Morphology of Hermatomyces colonies in culture. A-C. H. bifurcatus (CCF 5899). D-F. H. constrictus (CCF 5904). G-I. H. megasporus (CCF 5898). J-L. H. reticulatus (CCF 5893). M-O. H. sphaericoides (CCF 5908). P-R. H. sphaericus (CCF 5894). S-U. H. tucumanensis (CCF 5913). V-X. H. verrucosus (CCF 5903). All after 7 d growth at 25 ºC on MEA, PCA or PDA (from left to right). 

Morphology of Hermatomyces colonies in culture. A-C. H. bifurcatus (CCF 5899). D-F. H. constrictus (CCF 5904). G-I. H. megasporus (CCF 5898). J-L. H. reticulatus (CCF 5893). M-O. H. sphaericoides (CCF 5908). P-R. H. sphaericus (CCF 5894). S-U. H. tucumanensis (CCF 5913). V-X. H. verrucosus (CCF 5903). All after 7 d growth at 25 ºC on MEA, PCA or PDA (from left to right). 

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Five new species belonging to Hermatomyces (Hermatomycetaceae, Pleosporales) are described based on morphological investigations of specimens collected on rotten twigs and stems of various plants in Panama as well as phylogenetic analyses of sequence data of nuclear ribosomal and protein coding genes (EF1-α, RPB2, β-TUB). The new species are descri...

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... ( Fig. 3G-I): Colonies moderately slow growing reaching on MEA 10-14 mm diam, on PCA 10-12 mm, and on PDA 17-18 mm after 7 d at 25 °C. No sporulation observed after 4 m at 25 ...
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... Type: Panama: Chiriquí Province: west of Los Algarrobos, path through pastures along Majagua river, 8°29'47.82"N 82°26'19.721"W, 110 m a.s.l., on dry rotten twig of unknown tree, 14 Jul. 2017, O. Koukol KZP412 (PMA 116079 -holotype; CCF 5908 -ex-holotype living culture). Cultures (Fig. 3M-O verrucose, less compact and forming a dark gray subiculum around the fertile centre but combinations of some of these features were similarly found in H. sphaericus specimens. view, muriform, smooth or verruculose, central cells brown, dark brown to blackish brown, sometimes all cells brown and muriform septation visible, outer ring of ...
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... (Fig. 3P-R (Fig. 10B-D) are either smooth or inconspicuously verruculose. Sporulation in culture media, on the other hand, was rare with only one of 12 isolates able to produce conidia after 4 m of ...
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... (Fig. 3S-U): Colonies moderately slow growing reaching on MEA 12-13 mm diam, on PCA 14-16 mm, and on PDA14-16 mm after 7 d at 25 °C. No sporulation observed after 4 m at 25 ...
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... black, peripheral cells pale brown to brown, wide but sometimes narrow, broadly ellipsoidal to oblong in side view where two distinct adpressed halves can be recognized, each half seen laterally as a row of 4-7 cells, end cells subhyaline to pale brown, middle cells brown to blackish brown, 23-30(-39) îî ± P ± PP WKLFN /:: Sexual morph unknown. (Fig. 3V-X): Colonies moderately slow growing reaching on MEA 12-13 mm diam, on PCA 12 mm, and on PDA 13-15 mm after 7 d at 25 °C. No sporulation observed after 4 m at 25 ...
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... isolated from Panamanian specimens (Fig. 3) rarely sporulated and conidia were only seen in a very few strains of taxa with a single conidium type, such as H. sphaericus ( Fig. 9L-O) or H. sphaericoides (Fig. 8L-N). This is consistent with previously described species which failed to sporulate even after extended incubation periods (Tibpromma et al. 2016, Hashimoto et al. 2017) ...
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... (Fig. 3S-U fig. 65 that he considered the cylindrical conidia to be conidiophores ("sporophores"), bearing two melanized beaks with hanging lenticular conidia. This misinterpretation might have been caused by the limited quality of the optical equipment available at the time but also both types of conidia may give that impression when seen appressed together. Our three specimens match well the protologue of H. tucumanensis in the overall shape of conidia, but our measurements are smaller compared to those of Spegazzini (1911). He recorded lenticular conidia 35-40 × 30 µm and 15-20 µm thick and "sporophores" (cylindrical conidia) 20-40 µm long with a similar thickness of 10-15 µm. Leão et al. (2013) examined what they considered to be the holotype of H. tucumanensis (LPS15823) and also described larger conidia, the lenticular conidia being 31-42 × 14-16 μm and the cylindrical ones with two columns of 2-3 cells each and 31-46 × 17-28 μm. In view of similar differences between specimens of other taxa such as H. reticulatus, we suggest to consider them as the result of intraspecific ...
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... collectors after Spegazzini subsequently recorded "H. tucumanensis" but descriptions in the literature and our study of available specimens suggest that the name became rather an "umbrella" species and was largely attributed to numerous morphologically different two conidium type and one conidium type species recorded in various countries and on diverse hosts. Hughes (1953) mentioned collections of "H. tucumanensis" from dead branches of Averrhoa carambola, Alchornea cordifolia (Euphorbiaceae), Coffea liberica (Rubiaceae), and Elaeidis guineensis (Arecaceae) from Ghana and Sierra Leone. His species concept, based on examination of the type material from Argentina along with material in K(M)-IMI, was much broader than that of Spegazzini's. Hughes (1953) depicted one specimen with 2-3-celled cylindrical conidia in one column and another morphologically different specimen with six-celled cylindrical conidia in two columns, both with apical pigmentation. In our opinion the specimens studied and illustrated by Hughes ( fig. 43) belong to at least two different species, but not to H. tucumanensis. Further records are also questionable in the absence of morphological details. Mercado (1984) examined a specimen on the dead trunk of an unidentified tree in Cuba and described only lenticular conidia, but the picture also shows cylindrical conidia with 2-5 darker apical cells resembling H. amphisporus. Similarly, another collection from Cuba and revised in our study (PRM 842348) turned out to belong to H. sphaericus (Fig. 10). Matsushima (1993) reported "H. tucumanensis" from decaying petioles of a palm tree and a twig of a broad-leaved tree in Peru with 4-6-celled cylindrical conidia composed of globose, hyaline cells. His specimen is obviously not conspecific with H. tucumanensis and he most probably referred to a distinct, yet undescribed species of Hermatomyces. Delgado (2013) reported a specimen on rachides of dead leaves of Acoelorrhaphe wrightii (Arecaceae) in Florida without depicting cylindrical conidia. Unpublished morphological data along with unedited pictures were revised within the scope of this study and the Florida specimen seems to represent H. amphisporus. Prasher & Singh (2015) recorded a collection under the name "H. tucumanensis" from a branch in litter and bark of Mangifera indica (Anacardiaceae) in India. They mentioned "elliptical to almost round", smooth lenticular conidia 30-40 × 20-28 μm long and 13.5-15 μm thick. Cylindrical conidia were not described and the lenticular conidia in Fig. 1C match well those of H. reticulatus, particularly by the presence of a distinct constriction between halves, comparable dimensions and apparently verruculose wall ornamentation. Chang (1995) recorded a specimen on unknown decaying twigs from a stream in Taiwan having cylindrical conidia with two columns of 3-4 cells each and darkly pigmented apical cells which based on pigmentation and conidial shape does not seem to be conspecific with H. tucumanensis either. Finally, Castañeda & Heredia (2000) and Prasher & Sushma (2014) considered H. tucumanensis monomorphic following Ellis (1971) who described cylindrical conidia as 'setae'. Therefore, the currently known distribution of H. tucumanensis must be considered with caution and based on our revision is limited to the ...
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... ( Fig. 3G-I): Colonies moderately slow growing reaching on MEA 10-14 mm diam, on PCA 10-12 mm, and on PDA 17-18 mm after 7 d at 25 °C. No sporulation observed after 4 m at 25 ...
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... Differs from other species of Hermatomyces with one conidium type in having distinctly verrucose lenticular conidia and subicular hyphae. Description: Colonies on the natural substrate forming sporodochial, subiculate conidiomata, superficial, more or less circular or oval, scattered or crowded and confluent, brown-black, consisting of a velvety, annular, compact, dark brown, sterile mycelial outer zone enclosing a black, glistening sporulating centre, circular, oval or lobed by confluence, where conidia are easily liberated when touched, 250-700 μm diam when solitary, up to 1500 μm long when confluent. Mycelium superficial composed of compact network of repent, branched, septate, smooth or verruculose, pale brown to brown hyphae, 2-4 μm wide; subicular hyphae septate, branched, flexuous, undulate or irregularly geniculate once or few times, pale brown to brown, finely verruculose or finely spinulose to verruculose or locally verrucose, branching and anastomosing to form a dense network. Conidiophores micronematous, mononematous, cylindrical or slightly subulate, subhyaline to pale brown, smooth or finely verruculose, 6-10 × 2-4 μm, often corresponding to conidiogenous cells. Conidiogenous cells monoblastic, integrated, terminal, determinate, often arising directly on the superficial mycelium and closely packed together at the fertile centre, sphaerical, subsphaerical or ampulliform, pale brown to brown, smooth or finely verruculose, 4-8 × 2-5 μm. Conidia of one type, lenticular, globose, subglobose or disc-shaped in front view, sometimes somewhat irregular, solitary, dry, muriform, constricted at the septa or not, verrucose, central cells brown to dark brown or blackish brown to black, peripheral cells pale brown to brown, wide but sometimes narrow, broadly ellipsoidal to oblong in side view where two distinct adpressed halves can be recognized, each half seen laterally as a row of 4-7 cells, end cells subhyaline to pale brown, middle cells brown to blackish brown, 23-30(-39) × 21-29.5 μm, 14-22 μm thick, L/W = 1.12. Sexual morph unknown. (Fig. 3V-X): Colonies moderately slow growing reaching on MEA 12-13 mm diam, on PCA 12 mm, and on PDA 13-15 mm after 7 d at 25 °C. No sporulation observed after 4 m at 25 ...
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... isolated from Panamanian specimens (Fig. 3) rarely sporulated and conidia were only seen in a very few strains of taxa with a single conidium type, such as H. sphaericus ( Fig. 9L-O) or H. sphaericoides (Fig. 8L-N). This is consistent with previously described species which failed to sporulate even after extended incubation periods (Tibpromma et al. 2016, Hashimoto et al. 2017) and seems to be a regular tendency of Hermatomyces isolates. Matsushima (1993) was the only one who reported production of both types of conidia on Corn Meal agar for a putative new species misidentified as H. ...
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... (Fig. 3P-R (Fig. 10B-D) are either smooth or inconspicuously verruculose. Sporulation in culture media, on the other hand, was rare with only one of 12 isolates able to produce conidia after 4 m of ...

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... For instance, species represented by collection and isolate singletons dominate corticioid collections in North American pine and spruce forests (Rosenthal et al. 2017), leaves in moist tropical forest of Central Panama (Arnold et al. 2000), the phyllosphere of temperate Quercus macrocarpa (Jumpponen and Jones 2009), and the romaine lettuce phylloplane . Species of Hermatomyces (Pleosporales, Hermatomycetaceae) are often described based on single isolates or specimens collected from plant material (Koukol et al. 2018;Delgado et al. 2020). In some specific sampling locations, most macrofungi are known from a unique specimen in a single location (e.g., Malaysia; Mohammad et al. 2019). ...
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... Hashimoto et al. (2017) excluded Hermatomyces from Lophiotremataceae and resurrected the family Hermatomycetaceae based on their phylogenetic analyses using SSU, ITS, LSU, tef1-α and rpb2 sequences. Asexual morph of Hermatomyces species is characterized by sporodochial conidiomata and dimorphic conidia, i.e., cylindrical conidia and lenticular conidia (Ellis, 1971;Chang, 1995;Tibpromma et al., 2016;Doilom et al., 2017;Koukol et al., 2018;Ren et al., 2021). Its sexual morph was recently reported by de Silva et al. (2022), which has dark brown to black ascomata with central ostiole, 8-spored, bitunicate asci, and broadly fusiform, hyaline, 1-septate ascospores. ...
... Conidia dimorphic, (1) cylindrical conidia 51-67 × 16-24 μm (x = 58.5 × 21 μm, n = 30), hyaline to subhyaline, often with a distinct dark brown pigmentation from the top downwards or at rim of the conidia, straight or broadly curved, phragmoseptate or muriform, sometimes with oblique septa, constricted at the septa, consisting of two columns from one or two basal cells, rounded at the apex; (2) lenticular conidia 44-52 × 29-39 μm (x = 48 × 32 μm, n = 30), ellipsoidal in front view, central cells dark brown to blackish brown, peripheral cells subhyaline to pale olivaceous brown, forming a distinct ring, muriform, constricted at the septa, smooth-walled, side views not observed. (Chang, 1995;Tibpromma et al., 2016Tibpromma et al., , 2017Hashimoto et al., 2017;Koukol et al., 2018;Ren et al., 2021). Based on phylogenetic analyses, Hermatomyces hainanensis (GZCC 23-0592) is closely related to H. megasporus (CCF 5897 and CCF 5898) and H. reticulatus (CCF 5893 and MFLUCC 15-0843), although H. reticulatus only exhibits one type of conidia (Koukol et al., 2018). ...
... (Chang, 1995;Tibpromma et al., 2016Tibpromma et al., , 2017Hashimoto et al., 2017;Koukol et al., 2018;Ren et al., 2021). Based on phylogenetic analyses, Hermatomyces hainanensis (GZCC 23-0592) is closely related to H. megasporus (CCF 5897 and CCF 5898) and H. reticulatus (CCF 5893 and MFLUCC 15-0843), although H. reticulatus only exhibits one type of conidia (Koukol et al., 2018). Despite some overlap in the sizes of the cylindrical (51-67 × 16-24 μm vs. 49.5-60.5 × 18-28 μm) and lenticular (44-52 × 29-39 μm vs. 49-56 × 37-46 μm) conidia of Hermatomyces hainanensis and H. megasporus (Koukol et al., 2018), the phylogenetic analyses suggest that they are distinct species (Figure 2). ...
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... Therefore, we report [72] described only lenticular-type conidia and considered the cylindrical ones as conidiophores. However, with further morphological and phylogenetic investigations, the muriform, lenticular, hyaline or dematiaceous conidia of some Hermatomyces sp. were reported (previously referred to as monomorphic or dimorphic), and different species were mostly distinguished by cylindrical conidia [73,74]. Species of Hermatomyces seem to be limited by elevation and climate, and prefer to live on humid plant materials, including those found on immersed woody materials in water [73,75]. ...
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... An illustrative example of multiple taxonomic issues is the genus Hermatomyces, which has received much attention in recent years from both the Paleotropics and Neotropics. Recently, an alarming high number of imprecise species descriptions based on mixed phenotypic and molecular data together with insufficient understanding of previous species delimitations was revealed (Koukol et al. 2018;Koukol and Delgado 2019;Delgado et al. 2020). Six out of the 28 binomial names introduced in Hermatomyces (Index Fungorum http://www. ...
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Recent progress in the discovery of fungal diversity has been enabled by intensive mycological surveys in centres of global biodiversity. Descriptions of new fungal species have been almost routinely based on phenotypic studies coupled with single or multigene phylogenetic analyses of DNA sequence data. However, high accessibility of sequencing services together with an increasing amount of available molecular data are providing easier and less critical support for taxonomic novelties without carefully studying the phenotype, particularly morphology. As a result, the accelerated rate of species descriptions has been unfortunately accompanied by numerous cases of overlooking previously described and well documented species, some of them that have been known for more than a century. Here, we critically examined recent literature, phenotypic and molecular data, and detected multiple issues with putative novelties of asexual Ascomycota traditionally known as hyphomycetes. In order to fix these taxonomic problems, three new combinations within the genera Pleopunctum , Camposporium and Sporidesmium , and two new names in Camposporium are proposed. Moreover, three genera, Aquidictyomyces , Fusiconidium and Pseudohelminthosporium , together with nine species are reduced to synonymy. The examples outlined here clearly show the relevance of morphology in modern phylogenetic studies and the importance of more stringent ‘quality controls’ during biodiversity studies documenting the extensive fungal diversity in a speedy manner.
... Based on morphological comparisons and phylogenetic affinities, Koukol et al. (2018) revised Hermatomyces species and described five new species (viz. H. bifurcatus, H. constrictus, H. megasporus, H. sphaericoides and H. verrucosus) (Koukol et al. 2018). ...
... Based on morphological comparisons and phylogenetic affinities, Koukol et al. (2018) revised Hermatomyces species and described five new species (viz. H. bifurcatus, H. constrictus, H. megasporus, H. sphaericoides and H. verrucosus) (Koukol et al. 2018). These are probably species complexes that need more detailed study. ...
... Subsequent studies introduced H. bauhiniae, H. biconisporus, H. clematidis, H. trangensis and H. truncates into Hermatomyces Hyde et al. 2019;Koukol et al. 2019;Nuankaew et al. 2019;). Currently, 24 species are recognized in Hermatomyces (Koukol et al. 2018(Koukol et al. , 2019Nuankaew et al. 2019;Delgado et al. 2020;Phukhamsakda et al. 2020; Table 2). ...
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During our survey of the diversity of woody litter fungi in China and Thailand, three Hermatomyces species were collected from dead woody twigs of Dipterocarpus sp. (Dipterocarpaceae) and Ehretia acuminata (Boraginaceae). Both morphology and multigene analyses revealed two taxa as new species ( Hermatomyces turbinatus and H. jinghaensis ) and the remaining collections as new records of H. sphaericus . Hermatomyces turbinatus is characterized by 1) dimorphic conidia, having circular to oval lenticular conidia and 2) turbinate conidia consisting of two columns with two septa composed of 2–3 cells in each column. Hermatomyces jinghaensis is characterized by dimorphic conidia, having circular to oval lenticular conidia and clavate or subcylindrical to cylindrical conidia and consisting of one or two columns with 6–8 cells in each column. Phylogenetic analyses of combined LSU, ITS, tub 2, tef 1-α and rpb 2 sequence data supports the placement of these new taxa within Hermatomycetaceae with high statistical support.
... Based on morphological comparisons and phylogenetic affinities, Koukol et al. (2018) revised Hermatomyces species and described five new species (viz. H. bifurcatus, H. constrictus, H. megasporus, H. sphaericoides and H. verrucosus) (Koukol et al. 2018). ...
... Based on morphological comparisons and phylogenetic affinities, Koukol et al. (2018) revised Hermatomyces species and described five new species (viz. H. bifurcatus, H. constrictus, H. megasporus, H. sphaericoides and H. verrucosus) (Koukol et al. 2018). These are probably species complexes that need more detailed study. ...
... Subsequent studies introduced H. bauhiniae, H. biconisporus, H. clematidis, H. trangensis and H. truncates into Hermatomyces Hyde et al. 2019;Koukol et al. 2019;Nuankaew et al. 2019;). Currently, 24 species are recognized in Hermatomyces (Koukol et al. 2018(Koukol et al. , 2019Nuankaew et al. 2019;Delgado et al. 2020;Phukhamsakda et al. 2020; Table 2). ...
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Full-text available
During our survey of the diversity of woody litter fungi in China and Thailand, three Hermatomyces species were collected from dead woody twigs of Dipterocarpus sp. (Dipterocarpaceae) and Ehretia acuminata (Boraginaceae). Both morphology and multigene analyses revealed two taxa as new species (Hermatomyces turbinatus and H. jinghaensis) and the remaining collections as new records of H. sphaericus. Hermato-myces turbinatus is characterized by 1) dimorphic conidia, having circular to oval lenticular conidia and 2) turbinate conidia consisting of two columns with two septa composed of 2-3 cells in each column. Hermatomyces jinghaensis is characterized by dimorphic conidia, having circular to oval lenticular conidia and clavate or subcylindrical to cylindrical conidia and consisting of one or two columns with 6-8 cells in each column. Phylogenetic analyses of combined LSU, ITS, tub2, tef1-α and rpb2 sequence data supports the placement of these new taxa within Hermatomycetaceae with high statistical support.
... It was previously placed within Ascomycota as "family incertae sedis" (Wijayawardene et al. 2012), while Doilom et al. (2017) and Tibpromma et al. (2016a) suggested that it belongs in Lophiotremataceae based on phylogeny. Currently, Hermatomycetaceae is placed in Pleosporales as a monotypic family (Koukol et al. 2018;Tibpromma et al. 2018;Hongsanan et al. 2020). We follow and Mapook et al. (2020) and their latest treatments for Hermatomycetaceae. ...
... Conidia are cylindrical and comprise 1-4 columns with 2-11 cells and are irregularly pigmented (Castañeda Ruiz and Heredia 2000;Doilom et al. 2017;Hashimoto et al. 2017;Tibpromma et al. 2018). Hermatomyces has a worldwide distribution and they have been recorded from both temperate and tropical countries including Brazil, China, Cuba, Japan, Panama, Philipines, Thailand, Venezuela Hashimoto et al. 2017;Koukol et al. 2018;Tibpromma et al. 2018). To date, 28 Hermatomyces species are listed in Species Fungorum (2021). ...
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