Catenularia cupulifera (Berk. & Broome) Réblová & A.N. Mill.

Colonies effuse, hairy to velutinous, brown, dark brown to black, mycelium partly immersed, partly superficial; composed of conidiophores, capitate hyphae and sometimes ascomata. Anamorph. Conidiophores macronematous, mononematous, solitary or in tufts, with dark stromatic hyphal cells around the bases, erect, straight or flexuous, unbranched, brown to dark brown, thick-walled, paler and thinner-walled towards the apex. Capitate hyphae scattered among the conidiophores, occasionally absent, erect, brown, extending percurrently, paler towards the apex, apical cell sterile, thin-walled, subhyaline to hyaline, slightly swollen, broadly rounded with a hyaline mucilaginous cap that may disappear with age. Conidiogenous cells integrated, terminal, monophialidic, extending percurrently, cylindrical, subcylindrical or somewhat lageniform, brown, conidia produced successively; collarettes cup- or funnel-shaped, brown, smooth or slightly roughened, margin entire or frayed. Conidia cuneiform, obclavate, rounded-obconic to broadly obovoid in side view, with an angular outline when viewed from above with 3–6 blunt corners, broadly rounded to flattened at the apex, truncate at the distinctive, hyaline basal hilum, with a small, circular, thin-walled, pore-like area visible in the cell wall at each corner, sometimes with a visible central pore at the base, aseptate, hyaline when young, fuscous, fulvous, brown to dark brown at maturity, thick-walled, smooth; formed singly, adhered in basipetal chains, occasionally in clusters. Teleomorph. Ascomata perithecial, non-stromatic, superficial, globose, subglobose to conical, papillate, glabrous occasionally with a powdery layer that disappears with age, sometimes covered with conidiophores and capitate hyphae. Ostiolar canal periphysate. Ascomatal wall carbonaceous, two-layered. Paraphyses persistent, branching, anastomosing, hyaline, longer than the asci. Asci unitunicate, short-stipitate, apical annulus non-amyloid, with eight ascospores. Ascospores fusiform, transversely septate, hyaline, smooth, without mucilaginous sheath or appendages.

Saprobe on decaying bark, wood and bamboo culms of various hosts. Members of Catenularia have a worldwide distribution in temperate, subtropical and tropical geographic areas.

Hughes (1965) considered capitate hyphae to be an important diagnostic characteristic of Catenularia. These structures have long escaped attention, and mycologists began to notice them only after they were described by Hughes (1949). We studied holotype material of several species and original descriptions and illustrations to examine and trace this character in Catenularia. Capitate hyphae have not been mentioned in the original descriptions of C. cupulifera (Berkeley and Broome 1871; Richon 1877; Grove 1886). In studying collections of this species, we observed a variation in the presence of capitate hyphae. In some specimens, capitate hyphae are abundantly present, but may be scarce and difficult to find in others. Revision of the holotypes of C. cuneiformis var. minor (Holubová-Jechová 1983) and Ch. trianguloconidia (Réblová and Seifert 2003) not only revealed that both fungi are conspecific, but also led to the discovery of capitate hyphae, although they were not mentioned in the protologues of either species. They are scattered among conidiophores and easy to overlook. Phylogenetic analysis of several Catenularia representatives with capitate hyphae (C. cubensis and C. minor) and those without them (C. angulospora, C. catenulata) provided compelling evidence to consider these species congeneric.

In this study, we present a taxonomic circumscription of Catenularia using molecular and phenotypic data. The generic concept has been emended and species with and without capitate hyphae are accepted in Catenularia. We were unsuccessful in obtaining C. cupulifera into axenic culture from fresh material. The available non-type strain CBS 419.80 of this species is a contaminant (In the Blast search, ITS and 28S sequences derived from this strain showed 100% identity with sequences of various strains of Calycina citrina.). Eleven species are accepted in Catenularia and listed below, four of which have been verified with molecular DNA data. Other species are accepted based on morphological similarity, but have to be confirmed as members of Catenularia by molecular data. So far, the teleomorph has been observed in C. cubensis, C. cupulifera, C. minor and C. novae-zelandiae. Catenularia variegata (Li et al. 2017) is excluded from Catenularia and transferred to a new segregate genus Fuscocatenula in this study. Disposition of Catenularia and morphologically similar taxa previously attributed to the genus is presented in Table 1.

Haplochalara (Linder 1933) and Psiloniella (Costantin 1888) are accepted as generic synonyms of Catenularia. The systematic placement of H. pidoplitschkoi (Litvinov 1967) is unknown. The species was characterised by dematiaceous, erect, simple conidiophores producing ellipsoidal, hyaline conidia that accumulate in slimy droplets and formation of dark chlamydospores in culture. Based on these characteristics, the species shows affinity to Chloridium (Gams and Holubová-Jechová 1976) and would be better placed in this genus.

Colonies on natural substrate effuse, hairy to velutinous, dark brown to almost black. Anamorph. Conidiophores 77–220 × 4.5–6(–7) μm wide, 7–8 μm above the base, macronematous, solitary or arise in tufts, erect, straight or slightly flexuous, unbranched, dark brown, paler towards the apex, septate. Capitate hyphae absent. Conidiogenous cells 18–25 × 3.5–4.5 μm tapering to ca. 2.5 μm, integrated, terminal, monophialidic, extending percurrently, obclavate to subcylindrical or slightly lageniform, pale brown, paler towards the apex; collarettes 3–4 μm wide, 1.5(–2) μm deep, funnel-shaped, subhyaline, smooth, margin entire. Conidia 6–8(–9) μm long, 4.5–6(–7) μm wide at the apical end, ca. 2 μm wide at the basal hilum (mean ± SD = 7.4 ± 1.1 × 6.0 ± 1.2 μm × 2.0 ± 0.0 μm), rounded-obconic in side view, with three blunt corners when viewed from above, broadly rounded to flattened at the apex, truncate at the basal scar, pale brown to pale fuscous, thick-walled, smooth; formed singly, adhered in basipetal chains. Teleomorph. Unknown.

Figure 3. 

Catenularia angulospora (holotype FH 00965375) A vertical section of the wood with colony B–D conidiophores with conidia. Scale bars: 500 μm (A); 50 μm (B–D).

USA – Kentucky • near Louisville; on decaying beech log; 23 Mar. 1928; D.H. Linder (holotype of C. angulospora FH herbarium 00965375, as microscopic slides).

Saprobe on dead culms of Bambusa sp., decaying wood of Fagus sp. and other unknown hosts in freshwater and terrestrial habitats. It is known in China, India and the USA (Linder 1933; Sharma 1980; Luo et al. 2019 as C. cubensis).

For additional description and illustration, see Luo et al. (2019, as C. cubensis). Hughes (1965) revised the type material of H. angulospora, and despite the striking similarities to other Catenularia, he kept the species in Haplochalara due to the absence of capitate hyphae. Sharma (1980) described C. hughesii on dead bamboo culms in India with pale brown to brown conidia 6–8 × 4.5–5.8 μm and conidiophores up to 270 × 5–7 μm. Although the holotype of this species was not available for study, a detailed morphological comparison of its original description and illustration with C. angulospora suggests that they are conspecific. Luo et al. (2019) reported this species as C. cubensis (strain MFLUCC 18-1331) from China, characterised by the absence of capitate hyphae and cuneiform, greyish-brown to brown conidia 6–8 × 4–6 μm.

In the phylogenetic analysis, the strain of C. angulospora MFLUCC 18-1331 clustered as a sister to C. cubensis S.M.H. 3258, but their relationship is not statistically supported. Both species are, however, very similar. Catenularia cubensis (Holubová-Jechová 1982) differs from C. angulospora in brown to dark brown conidia, slightly narrower at the apical end (5.5–8.5 × 3.5–5.5 μm), and presence of capitate hyphae scattered among the conidiophores. The ITS sequence identity between C. cubensis and C. angulospora is 96.5% and supports our conclusion to treat them as separate species.

Saprobe on submerged wood, known only in China (Luo et al. 2019).

Catenularia catenulata is characterised by solitary conidiophores, absence of capitate hyphae and conidia 13–15 × 12–14 μm, greyish-brown, turbinate, triangular in side view with 3–4 corners when viewed from above (Luo et al. 2019). It resembles C. malabrica (Subramanian and Bhat 1989), but the latter species has larger conidia 12–18 × 18–21 μm with 4–5 corners.

Colonies on natural substrate effuse, hairy to velutinous, dark brown, mycelium partly immersed, partly superficial, brown; colonies composed of conidiophores, capitate hyphae and sometimes ascomata. Anamorph. Conidiophores 115–200 × 4–4.5 μm, 4.5–6 μm wide above the base, macronematous, solitary or arise in tufts, erect, straight or flexuous, unbranched, thick-walled, brown to dark brown, slightly paler towards the apex. Capitate hyphae 104–165 × 4–4.5 μm, 4–5.5 μm wide above the base, arise among the conidiophores, extending percurrently, erect, straight, brown to dark brown, paler towards the apex, apical cell sterile, thin-walled, subhyaline, slightly swollen, ca. 3.5 μm wide, broadly rounded, the hyaline gelatinous cap was not observed. Conidiogenous cells 22–38 × 3.5–4.5 μm tapering to 2–2.5 μm below the collarette, terminal, integrated, monophialidic, extending percurrently, cylindrical, pale brown to brown, producing conidia successively; collarettes 3.5–4 μm wide, 1–2 μm deep, shallow, funnel-shaped, pale brown, smooth, margin entire. Conidia 5.5–8.5 μm long, 3.5–5.5 μm wide at the apical end, 1.5–2 μm wide at the basal hilum (mean ± SD = 7.5 ± 0.7 × 4.3 ± 0.4 × 1.8 ± 0.2 μm), rounded-obconic to broadly obovoid in side view, with three bluntly rounded corners when viewed from above, broadly rounded to flattened at the apex, truncate at the basal scar, aseptate, brown to dark brown, thick-walled, smooth; formed singly, adhered in basipetal chains. Teleomorph. Ascomata 150–200 µm diam, 160–210 µm high, superficial, solitary or in groups, subglobose to conical, papillate, dark brown to black, covered with conidiophores and capitate hyphae. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 15–25 μm thick, two-layered. Outer layer consisting of brown, polyhedral cells with opaque walls. Inner layer consisting of several rows of thin-walled, hyaline cells. Paraphyses 2.5–3.5 μm wide, septate, hyaline, longer than the asci. Asci 62–84.5 × (6–)7–8.5 μm (mean ± SD = 72.2 ± 7.8 × 13.9 ± 0.9 µm), cylindrical-clavate, short-stipitate, apically rounded to obtuse, ascal apex with a non-amyloid apical annulus 2–2.5 μm wide, 1.5(–2) μm high. Ascospores 12–16(–17.5) × 3–4 μm (mean ± SD = 13.9 ± 0.9 × 3.5 ± 0.2 µm), fusiform, straight or slightly curved, hyaline, 3-septate, smooth, 2-seriate in the ascus.

Figure 4. 

Catenularia cubensis A ascomata accompanied by conidiophores and capitate hyphae B, J conidia C, D tufts of conidiophores with scattered capitate hyphae E–I conidiophores K–M asci with ascospores. Images: S.M.H. 3258 (A, H–J), PRM 825347 holotype (B–D, F, G, K–M); PRA-19884 (E); on natural substrate (A–G, K–M); in culture (H–J). Scale bars: 200 μm (A); 10 μm (B, E–M); 25 μm (C, D).

Costa Rica • Guanacaste, Liberia ACG, Sector Santa Maria, Estacion Biologica, trail to Bosque Encantado; 10.7647N, -85.3033W; alt. 750 m; on 5 cm diam branch on ground; 26 Jun. 1997; S.M. Huhndorf (S.M.H. 3258). Costa Rica • Alajuela, Cantón Upala, District Bijagua, Heliconias Station, Heliconias trail; 10.7081N, -85.0453W; on 25 cm diam log on ground; alt. 1190 m; 12 Jul. 2001; S.M. Huhndorf, F.A. Fernández, A.N. Miller & M. Darin (S.M.H. 4454). Cuba – Isla de la Juventud (Isla de Pinos) • Sierra de Casas, in a valley near El Abra, 2 km SW of Nueva Gerona; on dead trunk of Palmaceae; 22 Jan. 1981; V. Holubová-Jechová (holotype PRM 825347). Cuba – Isla de la Juventud (Isla de Pinos) • in forest near village Caryo Piedra; on wood of a trunk of a deciduous tree; 21 Jan. 1981; V. Holubová-Jechová (PRA-19884).

Saprobe on decaying wood of palm Euterpe oleracea and other hosts in Brazil, Cuba and Costa Rica (Holubová-Jechová 1982; De Castro et al. 2011; Miller and Huhndorf, unpubl.; this study), and on fallen leaves in India (Dubey and Pandey 2017).

The description is based on Cuban collections. In the Costa Rican material, conidia were 6–8.5 μm long, 3–5 µm wide at the widest point, 1.5–2 µm wide at the basal hilum, brown to dark brown, broadly obovoid or cuneiform, asci 60–80 × 7–9 µm, ascospores 12–20 × 3–5 µm, fusiform, 3-septate (Huhndorf and Miller, unpubl.). For additional details, see Holubová-Jechová (1982).

Catenularia cubensis closely resembles C. angulospora; for comparison see notes for the latter species. Catenularia minor can also be compared with C. cubensis, but differs in longer and wider conidia (6.5–)7.5–10.5(–13) × 6.5–11.5 μm with 3–5 blunt corners and conidiophores that form two distinct layers.

Colonies on natural substrate effuse, hairy or tufted, dark brown to black, mycelium partly immersed, partly superficial, brown; colonies composed of conidiophores, capitate hyphae and sometimes ascomata. Anamorph. Conidiophores 100–350(–530) × 6–7.5(–8) μm, 8.5–10.5 wide above the base, macronematous, solitary or in tufts, with dark brown stromatic hyphal cells around the bases, erect, straight or flexuous, unbranched, brown to dark brown, thick-walled, slightly paler towards the apex. Capitate hyphae 110–160 × 5.5–6 μm, 6.5–7 μm wide above the base, scattered among the conidiophores, erect, straight, brown to dark brown, paler towards the apex, apical cell sterile, thin-walled, subhyaline, slightly swollen, ca. 7 μm wide, broadly rounded with a hyaline gelatinous cap that disappears with age. Conidiogenous cells 40–59 × 5.5–6.5 μm, not tapering, terminal, integrated, monophialidic, extending percurrently, cylindrical, brown, producing conidia successively; collarettes 9.5–12.5 μm wide and 10–12.5 μm deep, funnel-shaped, brown, slightly roughened, with an irregularly frayed margin. Conidia (9–)10.5–13.5 μm long, 7–9.5 μm wide at the apical end, 3.5–4.5 μm wide at the basal hilum (mean ± SD = 11.8 ± 0.7 × 8.0 ± 0.6 μm × 4.0 ± 0.3 μm), cuneiform in side view, with (3–)4(–5) blunt corners when viewed from above, flattened to broadly rounded at the apex, truncate at the base, aseptate, fulvous, brown to dark brown, thick-walled, smooth; formed singly, adhered in basipetal chains.

Figure 5. 

Catenularia cupulifera A, B ascomata accompanied by conidiophores and capitate hyphae C colony composed of conidiophores and capitate hyphae D–F conidiophores G capitate hypha H–J upper parts of conidiophores with conidia K, L conidia M, N asci with ascospores. Images: W7972 (A, B, M); W7973 (C, D, H, I); PRA-19893 (E–G, J–L); JF 99018 (N); on natural substrate (A–N). Scale bars: 500 μm (A–C); 50 μm (D); 25 (E); 20 μm (F–L); 10 μm (M, N).

Ascomata 150–220 μm diam, 200–250 μm high, superficial with a base immersed, solitary or in groups or densely aggregated forming a crust, conical to subglobose, papillate, dark brown to black, rugose, sometimes covered with conidiophores and capitate hyphae or in a dense subiculum consisting of partly decumbent conidiophores. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 22–33 μm thick, two-layered. Outer layer consisting of brown, polyhedral to angular cells with opaque walls. Inner layer consisting of several rows of thin-walled, hyaline cells. Paraphyses 3–4 μm wide tapering to 2–2.5 μm, septate, hyaline, longer than the asci. Asci 110–140 × (8–)10–11(–12.5) μm (mean ± SD = 162.2 ± 11.1 × 10.5 ± 1.2 µm), cylindrical-clavate, short-stipitate, apically narrowly rounded to obtuse, ascal apex with a non-amyloid apical annulus 2–2.5(–3) μm wide, ca. 1.5 μm high. Ascospores 21–28.5 × 4.5–5.5 μm (mean ± SD = 25.3 ± 1.7 × 5.5 ± 0.4 µm), fusiform, straight or slightly curved, hyaline, 1–4-septate, smooth, 2-seriate in the ascus.

Belgium • West Flanders province, Adinkerke, Cabour; on decaying wood of Populus sp.; 21 Oct. 2007; B. Declerque (IFBL D0.16.23). Czech Republic – Moravia • Lanžhot, Ranšpurk National nature reserve; alt. 150 m; on decaying wood of Carpinus betulus; 14. Aug. 1979; V. Holubová-Jechová (PRA-19887) • Ibid.; on decaying wood of Populus alba, 28 Jul. 1970, V. Holubová-Jechová (PRA-19888) • Ibid.; on decaying wood Quercus robur, 28. Aug. 1976, V. Holubová-Jechová (PRA-19889). Czech Republic – Moravia • Bílé Karpaty, Velká Javořina Mt. near Kamenná Bouda; alt. 660 m; on decaying wood of a branch of Fagus sylvatica; 27 Jul. 1970; V. Holubová-Jechová (PRA-19886). France – Ariège • Pyreneés Mts., Rimont, Las Muros, alt. 480 m; on decaying wood of Fraxinus excelsior; 4 Feb. 1999; J. Fournier J.F. 99018 (PRA-19890). France – Ariège • Pyreneés Mts., Rimont, Las Muros, valley of the Peyrau brook, alt. 400 m; on decaying wood of Buxus sempervivens; 9 Nov. 1999, J. Fournier J.F. 99261 (PRA-19892) • Ibid.; on decaying wood of Salix caprea; 12 Mar. 2000; J. Fournier J.F. 00026 (PRA-19891). France – Ariège • Pyrénées Mts., Rimont, La Maille brook; alt. 550 m; on submerged wood; 28 May 2018; J. Fournier M.R. 4104 (PRA-19893). Slovak Republic • Brezová near Senica; on decaying wood of a trunk of Salix alba; 6 Aug. 1976; V. Holubová-Jechová (PRA-19885). United Kingdom – Somerset • Langridge, on decaying wood of roots of Ulmus sp.; Apr. 1869; C.E. Broome (holotype of S. cupulifera K(M) 57177). United Kingdom • on decaying wood; 14 Apr. 1873; ex Herbarium C.E. Broome 1886 (W 7972) • Ibid.; ex Herbarium C.E. Broome 1886, no. 366 (W 7973).

Saprobe on decaying wood of Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Hedera sp., Ilex sp., Quercus sp., Salix alba, Ulmus sp. and other unknown hosts. Most of the records originate from Europe in Belgium, Czech Republic, France, Slovak Republic and the United Kingdom (Berkeley and Broome 1871; Hughes 1965; Holubová-Jechová 1973; this study). Hughes (1965) suggested that C. cupulifera is apparently only known from Europe. However, findings of this species also come from other continents. Catenularia cupulifera has been reported from foam in a river in Venezuela (Fernández and Smits 2018), wood of Ulmus americana in the USA, Illinois (Shim 1969) and decaying leaves of Pandanus sp. in Mauritius (Whitton et al. 2012).

Our observations of the teleomorph-anamorph connection between Ch. cupulifera and C. cuneiformis agree with those of Berkeley and Broome (1871), De Seynes (1886) and Booth (1958). Although this relationship has not yet been verified experimentally, both morphs occur together in nature. Since the anamorph and teleomorph represent two different stages of the life cycle of one organism, we propose a new combination in Catenularia based on Sphaeria cupulifera with C. cuneiformis and C. simplex as synonyms.

Catenularia novae-zelandiae resembles C. cupulifera but differs in larger and rounded-obconic conidia, 11.5–17.5 μm long, 14.5–18.5 μm wide. Both species have conspicuous collarettes with a frayed margin, which is larger in C. novae-zelandiae, 19–27 μm wide and 12.2–19 μm deep, funnel- to cup-shaped.

On fossil wood, known only in the United Kingdom.

Catenularia elsikii was isolated from the material containing clay, charcoal and wood fragments present in the cracks of a large sample of fossil wood discovered in the United Kingdom (Pound et al. 2019). Thick-walled, dark brown conidia were the only structure that has been preserved in material dated to the Miocene. In the conidial characteristics, C. elsikii is remarkably similar to C. macrospora known from Canada and New Zealand and C. novae-zelandiae known only from New Zealand. These species share dark brown, rounded-obconic conidia with (3–)4–5 corners when viewed from above. In addition, C. elsikii and C. novae-zelandiae have a visible pore at the basal hilum. Conidia of C. elsikii (23.1–24.4 μm high, 20.8–23.9 μm wide with a basal scar 3–4 μm wide) are longer and wider than those of C. novae-zelandiae, but shorter than those of C. macrospora. For detailed comparison, see notes to the two latter species.

Saprobe on decaying wood, known only in China, India and Mexico (Subramanian and Bhat 1989; Heredia et al. 2004; Xia et al. 2013).

For descriptions and illustrations, refer to Subramanian and Bhat (1989) and Xia et al. (2013). Catenularia kalakadensis is unique among other species in conidia with six blunt corners when viewed from above. It resembles C. cubensis but differs in the absence of capitate hyphae and wider conidia (6–7 μm) with more corners at the apex (Subramanian and Bhat 1989).

Saprobe on decaying wood, known only in New Zealand (Hughes 1965).

Catenularia longispora is well recognisable by narrowly rounded-obconic, brown to dark brown conidia that are the longest in the genus, 27–45 μm long, 16.8–24 μm wide at the apical end, 7–10 μm wide at the basal hilum, with usually three blunt corners when viewed from above (Hughes 1965).

Saprobe on decaying bark and wood of Dacrydium cupressinum, Fuscospora cliffortioides, Vitex lucens and other unknown hosts, known in Canada and New Zealand (Hughes 1965).

Catenularia macrospora has broadly obovoid to rounded-obconic, brown to dark brown conidia, 21–28 μm long, 19–28 μm wide at the apical end and 4–7 μm wide at the basal hilum, with (3–)4(–5) blunt corners when seen from above (Hughes 1965). The conidial length is comparable with those of C. longispora and C. elsikii, but the former species differs in conidia narrowly rounded-obconic, narrower at the apical end (16.8–24 μm) with only (2–)3 corners. Although the length of conidia of C. elsikii and C. macrospora overlap and the number of corners is comparable, conidia of C. elsikii are slightly shorter and narrower in their upper range (23–24.5 × 21–24 μm) and narrower at the truncate base (3–4 μm) (Pound et al. 2019).

Saprobe on decaying wood of Magnolia liliifera and an unknown host, known only in India and Thailand (Subramanian and Bhat 1989; Kodsueb et al. 2008).

For descriptions and illustrations, see Subramanian and Bhat (1989). Catenularia malabrica produces one of the tallest conidiophores in the genus, 320–620 × 6–11 μm arising singly or in tufts. It resembles C. novae-zelandiae in dark brown conidia with 4–5 corners, but conidia of C. malabrica are wider (18–21 μm) and the capitate hyphae are absent.

Colonies on the natural substrate effuse, tufted or hairy, dark brown to black, mycelium partly immersed, partly semi-immersed, pale brown to brown; colonies composed of conidiophores, capitate hyphae and sometimes ascomata. Anamorph. Conidiophores macronematous, solitary or arise in tufts, with dark brown stromatic hyphal cells around the base, erect, straight or flexuous, unbranched, thick-walled, paler towards the apex, forming two layers. Conidiophores of the lower layer 95–212 × 3.5–4.5(–5) μm, 4.5–5.5(–8.5) μm wide above the base, pale brown to brown; conidiophores of the upper layer 260–527 × 4.5–7 μm long, 7.5–10 μm wide above the base, dark brown. Capitate hyphae 122–186 × 3.5–5 μm, 5–5.5 μm wide above the base, scattered among the conidiophores, erect, straight, brown, extending percurrently, paler towards the apex, apical cell subhyaline, slightly swollen, 3.5–4 μm wide, broadly rounded, thin-walled; the hyaline gelatinous cap was not observed. Conidiogenous cells 15–40 × 3.5–5.5 μm tapering to 2.5–3 μm below the collarette, integrated, terminal, monophialidic, extending percurrently, cylindrical to slightly lageniform, pale brown to brown, producing conidia successively; collarettes 3.5–5(–6) μm wide, 1.5–2.5 μm deep, shallow, funnel-shaped, pale brown to subhyaline, smooth, margin entire. Conidia (6.5–)7.5–10.5(–13) μm long, 6.5–11.5 μ wide at the apical end, 1.5–2.5 μm wide at the base (mean ± SD = 8.9 ± 0.9 × 9.0 ± 1.2 × 2.1 ± 0.2 μm), cuneiform to rounded-obconic to obtriangular in side view, with 3–5 blunt corners when viewed from above, broadly rounded to flattened at the apex, truncate at the basal scar with a central pore, aseptate, pale brown to dark brown, thick-walled, smooth; formed singly, adhered in basipetal chains or clusters. Teleomorph. Ascomata 230–250 μm diam, 250–275 μm high, superficial, solitary or densely aggregated, subglobose to globose, covered by a whitish-grey powder except for the black glabrous papilla; the powdery covering is ca. 5–15 μm thick, disappearing with age, leaving the perithecia dark and glabrous. Ascomata sparsely covered with conidiophores. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 30–37.5 μm thick, two-layered. Outer layer consisting of dark brown, opaque, thin-walled, polyhedral cells. Inner layer consisting of hyaline, thinner-walled, elongated, compressed cells. Paraphyses 3–4 μm wide, tapering to ca. 2 μm, branching, anastomosing, septate, hyaline, longer than asci. Asci 102–112 × 8–9(–9.5) μm (mean ± SD = 106 ± 1.6 × 8.9 ± 0.2 µm), cylindrical-clavate, short-stipitate, rounded apically, ascal apex with a non-amyloid apical annulus 3 μm diam, 1.5–2 μm high. Ascospores 25–29(–30) × (3.5–)4–4.5 μm (mean ± SD = 27 ± 0.5 × 4 ± 0.7 µm), fusiform, straight or curved, hyaline, 1–3-septate, smooth, 1–2-seriate in the ascus (adapted from Réblová and Seifert 2003).

Figure 6. 

Catenularia minor A, B colonies composed of ascomata, conidiophores and capitate hyphae C ascus with ascospores D–F conidiophores with capitate hyphae G–J upper parts of conidiophores with conidia in chains J capitate hypha K, L conidia (arrow indicates central pore in the basal scar). Images: PRM 828704 holotype of C. minor (D, E, G, K); PRM 900544 holotype of Ch. trianguloconidia (A–C, F, H–J, L); on natural substrate (A–L). Scale bars: 250 μm (A, B); 10 μm (C, G–L); 50 μm (D–F).

Cuba – Sancti Spiritus province • Soledad, Cienfuegos Province Botanical Garden; on decaying stem of Bambusa vulgaris; 19 Mar. 1981; M.A. Bondarceva & S. Herrera (holotype of C. cuneiformis var. minor PRM 828704). Thailand – Nakhon Nayok Province • Khao Yai National Park, trail to Haew Suwat waterfall, elev. 720 m; on decaying bamboo culm; 2 Sep. 2001; M. Réblová, Gary J. Samuels & R. Nasit M.R. 2186/TH 438 (holotype of Ch. trianguloconidia PRM 900544).

Saprobe on dead culms of bamboo, known in Cuba and Thailand (Holubová-Jechová 1983; Réblová and Seifert 2003).

For characteristics in culture, see Réblová and Seifert (2003). The apparent similarity of C. cuneiformis var. minor (Holubová-Jechová 1983) and Ch. trianguloconidia (Réblová and Seifert 2003) and their habitat on dead bamboo culms prompted a revision of both species. Examination of their holotypes revealed that they are conspecific. Additionally, we discovered capitate hyphae in the type material of both species, although they were not described in the protologues. They are scattered among the conidiophores and easy to overlook. The hyaline gelatinous cap around the swollen apex of the capitate hyphae was not observed. Conidia slightly vary in size and colour, and often smaller and pale brown conidia occur together with slightly larger and darker brown conidia.

Holubová-Jechová (1983) distinguished var. minor from var. cuneiformis (= C. cupulifera, this study) in shorter collarettes, smaller conidia and the absence of capitate hyphae. Based on their different morphology, a new combination for var. minor is proposed at the species level with Ch. trianguloconidia reduced to synonymy.

Catenularia angulospora is similar to C. minor, and it is challenging to distinguish both species, especially if capitate hyphae may rarely occur in some specimens of the latter species. Catenularia angulospora differs in fuscous to brown conidia that are narrower (4.5–6(–7)) μm and the lack of capitate hyphae. Catenularia cupulifera is comparable to C. minor but differs in larger collarettes (9.5–12.5 μm wide and 10–12.5 μm deep) with a frayed margin, and longer (10.5–13.5 μm) conidia that are wider (3.5–4.5 μm) at the basal hilum. Conidia of C. cupulifera are cuneiform in side view, whereas conidia of C. minor are more rounded-obconic to obtriangular.

Colonies on natural substrate effuse, tufted or velutinous, dark brown, mycelium partly immersed, partly superficial, brown; colonies composed of conidiophores, capitate hyphae and sometimes ascomata. Anamorph. Conidiophores 90–354 × 7.5–9.5 μm, 7–10.5 μm wide near the swollen base, macronematous, solitary or arise in tufts, with dark stromatic hyphal cells around the base, erect, straight or flexuous, unbranched, brown to dark brown, thick-walled. Capitate hyphae 95–215 × 5–7 μm, 6.5–9 μm wide above the base, 4.5–5.5 μm wide at the apex, solitary or in tufts, arise among the conidiophores, erect, straight to slightly flexuous, dark brown, paler towards the apex, apical cell pale brown to subhyaline, slightly swollen, broadly rounded, thin-walled, with a hyaline, mucilaginous cap that disintegrates with age. Conidiogenous cells 22.5–41(–65) × 7–11 μm, 7.5–9.5 μm wide below the collarette, terminal, integrated, monophialidic, extending percurrently, cylindrical, subcylindrical or slightly lageniform, brown, producing conidia successively; collarettes 19–27 μm wide and 12.2–19 μm deep, funnel-shaped or cup-shaped, brown to dark brown, roughened, with a frayed margin, the margin deteriorates, and the collarette becomes reduced in size 11.5–15.8 μm wide and 4.5–6 μm deep. Conidia 11.5–17.5 μm long, 14.5–18.5 μm wide at the apical end, 4–5.5 μm wide at the basal hilum, (mean ± SD = 15.8 ± 1.8 × 15.9 ± 1.3 × 5.5 ± 0.9 μm), cuneiform to rounded-obconic in side view, with 4–5 blunt corners when viewed from above, flattened to broadly rounded at the apex, truncate at the base, aseptate, brown to dark brown, thick-walled, smooth; formed singly, adhered in basipetal chains. Teleomorph. Ascomata 160–210 μm diam, 180–220 μm high, superficial, solitary or in small groups, subglobose to globose, papillate, dark brown, sometimes covered with capitate hyphae and conidiophores; capitate hyphae 80–130 × 5–5.5 μm, erect, simple, apical cell 6–6.5 μm wide, slightly inflated, broadly rounded apically, subhyaline, with a mucilaginous cap that disappears with age. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 17–22 μm thick, two-layered. Outer layer consisting of dark brown, polyhedral to angular cells with opaque walls. Inner layer consisting of rows of thin-walled, hyaline cells. Paraphyses 4–5 μm wide tapering to 1.5–2 μm, septate, hyaline, longer than the asci. Asci 102–130 × 11–13 μm (mean ± SD = 117.6 ± 9.8 × 12.3 ± 0.8 µm), 74–100(–110) μm in the sporiferous part (mean ± SD = 83.7 ± 12 µm), cylindrical-clavate, narrowly truncate apically, ascal apex with a non-amyloid apical annulus 3.5–4 μm wide, ca. 2 μm high. Ascospores 22–28(–30) × 4–5 μm (mean ± SD = 25.6 ± 1.6 × 4.7 ± 0.4 µm), fusiform, straight or slightly curved, hyaline, 3-septate, smooth, 2-seriate in the ascus.

Figure 7. 

Catenularia novae-zelandiae A ascomata B colony composed of conidiophores and capitate hyphae C, D, O, P conidiophores E capitate hypha arising among conidiophores F–I upper parts of conidiophores with conidia (arrow indicates central pore at the basal scar) J, K conidia in chain (arrow indicates appendages) L, M asci with ascospores and paraphyses N capitate hypha arising from the ascomal wall Q conidia. Images: PDD 81883 (A–C, F–I, L–Q); PDD 119362 (D, E, J, K); on natural substrate (A–N); in PCA culture after 2 wk (O–Q). Scale bars: 250 μm (A, B); 50 μm (C–E); 20 μm (F–Q).

On PCA: colonies 8–12 mm in 14d, circular, flat, margin entire, subsurface, aerial mycelium scarce, cobwebby to mucoid, beige-brown, reverse of the same colour. Sporulation was abundant, sporulating conidiophores developed from aerial mycelium and occasionally from immersed vegetative hyphae.

Colonies on PCA effuse, hairy, vegetative mycelium subhyaline to hyaline, 2–3 μm wide. Conidiophores, conidiogenous cells and conidia similar to those from nature. Conidiophores 31–120 × 6–7 μm, solitary or arise in tufts of 2–7, erect, straight, pale brown, 1–several-septate. Capitate hyphae absent. Conidiogenous cells 22–37 × 8–10 μm, tapering to ca. 7 μm below the collarette; collarettes 12.5–15 μm wide, 4–6(–8) μm deep, funnel-shaped, pale brown to dark brown, slightly roughened with a frayed to entire margin. Conidia (13–)14–18 μm long, 13–18 μm wide at the apical end, 4.5–6 μm wide basal hilum (mean ± SD = 15.2 ± 1.2 ×14.7 ± 1.4 µm × 5.5 ± 0.9 µm), broadly rounded-obconic in side view, aseptate, brown to grey-brown, thick-walled, smooth, formed singly, adhered in short basipetal chains.

New Zealand – Auckland region • Auckland district, Upper Piha Valley, Waitākere Ranges, Home track; on decaying wood of Metrosideros robusta; 9 Oct. 1963; J.M. Dingley (holotype PDD 21603, isotype DAOM 93575). New Zealand – West Coast region • Westland district, Otira, Kelly Shelter, Cockayane Nature Walk; on decaying wood; 16 Mar. 2003; M. Réblová MR 2846/NZ 362 (PDD 81883). New Zealand – West Coast region • Buller district, Victoria Forest Park, Reefton, Big River Inanganua track ca. 14 km; on decaying wood of Nothofagus sp.; 6 Mar. 2003; M. Réblová MR 2723/ NZ 224A (PDD 119362).

Saprobe on decaying wood of Coprosma lucida, Coprosma spp., Freycinetia banksii, Griselinia lucida, Leptospermum ericoides, Metrosideros robusta, Neopanax arboreum, Nothofagus sp., Olearia rani, Weinmannia racemosa and other unknown hosts, known only in New Zealand (Hughes 1965; this study).

The specimen PDD 81883 of C. novae-zelandiae was isolated in axenic culture (Fig. 7O–Q). In vitro, conidia were paler than those from nature and broadly rounded-obconic. Unfortunately, the isolate is no longer viable. The other collection PDD 119362 has conidia slightly larger 17.5–21 × 18–19 μm, 5–6 μm wide at the truncate base. In both specimens, we observed several conidia with minute hyaline appendages arising from the pale, circular, thin-walled areas in the cell wall (Fig. 7K).

Catenularia malabarica (Subramanian and Bhat 1989) is similar to C. novae-zelandiae in characters of conidia, but differs in the absence of capitate hyphae, longer conidiophores up to 620 μm long and conidiogenous cells with a shallow, funnel-shaped collarette without a frayed margin.

Colonies on natural substrate effuse, hairy, mycelium partly superficial, partly immersed; colonies composed of conidiophores and sometimes ascomata. Anamorph. Setae present, mostly associated with ascomata, simple, brown, apically rounded. Conidiophores mononematous, macronematous, solitary, erect, septate, unbranched, brown. Conidiogenous cells integrated, terminal, monophialidic, extending percurrently, cylindrical-lageniform to urceolate, brown; collarettes funnel-shaped, pale brown. Conidia obpyramidal, in side view cuneiform, obovoid to obtriangular, with angular outline when viewed from above, truncate at the basal scar, with a simple setula inserted apically at each corner, aseptate, hyaline, adhered in basipetal chains. Teleomorph. Ascomata non-stromatic, perithecial, papillate, dark brown, sparsely covered by setae and conidiophores. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, two-layered. Paraphyses persistent, septate, hyaline, longer than the asci. Asci unitunicate, 8-spored, cylindrical-clavate, ascal apex with a non-amyloid apical annulus. Ascospores fusiform, hyaline, transversely septate.

Saprobes on dead leaves of Freycinetia spp. (Pandanaceae) and decaying wood, known only in Australasia in New Caledonia and New Zealand (McKenzie 1991; this study).

The genus Chalarodes, typified with Cha. bisetis, was erected for dematiaceous hyphomycetes observed on leaf litter of Freycinetia spp. in New Zealand and New Caledonia (McKenzie 1991). It is characterised by mononematous, simple, dark brown conidiophores with terminal monophialidic conidiogenous cells extending percurrently and hyaline, aseptate, cuneiform, obconical to obtriangular conidia with setulae, adhered in short basipetal chains. In the protologue (McKenzie 1991), the conidia were described only in the side view with two simple setulae at the apical end. Based on the examination of newly collected material, the conidia have angular outline when viewed from above; they have (3–)4 corners with a setula inserted in each corner. Additionally, we observed sterile setae growing among the conidiophores or on the ascomatal wall. They resemble capitate hyphae of Catenularia, but the mucilaginous sheath around the apex was lacking.

To date, two species, Cha. bisetis and Cha. obconica, have been placed in Chalarodes (McKenzie 1991). A new species, Cha. obpyramidata, inhabiting decaying wood and originating from New Zealand is introduced below. The teleomorph-anamorph connection of Chalarodes is described for the first time. Based on the results of the phylogenetic study, Cha. obpyramidata is closely related to Catenularia.

Pyramidatus (L), pyramidal, prefix ob- (L), meaning reversely, inversely, referring to the conidial shape.

New Zealand – West Coast region • Westland district, Ross, Totara forest, Totara River valley; on decaying wood of a branch; 7 Mar. 2003; M. Réblová MR 2734/NZ 236 (holotype PDD 119363).

Colonies on natural substrate effuse, hairy, dark brown to black, mycelium partly superficial, partly immersed, brown; colonies composed of conidiophores and sometimes ascomata. Anamorph. Setae present, mostly associated with ascomata (see below). Conidiophores 195–360 × 5–7.5 μm, 7–8.5 μm wide above the base, mononematous, macronematous, solitary, erect, straight or flexuous, unbranched, thick-walled, dark brown, paler towards the apex. Conidiogenous cells 20–54 × 5–6.5(–8) μm tapering to 3.5–4.5 μm below the collarette, integrated, terminal, monophialidic, extending percurrently, cylindrical to cylindrical-lageniform, brown, producing conidia successively; collarettes 6–7.5 μm wide, 2.5–3(–4) μm deep, funnel-shaped, pale brown. Conidia 10.5–12 μm long, 8.5–12 μm wide, 2.5–3.5 μm wide at the basal hilum (mean ± SD = 11.2 ± 0.5 × 10.3 ± 1.0 × 2.9 ± 0.3 μm), obpyramidal, in side view cuneiform to obtriangular, with four corners when viewed from above, truncate at the basal scar, with straight or curved setulae inserted at each corner 5–8 μm long, aseptate, hyaline, thin-walled, smooth; formed singly, adhered in basipetal chains. Teleomorph. Ascomata 120–140 μm diam, 130–160 μm high, subglobose, dark brown to black, superficial, solitary or aggregated, subglobose, papillate, setose. Setae 37–157 × 3.5–5.5 μm, simple, straight, cylindrical, brown, pale brown towards the apex, extending percurrently, apical cell sterile, 3.5–4 μm wide, broadly rounded, pale brown to subhyaline, similar setae arise around ascomata on the substrate. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 20–24 μm thick, two-layered. Outer layer consisting of brown, polyhedral cells with opaque walls. Inner layer consisting of several rows of thin-walled, hyaline cells. Paraphyses 4–5 μm wide, tapering to ca. 2 μm, septate, hyaline, longer than the asci. Asci 95–114 × (9–)10–12 μm (mean ± SD = 103.5 ± 6.5 × 10.9 ± 1.1 µm), cylindrical-clavate, short-stipitate, apically narrowly rounded, ascal apex with a non-amyloid apical annulus ca. 3 μm wide, 2 μm high. Ascospores 18–22(–23) × 4–5 μm (mean ± SD = 20.4 ± 1.3 × 4.4 ± 0.4 µm), fusiform, hyaline, 1–3-septate, smooth, 2-seriate in the ascus.

Figure 8. 

Chalarodes obpyramidata A ascomata B colony composed of conidiophores C, H, O–Q conidiophores D–G, S, T conidia I–L, R upper parts of conidiophores with conidia M asci with ascospores N paraphyses. Images: PDD 119363 (A–L); PDD 119364 (M–T); on natural substrate (A–N); in PCA culture after 4 wk (O–T). Scale bars: 250 μm (A, B); 50 μm (C, H); 10 μm (D–G, M, N, R–T); 20 μm (I–L, O–Q).

On PCA: colonies 7–10 mm diam in 14d, circular, raised, margin entire, velvety-lanose, brown to dark grey-brown with whitish-grey conidial masses, reverse black. Sporulation abundant at the centre of the colony.

Colonies on PCA effuse, mycelium subhyaline to pale brown, 2–3 μm wide. Setae absent. Conidiophores, conidiogenous cells and conidia similar to those from nature. Conidiophores 74–141 × (4.5–)5–6 μm, 5.5–6.5 μm wide above the base, paler brown and less septate than those from nature, erect, straight. Conidiogenous cells 18–40 × 4.5–5.5 μm tapering to 3.5–4 μm below the collarette, cylindrical, pale brown; collarettes 5–6 μm wide, 3.5–4 μm deep, pale brown. Conidia 8–10(–11) μm long, 8–9(–10) μm wide, 2–2.5 μm wide at the hyaline basal hilum (mean ± SD = 9.7 ± 0.9 × 8.4 ± 0.6 × 2.0 ± 0.1 μm), cuneiform to obpyramidal, truncate at the basal scar, setulae not observed, aseptate, hyaline, thin-walled, smooth, formed basipetally in chains.

New Zealand – West Coast region • Buller district, Victoria Forest Park, Reefton, Big River Inanganua track; on decaying wood of Nothofagus sp. (associated with C. novae-zelandiae PDD 119362 and Zanclospora falcata PDD 119365); 6 Mar. 2003, M. Réblová MR 2724/ NZ 225 (PDD 119364).

Saprobe on decaying wood, known only in New Zealand.

In the size of conidia, our species appears intermediate between Cha. bisetis and Cha. obconica (McKenzie 1991). Chalarodes bisetis differs from Cha. obpyramidata in conidia longer and narrower at the apical end, (9.5–)12–14(–15) × 4.5–6(–9) μm, while Ch. obconica possesses conidia slightly shorter (8–)9–10.5(–11) μm and narrower at the basal hilum 1.75–2 μm.

Fuscus (L) dark, brown, dusky, catenula (L), a little chain, referring to pigmented conidia in chains.

Fuscocatenula submersa (Z.L. Luo, K.D. Hyde & H.Y. Su) Réblová & A.N. Mill.

Colonies effuse, hairy, brown, mycelium partly immersed, partly superficial. Anamorph. Conidiophores macronematous, mononematous, solitary, erect, unbranched, brown to dark brown, thick-walled, paler and thinner-walled towards the apex. Conidiogenous cells integrated, terminal, monophialidic, extending percurrently, cylindrical to lageniform, brown; collarettes funnel-shaped, brown. Conidia cuneiform to obovoid, broadly rounded apically, truncate at the base, aseptate, hyaline when young, pale brown at maturity, with protracted maturation, smooth, formed in a basipetal chain. Teleomorph. Unknown. (Description partly adapted from Li et al. 2017; Luo et al. 2019).

Members of the genus are saprobes on decaying plant matter in terrestrial and freshwater environments, known only in Asia in China.

Fuscocatenula is proposed as a segregate genus for fungi distantly related from Catenularia (Fig. 2), although morphologically similar. Conidia of Fuscocatenula are obovoid with a truncate base, lack an angular outline and small, circular, thin-walled pale areas in corners that are present in Catenularia. Conidia have a protracted maturation; at first they are hyaline and only later become pale brown, while still attached in a chain. Sometimes the chain consists of hyaline conidia with only one or a few mature pigmented conidia (Li et al. 2017: fig. 1; Luo et al. 2019: fig. 52). In Catenularia, conidia are also hyaline when young but mature soon and when released from the conidiogenous locus they are usually pigmented. Since Catenularia also includes species lacking capitate hyphae, this character alone is not reliable in the distinction of Fuscocatenula from Catenularia.

Two species are accepted in the genus. Li et al. (2017) introduced Catenularia variegata for a foliicolous species from China and Luo et al. (2019) described Chaetosphaeria submersa for a dematiaceous hyphomycete from submerged wood in Thailand. Both species are similar and reminiscent of Catenularia. In the phylogenetic analysis based on ITS-28S sequences, relationship of Ch. submersa and Catenularia was not supported. Molecular data of C. variegata are not available. Based on a detailed comparison of original descriptions and illustrations of both species we conclude that C. variegata is congeneric with Ch. submersa. Therefore, C. variegata is excluded from Catenularia and both species are transferred to the new genus Fuscocatenula.

Saprobe on submerged decaying wood in stream, known only in China (Luo et al. 2019).

The species is characterised by conidiophores 380–596(–691) μm × 15–21 μm and cuneiform, pale brown conidia 21–27 × 12–14 μm. The size of these structures clearly distinguishes F. submersa from the small-spored F. variegata with shorter conidiophores (Luo et al. 2019).

Saprobe on dead stems of an unidentified broadleaf tree, known only in China (Li et al. 2017).

Fuscocatenula variegata resembles F. submersa but differs in shorter conidia 8.5–11 × 5.5–7.5 μm and shorter conidiophores 150–270 × 4.5–8 μm (Li et al. 2017).

Antennatus (L) meaning ‘having antenna(s)’, referring to the presence of conidial appendages resembling insect antennas.

Thailand – Nakhon Nayok Province • Khao Yai National park, Phakrajai trail, on decaying wood and bark of a twig; 17 Aug. 2001; M. Réblová & N. Hywel-Jones M.R. 2056/TH 219 (PRA-20374).

Colonies on natural substrate effuse, hairy, dark brown, mycelium partly superficial, partly immersed, brown. Anamorph. Conidiophores forming two distinct layers; conidiophores of the upper layer 142–282 μm long, conidiophores of the lower layer 44–90 μm long, 5–6 μm wide, 6–8.5 wide above the base, basal cell bulbose with dark brown, thick-walled stromatic cells around the base, mononematous, macronematous, solitary or fasciculate in a group of 2–6, erect, straight or flexuous, unbranched, thick-walled, dark brown, paler towards the apex. Conidiogenous cells 19.5–29 × 5.5–7.5(–8) μm tapering to 3–5 μm below the collarette, integrated, terminal, monophialidic, extending percurrently, subcylindrical to lageniform, pale brown; collarettes 5.5–6.5 μm wide, 1.5–2.5 μm deep, funnel-shaped, pale brown. Conidia 14–17(–18) μm long, 11–14.5(–15.5) μm wide, 2.5–3.5 μm wide at the basal hilum (mean ± SD = 15.5 ± 1.2 × 12.9 ± 1.7 × 2.9 ± 0.3 μm), turbinate to obpyramidal, in side view cuneiform to obtriangular, truncate at the basal scar, flattened to slightly concave at the apical end, with (3–)4 corners when viewed from above, aseptate, hyaline, thin-walled, smooth, with simple setulae inserted at each corner, 17–43 μm long, 7.5–20 μm long when the ends are coiled, conidia accumulate in slimy droplets. Teleomorph. Not observed.

Figure 9. 

Nawawia antennata (PRA-20374 holotype) A, B colony C–G conidiophores H stromatic cells I–P conidia. Images: on natural substrate (A–P). Scale bars: 250 μm (A, B); 20 μm (C, F, G); 25 μm (D, E); 10 μm (H–N).

Saprobe on decaying wood, known only in Thailand.

We were unsuccessful in obtaining N. antennata in axenic culture. The species exhibits diagnostic characteristics of Nawawia such as pigmented, mononematous conidiophores with stromatic cells around the base, terminal monophialides extending percurrently and hyaline, aseptate, obtriangular conidia with an angular outline and several simple setulae at the apex. Conidia accumulate in a slimy head. Conidiophores forming two distinct layers were also documented in N. quadrisetulata (Goh et al. 2014: figs 2, 3).

Among Nawawia species, N. antennata is well distinguished by coiled appendages and the size of conidia. Nawawia quadrisetulata is similar to the new species in conidia with mostly four angles at the apex but differs in larger conidia (30–37.5 × 22.5–32.5 μm) with longer setulae (30–57.5 μm). Nawawia antennata resembles N. filiformis (Marvanová 1980) but the latter species has conidia wider at the apex (14–18 μm) and straight appendages.