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The Atlantic Forest has the greatest diversity of plants per hectare, and it is considered the main center of bamboo biodiversity in the world. Studies on bambusicolous fungi are scarce and concentrated in temperate regions. Cladosporium is one of the largest genera of dematiaceous hyphomycetes fungus and their adaptation to different environments allows them to have a broad lifestyle. Currently, this genus has been taxonomically and phylogenetically elucidating with the help of extensive polyphasic analyzes. This study aimed to characterize 17 Cladosporium isolates associated with decayed leaves of three bamboo species at a fragment of the Brazilian Atlantic Forest, namely “Parque Estadual da Serra do Brigadeiro”, Minas Gerais state, Brazil. Phylogenetic studies revealed that all isolates belong to the C. cladosporioides species complex. Six new Cladosporium species were found and named herein as C. aulonemiae, C. bambusicola, C. benschii, C. brigadeirensis, C. chusqueae and C. pseudotenuissimum. This study contributes with several taxonomic novelties for knowledge about the diversity of the C. cladosporioides species complex.

Cladosporium bambusicola P. P. Costa, A.W.C. Rosado & O.L. Pereira sp. nov. (Figure 3) MycoBank: MB824725 Systematic position: Fungi, Ascomycota, Pezizomycotina, Dothideomycetes, Dothideomycetidae, Cladosporiales, Cladosporiaceae Etymology:—Name refers to the host from which the species was isolated, bamboo. Colonies on PDA attaining 52 mm diam, dull green, greenish glaucous aerial mycelium, woolly, radially furrowed; sector formation, greenish glaucous, greenish-yellow at the margin, smoke grey aerial mycelium; reverse greenishgrey. Colonies on MEA attaining 50 mm diam, dark green to olivaceous grey, fluffy, woolly; reverse olivaceous black. Colonies on OA attaining 65 mm diam, smoke grey; pale olivaceous grey aerial mycelium, flat, wooly; reverse olivaceous grey. Colonies on SNA attaining 47 mm diam, flesh a rosy vinaceous, hazel, flat, sparse; reverse similar to surface. Mycelium scarce, 1.35–4.72µm wide, septate, sometimes very swollen and constricted at the septum, subhyaline to light brown, sometimes brown, irregularly rough-walled, stromatic hyphal aggregations, and numerous conidia. Conidiophores solitary, macronematous, 23.96–178.99 × 2.40–6.17 µm, arising terminal or laterally from hyphae, erect or flexuous, sometimes seceding at one of the lower septa, cylindrical attenuated towards the apex, sometimes filiform, non-nodulose, rarely geniculate towards the apex, pale brown to brown, sometimes becoming subhyaline towards the apex, 0–10 septate, more or less rough-walled, micronematous, lateral 5.56–21.08 × 2.01–2.78 µm, brown, narrowly cylindrical to filiform, 1–2 loci, 0–1 septum. Conidiogenous cells integrated, mostly terminal, cylindrical, cylindrical-filiform, sometimes geniculate with a small lateral shoulder at apex, 9.13–47.04 × 1.58–4.19µm, brown to pale brown, with 1–4 loci at apex, conidiogenous loci non-protuberant or only slightly so, thickened and darkened. Ramoconidia once or rarely formed, cylindrical-oblong attenuated to apex, 33.90–39.71 × 4.34–5.80µm aseptate, brown, truncate base, unthickened, refractive. Secondary ramoconidia ellipsoid, filiform, oblong-ellipsoid, sometimes subglobose, 1–4 distal hila, slightly thickened, 6.28–26.25 × 2.06–5.51 µm, 0–3 septate, subhyaline, pale brown to sometimes brown. Conidia numerous, catenate, acrogenous, aseptate or once septate, subhyaline to dark brown, intercalary conidia fusiform, ellipsoid-ovoid to subglobose 4.16–10.79 × 2.66–5.27 µm, terminal conidia subglobose to globose, sometimes obovoid, aseptate, 3.29–7.26 × 3.31–5.73 µm. Occurrence of microcyclic conidiogenesis. Type:— Brazil. Minas Gerais: Araponga, at “ Parque Estadual Serra do Brigadeiro ”, isolated from decayed leaf of Aulonemia amplissima (Poaceae), March 15, 2016, (holotype VIC 44237, ex-type culture COAD 2256). GenBank: MT373125 (ACT), MT680204 (TEF1-α), MZ318433 (rDNA-ITS) Additional specimen examined:— BRAZIL. Minas Gerais: Araponga at “Parque Estadual Serra do Brigadeiro”, isolated from decayed leaves of Aulonemia amplissima (Poaceae), 15 March 2016 (living culture COAD 2255). Notes: — Cladosporium bambusicola is a sister group of C. brigadeirensis, but the two species present morphological differences. Cladosporium bambusicola forms two types of intercalary conidia markedly different from each other. One smaller conidia type, fusiform, ellipsoid, ellipsoid-ovoid, ellipsoid-cylindrical, aseptate, subhyaline to pale brown, smooth. Another conidia type is predominantly larger, rarely catenate, subglobose, ellipsoid, limoniform, oblong, oblong-ellipsoid, brown to dark brown, wall rough, thickened. However, it is not possible to differentiate the conidiophore that originated these conidia as in Cladosporium herbaroides (C. herbarum species complex), which also have two morphological types of conidia. However, they are formed by two different conidiophores, macronematous and micronematous (Bensch et al. 2012). Cladosporium brigadeirensis does not have ramoconidia and its secondary ramoconidia is on average half the size of C. bambusicola. The conidial chains tend to be larger in the last species, while C. brigadeirensis form a cluster of conidia at the base of conidiogenous cells., Published as part of Costa, Pricila P., Rosado, André W. C. & Pereira, Olinto L., 2022, Six new species of Cladosporium associated with decayed leaves of native bamboo (Bambusoideae) in a fragment of Brazilian Atlantic Forest, pp. 1-29 in Phytotaxa 560 (1) on pages 15-17, DOI: 10.11646/phytotaxa.560.1.1, http://zenodo.org/record/7031143, {"references":["Bensch, K., Braun, U., Groenewald, J. Z. & Crous, P. W. (2012) The genus Cladosporium. Studies in Mycology 72: 1 - 401. https: // doi. org / 10.3114 / sim 0003"]}

Cladosporium chusqueae P. P. Costa, A.W.C. Rosado & O.L. Pereira sp. nov. (Figure 6) MycoBank: MB824724 Systematic position: Fungi, Ascomycota, Pezizomycotina, Dothideomycetes, Dothideomycetidae, Cladosporiales, Cladosporiaceae Etymology:—Name refers to the genus Chusquea, from which the species was isolated. Colonies on PDA attaining 52 mm diam, smoke grey, furrowed, submerged mycelium at the margin, velvety, sparse woolly mycelium central; reverse greenish grey, dull green at the margin. Colonies on MEA attaining 49 mm diam, grey olivaceous, velvety, sparse feathery mycelium central; reverse dull green. Colonies on OA attaining 53 mm diam, greenish-grey to glaucous grey, greenish glaucous at the border, fluffy, woolly on center, sector formation grey olivaceous, velvety; reverse dull green, with a light citrine to pure yellow, diffusible pigment. Colonies on SNA attaining 34 mm diam, greenish glaucous to olivaceous buff, flat and scares; reverse similar to surface. Mycelium loosely branched, 2.06–4.10 µm wide, septate, subhyaline to light brown, sometimes slightly swollen and darker brown near the origin of conidiophores, irregularly rough-walled. Conidiophores macronematous, solitary, 22.98–140.70 × 2.58–4.39 µm, arising terminal and laterally from hyphae, erect, flexuous, or bent, once dichotomously branched immediately before a septum at 90° angle, frequently cylindrical oblong, sometimes cylindrical attenuated towards the apex, some loci formed on the lateral shoulder or slight geniculate, sometimes slightly head-like, unilaterally swollen at the apex, brown or pale brown, unbranched, 0–7 septate, conspicuous at the base clearing towards the apex, more or less rough-walled, sometimes micronematous, lateral, 7.61–19.46 × 2.40–2.83 µm pale brown to brown, rarely subhyaline, narrower, 1–3 loci, 0–2 septate. Conidiogenous cells integrated, mainly terminal, sometimes geniculate towards the apex, rarely with loci formed on the minute lateral shoulder, 8.61–21.92 × 2.39–4.01µm, pale brown to brown, with 1–5 loci at apex, rarely 8, non-protuberant or only slightly so, somewhat thickened and darkened. Ramoconidia absent. Secondary ramoconidia ellipsoid to subcylindrical, 2–4 distal hila, 5.5–12.04 × 2.01–3.51 µm, aseptate, pale brown to brown, sometimes very similar to conidium, just a little bigger. Conidia numerous, catenate, acrogenous, fusiform, ellipsoid to narrowly ellipsoid-ovoid, aseptate, subhyaline to pale brown, intercalary conidia 3.81–7.34 × 1.81–2.53 µm, 1–2 distal hila, terminal conidia oblong-ellipsoid, 2.84–5.53 × 1.80–2.45 µm. Microcyclic conidiogenesis occurs. Type:— Brazil. Minas Gerais: Araponga, at “ Parque Estadual Serra do Brigadeiro ”, isolated from decayed leaf of Chusquea urelytra (Poaceae), March 15, 2016 (holotype VIC 44239, ex-type culture COAD 2258). GenBank: MT373122 (ACT), MT680201 (TEF1-α), MZ 318430 (rDNA-ITS) Additional specimens examined:— BRAZIL. Minas Gerais:Araponga, at “Parque Estadual da Serra do Brigadeiro”, isolated from decayed leaves of Chusquea urelytra (Poaceae), 15 March 2016, (living culture COAD 2261 and COAD 2262). Notes: — Cladosporium chusqueae is phylogenetically close to C. angulosum and C. xantochromaticum and has common features with both species. Similar to C. angulosum, C. chusqueae branches dichotomically near the base of the conidiophore, forming an angle of 90º. This branching occurs immediately before the septum, similar to the branches of the C. xantochromaticum. The unusual diffusible yellowish pigmentation in the culture medium was also observed in C. chusqueae as well as in C. xantochromaticum. However, this pigment was observed only in OA, whereas C. xantochromaticum was found only in the PDA medium. Unlike the two related species, ramoconidium was not observed for C. chusqueae., Published as part of Costa, Pricila P., Rosado, André W. C. & Pereira, Olinto L., 2022, Six new species of Cladosporium associated with decayed leaves of native bamboo (Bambusoideae) in a fragment of Brazilian Atlantic Forest, pp. 1-29 in Phytotaxa 560 (1) on page 21, DOI: 10.11646/phytotaxa.560.1.1, http://zenodo.org/record/7031143

Cladosporium pseudotenuissimum P. P. Costa, A.W.C. Rosado & O.L. Pereira sp. nov. (Figure 7) MycoBank: MB840952 Systematic position: Fungi, Ascomycota, Pezizomycotina, Dothideomycetes, Dothideomycetidae, Cladosporiales, Cladosporiaceae Etymology:—Name refers to phylogenetic proximity to C. tenuissimum. Colonies on PDA attaining 71 mm diam, dull green, flat, woolly; smoke grey aerial mycelium; reverse olivaceous grey to dull green. Colonies on MEA attaining 62 mm diam, grey olivaceous to smoke grey, fluffy, woolly; reverse dull green to iron grey. Colonies on OA attaining 34 mm diam, grey olivaceous to olivaceous, greenish glaucous at the border, velvet, streaked with grooves, sector formation dull green, reverse grey olivaceous. Colonies on SNA attaining 39 mm diam, isabelline to buff, very scarce, sector formation; reverse similar to surface. Mycelium diffuse, branched, 1.88–5.56 μm wide, septate, slightly swollen, pale brown to brown, sometimes irregularly rough-walled to verrucose, with some bubble-like swelling. Conidiophores numerous, solitary, macronematous, 43.38–299.42 × 2.77–4.60 μm, arising mainly laterally and rarely terminally from hyphae, erect or flexuous, cylindrical, cylindrical oblong, sometimes cylindrical attenuated towards the apex, or almost filiform, sometimes head-like swollen or occasionally with peg-like prolongations, some slightly geniculate with loci formed on the lateral shoulder, brown to dark brown, rarely dichotomically branched, 0–7 septate, septa darker and thicker toward the base, micronematous laterally, 20.7–24.7 × 4.04–5.06 μm, pale brown, cylindrical or cylindrical attenuated to apex, 1 apical locus, aseptate, reduced to conidiogenous cells. Conidiogenous cells integrated, mostly terminal, cylindrical or cylindrical attenuated to apex, sometimes terminal head-like swollen, 15.25–51.03 × 2.77–4.01 μm, rarely geniculate towards the apex with a single locus on the small lateral shoulder, brown to dark brown, sometimes pale brown, commonly 1–4 loci, rarely more at the apex, conidiogenous loci non-protuberant or only slightly so, sometimes lightly thickened and darkened. Ramoconidia occasionally formed, oblong-ellipsoid, cylindrical, 33.31–53.21 × 4.54–5.09 μm, aseptate, truncate base, brown to dark brown. Secondary ramoconidia ellipsoid, cylindrical-oblong, sometimes ovoid, 1–4 distal hila, 7.05–20.27 × 2.77–5.86 μm, 0–1 septate, pale brown to brown, darkened, slightly protuberant. Conidia numerous, sometimes catenate, acrogenous, fusiform, ellipsoid, ellipsoid-ovoid, ellipsoid-cylindrical, subglobose, limoniform, oblong, oblong-ellipsoid, aseptate, occasionally 1-septate, subhyaline, pale brown to dark brown, 4.27–6.95 × 2.24– 3.74 μm, terminal conidia ovoid to subglobose 3.09–4.90 × 1.96–3.59 μm. Microcyclic conidiogenesis occurs. Type:— Brazil. Minas Gerais: Araponga, at “ Parque Estadual Serra do Brigadeiro ”, isolated from decayed leaf of Chusquea urelytra (Poaceae), March 15, 2016 (holotype VIC 44422, ex-type culture COAD 2266). GenBank: MT373132 (ACT), MT680211 (TEF1-α), MZ318439 (rDNA-ITS) Additional specimens examined:— BRAZIL. Minas Gerais:Araponga, at “Parque Estadual da Serra do Brigadeiro”, isolated from decayed leaves of Chusquea urelytra (Poaceae), 15 March 2016, (living culture COAD 2264, COAD 2267 and COAD 2268), and isolated from decayed leaves of Chusquea anelytroides (Poaceae), 15 March 2016 (living culture COAD 2259). Notes:— Cladosporium pseudotenuissimum is phylogenetically close but distinct from C. tenuissimum. Moreover, some conspicuous morphological features distinguish the two species. Cladosporium tenuissimum have conidiophores solitary or in groups of 2–3, often are subnodulose or nodulose with apical and sometimes a few additional swellings on a lower level and have intercalary conidiogenous cells. While in C. pseudotenuissimum was only found solitary conidiophores, with swelling and conidiogenous cells restricted to the apex.Also, C. pseudotenuissimum has secondary ramoconidia, and its conidia are shorter (4.27–6.95 μm) than those described for C. tenuissimum [4–12(17)] μm. Cladosporium pseudotenuissimum also has bubble-like swelling on hyphae, which has not been described in any other species of Cladosporium until the moment., Published as part of Costa, Pricila P., Rosado, André W. C. & Pereira, Olinto L., 2022, Six new species of Cladosporium associated with decayed leaves of native bamboo (Bambusoideae) in a fragment of Brazilian Atlantic Forest, pp. 1-29 in Phytotaxa 560 (1) on pages 21-24, DOI: 10.11646/phytotaxa.560.1.1, http://zenodo.org/record/7031143

Cladosporium aulonemiae P.P. Costa, A.W.C. Rosado & O.L. Pereira sp. nov. (Figure 2) MycoBank: MB824721 Systematic position: Fungi, Ascomycota, Pezizomycotina, Dothideomycetes, Dothideomycetidae, Cladosporiales, Cladosporiaceae Etymology:—Name refers to the genus Aulonemia, from which the species was isolated. Colonies on PDA attaining 47 mm diam, dull green to dark herbage green, grey olivaceous to citrine scarce aerial mycelium, velvety, furrowed; reverse olivaceous grey. On MEA attaining 40 mm diam, grey olivaceous, greenish glaucous, pale olivaceous grey woolly central mycelium; velvety, slightly radially furrowed; reverse olivaceous grey. On OA attaining 45 mm diam, grey olivaceous, dull green at the margin; reverse greenish-grey, velvety, flat. On SNA attaining 19 mm diam, greenish glaucous to grey olivaceous, flat, and sparse; reverse similar to surface. Mycelium sparsely formed, 2.38–4.55 µm wide, septate, sometimes slightly swollen and constricted at the septum, medium to dark brown, irregularly rough-walled, sometimes walls thickened, usually covered by polysaccharide-like material, and formation of stromatic hyphal aggregations. Conidiophores macronematous, solitary, 37.14–228.51 × 2.31–4.53 µm, arising terminal and laterally from hyphae, erect or somewhat flexuous, sometimes bent, cylindrical, sometimes narrowed, several times distinctly geniculate-sinuous towards the apex due to accentuated zig-zag-like sympodial proliferation, loci then localized on small lateral shoulders, up to 37 µm long, frequently dichotomously branched towards the apex, occasionally with prolongations at the apex, brown at the base and becoming paler towards the apex, septum conspicuous, sometimes refractive,1–5 septate, walls irregularly rough-walled, thickened at the base, sometimes micronematous, lateral, 15.67–32.28 × 2.78–3.75 µm, short to medium scars 1–2 rarely 5, brown, loci somewhat protuberant, cylindrical, cylindrical attenuated. Conidiogenous cells integrated, terminal and intercalary, cylindrical, cylindrical-oblong, sometimes geniculate-sinuous, proliferation sympodial, 14.32–47.88 × 2.29–4.13 µm, conidiogenous loci at the apex (1–4) or in lateral shoulders (3–13), loci somewhat thickened and darkened. Ramoconidia rarely present, cylindrical, 19.08–27.7 × 3.08–3.51µm, 0–2 septa, pale brown, large base, somewhat thickened. Secondary ramoconidia oblong, oblong-ellipsoid, 2–5 distal hila, narrowed base, 8.26–17.98 × 2.53–4.23 µm, 0–1 septate, pale brown. Conidia numerous, catenate, acrogenous, branching in all directions, subglobose, ellipsoid, aseptate, rarely once septate, medium to dark brown, 4.59–7.9 × 2.77–4.96 µm, small terminal conidia, obovoid to globose, 3.63–5.59 × 2.90–4.06 µm, hila somewhat protuberant. Microcyclic conidiogenesis occurs. Type:— Brazil. Minas Gerais: Araponga, at “ Parque Estadual Serra do Brigadeiro ”, isolated from decayed leaf of Aulonemia amplissima (Poaceae), March 15, 2016 (holotype VIC 44413, ex-type culture COAD 2269). GenBank: MT373119 (ACT), MT680198 (TEF1-α), MZ318427 (rDNA-ITS) Additional specimens examined: — BRAZIL. Minas Gerais: Araponga, at “Parque Estadual Serra do Brigadeiro”, isolated from decayed leaves of Aulonemia amplissima (Poaceae), 15 March 2016 (living culture COAD 2270 and COAD 2271). Notes:— Cladosporium aulonemiae is phylogenetically close but distinct from C. rugulovarians. Although morphologically different, they have in common the presence of long conidiophores dichotomously branched towards the apex. Both belong to the C. cladosporioides species complex and form a basal sister clade in this complex. Perhaps this is the reason why C. aulonemiae possesses some features differentiated from their group and similar to species of C. herbarum species complex. Loci formation in close succession on small to large lateral shoulders results in a marked zig-zag-like growth. Similar conidiogenous cells are reported in C. arthropodii and C. sinuosum, belonging to the C. herbarum species complex. However, both species produce solitary and ornate conidia, characteristic of the C. herbarum species complex. In the sympodial loci of C. aulonemiae, besides the solitary and slightly rough conidia, short conidia chains or new conidiogenous cells with equal sympodial proliferation are also formed. Another unusual feature is the polysaccharide-like material produced in C. antarcticum and C. variabile (both belonging to the C. herbarum species complex). The material appears to adhere to the hypha in these species, enveloping these structures. However, in C. aulonemiae, it has more loosely adhered. It appears as a brownish smudge around the fungal structures and is also dispersed in the medium where the lamina was mounted., Published as part of Costa, Pricila P., Rosado, André W. C. & Pereira, Olinto L., 2022, Six new species of Cladosporium associated with decayed leaves of native bamboo (Bambusoideae) in a fragment of Brazilian Atlantic Forest, pp. 1-29 in Phytotaxa 560 (1) on pages 13-15, DOI: 10.11646/phytotaxa.560.1.1, http://zenodo.org/record/7031143