a clypeal thickening, the development of apex vary in a large degree, even to the occasional formation of a blackened internal clypeus, and sometimes apical cells become reddish or orange-brown (Barr 1990a). Barr (1990a) redefined the concept of Karstenula (sensu lato), which encompasses some species of Thyridium. In her concept, however, Barr (1990a) treated Karstenula as having trabeculate pseudoparaphyses and this is clearly not the case. In most cases the ascospores were brown with transverse septa and sparse longitudinal septa.
The ascomata of this species are similar to those found in Byssosphaeria and Herpotrichia, especially in the paler area around the ostiole and even in peridial structure and development under a subiculum. The numerous wide cellular pseudoparaphyses and cylindrical asci (in Herpotrichia) are also similar. The main difference of Karstenula from other two genera are the 3-septate ascospores with rare longitudinal septa (1-septate in Byssosphaeria and Herpotrichia).
Phylogenetic study
Karstenula forms a robust phylogenetic clade with Phaeodothis winteri (Niessl) Aptroot, Didymocrea sadasivanii, Bimuria novae-zelandiae, Montagnula opulenta, Curreya pityophila (J.C. Schmidt & Kunze) Arx & E. Müll. and some species of Letendraea and Paraphaeosphaeria (Kodsueb et al. 2006a; Zhang et al. 2009a). Consequently, Karstenula might be included in Montagnulaceae.
Concluding remarks
The description of the type of Karstenula here clearly excludes it from Melanommataceae as it has wide pseudoparaphyses. But its Montagnulaceae status can only be confirmed by more phylogenetic work including sequencing the generic type of Karstenula (K. rhodostoma).
Katumotoa Kaz. Tanaka & Y. Harada, Mycoscience 46: 313 (2005). (Lentitheciaceae)
Generic description
Habitat terrestrial or freshwater, saprobic. Ascomata small- to medium-sized, scattered or in small groups, immersed to erumpent, with a central protruding hairy papilla, subglobose. Peridium thin, comprising several layers of thin-walled compressed cells. Hamathecium of dense, cellular, filliform, embedded in mucilage, branching and anastomosing. Asci 8-spored, bitunicate, fissitunicate, clavate with short furcate pedicels. Ascospores apiosporous and hyaline when young, becoming 2-septate with reddish brown echinate central cell at maturity, with long gelatinous terminal appendages.
Anamorphs reported for genus: none.
Literature: Tanaka and Harada 2005b; Tanaka et al. 2009; Zhang et al. 2009a.
Type species
Katumotoa bambusicola Kaz. Tanaka & Y. Harada, Mycoscience 46: 313 (2005). (Fig. 41)
Some information for the following description is from Tanaka and Harada (2005).
Ascomata 240–330 μm high × 260–420 μm diam., scattered or in small groups, immersed, becoming erumpent, with a slightly protruding papilla covered with brown hyphae, subglobose (Fig. 41a). Peridium 13–30 μm thick, composed of a few layers of lightly pigmented, depressed cells. Hamathecium of dense, long cellular pseudoparaphyses, 1.5–3 μm broad, embedded in mucilage, branching and anastomosing. Asci 110–160 × 17.5–24 μm (\( \bar{x} = 139 \times 21\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, cylindro-clavate with a short furcate pedicel which is up to 25 μm long (Fig. 41b and d). Ascospores 39–50(−57) × 7–10 μm (\( \bar{x} = 45.8 \times 8.2\mu m \), n = 10), biseriate, fusoid to narrowly fusoid with acute ends, usually curved, apiosporus and hyaline when young, constricted at the primary septum, the upper cell longer and broader than the lower one, smooth, surrounded by a bipolar sheath which is up to 15 μm long, best seen in India ink, senescent ascospores yellowish brown, 2–4-septate (Fig. 41c).
Anamorph: none reported.
Material examined: JAPAN, Mt. Iwate, near Yakebashiri, Hirakasa, Nishine, Iwate, on culms of Oryza sativa L., 19 Oct. 2003, K. Tanaka (HHUF 28663, holotype).
Notes
Morphology
Katumotoa was formally established by Tanaka and Harada (2005b) to accommodate the monotypic species, K. bambusicola, which is characterized by immersed ascomata with a thin peridium comprising thin-walled compressed cells, cellular pseudoparaphyses, cylindro-clavate and fissitunicate asci and fusoid ascospores with an elongated bipolar mucilaginous sheath. Based on its immersed ascomata, psuedoparenchymatous peridium cells and cellular pseudoparaphyses, Katumotoa was assigned to Phaeosphaeriaceae (Tanaka and Harada 2005b; Tanaka et al. 2009), but this classification has been shown to be incorrect in subsequent phylogenetic studies (Tanaka et al. 2009; Zhang et al. 2009a).
Phylogenetic study
Phylogenetic analysis based on five genes (LSU, SSU, RPB1, RPB2 and EF1) indicates that Katumotoa bambusicola resides in Lentitheciaceae, and this receives high bootstrap support (Zhang et al. 2009a). In particular, K. bambusicola forms a robust clade with Ophiosphaerella sasicola (Nagas. & Y. Otani) Shoemaker & C.E. Babc., which has filliform ascospores (Shoemaker and Babcock 1989b).
Concluding remarks
The hyaline, apiosporous ascospores which become 2–4-celled with central reddish brown cells and large unraveling appendages are the most striking features of this species and readily distinguish it from other pleosporalean taxa. Both Katumotoa bambusicola and Ophiosphaerella sasicola are associated with bambusicolous hosts, which might indicate that host spectrum in this case, has greater phylogenetic significance than some morphological characters (Zhang et al. 2009a).
Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 582 (1919). (Lentitheciaceae)
Generic description
Habitat terrestrial or freshwater, saprobic. Ascomata small- to medium-sized, immersed, erumpent to nearly superficial, globose, papillate, ostiolate. Papilla covered by dark setae or small blackened cells. Peridium thick, composed of cells of pseudoparenchymatous and inner layer composed of pale cells. Hamathecium of dense, long pseudoparaphyses, rarely septate, anastomosing and branching. Asci 4- or 8-spored, bitunicate, fissitunicate, cylindro-clavate, with a furcate pedicel and a small ocular chamber. Ascospores hyaline to pale brown, ellipsoid to fusoid, 1-septate, constricted at the septum (Barr 1990a).
Anamorphs reported for genus: Dendrophoma (Bose 1961).
Literature: von Arx and Müller 1975; Bose 1961; Barr 1990a; Dennis 1978; Eriksson 1967a; von Höhnel 1919; Luttrell 1973; Munk 1957; Zhang et al. 2009a.
Type species
Keissleriella aesculi (Höhn.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 582 (1919). (Fig. 42)
≡ Pyrenochaeta aesculi Höhn., Ber. dt. bot. Ges. 35: 249 (1917).
Ascomata ca. 250 μm high × 450 μm diam., gregarious, immersed to erumpent, globose or subglobose, with a small black papilla, ca. 75 μm high and 110 μm broad, with short black external setae (Fig. 42a). Peridium ca. 25–40 μm wide laterally, up to 70 μm near the apex, thinner at the base, comprising two types of cells which merge in the middle; outer cells composed of small heavily pigmented thick-walled cells, cells ca. 4 μm diam., cell wall up to 4 μm thick, and thick near the apex and thinner laterally and absent in the immersed part of the ascoma, inner cells less pigmented, comprising lightly pigmented to hyaline cells, 5–7 μm thick (Fig. 42a). Hamathecium of dense, long pseudoparaphyses, 0.8–1.2 μm broad, rarely septate, anastomosing and branching, thicker near the base, ca. 2 μm, constricted near the septum (Fig. 42b). Asci 80–120 × 6–11 μm (\( \bar{x} = 101 \times 8.5\mu m \), n = 10), 4- or 8-spored, bitunicate, fissitunicate, cylindro-clavate, with a furcate pedicel which is up to 20–40 μm long, with a small ocular chamber (Fig. 42e and f). Ascospores 13–18 × 4–5.5 μm (\( \bar{x} = 14.5 \times 4.8\mu m \), n = 10), obliquely uniseriate and partially overlapping, fusoid with narrowly rounded ends, hyaline, 1-septate, constricted at the septum, smooth (Fig. 42c and d).
Anamorph: none reported.
Material examined: AUSTRIA, Brentenmaistal in the Viennese forest, Aesculus hippocastanum L., 1916, Höhnel (FH, holotype of Otthiella aesculi). (Note: only two slides; setae cannot be seen from the slides but could be seen from the drawings on the cover).
Notes
Morphology
Keissleriella is characterized by ascomata with setae in and over the papilla, asci are cylindrical and ascospores are hyaline, 1-septate. Based on the morphological characters, K. aesculi was regarded as conspecific with K. sambucina; as an earlier epithet, K. sambusina typifies the genus (see comments by Barr 1990a). Munk (1957) placed Trichometasphaeria and Keissleriella in Massarinaceae, and distinguished them by their substrates (Trichometasphaeria occurs on herbaceous plants and Keissleriella on woody substrates). Bose (1961) combined Trichometasphaeria under Keissleriella, which was followed by some workers (von Arx and Müller 1975; Dennis 1978; Eriksson 1967a; Luttrell 1973). Barr (1990a), however, maintained these as distinct genera based on the differences of peridium structure and pseudoparaphyses.
Phylogenetic study
The phylogeny of Keissleriella is poorly studied. Limited phylogenetic information indicates that K. cladophila forms a robust clade with other species of Lentitheciaceae (Zhang et al. 2009a).
Concluding remarks
The presence of black setae on the surface of papilla is a striking character of Keissleriella, but phylogenetic significance of setae is undetermined yet.
Lentithecium K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Lentitheciaceae)
= Tingoldiago K. Hirayama & Kaz. Tanaka, Mycologia 102: 740 (2010) syn. nov.
Generic description
Habitat freshwater, saprobic. Ascomata small, scattered or gregarious, immersed, slightly erumpent, depressed spherical to lenticular, ostiolate, papillate or epapillate. Peridium thin. Hamathecium of cellular pseudoparaphyses. Asci 8-ascospored, bitunicate, fissitunicate, clavate, short-stipitate. Ascospores broadly fusoid with broadly rounded ends, 1-septate, constricted, hyaline, usually with sheath.
Anamorphs reported for genus: none.
Literature: Shearer et al. 2009; Zhang et al. 2009a, b.
Type species
Lentithecium fluviatile (Aptroot & Van Ryck.) K.D. Hyde, J. Fourn. & Yin. Zhang, Fungal Divers. 38: 234 (2009). (Fig. 43)
≡ Massarina fluviatilis Aptroot & Van Ryck., Nova Hedwigia 73: 162 (2001).
Ascomata 230–260 μm high × 280–325 μm diam., scattered or gregarious, immersed, slightly erumpent, subglobose to depressed spherical, under a small black pseudostroma originating from the apical part of the peridium, apex slightly papillate, ostiole rounded, 60–70 μm diam. (Fig. 43a and b). Peridium 15–20 μm thick at sides and at base, comprising 4–5 layers of angular cells more thick-walled outwards, 50–55 μm thick at apex, of small very thick-walled cells. Hamathecium of cellular pseudoparaphyses, 2–2.5 μm broad (Fig. 43c and d). Asci 89–100 × 19–21 μm, 8-spored, bitunicate, fissitunicate, clavate, bumpy, short-stipitate, apex without obvious apical chamber (Fig. 43e). Ascospores 27–35 × 8.5–9.4 μm,, 2-3-seriate, broadly fusoid with broadly rounded ends, straight to slightly curved, 1-septate, slightly constricted, with four large guttules, hyaline, smooth-walled, a very thin mucilaginous sheath can be occasionally observed in India ink but in most cases no sheath can be observed (Fig. 43f and g).
Anamorph: none reported.
Material examined: FRANCE, Haute Garonne: Avignonet, Lac de Rosel, artificial lake, on bark and wood of a submerged branch Populus sp., 23 Nov. 2006, leg. Michel Delpont, det. Jacques Fournier (IFRD 2039, holotype).
Notes
Morphology
Lentithecium was introduced to accommodate some freshwater fungi previous assigned under Massarina, such as M. arundinacea (Sowerby) Leuchtm. and M. fluviatilis (Zhang et al. 2009a). It is characterized by its immersed and lenticular ascomata, thin peridium which is almost equal in thickness, short pedicellate asci and fusoid or filliform, hyaline or rarely lightly pigmented, 1- to multi-septate ascospores (Zhang et al. 2009b). Lentitheciaceae was introduced to accommodate Lentithecium and some other related taxa (Zhang et al. 2009a).
Phylogenetic study
The clade of Lentitheciaceae comprises the generic type Lentithecium fluviatile, as well as L. arundinaceum (Sowerby) K.D. Hyde, J. Fourn. & Yin. Zhang, Stagonospora macropycnidia, Wettsteinina lacustris (Fuckel) Shoemaker & C.E. Babc., Keissleriella cladophila, and the bambusicolous species Katumotoa bambusicola and Ophiosphaerella sasicola, which receive high bootstrap support (Zhang et al. 2009a).
Concluding remarks
Tingoldiago graminicola K. Hirayama & Kaz. Tanaka form a robust clade with species of Lentithecium (Shearer et al. 2009). Tingoldiago has lenticular immersed to erumpent ascomata, numerous and septate pseudoparaphyses, cylindro-clavate asci and hyaline, 1-septate ascospores with sheath. All of these characters fit Lentithecium well. We treat Tingoldiago as a synonym of Lentithecium.
Leptosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1: 234 (1863). (Leptosphaeriaceae)
Generic description
Habitat terrestrial, saprobic or parasitic. Ascomata small- to medium-sized, solitary, scattered or in small groups, erumpent to superficial, subglobose, broadly or narrowly conical, papillate, ostiolate. Peridium thick, comprising layers of cells of textura angularis. Hamathecium of dense cellular pseudoparaphyses, embedded in mucilage, anastomosing and branching. Asci 8-spored, bitunicate, fissitunicate unknown, cylindrical with a furcate pedicel and a large ocular chamber. Ascospores fusoid or narrowly fusoid, brown or reddish brown, 3-septate, constricted at each septum.
Anamorphs reported for genus: Coniothyrium and Phoma (Hyde et al. 2011; Sivanesan 1984).
Literature: von Arx and Müller 1975; Barr 1987a, b; Cesati and de Notaris 1863; Crane and Shearer 1991; Dong et al. 1998; Eriksson 1967a; Eriksson and Hawksworth 1986, 1991; de Greuter et al. 1988; Hedjaroude 1969; von Höhnel 1907; Holm 1957, 1975; Huhndorf et al. 1990; Luttrell 1973; Müller 1950; Munk 1957; Saccardo 1878b, 1883, 1891, 1895; Schoch et al. 2009; Shearer 1993; Shearer et al. 1990; Shoemaker 1984a; Sivanesan 1984; Zhang et al. 2009a.
Type species
Leptosphaeria doliolum Ces. & De Not., Comm. Soc. crittog. Ital. 1: 234 (1863). (Fig. 44)
≡ Sphaeria doliolum Pers., Icon. Desc. Fung. Min. Cognit. (Leipzig) 2: 39 (1800).
Ascomata 340–450 μm high × 380–500 μm diam., solitary, scattered or in small groups, superficial, subglobose, broadly or narrowly conical, with a flattened base on the host surface, black, usually with 2–4 ring-like ridges surrounding the ascomata surface, apex with a conical, usually shiny papilla (Fig. 44a). Peridium 85–110 μm wide at sides, thinner at the apex, comprising two types of cells, outer layer composed of small thick-walled cells of textura angularis, cells <2 μm diam., cell wall up to 8 μm thick, surface heavily pigmented and inner lightly pigmented, apex cells smaller, walls thicker, and cells more heavily pigmented, inner layer composed of subhyaline relatively thin-walled cells of textura angularis, 3–6 μm diam., wall up to 5 μm, cells near the base larger and wall thinner and paler (Fig. 44b). Hamathecium of dense, long cellular pseudoparaphyses, 1.5–3 μm broad, embedded in mucilage, anastomosing and branching. Asci 110–150 × 7–9(−10) μm (\( \bar{x} = 130.6 \times 8.5\mu m \), n = 10), 8-spored, bitunicate, fissitunicate unknown, cylindrical, furcate pedicel which is usually less than 25 μm long, with a large ocular chamber (Fig. 44c, d and e). Ascospores 25–31 × 4.5–6 μm (\( \bar{x} = 27.7 \times 5.3\mu m \), n = 10), uniseriate and somewhat partially overlapping, narrowly fusoid with sharp to narrowly rounded ends, reddish brown, 3-septate, constricted at each septum, smooth (Fig. 44f).
Anamorph: Phoma hoehnelii (Sivanesan 1984).
Material examined: Herb., Persoon 910270–650 (L, lectotype).
Notes
Morphology
Leptosphaeria was first established by Cesati and de Notaris (1863) with 26 species included; L. doliolum (Pers.:Fr.) Ces. & De Not. was subsequently selected as the lectotype species (de Greuter et al. 1988; Holm 1975; Shearer et al. 1990). Leptosphaeria was originally defined based mainly on the characters of ascospores being ellipsoid or fusoid, one to many septa, hyaline to dark brown. These few common characters meant that Leptosphaeria comprised many species, and some of them should be assigned to either Euascomycetes or Loculoascomycetes (Crane and Shearer 1991). Leptosphaeria had been divided based on host and habitat (Saccardo 1878b, 1891, 1895) as well as the pseudothecium (glabrous, hairy, setose) and ascospore septation (see comments by Crane and Shearer 1991). von Höhnel (1907) used centrum structure in the classification of Leptosphaeria, and divided Leptosphaeria into three genera, viz. Leptosphaeria, Scleropleella and Nodulosphaeria. Müller (1950) subdivided Leptosphaeria into four sections based on pseudothecial and centrum structure as well as ascospore characters. This classification was modified by Munk (1957), who named these four sections as section I (Eu-Leptosphaeria), section II (Para-Leptosphaeria), section III (Scleropleella) and section IV (Nodulosphaeria). Holm (1957) used a relatively narrow concept for Leptosphaeria, which included species closely related to the generic type, L. doliolum. This viewpoint was accepted by some workers (Eriksson 1967a; Hedjaroude 1969; Shoemaker 1984a). Nevertheless, it still seems a heterogeneous group of fungi (see comments by Crane and Shearer 1991). Its position among the Loculoascomycetes is also debated. It has been placed in the Pleosporaceae (von Arx and Müller 1975; Luttrell 1973; Sivanesan 1984) or Leptosphaeriaceae (Barr 1987a, b; Eriksson and Hawksworth 1991) or Phaeosphaeriaceae (Eriksson and Hawksworth 1986).
Phylogenetic study
Molecular phylogenetic analysis based on multigenes indicated that species of Leptosphaeria (including the generic type L. doliolum) and Neophaeosphaeria form a paraphyletic clade with moderate bootstrap support (Dong et al. 1998; Schoch et al. 2009; Zhang et al. 2009a), which is sister to other families of Pleosporales (Zhang et al. 2009a). Thus the familial rank of the Leptosphaeriaceae could be temporarily verified, but further molecular phylogenetic study is needed in which more related taxa should be included.
Concluding remarks
Morphologically, Leptosphaeria is mostly comparable with Amarenomyces, Bricookea, Diapleella, Entodesmium, Melanomma, Nodulosphaeria, Paraphaeosphaeria, Passeriniella, Phaeosphaeria and Trematosphaeria. While it prefers non-woody parts of dicotyledonous hosts, its cylindrical ascus with short pedicel and smooth, fusoid and multi-septate ascospores make it readily distinguishable from all other genera (Shoemaker 1984a).
Leptosphaerulina McAlpine, Fungus diseases of stone-fruit trees in Australia and their treatment: 103 (1902). (Didymellaceae)
Generic description
Habitat terrestrial, parasitic or saprobic. Ascomata small, scattered, immersed, globose to subglobose, with a small, slightly protruding papilla, ostiolate. Peridium thin. Hamathecium of rare or decomposing cellular pseudoparaphyses. Asci bitunicate, obpyriform. Ascospores broadly clavate or cylindrical, hyaline, turning pale brown when old, asymmetrical, multi-septate, smooth-walled.
Anamorphs reported for genus: Pithoascus and Pithomyces (Hyde et al. 2011).
Literature: Barr 1972; Chlebicki 2002; Crivelli 1983; Kodsueb et al. 2006a; Zhang et al. 2009a.
Type species
Leptosphaerulina australis McAlpine, Fungus diseases of stone-fruit trees in Australia and their treatment: 103 (1902). (Fig. 45)
Ascomata 140–170 μm diam., scattered, immersed, globose to subglobose, with a small slightly protruding papilla, ostiolate (Fig. 45a). Peridium thin, composed of one or two layers of large cells of textura angularis, pale brown (Fig. 45a). Hamathecium of rare or decomposing cellular pseudoparaphyses, up to 5 μm broad, filling the gaps between the asci. Asci 38–53 × 55–75 μm (\( \bar{x} = 67.5 \times 43.3\mu m \), n = 10), 8-spored, without pedicel, bitunicate, fissitunicate dehiscence not observed, obpyriform, with a large ocular chamber and apical ring (Fig. 45b and c). Ascospores 30–40(-47) × 11–14 μm (\( \bar{x} = 36.5 \times 13\mu m \), n = 10), broadly clavate, hyaline, turning pale brown when old, asymmetrical, upper hemisphere usually with one transverse septum and with a somewhat narrowly rounded end, lower hemisphere usually with two transverse septa and with broadly rounded ends, slighted constricted at the primary septum, mostly with one vertical septum in each central cell, smooth, with thin gelatinous sheath when young, 2–3 μm thick (Fig. 45d and e).
Anamorph: none reported.
Material examined: USA, Kansas, Kansas State College, on Poa pratensis L. Grass plots, 2 Jul. 1953, leg. T. Rogerson, det. L.E. Wehmeyer (NY, C.T. Rogerson 3836).
Notes
Morphology
Leptosphaerulina, introduced by McAlpine (1902), is characterized by small immersed ascomata, obpyriform asci with a large ocular chamber and apical ring as well as muriformly septate ascospores which may be hyaline or pigmented. Species of Leptosphaerulina may occur on monocotyledons or dicotyledons. Leptosphaerulina is most comparable with Pleospora, and the only difference between them is that Leptosphaerulina has smaller ascomata and hyaline ascospores that only become pigmented after discharge, whereas the ascospores of Pleospora become brown within the asci. Currently, about 60 names are accepted in this genus, and some even reported from marine environments, e.g. L. mangrovei (Inderbitzin et al. 2000).
Phylogenetic study
Based on multigene phylogenetic analysis, two putative strains of Leptosphaerulina australis, the generic type of Leptosphaerulina, from Switzerland (CBS 311.51) and Indonesia (CBS 317.83) resided within Didymellaceae (de Gruyter et al. 2009; Zhang et al. 2009a).
Concluding remarks
Because of its morphological confusion with Pleospora and the diversity of habitats within the genus, Leptosphaerulina sensu lato is likely to be polyphyletic. Fresh collections of this species are needed from Australia to epitypify this taxon and define the genus in a strict sense. The specimen described here is a collection from USA and therefore may not represent the type.
Lewia M.E. Barr & E.G. Simmons, Mycotaxon 25: 289 (1986). (Pleosporaceae)
Generic description
Habitat terrestrial, parasitic or saprobic? Ascomata small, scattered, erumpent to nearly superficial at maturity, subglobose to globose, black, smooth, papillate, ostiolate. Papilla short, blunt. Peridium thin. Hamathecium of pseudoparaphyses. Asci (4–6-)8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel. Ascospores muriform, ellipsoid to fusoid.
Anamorphs reported for genus: Alternaria (Simmons 1986).
Literature: Kwasna and Kosiak 2003; Kwasna et al. 2006; Simmons 1986, 2007; Vieira and Barreto 2006.
Type species
Lewia scrophulariae (Desm.) M.E. Barr & E.G. Simmons, Mycotaxon 25: 294 (1986). (Fig. 46)
≡ Sphaeria scrophulariae Desm., Plantes cryptogames du Nord de la France, ed. 1 fasc. 15:no. 718 (1834).
Ascomata ca. 150–200 μm diam., scattered, erumpent to nearly superficial at maturity, subglobose to globose, black, smooth, papillate. Papilla short, blunt. Peridium thin. Hamathecium of septate pseudoparaphyses, ca. 2–2.5 μm broad, anastomosing or branching not observed. Asci 100–140 × 13–17 μm, (4–6-)8-spored, bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel, ocular chamber unknown (Fig. 46a). Ascospores ellipsoid, 5 (rarely 6 or 7) transversal septa and one longitudinal septum mostly through the central cells, yellowish brown to gold-brown, 20–24 × 8–10 μm (\( \bar{x} = 21.5 \times 9.1\mu m \), n = 10), constricted at median septum, smooth or verruculose (Fig. 46b, e and f).
Anamorph: Alternaria conjuncta (Simmons 1986).
Primary conidiophore simple with a single conidiogenous locus; conidia produced in chains, the first conidia in chain is larger, 30–45 × 10–12 μm, 7 transverse septa, 1–2 longitudinal or oblique septa in lower cells. Secondary conidiophore with 5–7 conidiogenous loci, sometimes branched; sporulation in chains, rarely branched.
Material examined: (FH, slide from lectotype).
Note: The specimen contains only a slide, so limited structures could be observed e.g. ascospores. The information about ascomata, peridium and whole asci is referred to Simmons (1986).
Notes
Morphology
Lewia has “Pleospora-like” teleomorphs, while it has Alternaria anamorphs, which are characterized by the beakless conidia connected together with secondary conidiophore (Simmons 1986). Based on these characters, more species under this genus were subsequently reported, i.e. Lewia avenicola Kosiak & Kwaśna (Kwasna and Kosiak 2003); L. chlamidosporiformans B.S. Vieira & R.W. Barreto (Vieira and Barreto 2006); L. alternarina (M.D. Whitehead & J.G. Dicks.) E.G. Simmons and L. daucicaulis E.G. Simmons (Simmons 2007). Currently Lewia comprises 15 species (http://www.mycobank.org, 24-02-2009).
Phylogenetic study
Phylogenetic analysis based either on SSU rDNA sequences or on multigenes indicated that Lewia species (Allewia eureka (E.G. Simmons) E.G. Simmons = L. eureka) form a robust clade with other members of Pleosporaceae (Schoch et al. 2006; Schoch et al. 2009; Zhang et al. 2009a).
Concluding remarks
Its position in Pleosporaceae is confirmed.
Lichenopyrenis Calat., Sanz & Aptroot, Mycol. Res. 105: 634 (2001). (?Pleomassariaceae)
Generic description
Habitat terrestrial, parasitic on lichens. Ascomata medium-sized, globose or subglobose. Hamathecium of dense, filliform, branching, septate pseudoparaphyses. Asci bitunicate, fissitunicate, clavate, with a short sometimes furcate pedicel. Ascospores ellipsoidal with broadly rounded ends, pale orange-brown, 1-distoseptate.
Anamorphs reported for genus: see below.
Literature: Calatayud et al. 2001.
Type species
Lichenopyrenis galligena Calat., Sanz & Aptroot, Mycol. Res. 105: 636 (2001). (Fig. 47)
Ascomata 140–260 μm high × 140–250 μm diam., gregarious, initially immersed in galls, later becoming erumpent, globose or subglobose, black, roughened (Fig. 47a and b). Peridium 18–25 μm wide, composed of 2–5 layers of heavily pigmented cells of textura angularis to compressed, cells 6–11 μm diam., cell wall 1–3 μm thick (Fig. 47c, d and e). Hamathecium of dense, long filamentous pseudoparaphyses, 2.5–4 μm broad, branching, septate. Asci 65–85 × 15–20 μm (\( \bar{x} = 74 \times 18\mu m \), n = 10), 8-spored, bitunicate, fissitunicate, broadly clavate, with a short, thick, sometimes furcate pedicel, up to 13 μm long, ocular chamber not observed (Fig. 47f, g, h and k). Ascospores 16–20 × 9–11 μm (\( \bar{x} = 18 \times 10\mu m \), n = 10), biseriate, ellipsoidal, pale orange-brown, 1-distoseptate, with prominent swelling at the septum, containing refractive globules, smooth (Fig. 47i, j and l).
Anamorph: The following description is from Calatayud et al. (2001).
Conidiomata pycnidial, arising in galls together with the ascomata, immersed, ca. 100–200 μm diam.; wall dark brown throughout, composed of 2–5 layers of angular to laterally compressed cells; cells relatively large, ca. 8–16 μm diam. in superficial view. Conidiophores formed by 1–3 cells, frequently branched and with the uppermost cells bearing 1–4 conidiogenous cells; cells ± cylindrical, hyaline except at the base, which are sometimes pale brown, 7–15 × 3–4 μm. Conidiogenous cells tapered towards the apex, 14–18 × 3–4 μm. Conidia 5–7 × 1.5–2 μm. Vegetative hyphae hyaline.
Material examined: SPAIN, Andalucía, Province, Jaén, Andújar, lichenicolous on Leptochidium albociliatum (Desm.) M. Choisy on acid volcanic rock, 19 Apr. 2000, V. Calatayud (MA-Lichen 12715, holotype).
Notes
Morphology
Lichenopyrenis was formally established by Calatayud et al. (2001) based on its “perithecioid ascomata with peridium comprising compressed cells, fissitunicate and J- asci, wide hamathecium filaments, and 1-septate pale orange-brown ascospores with distoseptate thickenings at maturity”, and is monotypic with L. galligena. The genus was temporarily assigned to Pleomassariaceae. Lichenopyrenis galligena is a parasite of lichens, occurring in galls in the thallus of the host (Calatayud et al. 2001).
Phylogenetic study
None.
Concluding remarks
This is one of the few species that are parasitic on lichens. The most comparable species are Parapyrenis lichenicola Aptroot & Diederich and Lacrymospora parasitica Aptroot (both in Requienellaceae, Pyrenulales) as well as some species from Dacampiaceae. The peridium structure, cellular pseudoparaphyses, distoseptate and smooth, orange-brown ascospores as well as the anamorphic stage of Lichenopyrenis can easily distinguish from all of them (Calatayud et al. 2001).
Lineolata Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 687 (1990). (Pleosporales, genera incertae sedis)
Generic description
Habitat marine, saprobic (or perthophytic?). Ascomata medium-sized, gregarious, immersed to erumpent, obpyriform, ostiolate, papillate. Peridium thin, comprising two types of cells; outer cells thick stratum pseudostromatic, inner stratum thin, composed of a few layers of hyaline cells of textura angularis. Hamathecium of dense, long trabeculate pseudoparaphyses, embedded in mucilage, anastomosing and septate. Asci 8-spored, bitunicate, cylindrical, with short pedicels, with an ocular chamber. Ascospores uniseriate to partially overlapping, ellipsoidal, dark brown, 1-septate.
Anamorphs reported for genus: none.
Literature: Kohlmeyer and Kohlmeyer 1966; Kohlmeyer and Volkmann-Kohlmeyer 1990.
Type species
Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: 688 (1990). (Fig. 48)
≡ Didymosphaeria rhizophorae Kohlm. & E. Kohlm. Icones Fungorum Maris (Lehre) 4 & 5: tab. 62a (1967).
Ascomata 300–490 μm high × 200–360 μm diam., gregarious, immersed to erumpent, obpyriform, ostiolate, papillate, subcarbonaceous to subcoriaceous, blackish brown (Fig. 48a). Peridium 37–45 μm thick, comprising two types of cells; outer cells thick stratum pseudostromatic, composed of irregular or roundish, dark brown cells, on the outside with a more or less recognizable hyphal structure, enclosing some decaying cells of the host, inner stratum thin, composed of four or five layers of hyaline, polygonal, elongate, thin-walled cells with large lumina, merging into the pseudoparaphyses. Hamathecium of dense, long trabeculate pseudoparaphyses, 1–1.5 μm broad, embedded in mucilage, anastomosing and septate. Asci 150–175 × 14–17.5 μm, 8-spored, bitunicate, cylindrical, with short pedicels, with an ocular chamber (Fig. 48b). Ascospores 23–32(−33) × 9–12 μm, uniseriate to partially overlapping, ellipsoid, dark brown, 1-septate, not or slightly constricted at the septum, striate by delicate costae that run parallel or in a slight angle to the longitudinal axis of the ascospore (Fig. 48c, d, e and f) (adapted from Kohlmeyer and Kohlmeyer 1979).
Anamorph: none reported.
Material examined: US, Florida, Middle Torch Key, on Rhizophora mangle, 21 Nov. 1965, J. Kohlmeyer (Herb. J. Kohlmeyer No. 2390b, isotype); Pirate Grove Key, on R. mangle, 5 Jan. 1964 (Herb. J. Kohlmeyer No. 1721 paratype); Florida, Virginia Key, on R. mangle, 1 Jan. 1964, leg. E. Kohlmeyer (Herb. J. Kohlmeyer No. 1751 paratype); Florida, Torch Key, on R. mangle, 20 Nov. 1965, leg. J. Kohlmeyer (Herb. J. Kohlmeyer No. 2423 paratype).
Notes
Morphology
Lineolata was monotypified by L. rhizophorae, which was originally introduced by Kohlmeyer and Kohlmeyer (1966) as a species of Didymosphaeria (as D. rhizophorae). Based on the morphology of ascomata and asci, Barr (1990a) assigned it under Lojkania (as L. rhizophorae). Kohlmeyer and Volkmann-Kohlmeyer (1990) restudied this species and noticed that the absence of clypeus, almost superficial ascomata, coloured peridium, a hamathecium with gelatinous matrix, asci with apical ring-like structure and the ornamented ascospores are quite different from the modified concept of Didymosphaeria. Thus they introduced Lineolata to accommodate D. rhizophorae (Kohlmeyer and Volkmann-Kohlmeyer 1990).
Phylogenetic study
Three isolates of Lineolata rhizophorae from varied geographic localities were analyzed by Suetrong et al. (2009) and shown to be related to Caryosporella rhizophorae in Dothideomycetidae and excluded from Pleosporomycetidae and Pleosporales.
Concluding remarks
Based on initial molecular work it is likely that this species does not belong to Pleosporales in spite of its dense pseudoparaphyses and other characters shared with the order.
Loculohypoxylon M.E. Barr, Mycotaxon 3: 326 (1976). (Teichosporaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata relatively small, gregarious, immersed to erumpent, globose or subglobose, forming under a clypeus, papillate, ostiolate. Peridium thin, a single layer comprising hyaline thin-walled cells of textura angularis or textura prismatica. Hamathecium of septate pseudoparaphyses. Asci (2–4-)8-spored, bitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel, and wide ocular chamber. Ascospores broadly elliptic to subglobose, often apiculate at both ends, pale to dark brown, aseptate, with a germ slit.
Anamorphs reported for genus: none.
Literature: von Arx and Müller 1975; Barr 1976.
Type species
Loculohypoxylon grandineum (Berk. & Rav.) Barr, Mycotaxon 3: 326 (1976). (Fig. 49)
≡ Diatrype grandinea Berk. & Rav., in Berkeley, Grevillea 4: 95 (1876).
Ascomata 85–130 μm high × 75–145 μm diam., gregarious, immersed to widely erumpent, globose or subglobose, under a reddish brown to black clypeus, papillate, ostiolate (Fig. 49a and b). Peridium 18–30 μm thick laterally, 1-layered, composed of hyaline thin-walled cells of textura angularis to prismatica, cells up to 5 × 9 μm diam., cell wall 0.5–1 μm thick, apex cells smaller and walls thicker (Fig. 49c). Hamathecium comprising 2–3 μm broad, septate pseudoparaphyses. Asci 70–90 × 10–12.5 μm (\( \bar{x} = 76.5 \times 10.9\mu m \), n = 10), (2–4-)8-spored, bitunicate, cylindrical to cylindro-clavate, with a short, furcate pedicel, up to 25 μm long, with a wide ocular chamber (Fig. 49f, g, and h). Ascospores 7.5–10 × 5–7 μm (\( \bar{x} = 8.3 \times 5.9\mu m \), n = 10), uniseriate to partially overlapping at the upper part, broadly elliptic to subglobose, often apiculate at both ends, pale to dark brown, aseptate, with a germ slit (Fig. 49d and e).
Anamorph: none reported.
Material examined: USA, New Jersey, Newfield, on bark of Quercus coccinea, Sept. 1878, as Diatrype grandinea, Ellis N.A.F. 494 (NY, MASS); on Quercus sp. wood, Nov. 1893, as Anthostoma grandinea B. & Rav., Ellis & Everhart, N.A.F. 494 (NY); Newfield, Oct. 1881, as Diatrype grandinea (NY); Newfield, Jan. 1882, on Quercus coccinea, as Diatrype grandinea B. & Rav, Ex Herb Ellis (NY); Newfield, Nov. 1893, as Anthostoma grandinea, on bark of fallen trunks of Quercus coccinea (NY).
Notes
Morphology
Loculohypoxylon grandineum is one of the rare pleosporalean species having aseptate ascospores. When emphasis is given to ascospore morphology, Semidelitschia (monotypified by S. agasmatica Cain & Luck-Allen) is the most comparable genus. The large ascomata and ascospores, the mucilaginous sheath surrounding the ascospores as well as the coprophilous habitat of S. agasmatica differ from L. grandineum greatly. Thus Loculohypoxylon was introduced as a new genus.
Phylogenetic study
None.
Concluding remarks
Aseptate ascospores are rare in Pleosporales, and the position of this fungus needs further verification. The familial status of Loculohypoxylon in Teichosporaceae is questionable, as it is simply based on the similarity of living habitat, ascomata and asci with Immotthia and Teichospora (Barr 2002).
Lophionema Sacc., Syll. fung. (Abellini) 2: 717 (1883). (Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic? Ascomata solitary, scattered or in small groups, immersed to erumpent, globose to subglobose, with a flattened base, wall black, papillate, ostiolate. Peridium comprising two types of cells which merge in the middle. Hamathecium of trabeculate pseudoparaphyses, septate, rarely anastomosing and branching. Asci 8-spored, bitunicate, fissitunicate unknown, clavate to cylindro-clavate, with a short and furcate pedicel and a small inconspicuous ocular chamber. Ascospores filliform, hyaline to pale yellow, multi-septate, slightly constricted at each septum.
Anamorphs reported for genus: none.
Literature: Barr 1992b; Chesters and Bell 1970; Ellis and Everhart 1892; Höhnel 1909; Solheim 1949.
Type species
Lophionema vermisporum (Ellis) Sacc., Syll. fung. (Abellini) 2: 717 (1883). (Fig. 50)
≡ Lophiostoma vermispora Ellis, Bull. Torrey bot. Club 9: 19 (1882).
Ascomata 320–430 μm high × 280–350 μm diam., solitary, scattered or in small groups of 2–3, immersed to erumpent, globose to subglobose, black, papillate, ostiolate. Papilla 80–120 μm high, up to 150 μm broad, cylindrical to somewhat vertically flattened neck; mostly with a short slot-like ostiole, periphysate (Fig. 50a). Peridium 30–45 μm wide at the sides and slightly thicker at the apex, 2-layered, lateral walls and wall adjacent to neck comprising two types of cells which merge in the middle; outer cells small heavily pigmented thick-walled cells of textura angularis, cells 4–7 μm diam., cell wall 3.5–5 μm thick, inner cells less pigmented, comprising thin-walled compressed cells; apical wall cells smaller and walls thicker, basal wall thinner (ca. 15 μm wide), composed of lightly pigmented thin-walled compressed cells (Fig. 50b and c). Hamathecium of trabeculate pseudoparaphyses, 1–2 μm broad, septate, anastomosing and branching rarely between and mostly above the asci. Asci 105–130(−150) × 10–15 μm (\( \bar{x} = 123 \times 12\mu m \), n = 10), 8-spored, bitunicate, fissitunicate dehiscence not observed, clavate to cylindro-clavate, with a short, narrow, furcate pedicel which is 10–25 μm long, and a small inconspicuous ocular chamber (to 1.5 μm wide × 1 μm high) (Fig. 50d and e). Ascospores (80-)90–115 × 3–5 μm (\( \bar{x} = 95 \times 3.5\mu m \), n = 10), filliform, gradually tapering towards the base, hyaline to light yellow, (6-)7(−8)-septate, slightly constricted at each septum, smooth (Fig. 50f).
Anamorph: none reported.
Material examined: USA, New Jersey, Newfield, on dead stems of Oenothera biennis, Aug. 1881, Ellis (NY 643, holotype, NY 885, isotype).
Notes
Morphology
Lophionema is a relatively poorly studied genus, which was formally established by Saccardo (1883) as a monotypic genus represented by L. vermisporum based on its “globose ascomata, compressed ostiole, cylindrical to clavate ascus, and filamentous, septate, subhyaline to lightly pigmented ascospores”. Lophionema vermisporum was consequently listed as the generic type (Clements and Shear 1931). Berlese (1890) placed the genus in Lophiostomataceae but mentioned that the genus was similar to Ophiobolus according to the variable apex, and Shoemaker (1976) transferred Lophionema vermisporum to Ophiobolus sensu lato. Chesters and Bell (1970) however, had regarded Lophionema as related to Lophiostoma despite the distinct ascospore morphology. Barr (1992b) assigned Lophionema to Entodesmium based on the morphology of ascomata, papilla, peridium structure, pseudoparaphyses as well as the hyaline or slightly yellowish ascospores with a terminal appendage (not observed here). Species of Entodesmium, however, exclusively occur on legumes, but Lophionema vermisporum does not. We also note that the filliform ascospores, bitunicate asci, pseudoparaphyses and nature of the peridium may also be considered as typical of genera in the Tubeufiaceae (Barr 1980; Kodsueb et al. 2006b).
Phylogenetic study
None.
Concluding remarks
The immersed to erumpent ascomata, trabeculate pseudoparaphyses and laterally flattened papilla and periphysate ostioles indicate that this genus should be included in Lophiostomataceae. We do not accept the above proposals and, consider that Lophionema should be maintained as a separate genus with filliform ascospores in Lophiostomataceae until representative taxa can be sequenced and analyzed. Currently Lophionema comprises 10 species (http://www.mycobank.org, 08-01-2009). However, many of these are poorly studied and obscure.
Lophiostoma Ces. & De Not., Comm. Soc. crittog. Ital. 1: 219 (1863). (Lophiostomataceae)
Generic description
Habitat terrestrial, saprobic. Ascomata immersed to erumpent, usually with a distinct depressed papilla and a slot-like ostiole. Hamathecium of dense, long, septate pseudoparaphyses, embedded in mucilage, anastomosing and branching between and above the asci. Peridium unequal in thickness, thicker near the apex and thinner at base. Asci usually clavate. Ascospores 1-septate, multi-septate or even muriform, hyaline to deep brown, usually with terminal appendages.
Anamorphs reported for genus: Pleuorphomopsis-like (Hyde et al. 2011).
Literature: Barr 1990a; Chesters and Bell 1970; Holm and Holm 1988; Hyde and Aptroot 1998; Hyde et al. 2002; Tanaka and Harada 2003b; Yuan and Zhao 1994.
Type species
Lophiostoma macrostomum (Tode) Ces. & De Not., Comm. Soc. crittog. Ital. 1: 219 (1863). (Fig. 51)
≡ Sphaeria macrostoma Tode, Fung. mecklenb. sel. (Lüneburg) 2: 12 (1791).
Ascomata 400–600 μm high × 420–560 μm diam., densely scattered to gregarious, semi-immersed to erumpent, globose or subglobose, with a small to large flattened crest-like raised area above the ascomata which is variable in shape, up to 300 μm high and 480 μm wide, with a slit-like ostiole along the full length of the crest (Fig. 51a and b). Peridium 30–45 μm thick at the sides, thicker at the apex and thinner at the base, composed of one cell type of small lightly pigmented thin-walled cells of textura prismatica, cells ca. 6–9 × 3–4 μm diam., apex composed of pseudoparenchymatous cells (Fig. 51b). Hamathecium of dense, filliform, up to 3 μm near the base and less than 1.5 μm broad in the upper place, septate pseudoparaphyses, embedded in mucilage, anastomosing and branching between and above the asci (Fig. 51f). Asci 110–145 × 10–15 μm (\( \bar{x} = 127.5 \times 13\mu m \), n = 10), 8-spored, bitunicate, fissitunicate (ectotunica no constriction), cylindro-clavate, with a furcate pedicel and a small ocular chamber (to 1.5 μm wide × 2 μm high) (J-) (Fig. 51c, d and e). Ascospores 27–38(−43) × 5–7.5 μm (\( \bar{x} = 31.2 \times 6.4\mu m \), n = 10), biseriate, fusoid, curved, hyaline, usually 1-septate, with 3–5 septa and faintly brown when old, with (2-)3(−4) distinct oil drops in each cell and short terminal appendage at ends (Fig. 51h, i and j), and ornamented with warts when spores are senescent (Fig. 51g).
Anamorph: none reported.
Material examined: SWEDEN, Smaland, Femsjö par., Femsjö, on Prunus, 2006, Elias Fries, det. Geir Mathiassen (UPS, lectotype, as Sphaeria macrostoma Fr.). FRANCE, Ariège, Rimont, Las Muros, on dead stems of Vitis vinifera, 2 Sept. 1996 (IFRD2005).
Notes
Morphology
Lophiostoma is morphologically a well studied genus (Barr 1990a; Chesters and Bell 1970; Holm and Holm 1988; Mugambi and Huhndorf 2009b; Yuan and Zhao 1994), and currently it comprises about 30 species (Tanaka and Harada 2003b). The genus was characterized as having immersed to erumpent ascomata with a cylindrical or crest-like papilla and full length, slit-like ostiole; a peridium of unequal thickness, which was broader near the base (Lophiostoma-type); mostly clavate, bitunicate asci and 1- to several septate, hyaline to pigmented ascospores with terminal appendages or surrounded by a mucilaginous sheath (Holm and Holm 1988). This definition was followed by Barr (1990a), Yuan and Zhao (1994) and Hyde et al. (2002).
The crest-like papilla has been regarded as a prominent morphological character of Lophiostoma macrostomum (Chesters and Bell 1970; Holm and Holm 1988). In the lectotype specimen, the raised area above the ascomata is up to 300 μm high and 480 μm long, and seen as a flattened or even Y-shaped crest (Fig. 51a). In Lophiostoma curtum (Fr.) De Not. and Lophiotrema boreale Math. the raised area above the ascomata varies considerably in height or is even lacking (Holm and Holm 1988). Thus the variable “crest-like raised area in Lophiostomataceae” was explained as an evolutionarily adaptation to the hard substrate within which the ascomata develop (Holm and Holm 1988). The ascospores of L. macrostomum usually turn reddish brown when mature, and minutely verrucose ornamentation was also found on the surface of the pigmented ascospores. Hyaline ascospores that became pigmented with age are common in Lophiostoma, such as in L. appendiculatum Fuckel, L. massarioides (Sacc.) L. Holm & K. Holm, L. semiliberum, L. subcorticale Fuckel and L. winteri (Holm and Holm 1988; Tanaka and Harada 2003b). The phylogenetic significance of this character should be observed carefully in the future but at present its phylogenetic significance is unclear as this also occurs in some Lophiotrema species.
Phylogenetic study
Phylogenetic affinity with some Massarina species has been reported by Liew et al. (2002), and several Massarina species were transferred into Lophiostoma. In a systematic study of Lophiostoma- and Massarina-related fungi conducted by Zhang et al. (2009b), Lophiostoma taxa clustered into two groups; one includes the type species L. macrostomum with crest-like ostioles, L. rugulosum Yin. Zhang, J. Fourn. & K.D. Hyde with a wide, umbilicate pore surrounded by 4–6 radial ridges, and L. glabro-tunicatus with small ostiolar pores; the other cluster comprises Lophiostoma-like taxa with slot-like ostioles lacking raised crests, which includes L. arundinis (Pers.) Ces. & De Not., L. caulium, L. compressum (Pers.) Ces. & De Not., L. crenatum (Pers.) Fuckel, L. fuckelii (Sacc.) Sacc., L. macrostomoides, L. semiliberum and L. viridarium Cooke, which seems to represent a natural group at the family level. This conclusion is tentative until verified sequences of L. macrostomum are included in analyses (see comments of Zhang et al. 2009a).
Concluding remarks
We tend to accept a narrow concept of Lophiostomataceae, which only comprises species of Lophiostoma sensu stricto (Zhang et al. 2009a).
Lophiotrema Sacc., Michelia 1: 338 (1878). (Pleosporales, genera incertae sedis)
Generic description
Habitat terrestrial, saprobic. Ascomata small- to medium-sized, with or without short papilla. Hamathecium of dense, long, septate pseudoparaphyses, anastomosing and branching between and above asci. Asci cylindrical to cylindro-clavate. Ascospores hyaline, 1–3-septate, usually with mucilaginous sheath.
Anamorphs reported for genus: none.
Literature: Barr 1990a; Chesters and Bell 1970; Holm and Holm 1988; Saccardo 1878a; Tanaka and Harada 2003c; Tang et al. 2003; Yuan and Zhao 1994.
Type species
Lophiotrema nucula (Fr.) Sacc., Michelia 1: 338 (1878). (Fig. 52)
≡ Sphaeria nucula Fr., Syst. mycol. (Lundae) 2: 466 (1823).
Ascomata 200–240 μm high × 200–280 μm diam., scattered, erumpent to nearly superficial, with basal wall remaining immersed in host tissue, globose to subglobose, often laterally flattened, with a flattened base not easily removed from the substrate, black, roughened; with a cylindrical or slightly compressed papilla. Papilla to 120 μm long and 150 μm high, protruding, with a pore-like ostiole (Fig. 52a). Peridium 25–30 μm wide, very thin at the base, composed of heavily pigmented pseudoparenchymatous cells near the apex, cells 2–2 × 6 μm diam., wall 1–3(−4) μm thick, lower sides composed of pigmented cells of textura angularis, 3–5 μm diam., wall 0.8–1.5 μm thick, ostiole wall composed of heavily pigmented and thick-walled small cells (Fig. 52b and c). Hamathecium of dense, long, septate pseudoparaphyses, 1–2 μm broad, anastomosing and branching between and above asci, embedded in mucilage (Fig. 52i). Asci 90–115 × 9–11.5 μm (\( \bar{x} = 99.5 \times 11.5\mu m \); n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, with a short, narrowed, furcate pedicel which is up to 10 μm long, with a small ocular chamber (ca. 1.5 μm wide × 1 μm high) (J-) (Fig. 52d, e, f and h). Ascospores 17–21(−25) × (4-)5–6.5 μm (\( \bar{x} = 19.5 \times 5.5\mu m \), n = 10), obliquely uniseriate and partially overlapping to biseriate, broadly fusoid to fusoid, with narrowly rounded ends, hyaline and lightly pigmented on very rare occasions when senescent, 1-septate, 3-septate when old, constricted at the median septum, the upper cell often broader than the lower one (Fig. 52g).
Anamorph: none reported.
Material examined: on decaying wood (UPS, lectotype as Sphaeria nucula Fr.).
Notes
Morphology
Holm and Holm (1988) provided a relatively strict definition for Lophiotrema after they examined several specimens including the type material which they lectotypified. Lophiotrema was mainly defined on the unique characters of small to medium ascomata, a “Lophiotrema-type” peridium and 1-septate ascospores. In Lophiotrema, Holm and Holm (1988) considered the ascomata to be small- to medium-sized, ca. pyriform but neck often reduced, even lacking and sometimes cylindrical. The peridium was of approximately equal thickness, 20–30 μm, composed of an outer textura angularis of uniformly pigmented cells, up to 12 μm, and an inner layer of very small hyaline cells, with somewhat thickened walls. Asci are cylindrical, spores hyaline, at first 1-septate, becoming 3-septate, with distinct guttules, often with a mucilaginous sheath. Much emphasis was given to the 1-septate ascospores by Holm and Holm (1988) when they described and distinguished the three Lophiotrema species: L. boreale, L. nucula, L. vagabundum (Sacc.) Sacc. and two other unnamed species. This concept was widely accepted by later workers (Kirk et al. 2001; Yuan and Zhao 1994). Tanaka and Harada (2003c) considered the peridium and asci to distinguish Lophiotrema from Lophiostoma, while Tang et al. (2003) introduced a new Lophiotrema species with elongated slit-like ostiole stating that the main difference between Lophiotrema and Lophiostoma were size of ascomata, structure of peridium, shape of asci and sheath of ascospores. This peridium concept however, is not supported by the lectotype specimen we examined here, which has a flattened thin-walled base. Thus the “Lophiotrema-like peridium” sensu Holm and Holm (1988) should not serve as a diagnostic character of Lophiotrema, while the ostiole, asci and ascospores might have some phylogenetic significance (Zhang et al. 2009b). No anamorph is yet known for Lophiotrema. Although the ascospores was reported by Holm and Holm (1988) to be verruculose this could not be observed in the lectotype examined under light microscope (1000 ×) in the present study.
Phylogenetic study
In the phylogenetic study of Lophiostoma, Massarina and related genera (Zhang et al. 2009b), Lophiotrema nucula formed a consistent and robust clade with three other Lophiotrema species: L. lignicola Yin. Zhang, J. Fourn. & K.D. Hyde, L. brunneosporum Yin. Zhang, J. Fourn. & K.D. Hyde and L. vagabundum, separate from other members of Lophiostoma and Massarina sensu stricto. This clade might represent Lophiotrema sensu stricto, however, the correctness of strains of L. vagabundum (CBS 628.86) and L. nucula (CBS 627.86) used in the phylogenetic study are not verified and warrant further study.
Concluding remarks
Holm and Holm (1988) distinguished Lophiostoma from Lophiotrema based on the smaller ascomata, 1-septate versus multi-septate ascospores, and peridial wall structure. However, we doubt that these distinguishing characters (size of ascomata, number of septa of ascospores) can be confidently used to separate these genera and we could not establish any characters that could reliably distinguish between these two genera. The molecular data, however, does separate Lophiostoma macrostomum and Lophiotrema nucula into separate clades and provides some support that these are separate genera. Although the strain of L. nucula (CBS 627.86) was isolated by K. & L. Holm, who had examined the type specimen of L. nucula (Holm and Holm 1988), the culture of Lophiostoma macrostomum used in the analysis are unverified (see comment by Zhang et al. 2009b). For the purpose of this monograph we tentatively maintain Lophiotrema as distinct from Lophiostoma.
Macroventuria Aa, Persoonia 6: 359 (1971). (Didymellaceae)
Generic description
Habitat terrestrial, saprobic. Ascomata small, solitary, scattered, or in groups, initially immersed, becoming erumpent, to nearly superficial, globose to subglobose, roughened with cylindrical setae erect from apex. Peridium thin, membranous. Hamathecium of cellular pseudoparaphyses, seems to easily disappear when mature. Asci bitunicate, somewhat obclavate to fusoid. Ascospores fusoid with broadly to narrowly rounded ends, hyaline, 1-septate, constricted at the septum.
Anamorphs reported for genus: none.
Literature: van der Aa 1971; von Arx and Müller 1975; Barr 1987a.
Type species
Macroventuria wentii Aa, Persoonia 6: 361 (1971). (Fig. 53)
Ascomata 135–180 μm diam., rarely more than 200 μm diam., solitary, scattered or in groups, initially immersed, becoming erumpent, to nearly superficial, with basal wall remaining immersed in host tissue, globose to subglobose, broadly or narrowly conical, setae erect from the apical region of the ascomata, cylindrical or tapering to the rounded or pointed tip, brown, up to 90 μm long, 5–7.5 μm thick (Fig. 53a). Peridium, 25–35 μm thick, 2-layered, out layer composed of relatively thick-walled cells of textura angularis, cell wall up to 3 μm thick; inner layer cells with a thinner wall and subhyaline; near apex cells smaller (Fig. 53a). Hamathecium of cellular pseudoparaphyses, 1–2 μm thick, evanescing not sure. Asci 75–93 × 24–30 μm, 8-spored, without pedicel, bitunicate, somewhat obclavate to fusoid (Fig. 53b). Ascospores 22–32 × 8–14 μm, 1–3 seriate, fusoid with broadly to narrowly rounded ends, hyaline, 1-septate