Material examination
Specimens were loaned from BPI, BRIP, F, G, GZU, IMI, NBRC, NY, PC, PDD, PRM, S, TRTC and UPS herbaria. These samples were observed in pure water and/or 5% KOH for sectioning. Specimens were examined using a Motic SMZ 168 stereomicroscope. Micromorphological characters were examined using a Nikon ECLIPSE 80i compound microscope and images were captured with a Canon EOS 550D and/or a Canon EOS 600D digital camera. Taro soft ® Image Framework program version 0.9.0.7 was used for measurements. Some of the type specimens were in poor condition or could not be loaned. For these taxa, we redrew the morphological features based on the original publications. The images used for photoplates were processed with Adobe Photoshop CS6 software (Adobe Systems, USA).
Phylogenetic analyses
Sequences were obtained from GenBank following previous publications and aligned using the default settings of MAFFT version 7 (http://mafft.cbrc.jp/alignment/server/index.html) (Katoh and Standley 2013). These datasets were manually corrected using Bioedit 7.0.9.0 (Hall 1999, Dissanayake et al. 2020). The phylogenetic analyses of the combined gene regions were performed using maximum-likelihood (ML) and/or Bayesian inference (BI). The evolutionary model was obtained using jModeltest 2.1.1 (Darriba et al. 2012) or ModelFinder (Kalyaanamoorthy et al. 2017). The TIM1, TIM2, TIM3, TrN and TrNef substitution models were replaced by the GTR model (Lecocq et al. 2013). The ML analyses were run with IQ-TREE (Nguyen et al. 2015; Chernomor et al. 2016), using models for each partitioned gene, with 1000 rapid bootstrap replicates. Bayesian inference was implemented by MrBayes v.3.0b4 (Ronquist and Huelsenbeck 2003) with the best-fit model of sequence evolution. Phylogenetic trees were viewed with FigTree v1.4.0 (http://tree.bio.ed.ac.uk/software/figtree/) and processed by Adobe Illustrator CS5. The final alignments and trees were deposited in TreeBASE.
Table of contents
Arrangement of genera in Boliniaceae, Calosphaeriaceae, Helminthosphaeriaceae, Jobellisiaceae, Lasiosphaeriaceae, Pleurostomataceae, Podosporaceae andSordariaceae. Those not in agreement with Wijayawardene et al. (2020) are marked with an asterisk.
Phylum Ascomycota Caval.-Sm.
ClassSordariomycetes O.E. Erikss. & Winka
SubclassDiaporthomycetidae Senan., Maharachch. & K.D. Hyde
Calosphaeriales M.E. Barr
Calosphaeriaceae Munk
Calosphaeria Tul. & C Tul.
Flabellascus Réblová
Jattaea Berl.
Togniniella Réblová et al.
Pleurostomataceae Réblová et al.
Pleurostoma Tul. & C. Tul
Calosphaeriales genus incertae sedis
Kacosphaeria Speg.
Jobellisiales M.J. D'souza & K.D. Hyde*
Jobellisiaceae Réblová
Jobellisia M.E. Barr
Other genera in Diaporthomycetidae
Ophiostomatales Benny & Kimbr.
Ophiostomataceae Nannf.
Fragosphaeria Shear
SubclassHypocreomycetidae O.E. Erikss. & Winka
Genera transferred to Hypocreomycetidae
Coronophorales genus incertae sedis
Tengiomyces Réblová*
Hypocreales genus incertae sedis
Sulcatistroma A.W. Ramaley*
SubclassSordariomycetidae O.E. Erikss. & Winka
Boliniales P.F. Cannon
Boliniaceae Rick
Apiocamarops Samuels & J.D. Rogers
Apiorhynchostoma Petr.
Camaropella Lar.N. Vassiljeva
Camarops P. Karst.
= Bolinia (Nitschke) Sacc.
Cornipulvina Huhndorf et al.
Endoxyla Fuckel
Mollicamarops Lar.N. Vassiljeva
Neohypodiscus J.D. Rogers et al.
Pseudovalsaria Spooner
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández
Helminthosphaeriaceae Samuels, Cand. & Magni
Echinosphaeria A.N. Mill. & Huhndorf
Helminthosphaeria Fuckel
Hilberina Huhndorf & A.N. Mill.
Ruzenia O. Hilber
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss
Bombardiaceae S.K. Huang & K.D. Hyde*
Apodospora Cain & J.H. Mirza*
Bombardia (Fr.) P. Karst.*
Bombardioidea C. Moreau ex N. Lundqv.*
Fimetariella N. Lundq.*
Lasiosphaeriaceae Nannf.
Anopodium Lundq.
Bellojisia Réblová
Corylomyces Stchigel et al.
Lasiosphaeria Ces. & De Not.
Mammaria Ces. ex Rabenh.
Zopfiella G. Winter
Lasiosphaeridaceae S.K. Huang & K.D. Hyde*
Lasiosphaeris Clem.*
Podosporaceae X. Wei Wang & Houbraken
Cladorrhinum Sacc. & Marchal
Podospora Ces.
= Schizothecium Corda*
Triangularia Boedijn
= Apiosordaria Arx & W. Gams*
Sordariaceae G. Winter
Boothiella Lodhi & Mirza*
Guilliermondia Boud.
Neurospora Shear & B.O. Dodge
Pseudoneurospora Dania García et al.
Sordaria Ces & De Not.
Strattoniaceae S.K. Huang & K.D. Hyde*
Strattonia Cif.*
Zygospermellaceae S.K. Huang & K.D. Hyde*
Episternus Górz & Boroń*
Zygospermella Cain*
Other genera in Sordariales
Chaetomiaceae G. Winter
Stellatospora T. Ito & Nakagiri*
Neoschizotheciaceae S.K. Huang & K.D. Hyde*
Apodus Malloch & Cain*
Cercophora Fuckel*
Echria (N. Lundq.) Kruys et al.*
Immersiella A.N. Mill. & Huhndorf*
Jugulospora N. Lundq.*
Neoschizothecium S.K. Huang & K.D. Hyde*
Rinaldiella Deanna A. Sutton et al.*
Zygopleurage Boedijn*
Sordariales genera incertae sedis
Arnium Nitschke ex G. Winter*
Biconiosporella Schaumann*
Camptosphaeria Fuckel*
Diffractella Guarro et al.*
Emblemospora Jeng & J.C. Krug*
Eosphaeria Höhn.*
Isia D. Hawksw. & Manohar.
Lockerbia K.D. Hyde
Periamphispora J.C. Krug*
Ramophialophora M. Calduch et al.*
Reconditella Matzer & Hafellner
Roselliniopsis Matzer & Hafellner
Synaptospora Cain*
Tripterosporella Subram. & Lodha*
Other genera studied
SubclassXylariomycetidae O.E. Erikss. & Winka
Xylariales Nannf.
Diatrypaceae Nitschke
Monosporascus Pollack & Uecker
Sordariomycetes genus incertae sedis
Conidiotheca Réblová & L Mostert *
Ascomycota genera incertae sedis
Copromyces N. Lundq.*
Effetia Bartoli et al.*
Endophragmiella B. Sutton*
Tulipispora Révay & J. Gönczöl *
Table 2. Change of taxa placement from Wijayawardene et al. 2020 to present
Taxa
|
Taxa placement in Wijayawardene et al. 2020
|
Present placement
|
Jobellisiales M.J. D'souza & K.D. Hyde
|
Hypocreomycetidae O.E. Erikss. & Winka
|
Diaporthomycetidae Senan., Maharachch. & K.D. Hyde
|
Apodospora Cain & J.H. Mirza
|
Lasiosphaeriaceae Nannf.
|
Bombardiaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Apodus Malloch & Cain
|
Lasiosphaeriaceae Nannf.
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Arnium Nitschke ex G. Winter
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Biconiosporella Schaumann
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Bombardia (Fr.) P. Karst.
|
Lasiosphaeriaceae Nannf.
|
Bombardiaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Bombardioidea C. Moreau ex N. Lundqv.
|
Lasiosphaeriaceae Nannf.
|
Bombardiaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Boothiella Lodhi & Mirza
|
Sordariomycetes genera incertae sedis
|
Sordariaceae G. Winter
|
Camptosphaeria Fuckel
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Cercophora Fuckel
|
Lasiosphaeriaceae Nannf.
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Conidiotheca Réblová & L Mostert
|
Togniniaceae Réblová, L. Mostert, W. Gams & Crous
|
Sordariomycetes genera incertae sedis
|
Copromyces N. Lundq.
|
Sordariaceae G. Winter
|
Ascomycota genera incertae sedis
|
Diffractella Guarro et al.
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Echria (N. Lundq.) Kruys et al.
|
Not mentioned
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Effetia Bartoli et al.
|
Sordariaceae G. Winter
|
Ascomycota genera incertae sedis
|
Emblemospora Jeng & J.C. Krug
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Endophragmiella B. Sutton
|
Helminthosphaeriaceae Samuels, Cand. & Magni.
|
Ascomycota genera incertae sedis
|
Eosphaeria Höhn.
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Episternus Górz & Boroń
|
Lasiosphaeriaceae Nannf.
|
Zygospermellaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Fimetariella N. Lundq.
|
Lasiosphaeriaceae Nannf.
|
Bombardiaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Immersiella A.N. Mill. & Huhndorf
|
Lasiosphaeriaceae Nannf.
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Jugulospora N. Lundq.
|
Lasiosphaeriaceae Nannf.
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Lasiosphaeris Clem.
|
Sordariales genera incertae sedis
|
Lasiosphaeridaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Periamphispora J.C. Krug
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Ramophialophora M. Calduch et al.
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Rinaldiella Deanna A. Sutton et al.
|
Lasiosphaeriaceae Nannf.
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Stellatospora T. Ito & Nakagiri
|
Sordariaceae G. Winter
|
Chaetomiaceae G. Winter
|
Strattonia Cif.
|
Lasiosphaeriaceae Nannf.
|
Strattoniaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Sulcatistroma A.W. Ramaley
|
Calosphaeriales genera incertae sedis
|
Hypocreales genera incertae sedis
|
Synaptospora Cain
|
Helminthosphaeriaceae Samuels, Cand. & Magni.
|
Sordariales genera incertae sedis
|
Tengiomyces Réblová
|
Helminthosphaeriaceae Samuels, Cand. & Magni.
|
Coronophorales genera incertae sedis
|
Tripterosporella Subram. & Lodha
|
Lasiosphaeriaceae Nannf.
|
Sordariales genera incertae sedis
|
Tulipispora Révay & J. Gönczöl
|
Sordariomycetes genera incertae sedis
|
Ascomycota genera incertae sedis
|
Zygopleurage Boedijn
|
Lasiosphaeriaceae Nannf.
|
Neoschizotheciaceae S.K. Huang & K.D. Hyde
|
Zygospermella Cain
|
Lasiosphaeriaceae Nannf.
|
Zygospermellaceae S.K. Huang, Maharachch. & K.D. Hyde
|
Diaporthomycetidae Senan., Maharachch. & K.D. Hyde, Fungal Diversity 72: 208 (2015)
Maharachchikumbura et al. (2015) introduced the subclass Diaporthomycetidae, which is related to Sordariomycetidae based on phylogenetic analyses (Hongsanan et al. 2017; Hyde et al. 2017, 2020; this study). There are 15 orders in this subclass, viz. Annulatascales, Atractosporales, Calosphaeriales, Diaporthales, Distoseptisporales, Jobellisiales, Magnaporthales, Myrmecridiales, Ophiostomatales, Pararamichloridiales, Phomatosporales, Sporidesmiales, Tirisporellales, Togniniales and Xenospadicoidales (Maharachchikumbura et al. 2015; Hyde et al. 2020; this study, Fig. 1) and they are widely distributed as saprobes, pathogens, or endophytes in aquatic or/and terrestrial habitats (Hongsanan et al. 2017; Hyde et al. 2017, 2020). The known asexual morphs in this subclass have been linked to hyphomycetous or coelomycetous fungi generally forming phialidic conidiogenesis and holoblastic conidiogenous cells (Zhang and Blackwell 2001; Vijaykrishna et al. 2004; Réblová et al. 2004, 2015; Najwa et al. 2012; Senanayake et al. 2017).
Calosphaeriales M.E. Barr, Mycologia 75(1): 11 (1983)
The genera in Calosphaeriales are characterized by black ascomata with an eccentric papilla or long neck, with conspicuously septate, tapered paraphyses, numerous, clavate or subglobose asci borne from fascicled or spicate ascogenous hyphae, hyaline, allantoid to subcylindrical ascospores and phialidic hyphomycetous asexual morphs (Barr 1985; Réblová et al. 2004, 2015; Réblová and Mostert 2007). Calosphaeriales and Togniniales are similar in morphology, but are phylogenetically distant (Réblová et al. 2015; this study, Fig. 1). Ascogenous hypha, on which the asci form, are conspicuous character of members in Calosphaeriales and Togniniales, and is an important feature to distinguish the genera in these two orders (Barr 1985; Mostert et al. 2006; Damm et al. 2008; Réblová et al. 2015). Two families are included in this order, Calosphaeriaceae with four genera, and the monotypic Pleurostomataceae (Wijayawardene et al. 2020; this study). Jobellisiaceae was transferred to Calosphaeriales based on multigene analysis (Hongsanan et al. 2017). However, Hongsanan et al. (2017) mentioned that Jobellisiaceae is an unstable group and it is closely related to Calosphaeriales, Diaporthales and Togniniales (Hyde et al. 2017; this study, Fig. 1). We place Jobellisiaceae under Jobellisiales (see notes for Jobellisiales). Calosphaeriales is sister to Togniniales in Diaporthomycetidae (Fig. 1).
Calosphaeriaceae Munk, Dansk bot. Ark. 17(no. 1): 278 (1957)
Saprobic on dead wood or leaves in aquatic and/or terrestrial habitats, sometimes fungicolous. Sexual morph: Ascostromata semi-immersed to erumpent through the surface of the host. Ascomata perithecial, gregarious or scattered or solitary, immersed, semi-immersed or superficial, globose to subglobose, brown to black, coriaceous to membranaceous, tuberculate or smooth or with hyphal coating, ostiolate, with long or short necks. Peridium coriaceous or membranaceous, composed of hyaline to brown cells of textura angularis to textura prismatica. Paraphyses numerous, broad, cylindrical, septate, tapered. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with obovoid to obpyriform cells formed in sympodial succession, growing in dense clusters and each giving rise to an ascus. Asci 8-spored, unitunicate, clavate to subglobose, pedicellate, apex rounded or truncate, with or without an apical ring. Ascospores bi-seriate to overlapping, allantoid to ellipsoidal, straight to slightly curved, mostly aseptate, sometimes multi-septate and producing numerous ascoconidia when mature, hyaline to pale brown, smooth-walled, sometimes with guttules. Ascoconidia hyaline, allantoid to ellipsoidal, aseptate, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Conidiophores micronematous or mononematous, hyaline, straight or flexuous, septate, branched or unbranched. Phialides enteroblastic or holoblastic, monophialidic, hyaline, smooth-walled. Conidia aggregated, hyaline, aseptate, cylindrical to allantoid, with or without guttules (adapted from Réblová et al. 2004, 2015, 2016; Maharachchikumbura et al. 2016).
Type genus: Calosphaeria Tul. & C. Tul. 1863
Notes: Calosphaeria, Conidiotheca, Jattaea, Kacosphaeria, Sulcatistroma, Togniniella and Tulipisopora have traditionally been accommodated Calosphaeriaceae (Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2018). Most species are wood inhabitants and are mostly reported from Asia and Europe (Barr 1985; Damm et al. 2008; Réblová et al. 2015). Réblová et al. (2004) established Calosphaeriophora as a member of Calosphaeriaceae, typified by Cal. pulchella. Subsequently, according to nomenclatural priority, Réblová et al. (2016) recommended that Calosphaeriophora should be synonymized under Calosphaeria (Réblová et al. 2004, 2015, 2016). Vu et al. (2019) sequenced the ex-type strain of Sulcatistroma and found that it is more closely related to Allantonectria (Nectriaceae) (Ramaley 2005; Hirooka et al. 2012; this study). Kacosphaeria was accepted as a member of Calosphaeriaceae based on the similarity of clavate asci with Calosphaeria (Spegazzini 1888). We were unable to obtain authentic material of Kacosphaeria and its description lacks sufficient detail to prove that this genus belongs to Calosphaeriaceae. Therefore, we propose to transfer it to Calosphaeriales genera incertae sedis. The Tulipispora has been placed in Calosphaeriaceae without molecular data (Maharachchikumbura et al. 2015, 2016). This hyphomycetous genus has multi-septate conidia which is different from other members of Calosphaeriaceae. Genera such as Triscelophorus and Triramulispora have multi-septate conidia which are similar to Tulipispora (Révay et al. 2009). Thus, we place Tulipispora in Ascomycota genera incertae sedis. The monotypic Conidiotheca is similar to Jattaea and Pleonectria (Nectriaceae) in having ellipsoidal to cylindrical ascospores producing ascoconidia (a conidium formed directly from an ascospore, especially when still within the ascus) (Hirooka et al. 2012). Molecular data of Conidiotheca is unavailable and it has black ascomata and muriform ascospores, instead of red ascomata in Pleonectria or transverse-septate ascospores in Jattaea. Therefore, we place this genus in Sordariomycetes genera incertae sedis pending further studies. Réblová et al. (2015) introduced Flabellascus as a member of Calosphaeriaceae based on morphological and phylogenetic analyses. Available phylogenetic data for Calosphaeriaceae, including Calosphaeria, Flabellascus, Jattaea and Togniniella, are shown in Fig. 2 and forms a sister clade to Pleurostomaceae in Calosphaeriales (Fig. 1).
Calosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 108 (1863)
Saprobic on dead wood. Sexual morph: Ascomata perithecial, gregarious or solitary, immersed to semi-immersed, globose to subglobose, brown to black, coriaceous to membranaceous, tuberculate or smooth or with hyphal coating. Necks long or short, oblong to subglobose, ostiolate, periphysate. Peridium comprising two layers, outer layer coriaceous, composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses numerous, broad, cylindrical, septate, longer than the asci. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with ovoid to ellipsoidal cells, growing in sympodial succession, in dense clusters and each giving rise to an ascus. Asci 8-spored, unitunicate, clavate, pedicellate, apex rounded or truncate, with or without J- apical ring. Ascospores bi-seriate to overlapping in the upper part of ascus, allantoid to ellipsoidal, straight to slightly curved, aseptate, hyaline, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Conidiophores micronematous, hyaline, straight or flexuous, septate, branched. Phialides enteroblastic or holoblastic, monophialidic, elongate ampulliform to subcylindrical, hyaline, smooth-walled. Conidia aggregated, hyaline, aseptate, cylindrical to allantoid, with or without guttules (adapted from Barr et al. 1993; Réblová et al. 2004).
Notes: Calosphaeria has been included in various families, such as Diatrypaceae and Gnomoniaceae, based on the tiny asci and allantoid ascospores (Barr et al. 1993; Réblová et al. 2004). Tulasne and Tulasne (1863) established Calosphaeria, typified by C. princeps. Over 80 species of Calosphaeria have been found on wood in freshwater and/or terrestrial habitats in America, Asia, Australia and Europe (Barr 1985; Réblová et al. 2004; Damm et al. 2008), but few have molecular data. Damm et al. (2008) and Réblová et al. (2004) sequenced C. africana and C. pulchella, and these Calosphaeria species form a sister clade to Jattaea in Calosphaeriaceae based on multi-gene analysis (0.98BY, Fig. 2).
Type species: Calosphaeria princeps Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 109 (1863)
Basionym: Sphaeria pulchella Pers., Neues Mag. Bot. 1: 83 (1794)
Facesoffungi number: *****??; Fig. 3
Saprobic on dead wood. Sexual morph: Ascomata 300–450 µm (n = 10) diam. at the venter, perithecial, gregarious, immersed, the long necks converging radially and erumpent through the bark of the host, globose, brown to dark brown, coriaceous, glabrous. Necks 1.5–2.5 × 0.1–0.2 mm (x̄ = 2 × 0.15 mm, n = 10), coriaceous, central, fasciculate, dark brown to black, smooth, wall composed of brown to hyaline cells of textura epidermoidea to textura prismatica (45–75 µm wide, n = 20), ostiole with hyaline periphyses. Peridium 40–80 µm (x̄ = 55 µm, n = 30) wide at venter-walled, comprising two layers, outer layer coriaceous, composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses 2–4.5 µm (x̄ = 3.5 µm, n = 30) wide, numerous, cylindrical, septate, slightly constricted at the septa, branched, apically rounded, tapering, longer than the asci. Ascogenous hyphae discrete, elongate, hyaline, smooth-walled, branched, with hyaline, ovoid to ellipsoidal cells, 10–25 × 1.5–4 µm (x̄ = 15.5 × 3 µm, n = 20), growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci 30–45(–50) × 3–5 µm (x̄ = 40 × 4 µm, n = 50), 8-spored, unitunicate, clavate, very long pedicellate, apex blunt, without an apical ring. Ascospores 3.5–5.5 × 0.5–1.5 µm (x̄ = 4 × 1 µm, n = 50), overlapping near the apex, hyaline, allantoid to ellipsoidal, straight to slightly curved, aseptate, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Conidiophores micronematous, hyaline, straight or flexuous, septate. Phialides terminal or intercalary, monophialidic, ampulliform to subcylindrical, hyaline, smooth-walled. Conidia hyaline, aseptate, cylindrical to allantoid, with guttules (adapted from Réblová et al. 2004).
Material examined: USA, Kansas, Stockton, on the inner bark of Prunus cerasus, 5 October 1906, E. Bartholomew (S-F263455); USA, Missouri, Emma, on dead trunks of Persica vulgaris, November 1903, C.H. Demetrio (S-F263457); France, 86 Vienne, Poitiers, Pictavii, on trunks of cherry tree, 1840 (PC-MNHN-PC-PC0167696); France, 49 Maine-et-Loire, Angers, on trunks of cherry tree, J.P. Guépin (PC-MNHN-PC-PC0167711).
Known hosts and distribution: On decayed wood of Betula, Cerasus and Prunus in France (type locality), Germany and North America (Persoon 1794; Fries 1823a; Tulasne and Tulasne 1863; Schröter 1897; Barr 1985); on branches of Solanum auriculatum in South Africa (Doidge 1950).
Notes: Persoon (1794) introduced Sphaeria pulchella collected on decayed wood of Cerasus and Prunus and Fries (1823a) made further collections on Betula and Prunus in America. Tulasne and Tulasne (1863) established Calosphaeria and transferred Sphaeria pulchella to Calosphaeria princeps as the generic type. Schröter (1897) subsequently re-named C. princeps as Calosphaeria pulchella as type species, but this view was refuted by Clements and Shear (1931) and Barr (1985). Damm et al. (2008) reported that the ascospore size of ‘Calosphaeria princeps’ differs from ‘Calosphaeria pulchella’. In this study, based on priority, the oldest name Calosphaeria princeps is recommended for use. The type material could not be obtained. Therefore, we re-examined the authentic samples collected from France and the USA. They are morphologically similar to C. princeps as described by Tulasne and Tulasne (1863). Phylogenetically, in this study, C. pulchella (strain CBS 115999) is considered as C. princeps and is sister to C. africana (100%ML/1.00BY, Fig. 2). However, more fresh collections and phylogenetic information are expected to determine the affinities of Calosphaeria species.
Flabellascus Réblová, in Réblová, Jaklitsch, Réblová& Štěpánek, PLoS ONE 10(12): e0144616, 15 (2015)
Saprobic on wood. Sexual morph: Ascomata perithecial, immersed to semi-immersed, subglobose, ostiolate, with elongate necks, periphysate. Peridium coriaceous to membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, broad, cylindrical, septate, tapering, longer than the asci. Ascogenous hyphae discrete, hyaline, smooth-walled, elongate, branched, with ellipsoidal to obpyriform cells, growing in sympodial succession, in dense clusters and each giving rise to an ascus. Asci 8-spored, unitunicate, clavate, pedicellate, thickened at the apex. Ascospores bi-seriate to overlapping in the upper part of ascus, hyaline, allantoid to ellipsoidal, smooth-walled, aseptate. Asexual morph: Hyphomycetous. Conidiophores macronematous to semi-macronematous, brown, unbranched or branched, with terminate in a single or multiple verticillate phialide(s). Phialides ampulliform to elongate-ampulliform, hyaline to pale brown, tapering. Conidia aggregated, hyaline, aseptate, allantoid to reniform (adapted from Réblová et al. 2015).
Notes: Flabellascus is monotypic. Its sexual morph is similar to Jattaea and different asexual morphs distinguish them. The former has macronematous and branched conidiophores, whereas the latter generally has micronematous and unbranched conidiophores (Réblová et al. 2015). In our study, Flabellascus is sister to Togniniella in Calosphaeriaceae (100%ML/1.00BY, Fig. 2).
Type species: Flabellascus tenuirostris Réblová, in Réblová, Jaklitsch, Réblová & Štěpánek, PLoS ONE 10(12): e0144616, 15 (2015)
Fig. 4 a–b
Saprobic on wood. Sexual morph: Ascomata perithecial, immersed, the necks converging radially and erumpent through the bark of the host, subglobose, ostiolate, with elongate necks, periphysate. Peridium coriaceous to membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, broad, cylindrical, septate, tapering, longer than the asci. Ascogenous hyphae discrete, hyaline, smooth-walled, elongate, branched, with ellipsoidal to obpyriform cells, growing in sympodial succession, in dense clusters and each giving rise to an ascus. Asci 8-spored, unitunicate, clavate, pedicellate, conspicuously thickened at the apex. Ascospores bi-seriate to overlapping in the upper part of ascus, hyaline, allantoid to ellipsoidal, smooth-walled, aseptate. Asexual morph: Hyphomycetous. Conidiophores macronematous to semi-macronematous, brown, unbranched or branched, with terminate in a single phialide or 2–3 verticillate phialides. Phialides ampulliform to elongate-ampulliform, hyaline to pale brown, gradually tapering. Conidia aggregated, hyaline, aseptate, allantoid to reniform (adapted from Réblová et al. 2015).
Known hosts and distribution: On wood of Fagus sylvatica and Quercus cerris in Czech Republic (type locality) (Réblová et al. 2015).
Notes: Réblová et al. (2015) introduced the characteristics of Flabellascus tenuirostris in detail, and analyzed four strains with ITS, LSU, TUB and RPB2 regions. The drawing is provided for the type species (Fig. 4 a–b), based on Réblová et al. (2015).
Jattaea Berl., Icon. fung. (Abellini) 3(1-2): 6 (1900)
Saprobic on wood. Sexual morph: Ascomata perithecial, aggregated to scattered, immersed to semi-immersed, often associated with other dead ascomata, brown to dark brown, globose to subglobose, membranaceous, glabrous or covered with hyphae, ostiolate, with long or short necks, periphysate. Peridium comprising two layers, outer layer composed of brown to pale brown cells of textura angularis to textura prismatica; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, broad, cylindrical, septate. Ascogenous hyphae discrete, hyaline, smooth-walled, with ellipsoidal to obpyriform cells, growing in sympodial succession, in dense clusters and each giving rise to an ascus. Asci 8-spored, unitunicate, clavate, pedicellate, apex obtuse. Ascospores bi-seriate to overlapping, hyaline, allantoid to ellipsoidal, 0–multi-septate, smooth-walled, sometimes with guttules, sometimes producing numerous ascoconidia when mature. Ascoconidia hyaline, allantoid to ellipsoidal, aseptate, smooth-walled, guttulate. Asexual morph: Hyphomycetous, Conidiophores hyaline to pale brown, septate, unbranched to branched. Phialides monophialidic, elongate to ampulliform, fusiform, smooth-walled. Conidia hyaline, ovoid to suballantoid, straight or slightly curved, aseptate, smooth-walled (adapted from Berlese 1900; Réblová 2011).
Notes: Jattaea and Wegelina were proposed as independent genera (Clements and Shear 1931). Subsequently, Réblová (2011) showed that the type Wegelina discreta and Jattaea species are congeneric based on DNA sequence analyses, and reduced Wegelina to a synonym of Jattaea (Réblová 2011). Jattaea resembles Calosphaeria, Flabellascus, Pleurostoma and Togniniella in having clavate asci and allantoid ascospores (Réblová 2011; Huang et al. 2018). Jattaea species are aggregated and sister to Calosphaeria (0.98BY) and distinct from Flabellascus, Pleurostoma and Togniniella (Fig. 2).
Type species: Jattaea algeriensis Berl., Icon. fung. (Abellini) 3(1-2): 6 (1900)
Saprobic on wood. Sexual morph: Ascomata perithecial, aggregated to scattered, immersed, black, glabrous, ostiolate, with necks. Asci 8-spored, unitunicate, cylindrical. Ascospores hyaline, allantoid, aseptate, smooth-walled. Asexual morph: Hyphomycetous, phialophora-like. Conidia hyaline, ovoid to suballantoid, aseptate, smooth-walled (adapted from Berlese 1900; Damm et al. 2008).
Known hosts and distribution: On decayed stems or wood of Rubus in Algeria (type locality) (Berlese 1900).
Notes: Jattaea algeriensis was reported on the sarmentum of Rubus fruticosus in Algeria (Berlese 1900). Réblová (2011) used CBS 120871 to stand for J. algeriensis. However, strain CBS 120871 was sequenced from type material of Jattaea prunicola (Damm et al. 2008). The asci of J. algeriensis is clavate with long pedicel which is different from J. prunicola with cylindrical asci (Damm et al. 2008). We were unable to obtain authentic materials or find a reliable illustration. Therefore, we re-examined lectotype of Jattaea discreta, which is similar to J. algeriensis (Damm et al. 2008; Réblová 2011), and provide an illustration. Fresh collections and phylogenetic data of J. algeriensis are needed to determine its taxonomic placement.
Jattaea discreta (Berl.) Réblová, Fungal Diversity 49: 179 (2011)
Basionym: Wegelina discreta Berl., Icon. fung. (Abellini) 3(1-2): 8 (1900)
Facesoffungi number: *****; Fig. 5
Saprobic on wood. Sexual morph: Ascomata 200–350 µm (n = 10) diam. at the venter, perithecial, aggregated to scattered, immersed, the elongate necks converging radially and erumpent through the bark of the host, often associated with other dead ascomata, brown to dark brown, globose to subglobose, glabrous. Necks 100–150 × 400–700 μm (x̄ = 130 × 550 µm, n = 10), central, ostiolate, periphysate, straight or slightly flexuous, rounded to slightly tapering at the apex, the wall composed of dark brown to brown cells of textura epidermoidea to textura prismatica. Peridium 20–55 µm (x̄ = 35 µm, n = 30) wide at venter-walled, comprising two layers, outer layer composed of brown to pale brown cells of textura angularis to textura prismatica; inner layer composed of hyaline cells of textura prismatica. Paraphyses 1–2 µm wide, numerous, broad, cylindrical, septate, slightly constricted at the septa, unbranched, apex rounded. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with hyaline, ellipsoidal to obpyriform cells, 3–4.5 × 1.5–2.5 µm (x̄ = 3.5 × 2 µm, n = 20), growing in sympodial succession, in dense clusters and each giving rise to an ascus. Asci (15–)17–23(–30) × 2–4 µm (x̄ = 20 × 3 µm, n = 50), 8-spored, unitunicate, clavate, apex obtuse, without apical ring, long pedicellate. Ascospores 4–6 × 1–2 µm (x̄ = 5 × 1.5 µm, n = 50), bi-seriate to overlapping, hyaline, allantoid to ellipsoidal, slightly curved, aseptate, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Conidiophores hyaline to pale brown, erect, septate, unbranched or branched. Phialides monophialidic, terminal to intercalary, elongate to ampulliform, fusiform, narrowed at the apex, smooth-walled, hyaline, with cylindrical to ampulliform adelophialides. Conidia hyaline, ovoid to suballantoid, straight or slightly curved, aseptate, smooth-walled, aggregated, accumulating in slimy masses on the apex of the phialides (adapted from Réblová 2011).
Material examined: Italy, Veneto, Belluno, on decaying wood of a branch of Acer pseudoplatanus, Autumn 1879, (NY-00912077, lectotype).
Known hosts and distribution: On decayed wood of Crataegus, Acer, Quercus, Vaccinium in France, Italy (type locality) and USA (Berlese 1900; Réblová 2006, 2007; 2011).
Notes: Wegelina species have been transferred to Barbatosphaeria, Ceratostomella and Jattaea (Kummer et al. 2005; Réblová 2006, 2007, 2011). Jattaea and Wegelina have similar characters with clavate asci and hyaline, allantoid ascospores, but differ in the length of the ascomata’s neck (Berlese 1900). Réblová (2011) confirmed that Wegelina discreta is a member of Jattaea. This sample (NY-00912077) was initially determined as Calosphaeria wahlenbergii, then identified as an isotype of Wegelina discreta by Barr in 1984 (mentioned in the label of material), and designated as a lectotype of J. discreta by Réblová (2011).
Togniniella Réblová, L. Mostert, W. Gams & Crous, Stud. Mycol. 50(2): 543 (2004)
Saprobic on wood. Sexual morph: Ascomata perithecial, gregarious or solitary, immersed to semi-immersed, globose to subglobose, glabrous, brown to black, carbonaceous to coriaceous, immersed, the long necks erumpent through the bark of the host. Necks central, cylindrical or slightly flexuous, black, smooth, apex broadly rounded, ostiolate, periphysate. Peridium comprising two layers, outer layer carbonaceous or coriaceous, composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses unbranched, broad, cylindrical, septate, slightly constricted at the septa, tapering, with a rounded apex, longer than asci. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with hyaline, obovoid to ellipsoidal cells, growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci 8-spored, unitunicate, clavate, pedicellate, apex obtuse, without apical ring. Ascospores overlapping near the apex, allantoid to suballantoid, straight to slightly curved, aseptate, hyaline, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Conidiophores macronematous or micronematous, usually arising from aerial hyphae, erect, hyaline to pale brown, unbranched or branched, straight or flexuous, septate, constricted at the septa. Phialides monophialidic or polyphialidic, terminal or intercalary, subcylindrical to ampulliform, tapering, smooth-walled, hyaline, with cylindrical adelophialides. Conidia hyaline, obovoid to reniform, straight or slightly curved, aseptate, smooth-walled, accumulating in slimy masses on the apex of the phialides (adapted from Réblová et al. 2004).
Notes: The monotypic Togniniella initially typified by T. acerosa, and its asexual morph was introduced as Phaeocrella acerosa (Réblová et al. 2004). Subsequently, the name T. acerosa was corrected as T. microspora (Réblová 2011; Réblová et al. 2015). Réblová et al. (2016) recommended that Phaeocrella should be synonymized under Togniniella, according to nomenclatural priority. This genus is characterized by clavate, tiny asci, allantoid ascospores and verrucous phialides with reniform conidia (Réblová et al. 2004, 2015, 2016; Réblová 2011), and phylogenetic information is available (Réblová et al. 2004; Damm et al. 2008; Vu et al. 2019). In this study, Togniniella is sister to Flabellascus (100%ML/1.00BY, Fig. 2).
Type species: Togniniella microspora (Ellis & Everh.) Réblová, Fungal Diversity 49: 193 (2011)
Basionym: Ceratostomella microspora Ellis & Everh., Proc. Acad. nat. Sci. Philad. 45: 444 (1894)
Facesoffungi number: *****??; Fig. 6
Saprobic on wood. Sexual morph: Ascomata 200–400 µm (250 µm, n = 10) diam. at the venter, perithecial, gregarious or solitary, immersed to semi-immersed, the long necks erumpent through the bark of the host, globose to subglobose, glabrous, brown to black, carbonaceous to coriaceous. Necks 550–1000 × 55–75 µm (x̄ = 755 × 65 µm, n = 10), carbonaceous, central, cylindrical or slightly flexuous, black, smooth, apex broadly rounded, ostiole with hyaline periphyses. Peridium 20–30 µm (x̄ = 25 µm, n = 30) wide at venter-walled, comprising two layers, outer layer carbonaceous to coriaceous, composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses 7–8 µm (x̄ = 7.5 µm, n = 30) wide near the base, 1.5–5 µm (x̄ = 2.5 µm, n = 30) wide near the apex, unbranched, broad, cylindrical, septate, slightly constricted at the septa, tapering, with a rounded apex, longer than asci. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with hyaline, obovoid to ellipsoidal cells, 3–4.5 × 1.5–2.5 µm (x̄ = 3.5 × 2 µm, n = 20), growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci 10–16(–20) × 2–5 µm (x̄ = 15 × 3 µm, n = 50), 8-spored, unitunicate, clavate, long pedicellate, apex obtuse, apical ring absent. Ascospores 2.5–4 × 0.2–0.5 µm (x̄ = 3.5 × 0.3 µm, n = 50), overlapping near the apex, allantoid to suballantoid, straight to slightly curved, aseptate, hyaline, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Conidiophores macronematous or micronematous, usually arising from aerial hyphae, erect, hyaline to pale brown, unbranched or branched, straight or flexuous, 1–4-septate, constricted at the septa. Phialides monophialidic or polyphialidic, terminal to intercalary, subcylindrical to ampulliform, tapering, smooth-walled, hyaline, with cylindrical adelophialides. Conidia hyaline, obovoid to reniform, straight or slightly curved, aseptate, smooth-walled, aggregated into a round at the apex of phialides (adapted from Réblová et al. 2004).
Material examined: New Zealand, South Island, Tasman Province, Nelson Lake National Park, St. Arnaud, Lake Rotoiti, Lakehead track ca. 1.5 km SE of NP Headquarters, on decayed wood of a trunk of Nothofagus sp., 22 February 2003, M. Réblová (PDD-81431, holotype); New Zealand, West Coast Province, Harihari 73 km SW of Hokitika, Saltwater Forest, Poerua River Valley, on decayed wood, 12 March 2003, M. Réblová (PDD-81432).
Known hosts and distribution: On decayed wood of Nothofagus, Populus in Canada (type locality) and New Zealand (Réblová et al. 2004; Réblová 2011).
Notes: Togniniella microspora is sister to Flabellascus tenuirostris (Réblová et al. 2004; Damm et al. 2008; Vu et al. 2019; 100%ML/1.00BY, Fig. 2). Ascomata of 81431 and 81432 are fragile, almost no ascoma with a long intact neck is seen, and most features are evanescent. We provided photo of observable characters and complete the description through Réblová et al. (2004).
Pleurostomataceae Réblová, L. Mostert, W. Gams & Crous, Stud. Mycol. 50(2): 540 (2004)
Saprobic on wood and soil; parasitic in humans. Sexual morph: Ascomata perithecial, gregarious or solitary, immersed or superficial, globose to subglobose, dark brown to black, glabrous, ostiolate, with papilla, periphysate. Peridium outer layer coriaceous, composed of dark brown to pale brown cells of textura intricata to textura epidermoidea to textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses not observed. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with hyaline, ellipsoidal cells, growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci multi-spored, unitunicate, reniform to ellipsoidal, with blunt or rounded, thickened apex. Ascospores overlapping, cylindrical to allantoid, straight to curved, aseptate, hyaline, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Mycelium composed of branched, septate, tuberculate or smooth-walled, hyaline to brown hyphae. Conidiophores usually arising from aerial or submerged hyphae, erect, septate, straight or flexuous, hyaline to brown. Phialides monophialidic or polyphialidic, with inconspicuous or flared collarettes, smooth-walled, hyaline, cylindrical. Conidia hyaline, ovoid to suballantoid, straight or slightly curved, aseptate, smooth-walled, accumulating in slimy masses on the apex of the phialides (adapted from Vijaykrishna et al. 2004).
Type genus: Pleurostoma Tul. & C. Tul. 1863
Notes: Tulasne and Tulasne (1863) established Pleurostoma candollei as the generic type. Vijaykrishna et al. (2004) introduced a second species P. ootheca and its asexual morph Pleurostomophora ootheca. Subsequently, three more species, Pleurostomophora ochraceum (human pathogen), Ple. repens and Ple. richardsiae, were confirmed with molecular data (Vijaykrishna et al. 2004; Najwa et al. 2012). Pleurostomataceae was proposed as an independent family in Calosphaeriales with two genera Pleurostoma and Pleurostomophora based on morphological and molecular analysis (Réblová et al. 2004). Réblová et al. (2015, 2016) recommended that the asexual Pleurostomophora be reduced as a synonym to Pleurostoma based on phylogenetic analysis and to maintain nomenclatural stability. The monotypic nature of Pleurostomataceae is characterized by branched ascogenous hyphae, allantoid ascospores and ovoid conidia, which is similar to Calosphaeriaceae, but the former has short-necked ascomata and reniform asci, and the latter has long-necked ascomata and clavate asci. Pleurostomataceae was revised based on morphology and phylogenetic analysis and it is confirmed as a sister family to Calosphaeriaceae in Calosphaeriales (100%ML/1.00BY, Fig. 2).
Pleurostoma Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 247 (1863)
Saprobic on wood and soil; parasitic in humans. Sexual morph: Ascomata perithecial, gregarious or solitary, at first immersed, then becoming superficial, globose to subglobose, dark brown to black, coriaceous, glabrous, ostiolate, with papilla, periphysate. Peridium comprising three layers, outer layer coriaceous, composed of dark brown to pale brown cells of textura intricata to textura epidermoidea; middle layer coriaceous, composed of brown cells of textura epidermoidea to textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses not observed. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with hyaline, ellipsoidal cells, growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci multi-spored, unitunicate, reniform to ellipsoidal, producing bristle-like pedicel when separating the ascogenous hyphae, with blunt or rounded and thickened apex. Ascospores overlapping, cylindrical to allantoid, straight to curved, aseptate, hyaline, smooth-walled, sometimes with guttules. Asexual morph: Hyphomycetous. Mycelium composed of branched, septate, tuberculate or smooth-walled, hyaline to brown hyphae. Conidiophores usually arising from aerial or submerged hyphae, erect, septate, straight or flexuous, hyaline to brown. Phialides monophialidic or polyphialidic, with inconspicuous or flared collarettes, smooth-walled, hyaline, cylindrical. Conidia hyaline, ovoid to suballantoid, straight or slightly curved, aseptate, smooth-walled, accumulating in slimy masses on the apex of the phialides (adapted from Tulasne and Tulasne 1863; Vijaykrishna et al. 2004).
Notes: Pleurostoma is characterized by glabrous, black ascomata, reniform to ellipsoidal asci with a thickened apex, cylindrical to allantoid ascospores and a phialophora-like asexual morph (Réblová et al. 2004, 2015, 2016; Vijaykrishna et al. 2004; Najwa et al. 2012). Two asexual species, Pleurostoma minimum and P. vibratile, were transferred to Phaeoacremonium based on phylogenetic analysis (Gramaje et al. 2015). Thus, Pleurostoma accommodates five species, three have been reported as asexual (P. repens, P. richardsiae and P. ochraceum), one sexual (P. candollei) and one holomorphic species (P. ootheca) (Tulasne and Tulasne 1863; Vijaykrishna et al. 2004; Najwa et al. 2012; Réblová et al. 2015).
Type species: Pleurostoma candollei Tul. & C. Tul. [as 'candollii'], Select. fung. carpol. (Paris) 2: 247 (1863)
Facesoffungi number: *****??; Fig. 7
Saprobic on decorticated wood. Sexual morph: Ascomata 470–520 × 370–420 µm (x̄ = 500 × 400 µm, n = 10), perithecial, gregarious or solitary, semi-immersed or superficial, globose to subglobose, dark brown to black, coriaceous, glabrous, ostiolate papilla, central, periphysate. Peridium 35–95 µm (x̄ = 60 µm, n = 30) wide, comprising three layers, outer layer coriaceous, composed of dark brown to pale brown cells of textura intricata to textura epidermoidea; middle layer coriaceous, composed of brown cells of textura epidermoidea to textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses not observed. Ascogenous hyphae discrete, hyaline, smooth-walled, branched, with hyaline, ellipsoidal cells, 1.5–4.5 × 1.5–3 µm (x̄ = 2.5 × 2 µm, n = 20), growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci (15–)18–25(–35) × 6.5–11.5 µm (x̄ = 23 × 8.5 µm, n = 50), multi-spored, unitunicate, reniform to ellipsoidal, producing bristle-like pedicel when separating the ascogenous hyphae, with rounded and asymmetrically thickened apex. Ascospores 2.5–4 × 0.5–1.5 µm (x̄ = 3 × 1 µm, n = 50), overlapping, cylindrical to allantoid, straight to curved, aseptate, hyaline, smooth-walled, sometimes with guttules. Asexual morph: Undetermined.
Material examined: France, 92 Hauts-de-Seine, Chaville, on dead wood of Quercus sp., 8 March 1859 (PC-MNHN-PC-PC0167640, holotype); USA, Massachusetts, Medford, on inner surface of bark of standing beech tree, 20 April 1936, G.D. Darker (S-F21567).
Known hosts and distribution: On decayed and decorticated wood of Quercus in France (type locality), North America (Tulasne and Tulasne 1863; Bates et al. 2018).
Notes: Tulasne and Tulasne (1863) examined specimens of rotting bark-less oak collected in Chaville, France in the winter and introduced Pleurostoma candollei. We re-examined the specimen, PC0167640, which was collected on decorticated wood of Quercus at Chaville in March 1859. This sample was named Sphaeria latericollis DC. and Sphaeria spermoides, and finally determined as Pleurostoma candollei (mentioned in the label of material). This information is similar to the history of type material of P. candollei mentioned in Tulasne and Tulasne (1863). Thus, we re-examined the authentic material of PC0167640 and designated that it is the holotype of P. candollei.
Tulasne and Tulasne (1863) also found a kind of villiform, brown to greyish mycelium with branched conidiophores and aggregated, terminal, slender, straight or curved conidia in nature on the same specimen (Fig. 7 r). However, cultural characteristics and phylogenetic analysis have not confirmed the relationship between this hyphomycetous fungus and P. candollei. We could not find any asexual taxon on the materials. The hand-drawn provided for this hyphomycete (Fig. 7 r) is based on Tulasne and Tulasne (1863).
Calosphaeriales genera incertae sedis
Kacosphaeria Speg., Boln Acad. nac. Cienc. Córdoba 11(2): 214 (1887) [1888]
Saprobic on wood. Sexual morph Ascomata perithecial, aggregated, discoid, immersed to erumpent through the bark of the host, black, glabrous, thick-walled, with short necks. Asci 8-spored, unitunicate, clavate, long pedicellate, apex obtuse, with long paraphyses. Ascospores bi-seriate, allantoid, 1-septate, hyaline. Asexual morph Undetermined (adapted from Spegazzini 1888).
Notes: The monotypic Kacosphaeria with clavate asci and allantoid ascospores that is similar to some species of Calosphaeria (Spegazzini 1888). The specific ascogenous hyphae of Calosphaeriales is not mentioned in the description (Spegazzini 1888), nor is it found in the drawings of Spegazzini (Arambarri et al. 2020). Thus, we propose that Kacosphaeria be placed in Calosphaeriales genera incertae sedis, and await more fresh samples pending.
Type species: Kacosphaeria antarctica Speg., Boln Acad. nac. Cienc. Córdoba 11(2): 214 (1887) [1888]
Fig. 4 c–e
Saprobic on Ribisma gellanicae. Sexual morph Ascomata 280–320 µm, perithecial, 4–9 ascomata aggregated, discoid, immersed to erumpent through the bark of the host, black, glabrous, with short necks. Asci 40–50 × 4–5 µm, 8-spored, unitunicate, clavate, long pedicellate, obtuse apex, with long paraphyses. Ascospores 9–11 × 1.5 µm, bi-seriate, allantoid, 1-septate, hyaline. Asexual morph Undetermined (adapted from Spegazzini 1888).
Known hosts and distribution: On decayed wood of Ribisma gellanicae in Argentina (type locality) (Spegazzini 1888).
Notes: Kacosphaeria antarctica was found on dead branches of Ribisma gellanicae in Ushuaia, Argentina, in June 1882 (Spegazzini 1888). We were unable to obtain authentic material of K. antarctica, but provide a drawing (Fig. 4 c–e) based on Spegazzini's drawings from Arambarri et al. (2020).
Jobellisiales M.J. D'souza & K.D. Hyde 2015
Réblová (2008) and Liu et al. (2012) showed that Jobellisia species were closely related to members of Calosphaeriales, Diaporthales and Togniniaceae based on LSU sequence analysis. Maharachchikumbura et al. (2015) established the monotypic order Jobellisiales in Diaporthomycetidae based on the multi-gene analysis. The phylogenetic position of this order is unstable and was previously thought to be related to Calosphaeriales and Togniniales (Hongsanan et al. 2017; Hyde et al. 2017). We find that Jobellisiales is basal to Calosphaeriales, Togniniales, Diaporthales and Tirisporellales (Fig. 1). Jobellisiales is different from Calosphaeriales and Togniniales with clavate asci and hyaline allantoid ascospores, and different from Tirisporellales with cylindrical asci and fusoid, falcate to lunate ascospores (Hyde et al. 2020). It is similar to Diaporthales, in having cylindrical to ellipsoidal asci and brown, subglobose ascospores; however, they are distantly related in phylogenetic analyses (Hyde et al. 2020; this study, Fig. 1).
Jobellisiaceae Réblová, Mycologia 100(6): 899 (2008)
Saprobic on wood. Sexual morph: Ascostromata crustose. Ascomata perithecial, gregarious, semi-immersed to superficial, globose to subglobose, black to dark brown, rough, ostiolate, with papilla, periphysate. Peridium membranaceous, composed of blue-green or brown or yellow to hyaline cells of textura angularis or textura globosa to textura prismatica. Paraphyses numerous, filiform, septate, unbranched, evanescent. Asci unitunicate, cylindrical, long pedicellate, apex obtuse, with J- apical ring. Ascospores 8-spored, ellipsoidal to subglobose, pale brown to dark brown, 0–1-septate, smooth-walled, with guttules and a germ pore at one or each end, without gelatinous appendages. Asexual morph: Undetermined (adapted from Barr 1993; Huhndorf et al. 1999a).
Type genus: Jobellisia M.E. Barr 1993
Notes: Jobellisia was established based on Letendraea luteola as a member of Clypeosphaeriaceae (Barr 1993). Subsequently, other species of Jobellisia which have been found in aquatic and/or terrestrial habitats in Asia, Europe and the USA, were reported as resembling J. luteola, in having ascomata with necks and brown, 1-septate ascospores with germ pore(s) (Barr 1994; Huhndorf et al. 1999a; Ranghoo et al. 2001; Leroy 2006; Untereiner et al. 2013). Huhndorf et al. (2004b) analyzed LSU sequence data for J. fraterna and J. luteola and found that they located in Diaporthales. However, Réblová (2008) transferred J. rhynchostoma to Bellojisia (Lasiosphaeriaceae) and established Jobellisiaceae as an independent family in Sordariomycetes based on LSU sequence data. Liu et al. (2012) introduced J. guangdongensis and found that Jobellisiaceae is basal to Calosphaeriales and Diaporthales. Maharachchikumbura et al. (2015, 2016) introduced the monotypic Jobellisiales, which is closely related to Calosphaeriales based on the multi-gene analysis. Hyde et al. (2017) found that Jobellisiales is sister to Togniniales with a divergence of ca. 130–188 MYA in the MCC tree. Hongsanan et al. (2017) and Wijayawardene et al. (2018) suggested that Jobellisiales should be reduced as a family in Calosphaeriales. Hyde et al. (2020) and Wijayawardene et al. (2020) listed Jobellisiales in Hypocreomycetidae. However, species of Jobellisiales have cylindrical asci with rounded apices and ellipsoidal, brown, 1-septate ascospores similar to those of Diaporthales (Maharachchikumbura et al. 2016; Hyde et al. 2020). The clade characterising Jobellisiales is phylogenetically distinct from other orders of Diaporthomycetidae (Fig. 1).
Jobellisia M.E. Barr, Mycotaxon 46: 60 (1993)
Saprobic on wood. Sexual morph: Ascomata perithecial, gregarious, semi-immersed to superficial, obpyriform to subglobose, collapsing when dry or not, blue-green, yellow, brown to black, rough, ostiolate, with beaked to conical necks, periphysate. Peridium membranaceous, comprising three layers, outer layer composed of brown or blue-green or orange cells of textura angularis-globosa; middle layer composed of brown to yellow cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, filiform, septate, unbranched, evanescent. Asci unitunicate, cylindrical, long pedicellate, apex obtuse, with J- apical ring. Ascospores 8-spored, ellipsoidal to subglobose, pale brown to dark brown, straight to slightly curved, 0–1-septate, smooth-walled, with guttules and a germ pore at one or each end, without gelatinous appendages. Asexual morph: Undetermined (adapted from Barr 1993; Huhndorf et al. 1999a).
Notes: Barr (1993) established Jobellisia based on Letendraea luteola. Eight species have been accommodated in Jobellisia; however, only three (Jobellisia fraterna SMH2863, J. guangdongensis HMAS 251240 and J. luteola SMH2753) have molecular data (Huhndorf et al. 1999a; Réblová 2008; Liu et al. 2012). More phylogenetic information is needed to determine their taxonomic placement.
Type species: Jobellisia luteola (Ellis & Everh.) M.E. Barr, Mycotaxon 46: 61 (1993)
Basionym: Letendraea luteola Ellis & Everh., Proc. Acad. nat. Sci. Philad. 47: 415 (1895)
Facesoffungi number: *****??; Fig. 8
Saprobic on wood. Sexual morph: Ascostromata crustose. Ascomata 460–570 × 515–620 µm (x̄ = 500 × 560 µm, n = 10), perithecial, gregarious, superficial, obpyriform to subglobose, collapsing when dry, black to dark brown, rough, ostiolate, with conical necks, periphysate. Peridium 40–75 µm (x̄ = 55 µm, n = 30) wide, membranaceous, comprising three layers, outer layer composed of brown to orange cells of textura angularis-globosa; middle layer composed of brown to pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, filiform, septate, unbranched, evanescent. Asci unitunicate, cylindrical, long pedicellate, apex obtuse, with J- apical ring. Ascospores 12–14 × 4.5–5.5 µm (x̄ = 13.5 × 5 µm, n = 50), 8-spored, ellipsoidal to subglobose, pale brown to dark brown, straight to slightly curved, 1-septate, smooth-walled, with guttules and a germ pore at each end, without gelatinous appendages. Asexual morph: Undetermined (adapted from Barr 1993; Huhndorf et al. 1999a).
Material examined: USA, Michigan, Marquette, Huron Mountain Club, end of road at Mountain Lake, 45º00’ N / 87°0’ W, on dead wood over the water, 17 August 1997, S.M. Huhndorf (F-SMH 3349).
Known hosts and distribution: On decayed wood in Ohio, USA (type locality) (Ellis and Everhart 1895); on decayed decorticated wood in Iowa, Michigan, North Carolina, New York and Ontario, USA (Barr 1993; Huhndorf et al. 1999a).
Notes: Jobellisia luteola has been collected in temperate regions (Barr 1993; Huhndorf et al. 1999a). Maharachchikumbura et al. (2016) re-examined the holotype (NY00914363) collected by J.P. Morgan in Ohio, USA. Huhndorf et al. (1999a) also introduced two new specimens of J. luteola, SMH 2753 and SMH 3349. Subsequently, Huhndorf et al. (2004b) analyzed LSU sequence data for SMH 2753 which is widely used to represent J. luteola. In this study, the authentic material of SMH 3349 was re-examined.
Other genera in Diaporthomycetidae
Ophiostomatales Benny & Kimbr.
Ophiostomataceae Nannf.
Fragosphaeria Shear, Mycologia 15(3): 124 (1923)
Saprobic on wood. Sexual morph: Ascomata solitary, superficial, carbonaceous or coriaceous, globose, black, glabrous or surrounded by hyaline hyphae. Asci 8-spored, unitunicate, globose. Ascospores ovoid to reniform, aseptate, guttulate, smooth-walled. Asexual morph: Hyphomycetous. Conidiophores hyaline, branched. Conidia hyaline, aseptate, suballantoid to ellipsoidal (adapted from Saccardo 1881; Shear 1923).
Notes: Fragosphaeria is typified by F. purpurea and has cleistothecial ascomata, globose asci and reniform ascospores (Shear 1923). Chesters (1935) transferred this genus to Cephalotheca based on similar morphologies. In this study, we find that the sexual characteristics of Fragosphaeria are similar to members of Eurotiales (Eurotiomycetes), but the asexual morph is unknown. However, Suh and Blackwell (1999) found that F. purpurea is closely related to Ophiostoma (Ophiostomatales) based on LSU and SSU sequence analyses. Hence, Fragosphaeria is accepted as a genus in Ophiostomataceae (Diaporthomycetidae) (Hyde et al. 2020).
Type species: Fragosphaeria purpurea Shear, Mycologia 15(3): 124 (1923)
Facesoffungi number: *****??; Fig. 9
Saprobic on wood. Sexual morph: Ascomata 90–130 µm (x̄ = 120 µm, n = 10) diam., cleistothecial, solitary, superficial, carbonaceous, globose, dark brown to black. Peridium brittle, composed of carbonaceous, brown to reddish brown cells of textura prismatica, squashed fragments pentagonal. Asci 4–6 µm (x̄ = 5 µm, n = 20) diam., 8-spored, unitunicate, globose. Ascospores 2.5–3 × 1.5–2.8 µm (x̄ = 2.8 × 2 µm, n = 20), overlapping, reniform, aseptate, hyaline when young, becoming pale brown to olivaceous when mature, guttulate, smooth-walled. Asexual morph: Hyphomycetous. Conidiophores hyaline, branched. Conidia hyaline, aseptate, suballantoid to ellipsoidal (adapted from Shear 1923).
Material examined: USA, Virginia, on culture of Pilacre sp., July 1920, C.L. Shear (BPI-566225, type); Sweden, Göteborg, Nature Park of Botanical Garden, on dead deciduous wood, 15 October 1970, U. Eliasson (BPI-566226).
Known hosts and distribution: On culture of Pilacre petersii in USA (type locality) (Shear 1923); on dead deciduous wood in UK, Sweden (Eliasson 1971; Yaguchi et al. 2006).
Notes: Shear (1923) introduced Fragosphaeria purpurea, which was found in the culture of Pilacre petersii in Virginia, September 1920, as the generic type. In this study, we re-examined type material (566225), which was isolated from a culture of Pilacre collected in Virginia, July 1920. This specimen is a dry fragmented culture, and we were unable to find any structure for observation. We also obtained authentic material, 566226 collected and determined by Eliasson (1971).
Genera transferred to Hypocreomycetidae
Coronophorales genera incertae sedis
Tengiomyces Réblová, Mycotaxon 70: 408 (1999)
Saprobic on wood. Sexual morph: Ascomata perithecial, superficial, solitary to gregarious, globose to subglobose, coriaceous, dark brown to black, surrounded by setae, seated on sparse, brown, septate, branched hyphae, ostiolate, with papilla. Setae brown to dark brown, spine-like, straight, acute, aseptate, opaque, unbranched. Peridium membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses septate, evanescent. Asci 8-spored, unitunicate, cylindrical to clavate, broadly rounded to truncate and thickened at the apex, pedicellate. Ascospores ellipsoidal to clavate, with broadly rounded ends, straight to slightly curved, septate, middle cell(s) pale brown to brown, end cell(s) hyaline to subhyaline, slightly constricted at the septa, smooth-walled. Asexual morph (associated): Hyphomycetous, spadicoides-like. Mycelium semi-immersed to superficial. Conidiophores arising from superficial hyphae growing from the base of the perithecial initials and mature ascomata, macronermatous, mononematous, unbranched or branched in the lower parts, cylindrical, pale brown, straight, flexuous in the lower parts, septate, not constricted or slightly constricted at the septa, apical cells rounded. Conidiogenous cells polytretic, integrated, intercalary and terminal. Conidia obpyriform to clavate, broadly rounded at the apex, septate, not constricted or slightly constricted at the septa, brown or the pigment fading from apex to bottom (adapted from Réblová 1999a).
Notes: Réblová (1999a) transferred Chaetosphaerella indica to Tengiomyces and established a monotypic genus. The black ascomata with setae on T. indicus are similar to Helminthosphaeria and Chaetosphaerella, and the ascospores comprise two brown middle cells and one hyaline cell at each end and are similar to Crassochaeta and Chaetosphaerella (Chaetosphaerellaceae, Coronophorales) (Réblová 1999a; Huhndorf et al. 2004a). A hyphomycetous spadicoides-like form was associated with T. indicus on the natural substratum (Réblová 1999a). Spadicoides is polyphyletic in Pleosporales (Dothideomycetes) (Ma et al. 2016; Réblová et al. 2018). However, we could not determine the relationship between Tengiomyces and Spadicoides without appropriate cultural characteristics and phylogenetic analysis. In this study, Tengiomyces with black, bristly ascomata and ellipsoidal ascospores comprise brown and hyaline cells is similar to Chaetosphaerella rather than Helminthosphaeria. Thus, we place Tengiomyces in Coronophorales genera incertae sedis pending further studies with molecular data.
Type species: Tengiomyces indicus (Varghese & V.G. Rao) Réblová, Mycotaxon 70: 408 (1999)
Basionym: Chaetosphaerella indica Varghese & V.G. Rao, Biovigyanam 5(1): 2 (1979)
Facesoffungi number: ??; Fig. 10
Saprobic on wood. Sexual morph: Ascomata 200–250 µm (x̄ = 230 µm, n = 5) diam., perithecial, superficial, solitary to gregarious, globose to subglobose, coriaceous, dark brown to black, surrounded by setae, seated on sparse, brown, septate, branched hyphae 3–8 µm wide, ostiolate, with papilla. Setae 3.5–11 µm wide, brown to dark brown, spine-like, straight, acute, aseptate, opaque, unbranched. Peridium 20–26 µm wide, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses septate, evanescent. Asci 60–70 × 10–15 µm, 8-spored, unitunicate, cylindrical to clavate, broadly rounded to truncate and thickened at the apex, with short pedicel. Ascospores 10–15 × 4–5.5 µm (x̄ = 13 × 5 µm, n = 20), bi-seriate, ellipsoidal to clavate, with broadly rounded ends, straight to slightly curved, 3-septate, two middle cells pale brown to brown, two end cells hyaline to subhyaline, slightly constricted at the septa, smooth-walled. Asexual morph (associated): Hyphomycetous, spadicoides-like. Mycelium semi-immersed to superficial. Conidiophores 3.5–8 µm wide, arising from superficial hyphae growing from the base of the perithecial initials and mature ascomata, macronematous, mononematous, unbranched or branched in the lower parts, cylindrical, pale brown, straight, flexuous in the lower parts, septate, not constricted or slightly constricted at the septa, apical cells rounded. Conidiogenous cells 4–6 × 5–6 µm, polytretic, integrated, intercalary and terminal. Conidia 14–18 × 5.5–10 µm (x̄ = 15 × 7.5 µm, n = 20), obpyriform to clavate, broadly rounded at the apex, 2–3-septate, not constricted or slightly constricted at the septa, brown or the pigment fades from apex to bottom (adapted from Réblová 1999a).
Material examined: China, AnHui, LangYa Shan, on dead wood of a deciduous tree, 20 June 1933, S.Q. Deng (BPI-622100); Guiana, Saint Laurent, Piste Balate, 12km from Saint Laurent, 20 m elev, on dead branch, A. Rossman, C. Feuillet and L. Skog, 19 November 1986 (BPI-622098).
Known hosts and distribution: On dead stems of a dicotyledoneous plant in India (type locality) (Varghese and Rao 1979); on dead wood of a deciduous tree in China (Réblová 1999a); on dead branch in Guiana.
Notes: Réblová (1999a) introduced Tengiomyces indicus after examining the holotype of Chaetosphaerella indica (AMH 3871), collected by Varghese in India (Varghese and Rao 1979) and a sample BPI 622100 collected by Deng in China. We re-examined two authentic specimens (622100 and 622098) which were determined by Réblová (mentioned in the label of material) and found that they were dry and observed only ascomata and evanescent asci and ascospores. We also found a spadicoides-like taxon by T. indicus on the natural substrate (Fig. 10 h–i).
Hypocreales genera incertae sedis
Sulcatistroma A.W. Ramaley, Mycotaxon 93: 140 (2005)
Saprobic on leaves. Sexual morph: Ascostromata discrete, multiloculate, erumpent through leaf epidermis, ellipsoidal, rarely circular. Ascomata perithecial, immersed, formed in stroma, subglobose, dark brown, ostiole with hyaline periphyses. Peridium membranaceous, composed of hyaline to brown cells of textura angularis. Paraphyses numerous, septate, slightly constricted at the septa, unbranched, tapering to a rounded apex. Asci 8-spored, unitunicate, clavate, short pedicellate. Ascospores bi-seriate, hyaline, allantoid with rounded ends, aseptate, smooth-walled. Asexual morph: Hyphomycetous. Conidiophores arising from aerial hyphae, erect, inflated at the base. Phialides monophialidic, smooth-walled, hyaline, cylindrical, unbranched or with sympodial conidiogenous cells. Conidia hyaline to pale orange, ellipsoidal to allantoid, aseptate, smooth-walled, with rounded ends (adapted from Ramaley 2005).
Notes: The monotypic genus Sulcatistroma has ellipsoidal ascostromata, clavate asci, allantoid ascospores and ellipsoidal to allantoid conidia (Ramaley 2005). This genus is similar to the sexual morph of Diatrype (Xylariales), differing from Sulcatistroma in having sympodial conidiogenous cells, whereas Diatrype has percurrently proliferating conidiogenous cells (Ramaley 2005). Initially, Sulcatistroma was accepted as a member of Calosphaeriales based on its spicate clusters of asci (Ramaley 2005), and later it was included in Calosphaeriaceae (Lumbsch and Huhndorf 2010; Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2018). Vu et al. (2019) analyzed ITS sequence data for ex-type strain Sulcatistroma nolinae (CBS 117709). This strain is more closely related to members of Nectriaceae (Hypocreales) (Fig. 1). Sulcatistroma has similar features to Nectriaceae species, such as clavate asci, aseptate, allantoid ascospores and phialophora-like asexual morphs, but does not have the red ascomata which is characteristic of Nectriaceae (Hirooka et al. 2012). We place Sulcatistroma in Hypocreales genera incertae sedis pending phylogenetic data.
Type species: Sulcatistroma nolinae A.W. Ramaley, Mycotaxon 93: 140 (2005)
Facesoffungi number: ??; Fig. 11
Saprobic on leaves. Sexual morph: Ascostromata discrete, multiloculate, erumpent through leaf epidermis, ellipsoidal, rarely circular, with long axis parallel to length of leaf, 900–1500 × 350–850 µm (x̄ = 1200 × 450 µm, n = 5). Ascomata 120–250 µm diam., perithecial, immersed, subglobose, brown to dark brown, ostiole with hyaline periphyses. Peridium 7–25 µm (x̄ = 16.5 µm, n = 20) wide, membranaceous, composed of hyaline to pale brown cells of textura angularis. Paraphyses numerous, septate, slightly constricted at the septa, unbranched, tapering to the rounded apex. Asci 8-spored, unitunicate, clavate, short pedicellate, apical ring not apparent. Ascospores bi-seriate, hyaline, allantoid with rounded ends, aseptate, smooth-walled. Asexual morph: Hyphomycetous, phialophora-like. Conidiophores arising from aerial hyphae, erect, inflated at the base. Phialides monophialidic, smooth-walled, hyaline, cylindrical, unbranched or an axis with 1–4 sympodial conidiogenous cells. Conidia hyaline to pale orange, ellipsoidal to allantoid, aseptate, smooth-walled, with rounded ends (adapted from Ramaley 2005).
Material examined: USA, New Mexico, Lincoln Co., Valley of Fires, roadside, on leaves of Nolina micrantha, A.W. Ramaley, 9 October 2002 (BPI-864276, holotype).
Known hosts and distribution: On decayed leaves of Nolina micrantha in the USA (type locality) (Ramaley 2005).
Notes: We re-examined the holotype (864276), but the ascomata contained only some filament-like supporting structures (Fig. 11 e–g). Therefore, we provided hand drawings of other characteristics (Fig. 11 a, h–i) based on Ramaley (2005).
Sordariomycetidae O.E. Erikss. & Winka
The subclass Sordariomycetidae was introduced by Eriksson and Winka (1997) and previously comprised six orders. Eight orders with 23 families are now included based on phylogenetic and morphological studies (Maharachchikumbura et al. 2015; Hongsanan et al. 2017; Hyde et al. 2017, 2020; Marin-Felix et al. 2020; Wijayawardene et al. 2020). Sordariomycetidae members are widespread and usually as saprobes, pathogens or endophytes (Maharachchikumbura et al. 2015, 2016; Hyde et al. 2020). Fig. 1 reveals that this subclass is sister to Diaporthomycetidae (68%ML) in Sordariomycetes.
Boliniales P.F. Cannon, in Kirk et al., Dictionary of Fungi 9 (2001)
The saprotrophic Boliniales was introduced by Kirk et al. (2001) and is characterized by black ascomata, cylindrical asci and fusiform to subglobose ascospores (Maharachchikumbura et al. 2015, 2016; Hyde et al. 2020). Some members of this order with carbonaceous ascomata were traditionally linked to Xylariales, e.g. Camarops (Karsten 1873). Boliniales was clarified by Maharachchikumbura et al. (2015) and currently contains nine genera based on morphology and phylogenetic analyses (Hyde et al. 2020; Wijayawardene et al. 2020). This order is sister to Phyllachorales (Maharachchikumbura et al. 2016; Hongsanan et al. 2017; Hyde et al. 2017; this study, 83%ML, Fig. 1).
Boliniaceae Rick, Brotéria, sér. bot. 25(2-3): 65 (1931)
Saprobic on wood in terrestrial habitats. Sexual morph: Ascostromata immersed, erumpent or superficial, pale brown to black, irregular in shape, surface furfuraceous or surrounded by hyaline to brown hyphae. Ascomata perithecial, solitary or gregarious, immersed, erumpent or superficial, globose to obpyriform, brown to black, papillate. Peridium carbonaceous or coriaceous or membranaceous, composed of brown to dark brown cells of textura intricata or textura prismatica. Paraphyses abundant, filamentous, tapering, septate, branched. Asci 8-spored, unitunicate, cylindrical to clavate, apex rounded to blunt, pedicellate. Ascospores 2–3-seriate, normally cells in the center brown to dark brown, and the terminal cell(s) hyaline to pale brown, subglobose to ellipsoidal, 0–2-septate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Rick 1931; Romero and Samuels 1991; Maharachchikumbura et al. 2016).
Type genus: Camarops P. Karst. 1873 (= Bolinia (Nitschke) Sacc. 1882)
Notes: Boliniaceae, introduced by Rick (1931), is characterized by black ascostromata and cylindrical asci with smooth-walled ascospores. Earlier, this family was placed in Xylariales based on morphological similarities, such as carbonaceous ascomata (Barr 1990; Romero and Samuels 1991). Andersson et al. (1995) relied on analyzing SSU sequence data and concluded that Boliniaceae was more closely related to Sordariales. This family was subsequently used to establish Boliniales by Kirk et al. (2001). Thus, Boliniaceae, based on Bolinia is typified by B. tubulina, is the only member of Boliniales (Maharachchikumbura et al. 2016; Kirk et al. 2001). Nannfeldt (1972) recommended Bolinia should be synonymized as Camarops which is typified by Camarops hypoxyloides (Karsten 1873) as they have similar characters. Phylogenetically, Boliniales is closely related to Phyllachorales and Sordariales in Sordariomycetidae (Huhndorf et al. 2004b; Miller and Huhndorf 2005; Hongsanan et al. 2017; Hyde et al. 2017; this study, Fig. 1). However, since both Boliniales and Boliniaceae were established based on Bolinia tubulina, even if B. tubulina was transferred to Camarops tubulina, this species is still the type of its order and family (Maharachchikumbura et al. 2016). We accept Camarops tubulina as type species of Boliniales and Boliniaceae, and Camarops hypoxyloides as the type of Camarops, as established (Karsten 1873; Rick 1931; Kirk et al. 2001); and more characteristics and phylogenetic information are expected to determine the affinities of Camarops and other genera in the Boliniaceae. Nine genera are accepted into this family, six of which have available molecular data (viz. Apiorhynchostoma, Camaropella, Camarops, Cornipulvina, Endoxyla, Pseudovalsaria) (Huhndorf and Miller 2008; Raja et al. 2011; Untereiner et al. 2013; Maharachchikumbura et al. 2015, 2016; Vu et al. 2019; Hyde et al. 2020; Wijayawardene et al. 2020), and Apiocamarops, Mollicamarops, Neohypodiscus were included because of their morphology similar to Camarops (Andersson et al. 1995; Rogers and Ju 2003; Vasilyeva 2007). In this study, we find that some species of ascospores are not arranged in a uniform direction in the ascus. For example, in Apiocamarops, ascospore is composed of a large brown cell and a small hyaline cell (Samuels and Rogers 1987). Ascospores partly are oriented with the hyaline cell to the ascus apex; and partly in the opposite direction to the ascus base.
Apiocamarops Samuels & J.D. Rogers, Mycotaxon 28(1): 54 (1987)
Saprobic on wood. Sexual morph: Ascostromata superficial or semi-immersed, brown to black, irregular in shape, surface furfuraceous or glabrous. Ascomata perithecial, gregarious, superficial or semi-immersed, globose to subglobose, membranaceous, orange, brown to black, ostiolate, with periphyses. Peridium membranaceous, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- apical ring. Ascospores uniseriate, 0–1-septate, unequal, oblong to ellipsoidal, smooth-walled, hyaline when young, composed of a large, brown cell with or without a terminal pore and a small, hyaline cell when mature, guttulate. Asexual morph: Undetermined (adapted from Samuels and Rogers 1987; Rogers 1988; Reagan and Waide 1996; Rogers and Ju 2003).
Notes: Apiocamarops was established by Samuels and Rogers (1987) and resembles Camarops. Both genera have similar cylindrical asci and ellipsoidal ascospores with germ pore(s), but the ascospores in Apiocamarops are 1-septate and apicosporous, while those of Camarops lack septum (Samuels and Rogers 1987; Rogers 1988; Reagan and Waide 1996; Rogers and Ju 2003). Four species are accommodated in this genus, but molecular data is lacking.
Type species: Apiocamarops alba Samuels & J.D. Rogers, Mycotaxon 28(1): 54 (1987)
Facesoffungi number: *****??; Fig. 13
Saprobic on decorticated wood. Sexual morph: Ascostromata superficial, brown to black, irregular in shape, surface furfuraceous. Ascomata 450–750 × 350–700 µm (x̄ = 550 × 450 µm, n = 10), perithecial, gregarious, superficial, globose to subglobose, brown to black, papillate, ostiole central, with periphyses, covered by white to yellowish, furfuraceous layer. Peridium 35–60 µm (x̄ = 45 µm, n = 30) wide, membranaceous, comprising two layers, outer layer composed of dark brown to pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 1–3 µm wide, numerous, filiform, septate, branched, longer than the asci. Asci 75–95(–110) × 4–6 µm (x̄ = 90 × 5 µm, n = 50), 8-spored, unitunicate, cylindrical, pedicel short, apex rounded, with J- inconspicuous apical ring. Ascospores (4.5–)5.5–6.5(–8) × 1.5–3 µm (x̄ = 6 × 2.5 µm, n = 50), uniseriate, oblong to ellipsoidal, slightly curved, 1-septate, unequal, hyaline when young, composed of a large, brown cell with a minute terminal pore and a small, hyaline cell when mature, partly hyaline cell towards the ascus apex; partly in the opposite direction, smooth-walled, guttulate. Asexual morph: Undetermined.
Material examined: Brazil, Amazonas, Plateau of Serra Araca, N side of North Mountain, alt. 1250 m., 00° 57' N, 63° 21' W, cloud forest, on decorticated wood, 19–24 February 1984, G.J. Samuels, G.T. Prance and J. Pipoly (NY-00986029, holotype); ibid. (NY-00986030, paratype).
Known hosts and distribution: On decorticated wood in Brazil (type locality), Guiana, Venezuela (Samuels and Rogers 1987).
Notes: As noted by Samuels and Rogers (1987), we re-examined the type materials, 00986029 and 00986030, which are well preserved.
Apiorhynchostoma Petr., Annls mycol. 21(3/4): 185 (1923)
Saprobic on wood. Sexual morph: Ascomata perithecial, scattered, immersed to semi-immersed, globose to subglobose, dark brown to black, papillate, ostiole central, with periphyses. Peridium membranaceous, comprising two layers, outer layer composed of brown to pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci 8-spored, unitunicate, cylindrical, pedicellate, with J- apical ring. Ascospores uniseriate, oblong to ellipsoidal, straight to slightly curved, 0–2-septate, hyaline when young, becoming composed of 1 or 2 large, brown cell(s) with a minute pore at the end, other end with a small, hyaline to pale brown cell, smooth-walled or verrucose, guttulate. Asexual morph: Undetermined (adapted from Petrak 1923).
Notes: Currey (1859) introduced two species, named ‘Sphaeria apiculata’ and ‘Sphaeria curreyi Blox. MSS.’. Rabenhorst (1860) examined some materials and stated that Sphaeria apiculata Curr. is illegitimate as it has a homonym of Sphaeria apiculata Wallr. (Wallroth 1833). Thus, Rabenhorst (1860) re-named S. apiculata Curr. as Sphaeria curreyi Rabenh. which is a homonym of S. curreyi Blox. MSS. Due to the illegitimate species naming of ‘Sphaeria apiculata’ and ‘Sphaeria curreyi’, subsequent research has been controversial. Petrak (1923) established Apiorhynchostoma and named S. apiculata Curr. as Apiorhynchostoma apiculatum as the type species. Winter (1886) and Müller and von Arx (1962) recognized the former species related to ‘Sphaeria curreyi’ and transferred S. curreyi Rabenh. as Apiorhynchostoma curreyi as the type species of Apiorhynchostoma (Rogers et al. 1994a; Hyde et al. 1998; Stadler et al. 2013; Untereiner et al. 2013). In our study, we proposed a new epithet, Apiorhynchostoma apiosporum, based on isotype material of Sphaeria curreyi Rabenh. to avoid future confusion. Therefore, the illegitimate A. apiculatum (≡ S. apiculata Curr.) and A. curreyi (≡ S. curreyi Rabenh.) are replaced by A. apiosporum. Apiorhynchostoma is characterized by subglobose ascomata and ascospores which are composed of two brown cells and a hyaline terminal cell (Müller and von Arx 1962; Rappaz 1995; Untereiner et al. 2013). This genus has been included in Amphisphaeriaceae and Clypeosphaeriaceae based on their similar ascomata and ascospores (Hyde et al. 1998; Réblová 1998; Wang et al. 2004). Untereiner et al. (2013) analyzed ITS-LSU sequence data for ‘A. curreyi’ and showed that Apiorhynchostoma is closely related to Cornipulvina and Pseudovalsaria in Boliniales. In this study, the strain name of A. curreyi is replaced by Apiorhynchostoma apiosporum and it is sister to Pseudovalsaria ferruginea in Boliniales (72%ML, Fig. 12).
Type species: Apiorhynchostoma apiosporum S.K. Huang & K.D.Hyde, nom. nov.
Synonym: Sphaeria apiculata Curr., Trans. Linn. Soc. London 22: 326 (1859)
Sphaeria curreyi Rabenh., Fungi europ. exsicc. Klotzschii herbarii vivi mycologici continuatio, Edn nova. Series secunda, Cent. 3: no. 250 (1860)
Apiorhynchostoma apiculatum (Sacc.) Petr. [as 'apiculata'], Annls mycol. 21(3/4): 185 (1923)
Apiorhynchostoma curreyi E. Müll., in Müller & von Arx, Beitr. Kryptfl. Schweiz 11(no. 2): 706 (1962)
Facesoffungi number: *****??; Index Fungorum number: IF558201; Fig. 14
Etymology: apiosporum refers to the ellipsoidal ascospores with one hyaline cell markedly smaller than the other.
Saprobic on decorticated wood. Sexual morph: Ascomata 530–660 × 470–580 µm (x̄ = 590 × 530 µm, n = 10), perithecial, scattered, immersed to semi-immersed, globose to subglobose, dark brown to black, papillate, ostiole central, with periphyses. Peridium 15–65 µm (x̄ = 35 µm, n = 30) wide, membranaceous, comprising two layers, outer layer composed of brown to pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 2–4.5 µm wide, numerous, filiform, septate, longer than asci. Asci 190–200(–250) × 7.5–13 µm (x̄ = 198 × 10 µm, n = 50), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- inconspicuous apical ring. Ascospores 23.5–30 × 5.5–9 µm (x̄ = 25 × 7.5 µm, n = 50), uniseriate, oblong to ellipsoidal, straight to slightly curved, 1–2-septate, unequal, hyaline when young, becoming composed of 1 or 2 large, brown cell(s) with a minute pore and a hyaline, thick-walled at the end, other end with a small, hyaline to pale brown cell, smooth-walled, guttulate. Asexual morph: Undetermined.
Material examined: UK, Batheaston, on dead decorticated wood, F. Currey, Rabenhorst Fungi Europaei Exsiccati nr. 250 (S-F61087, lectotype).
Known hosts and distribution: on decayed decorticated wood of conifers in Austria, Canada, Germany, Sweden, UK (type locality) (Currey 1859; Untereiner et al. 2013).
Notes: Currey (1859) described samples as being collected on dry old wood of the South Western Railway, not far from the Weybridge Station. Rabenhorst (1860) mentioned that Currey collected material of Sphaeria curreyi Rabenh. in Batheaston in 1856 and 1857, and this material was determined as the isotype of S. curreyi Rabenh. (Untereiner et al. 2013). As noted in S-F61087, this material was collected by Currey in Batheaston and determined by Rabenhorst. Thus, we proposed this material as lectotype of Apiorhynchostoma apiosporum.
Camaropella Lar.N. Vassiljeva, Mikol. Fitopatol. 31(1): 6 (1997)
Saprobic on wood. Sexual morph: Ascostromata erect, immersed to semi-immersed, erumpent through bark of host, carbonaceous, black, glabrous, pulvinate, peltate, discoid or valsoid, hemispherical or spherical, sometimes with broad attachment to the substrate. Ascomata perithecial, gregarious, carbonaceous, cylindrical, black, ostiolate, with necks, periphysate. Asci 8-spored, unitunicate, cylindrical, apex rounded, pedicellate, with J- apical ring. Ascospores uniseriate, cylindrical to ellipsoidal, aseptate, pale brown or dark brown, smooth-walled, with guttules. Asexual morph: Undetermined (adapted from Vasilyeva 1997; Vasilyeva et al. 2007).
Notes: Vasilyeva (1997) transferred Camarops pugillus (≡ Sphaeria pugillus Schw.) to Camaropella as generic type. Camaropella species do not have a lopadostomoid type of stromata as found in Camarops, but has special eutypoid and eutypelloid or valsoid forms (Vasilyeva 1997). She also introduced a second species, Camaropella lutea (Vasilyeva et al. 2007). These two species are nested in Boliniales and distant from Camarops tubulina based on phylogenetic analysis (Huhndorf and Miller 2008; Untereiner et al. 2013; Vu et al. 2019). In this study, strains of Camarops amorpha, C. microspora and C. plana, which have pulvinate or peltate stromata (Miller and Huhndorf 2005; Spatafora et al. 2006; Raja et al. 2011), clustered with Camaropella species with 89%ML/0.93BY support (Fig. 12), and this clade is distant from Camarops clade (Untereiner et al. 2013; this study). Therefore, we recommend transferring these three Camarops species to Camaropella. However, Camaropella and Camarops are very similar in morphology, in the cylindrical asci, ellipsoidal ascospores and the arrangement type of the ascomata in stroma. The relationship between Camarops and Camaropella needs to be resolved with more collections and sequence data.
Type species: Camaropella pugillus (Schwein.) Lar.N. Vassiljeva, Mikol. Fitopatol. 31(1): 6 (1997)
Basionym: Sphaeria pugillus Schwein., Schr. naturf. Ges. Leipzig 1: 38 [12 of repr.] (1822)
Facesoffungi number: *****??; Fig. 15
Saprobic on wood. Sexual morph: Ascostromata 2–3 × 1.5–3 mm (x̄ = 2.2 × 2 mm, n = 5) immersed to semi-immersed, erumpent through bark of host, carbonaceous, valsoid, black, glabrous. Ascomata 1.5–2.5 × 0.3–0.5 mm (x̄ = 1.8 × 0.4 mm, n = 10), perithecial, gregarious, cylindrical, black, ostiolate, with necks, circular in apical view, periphysate. Peridium 25–60 µm (x̄ = 50 µm, n = 30) wide, comprising two layers, outer layer carbonaceous, composed of brown to black cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses 1.5–3.5 µm wide, numerous, septate, unbranched, filiform, guttulate. Asci (45–)47–53(–65) × 3.5–5.5 µm (x̄ = 50 × 4.5 µm, n = 50), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- apical ring inconspicuous. Ascospores 4.5–6.5 × 2–4 µm (x̄ = 6 × 3 µm, n = 50), uniseriate, cylindrical to ellipsoidal, aseptate, hyaline when young, becoming pale brown or dark grey, sometimes slightly contracted at middle of the cell, smooth-walled, 2 guttules, with a germ pore at the base. Asexual morph: Undetermined.
Material examined: France, Aquitaine, Pyrénées-Atlantiques, Pau 64, on dead wood of Quercus sp., 24 October 1998, F. Candoussau and G.J. Samuels (S-F24255); Sweden, Uppland, Sollentuna, 700 m NNW of Hägerstalund, on rotten log, 3 May 1984, N. Lundqvist (UPS-UPS:BOT:F-120952).
Known hosts and distribution: On dead wood of Acer, Liriodendron and Rhododendron in the USA (type locality) (Shear 1940); on dead wood of Alnus glutinosa in Slovakia (Vasilyeva 1997); on dead wood in Austria, Sweden, Russia (Vasilyeva 1997); on dead wood of Fraxinus excelsior in Czech Republic (del Valle Catania and Romero 2003); on decorticated wood of Podocarpus parlatorei in Argentina (del Valle Catania and Romero 2003); on dead wood of Quercus in France (del Valle Catania and Romero 2003).
Notes: We could not obtain the holotype, so we re-examined specimens F-120952 from UPS and F24255 from S herbaria based on information provided by del Valle Catania and Romero (2003). The strain of Camaropella pugillus nests in Camaropella (Walker et al. 2012; Vu et al. 2019) and related to Camaropella amorpha and Cam. microspora (Fig. 12).
New combinations:
Camaropella amorpha (Boedijin) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sarcostromella amorpha Boedijn, Persoonia 1(1): 17 (1959)
Synonym: Camarops amorpha (Boedijn) Nannf., Svensk bot. Tidskr. 66(4): 362 (1972)
Index Fungorum number: IF558202
Camaropella microspora (P. Karst.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Anthostoma microsporum P. Karst., Fungi Fenniae Exsiccati, Fasc. 9: no. 860 (1869)
Synonym: Camarops microspora (P. Karst.) Shear, Mycologia 30(5): 588 (1938)
Index Fungorum number: IF558249
Camaropella plana (Pouzar) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Camarops plana Pouzar, Česká Mykol. 40(4): 219 (1986)
Index Fungorum number: IF558250
Camarops P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 6, 53 (1873)
Synonym: Bolinia (Nitschke) Sacc., Syll. fung. (Abellini) 1: 352 (1882)
Saprobic on wood. Sexual morph: Ascostromata immersed, erumpent to superficial, dark brown to black, carbonaceous or coriaceous, pulvinate, peltate, discoid or valsoid, hemispherical, spherical or irregular, sometimes with broad attachment to the substrate. Ascomata perithecial, gregarious, globose to subglobose, brown to dark brown, ostiolate. Peridium comprising two layers, outer layer composed of brown to dark brown cells of textura intricata; inner layer composed of hyaline cells of textura prismatica. Paraphyses abundant, persistent, filamentous, septate, branched. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded to blunt, with J- apical ring. Ascospores 2–3-seriate, subglobose to ellipsoidal, aseptate, pale brown to dark brown, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Nannfeldt 1972).
Notes: Bolinia was introduced as a subgenus by Nitschke (1867) and raised to generic rank by Saccardo (1882), based on Sphaeria tubulina Alb. & Schw. Karsten (1873) had earlier introduced Camarops based on C. hypoxyloides P. Karst. Bolinia and Camarops, as very similar genera, have often been discussed together in subsequent studies. Munk (1957) and von Arx and Müller (1954) reviewed the taxonomy of these taxa and concluded that Camarops and Bolinia were distinct genera with similar characters, except the former has monostichous ascomata (ascomata arranged in one line or series in stroma, e.g. Fig. 16 b) and the latter has polystichous (ascomata arranged in multiple lines or groups in stroma, e.g. Fig. 17 d–e). Shear (1938) and Svrček (1969) recognized that Sphaeria tubulina Alb. & Schw. should be a species of Camarops, named Camarops tubulina. However, Nannfeldt (1972) re-examined species of Camarops and Bolinia, and suggested Bolinia should be merged with Camarops based on their similar characters. Nannfeldt (1972) also proposed that the structure of the ascospores, peridium and stroma should be emphasized in the remodeled Camarops, and the arrangement of ascomata in stroma and size, shape of stromata are considered to be the basis for distinguishing species (del Valle Catania and Romero 2003). Therefore, Bolinia was accepted as a synonym of Camarops under Boliniaceae (Lumbsch and Huhndorf 2010; Hyde et al. 2020; Wijayawardene et al. 2020). Vasilyeva (1997) established Camaropella based on Camarops pugillus. Untereiner et al. (2013) found that species of Camaropella and Camarops sensu lato form a cluster (see notes for Camaropella). In this study, we found that the previously proposed methods for distinguishing Camaropella and Camarops morphologically are inconsistent with phylogenetic analysis. Currently, we suggest using phylogenetic analysis as the main method to distinguish Camaropella and Camarops. The complex needs resolving based on more fresh collections.
Type species: Camarops hypoxyloides P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 53 (1873)
Facesoffungi number: *****??; Fig. 16
Saprobic on wood. Sexual morph: Ascostromata dark brown to black, carbonaceous, pulvinate, discoid, hemispherical or spherical, sometimes with broad attachment to the substrate. Ascomata perithecial, gregarious, globose to subglobose, brown to dark brown, monostichous, carbonaceous, ostiolate, with periphyses. Peridium 25–70 µm (x̄ = 40 µm, n = 20) wide, comprising two layers, outer layer carbonaceous to coriaceous, composed of brown to dark brown cells of textura intricata to textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Asci 8-spored, unitunicate, cylindrical, apex rounded, pedicellate. Ascospores 6–8 × 4–6 µm (x̄ = 6.5 × 4.5 µm, n = 20), uniseriate, subglobose to ellipsoidal, aseptate, hyaline when young, becoming pale brown, smooth-walled, with 2 large guttules, a germ pore at end. Asexual morph: Undetermined (adapted from Karsten 1873).
Material examined: USA, Louisiana, Ferriday, on dead wood of Gleditsia sp., 18 July 1931, Overholts and Siggers (BPI-797104).
Known hosts and distribution: On dead wood of Alnus glutinosa in UK (type locality) (Karsten 1873); on dead wood in tropical America and rare in Europe (Miller 1930); on decayed wood in Argentina (Mercuri 1972).
Notes: Camarops hypoxyloides was introduced by Karsten (1873). Miller (1930) proposed that C. hypoxyloides is closely related to Solenoplea peltata Lloyd from Porto Rico, and found that ‘The Montagne specimen of Hypoxylon polyspermum at Kew is, according to Mason in a letter, the same as his English specimen of Camarops which is equal to Karsten’s C. hypoxyloides’, and then introduced Camarops polysperma (≡ Hypoxylon polyspermum Mont.) as the current name of C. hypoxyloides (Miller 1930; Mercuri 1972). However, this statement is not generally recognized and C. hypoxyloides is widely accepted as the type species of Camarops (Untereiner et al. 2013; Hyde et al. 2020; Index Fungorum 2020). We also recommend C. hypoxyloides, the name first used to introduce Camarops, as the type species, and accept C. polysperma as an independent species similar to C. hypoxyloides, until there is sufficient morphological or molecular data to discern their relationship. Raja et al. (2011) sequenced molecular data for C. polysperma which was collected from Sweden. Camarops tubulina is sister to C. polysperma (95%ML/0.99BY, Fig. 12). Morphologically, the former has polystichous ascomata in stroma, whereas the latter has monostichous ascomata (Shear 1938; Mercuri 1972). In this study, we were unable to obtain the type specimen. Therefore, we re-examined an authentic material (BPI-797104) collected in America.
Camarops tubulina (Alb. & Schwein.) Shear, Mycologia 30(5): 585 (1938)
Basionym: Sphaeria tubulina Alb. & Schwein., Consp. fung. (Leipzig): 6 (1805)
Facesoffungi number: *****??; Fig. 17
Saprobic on wood. Sexual morph: Ascostromata 15–20 × 2–5 mm (x̄ = 18 × 4 mm, n = 5). carbonaceous to coriaceous, immersed to semi-immersed, dark brown, smooth, pulvinate, discoid, hemispherical or irregular, with broad attachment to the substrate. Ascomata 1–2.5 × 0.5–1 mm (x̄ = 1.5 × 0.7 mm, n = 10), perithecial, gregarious, polystichous, carbonaceous, globose to subglobose, dark brown to black, ostiolate, with necks. Necks carbonaceous, dark brown to black, the wall composed of brown to hyaline cells of textura intricata to textura prismatica, with hyaline periphyses. Peridium of venter 40–120 µm (x̄ = 90 µm, n = 30) wide, comprising two layers, outer layer coriaceous, composed of brown to dark brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses 0.8–1.8 µm wide, numerous, septate, branched, filiform. Asci 30–55 × 2–4.5 µm (x̄ = 40 × 3 µm, n = 50), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- apical ring. Ascospores 3–6 × 1.5–2.5 µm (x̄ = 4.5 × 2 µm, n = 50), uniseriate, subglobose to ellipsoidal, aseptate, hyaline when young, becoming dark brown or dark grey, with germ spore at base, smooth-walled, with 2 guttules. Asexual morph: Undetermined.
Material examined: Czech Republic, Novohradských Horách Mountains, Virgin forest of Žofínský prales, on the dead trunks of Picea sp., 18 October 1967, Z. Pouzar, determined by M. Svrček in January 1969 (PRM-PR647008).
Known hosts and distribution: On trunk of Abies in Upper Lusatia (type locality) (Albertini and Schweinitz 1805); on dead spruce stumps in moist places in the south of Sweden (Fries 1816); on decayed wood of Abies, rare Juglans and Alnus in Lusatia (Germany), Sweden and North America (Saccardo 1882); on dead wood of conifers (Picea, Abies) and rare broadleaved trees (Fagus) in Czech Republic (Holec 2005).
Notes: Saccardo (1882) transferred Sphaeria tubulina Alb. & Schw. to Bolinia as generic type (see notes for Boliniaceae). Shear (1938) and Svrček (1969) combined Sphaeria tubulina Alb. & Schw. under Camarops tubulina (see notes for Camarops). Thus, Bolinia was subsequently accepted as a synonym for Camarops, and its members were assigned to Camaropella and Camarops (Shear 1938; Nannfeldt 1972; Vasilyeva et al. 2007). Rehm (1904) and Theissen (1909) proposed C. tubulina is a synonym of C. hypoxyloides, but Miller (1930) thought that the correct samples were not examined. Thus, there is an issue with their assumption, that is, C. tubulina (≡ Sphaeria tubulina) collected by Mason has polystichous ascomata in stroma with long elongate necks, different from C. hypoxyloides as described by Karsten has monostichous ascomata. Huhndorf et al. (2004b) sequenced C. tubulina, which is sister to C. polysperma (Raja et al. 2011; this study, 95%ML/0.99BY, Fig. 12). In this study, we were unable to obtain the type material. Therefore, we re-examined an authentic sample determined by Svrček (mentioned in the label of material). Camarops hypoxyloides and C. tubulina are two vital species in Boliniaceae (Boliniales), and their relationships should be resolved following fresh collections and epitypification.
Cornipulvina Huhndorf, A.N. Mill., F.A. Fernández & Lodge 2005
Saprobic on decorticated wood. Sexual morph: Ascostromata superficial, irregular in shape, membranaceous, brown to black, glabrous, with fascicular arrangement, necks rostrate. Ascomata perithecial, subglobose, dark brown to black, ostiolate, with papilla, periphysate. Peridium composed of brown to dark brown cells of textura angularis. Paraphyses numerous, septate, unbranched, filiform, guttulate, evanescent. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- inconspicuous apical ring. Ascospores uniseriate, subglobose to ellipsoidal, aseptate, hyaline, smooth-walled, with 2 guttules, without gelatinous appendages. Asexual morph: Undetermined (adapted from Huhndorf et al. 2005).
Notes: The monotypic genus Cornipulvina is characterized by ascostromata with long rostrate necks and ellipsoidal to subglobose ascospores without germ pores (Huhndorf et al. 2005). Sequence data from Cornipulvina ellipsoides (SMH 1378) is basal to Apiorhynchostoma and Pseudovalsaria in Boliniaceae (Huhndorf et al. 2005; 70%ML, Fig. 12).
Type species: Cornipulvina ellipsoides Huhndorf, A.N. Mill., F.A. Fernández & Lodge, Fungal Diversity 20: 63 (2005)
Facesoffungi number: *****??; Fig. 18
Saprobic on wood. Sexual morph: Ascostromata 1.5–3.5 × 1–2.5 mm (x̄ = 2.5 × 1.5 mm, n = 5), superficial, irregular in shape, membranaceous, brown to black, glabrous, with fascicular arrangement, necks rostrate. Ascomata 450–900 × 250–600 µm (x̄ = 750 × 450 µm, n = 10), perithecial, subglobose, dark brown to black, ostiolate, with papilla, periphysate. Peridium 25–65 µm (x̄ = 50 µm, n = 30) wide, composed of brown to dark brown cells of textura angularis. Paraphyses 1–3.5 µm wide, numerous, septate, unbranched, filiform, guttulate, evanescent. Asci (50–)60–77(–85) × 3.5–7 µm (x̄ = 70 × 5.5 µm, n = 50), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- inconspicuous apical ring. Ascospores (5–)7–8.5(–10) × 3–5 µm (x̄ = 8 × 4.5 µm, n = 50), uniseriate, subglobose to ellipsoidal, aseptate, hyaline, smooth-walled, usually with two large guttules, without gelatinous appendages. Asexual morph: Undetermined.
Material examined: Puerto Rico, Luquillo Mountains, Bisley Watershed 3, 220m, 18º 19.00’ N / 65°0’ W, on dead decorticated wood, 8 May 1995, S.M. Huhndorf and D.J. Lodge (F-SMH 1378, holotype).
Known hosts and distribution: On decayed decorticated wood in Brazil, Puerto Rico (type locality) and Venezuela (Huhndorf et al. 2005).
Notes: We re-examined the holotype of Cornipulvina ellipsoides (SMH 1378), and it is sister to Apiorhynchostoma and Pseudovalsaria (70%ML, Fig. 12).
Endoxyla Fuckel, Jb. nassau. Ver. Naturk. 25-26: 321 (1871)
Saprobic on wood. Sexual morph: Ascostromata poorly developed or absent. Ascomata perithecial, gregarious or in bi-seriate linear groups, immersed to semi-immersed, globose to subglobose, black, ostiolate. Asci 8-spored, clavate, pedicellate. Ascospores ellipsoidal to cylindrical, hyaline to brown, slightly curvate, with a germ pore at one end. Asexual morph: Undetermined (adapted from Fuckel 1872).
Notes: Fuckel (1872) established Endoxyla based on globose ascomata with clavate asci and ellipsoidal ascospores, and while introducing E. macrostoma, mentioned that Sphaeria parallela Fr. and S. operculata Alb. & Schw. belong to Endoxyla. Saccardo (1882) accepted S. parallela as Endoxyla parallela and S. operculata as E. operculata. Subsequently, there was a controversy as to which species should be selected as the type of Endoxyla. Some mycologists have recognized S. operculata (≡ E. operculata) published earlier, as the type species (Clements and Shear 1931; Holm 1975). Others believed that Fuckel (1872) only described E. macrostoma in detail when establishing Endoxyla, so that, it should be regarded as the generic type (Untereiner 1993). Endoxyla macrostoma is widely accepted as the type species of Endoxyla (Untereiner 1993; Untereiner et al. 2013; Hyde et al. 2020). Untereiner et al. (2013) sequenced strains of E. macrostoma, E. mallochii and E. operculata which clustered with Endoxyla (98%ML, Fig. 12) in Boliniaceae. The Endoxyla species forms a sister clade to Camaropella (53%ML, Fig. 12).
Type species: Endoxyla macrostoma Fuckel, Jb. nassau. Ver. Naturk. 25-26: 322 (1871)
Fig. 20 l–m
Saprobic on decorticated wood. Sexual morph: Ascostromata poorly developed, brown, septate hyphae. Ascomata perithecial, gregarious or in bi- or uni-seriate linear groups, immersed to semi-immersed, ampulliform to subglobose, black, ostiolate, with papilla. Peridium comprising two layers, outer layer composed of dark brown to brown cells of textura intricata to textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses filiform, septate, sparse in mature ascomata. Asci 8-spored, clavate to cylindrical, pedicellate, with J- apical ring. Ascospores ellipsoidal to suballantoid, curved, 0–1-septate, unequal, composed of a large brown to pale brown cell with an apical germ pore and a tiny, conical, hyaline cell. Asexual morph: Undetermined (adapted from Fuckel 1872; Untereiner 1993).
Known hosts and distribution: On decayed, decorticated wood of Quercus in Germany (type locality) (Fuckel 1872); on decorticated wood of angiosperms and gymnosperms (Abies, Picea, Pinus) in Canada, Czech Republic, Germany and Sweden (Untereiner 1993; Untereiner et al. 2013).
Notes: Fuckel (1872) established Endoxyla and introduced E. macrostoma which was collected from Germany. Holm (1975) suggested E. operculata as the type species. However, Untereiner (1993), after reviewing the species of Endoxyla, proposed that E. macrostoma should be used as the type (see notes for Endoxyla). Untereiner et al. (2013) sequenced molecular data for E. macrostoma collected from Canada and this strain is basal to E. mallochii and E. operculata (98%ML, Fig. 12).
Endoxyla operculata (Alb. & Schw.) Sacc., Syll. fung. (Abellini) 1: 181 (1882)
Basionym: Sphaeria operculata Alb. & Schwein., Consp. fung. (Leipzig): 40 (1805)
Facesoffungi number: *****??; Fig. 19
Saprobic on wood. Sexual morph: Mycelium brown, septate hyphae 2–6.5 µm wide, surrounded the base of the ascoma. Ascostromata poorly developed or absent. Ascomata 800–900 × 500–565 µm (x̄ = 845 × 530 µm, n = 10), perithecial, solitary, immersed, with necks erumpent the rough wood of the surface, in linear groups, ampulliform to subglobose, black, ostiolate, with necks, periphysate. Peridium 40–100 µm (x̄ = 70 µm, n = 30) wide, comprising three layers, outer layer carbonaceous or coriaceous, composed of dark brown cells of textura intricata; middle layer membranaceous, composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses filiform to ventricose, septate, sparse in mature ascomata. Asci 85–90 × 6.5–12 µm (x̄ = 87.5 × 9.5 µm, n = 30), 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex blunt, with J- apical ring. Ascospores 9.5–16 × 4–6.5 µm (x̄ = 13.5 × 5 µm, n = 50), uni- or bi-seriate, ellipsoidal to cylindrical, aseptate, hyaline when young, becoming pale grey, smooth-walled, slightly curved, guttulate, sometimes with an apical germ pore. Asexual morph: Undetermined (adapted from Untereiner 1993).
Material examined: Sweden, Uppland, Östera Ryd par., Rydboholm. S. of the castle in deciduous forest, on naked wood of deciduous tree, 17 May 1983, N. Lundqvist (S-F 138814); Norway, Troms, Storfjord commune, Helligskogen in Skibotndalen Valley (26 km SE of Skibotn), subalpine birch forest, on decayed wood of Betula pubescens, 17 August 1992, N. Lundqvist (S-F 138817).
Known hosts and distribution: On decayed wood of Abies in Lusatia and Sweden (type locality) (Albertini and Schweinitz 1805); on decayed, decorticated wood of angiosperms (Acer, Fagus, Populus, Quercus) and gymnosperms (Alnus, Pinus) in Canada, Czech Republic, Finland, France, Germany, Sweden, USA (Untereiner 1993; Untereiner et al. 2013).
Notes: Untereiner (1993) introduced a lectotype for Endoxyla operculata which was collected from Sweden by Fries which has immersed ascomata with clavate to cylindrical asci and cylindrical ascospores; however, the stomata are poorly developed or absent. In this study, we could not obtain the type material. Therefore, we re-examined authentic specimens collected by Lundqvist in Sweden and Norway. These collections of E. operculata examined here, had prominent apical rings in the asci, and ascomata were not in a stroma (Fig. 19). These characters are not identical to Camarops tubulina, which is considered to be the current name of E. operculata (Index Fungorum 2020). Endoxyla is different from Camarops with crust-like stroma. Phylogenetically, the strain of E. operculata is sister to E. mallochii and is distant from C. tubulina (Untereiner et al. 2013; Réblová et al. 2018; this study, 84%ML, Fig. 12).
Mollicamarops Lar.N. Vassiljeva, Mycotaxon 99: 160 (2007)
Saprobic on decorticated wood. Sexual morph: Ascostromata semi-immersed, irregular in shape, widely effuse, ochreous or fulvous. Ascomata black, ostiolar region conspicuously stellate to furrowed. Paraphyses numerous, filiform, sinuous. Asci 8-spored, unitunicate, cylindrical, apical ring indistinct, long pedicellate. Ascospores ellipsoidal, light brown to brown, aseptate, smooth-walled, with guttules. Asexual morph: Undetermined (adapted from Vasilyeva 2007).
Notes: The monotypic genus Mollicamarops is characterized by stellate, ostiolate ascomata, cylindrical asci and ellipsoidal, aseptate ascospores. It is similar to Camarops flava in having yellow to brown ascostromata and can be distinguished as the latter has astellate ostioles and ornamented ascospores (Samuels and Rogers 1987; Vasilyeva 2007). Therefore, we place Mollicamarops in Boliniaceae.
Type species: Mollicamarops stellata Lar.N. Vassiljeva, Mycotaxon 99: 160 (2007)
Fig. 20 n–o
Saprobic on decorticated wood. Sexual morph: Ascostromata semi-immersed, irregular in shape, widely effuse, ochreous or fulvous. Ascomata black, ostiolar region conspicuously stellate to furrowed. Paraphyses numerous, narrow, sinuous. Asci 8-spored, unitunicate, cylindrical, apical ring indistinct. Ascospores ellipsoidal, light brown, aseptate, smooth-walled, with guttules. Asexual morph: Undetermined (adapted from Vasilyeva 2007).
Known hosts and distribution: On decayed, decorticated wood in Russia (type locality) (Vasilyeva 2007).
Notes: We were unable to obtain type material. Therefore, we draw the asci and ascospores of Mollicamarops stellata (Fig. 20 n–o) from Vasilyeva (2007).
Neohypodiscus J.D. Rogers, Y.M. Ju & Læssøe, Mycologia 86(5): 684 (1994)
Saprobic on decorticated wood. Sexual morph: Ascostromata subglobose to pulvinate to irregular, solitary or confluent, narrow to broad, carbonaceous, with longitudinal veins below and attachment to the substrate. Ascomata subglobose to ampulliform, brown to black, subglobose, carbonaceous, monostichous, ostiolate, rough-walled. Paraphyses numerous, mostly unbranched, septate. Asci 8-spored, unitunicate, cylindrical to clavate, with J- apical ring. Ascospores uniseriate, greenish to brown, ellipsoidal to crescent-shaped, 0–2-septate, smooth-walled, some with an apical thickening or acellular appendage at one end, with a germ pore at one or both end(s), guttulate. Asexual morph: Undetermined (adapted from Rogers et al. 1994b).
Notes: Rogers et al. (1994b) placed Neohypodiscus in Amphisphaeriaceae (Xylariomycetidae) based on its carbonaceous stromata, cylindrical asci and ellipsoidal to fusiform ascospores. Subsequently, Andersson et al. (1995) transferred it to Boliniaceae. No molecular data for this genus has been sequenced yet. In this study, we place Neohypodiscus in Boliniaceae because it has the same carbonaceous stromata and unequal ascospores with hyaline thickened end(s) as Camarops and Apiorhynchostoma.
Type species: Neohypodiscus rickii (Lloyd) J.D. Rogers, Y.M. Ju & Læssøe, Mycologia 86(5): 685 (1994)
Basionym: Hypodiscus rickii Lloyd, Mycol. Writ. 7(Letter 68): 1181 (1923)
Fig. 20 a–k
Saprobic on decorticated wood. Sexual morph: Ascostromata subglobose to irregular, confluent in mass, with narrow to broad connection to substrate, brown to black, with veins. Ascomata 350–1300 × 550–800 µm (x̄ = 1100 × 755 µm, n = 10), brown to black, subglobose, carbonaceous, with a short umbilicate ostiole, rough-walled. Paraphyses 1.5–10 µm wide, numerous, mostly unbranched, septate. Asci 200 × 11 µm, 8-spored, unitunicate, cylindrical, apical ring indistinct. Ascospores 26.5–32 × 7–9 µm, uniseriate, greenish to brown, ellipsoidal to crescentic, aseptate or 1-septate near one end or rarely 2-septate, unequal, normally composed of one small cell and one large cell with a germ pore at end, smooth-walled, some with an apical thickening or acellular appendage at one end, with guttules. Asexual morph: Undetermined (adapted from Rogers et al. 1994b).
Material examined: Puerto Rico, El Toro Trail, off Rt. 186, Luquillo Mountains, on dead wood, 24 December 1996, D.J. Lodge, G.J. Samuels and H.J. Strauss (BPI-737826); Puerto Rico, El Verde, Luquillo Mts., Trail to the Radiation Center, on dead wood, 23 March 1993, D.J. Lodge and M. Boyd (BPI-802543).
Known hosts and distribution: On decayed, decorticated wood in Brazil (type locality), Puerto Rico (Rogers et al. 1994b).
Notes: Lloyd (1923) described Hypodiscus rickii, and Rogers et al. (1994b) transferred this species as Neohypodiscus rickii as its generic type. Rogers et al. (1994b) examined two samples collected by Lloyd from Brazil and defined those collections as lectotype and paralectotype materials. In this study, we re-examined two authentic samples determined by Rogers (mentioned in the label of material) as we were unable to obtain type material. We were unable to find any structures in the samples except the ascomata, therefore, we provided a hand drawing of asci and ascospores (Fig. 20 j–k) following Rogers et al. (1994b).
Pseudovalsaria Spooner, Trans. Br. mycol. Soc. 86(3): 405 (1986)
Saprobic on wood. Sexual morph: Ascostromata immersed to erumpent through bark. Ascomata gregarious, brown to black, ellipsoidal to pyriform, immersed to semi-immersed in stroma. Asci 8-spored, unitunicate, cylindrical, apex rounded or blunt. Ascospores uniseriate, brown to dark brown, cylindrical to ellipsoidal, ends slightly tapered to rounded, 0–1-septate, smooth-walled, with a germ pore at the base, guttulate. Asexual morph: Undetermined (adapted from Spooner 1986).
Notes: Spooner (1986) established Pseudovalsaria typified by P. foedans, and Barr (1994) placed this genus in Clypeosphaeriaceae. Rappaz (1995) subsequently transferred it to Boliniaceae as it has similar characters to Camarops, such as cylindrical asci and ellipsoidal ascospores with germ pore. Untereiner et al. (2013) confirmed that Pseudovalsaria belong in Boliniaceae based on ITS-LSU sequence analyses of P. ferruginea. In this study, the strain of P. ferruginea is sister to Apiorhynchostoma apiosporum in Boliniaceae (72%/ML, Fig. 12).
Type species: Pseudovalsaria foedans (P. Karst.) Spooner, Trans. Br. mycol. Soc. 86(3): 405 (1986)
Basionym: Phaeosperma foedans P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 55 (1873)
Fig 20 p–q
Saprobic on wood of Pinus silvestris. Sexual morph: Ascostromata gregarious, immersed to erumpent through bark. Ascomata gregarious, immersed to semi-immersed in stroma, brown to black, ellipsoidal to pyriform, with long necks. Necks composed of pale brown to hyaline cells of textura porrecta, surrounded by stromatic tissue. Peridium of venter outer layer composed of dark brown, irregular patches forming opaque amorphous matter; middle layer composed of brown cells of textura angularis to textura prismatica; inner layer composed of hyaline cells of textura prismatica. Paraphyses mostly branched, filiform, septate. Asci 8-spored, unitunicate, cylindrical, apex rounded, with J- apical ring. Ascospores uniseriate, dark brown, cylindrical to ellipsoidal, slightly tapered to rounded ends, sometimes slightly curved, 1-septate, smooth-walled, with a terminal germ pore, guttulate. Asexual morph: Undetermined (adapted from Spooner 1986).
Known hosts and distribution: On decorticated wood of Viburnum sp. in China (Spooner 1986); on decayed wood of Betula sp., Castanea sativa, Pinus silvestris, Prunus laurocerasus, Quercus sp. in England (type locality) (Spooner 1986); on decayed wood of Alnus in Finland (Spooner 1986); on decayed wood of Alnus sp. in Canada (Untereiner et al. 2013).
Notes: Spooner (1986) established Pseudovalsaria based on Phaeosperma foedans. Untereiner (1993) proposed that Anthostoma ferrugineum, which has J- asci and brown, 1-septate ascospores with an apical germ pore, is similar to Pse. foedens. Rappaz (1995) subsequently synonymized A. ferrugineum under Pse. ferruginea and mentioned that ascospores of Pse. foedans are normally cylindrical and slightly curved which distinguishes it from Pse. ferruginea which has symmetrical, cylindrical ascospores. Ju et al. (1996) believed that the difference in ascospores between P. ferruginea and P. foedans were indistinct, and these species were probably conspecific. They also merged these two species and recognized P. ferruginea as the current name (Ju et al. 1996). However, this was not verified by phylogenetic analysis. Morphologically, Rappaz (1995) had considered Pse. ferrugine as having grey-brown tissue around the brown-red ascomata as different to Pse. foedans, which has dark brown tissue around the dark brown ascomata (Spooner 1986). The holotype of Pse. ferruginea and Pse. foedans were collected from Germany and England respectively (Nitschke 1867; Karsten 1873). Untereiner et al. (2013) analyzed the sequence data from a collection of Pse. ferruginea which was isolated from a decorticate branch of Populus in Canada. This study recommends that the generic type Pse. foedans (Spooner 1986) and Pse. ferruginea as two independent species until there are more characteristics and phylogenetic data to determine their classification. The strain of Pse. ferruginea is sister to Apiorhynchostoma apiosporum (72%/ML, Fig. 12), but no sequence is available for Pse. foedans. We were unable to examine the type taxon, therefore, we provided a hand drawing of asci and ascospores (Fig. 20 p–q) following Spooner (1986).
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández, Mycologia 96(2): 378 (2004)
Chaetosphaeriales was introduced by Huhndorf et al. (2004a) and four families (Chaetosphaeriaceae, Helminthosphaeriaceae, Leptosporellaceae and Linocarpaceae) are included in this order based on the morphology and phylogenetic analyses (Maharachchikumbura et al. 2015, 2016; Hernández-Restrepo et al. 2017; Hyde et al. 2020). Species of Chaetosphaeriales are mostly saprobes or/and pathogens in terrestrial and aquatic habitats, and can also be fungicolous (Maharachchikumbura et al. 2015, 2016).
Helminthosphaeriaceae Samuels, Cand. & Magni, Mycologia 89(1): 144 (1997)
Saprobic or parasitic on wood or decorticated branches or stems in terrestrial habitats, sometimes fungicolous. Sexual morph: Ascomata perithecial, solitary, scattered or gregarious, superficial or immersed, ovoid to subglobose, dark brown to black, carbonaceous or membranaceous, glabrous or tuberculate or with setae, papillate. Peridium carbonaceous or membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, evanescent, septate, unbranched or branched, filiform to cylindrical. Asci 8-spored, unitunicate, cylindrical to clavate, apex truncate, pedicellate, with J- apical ring. Ascospores normally bi-seriate, allantoid or clavate or cylindrical, 0–3-septate, hyaline to brown or grey or becoming dark colored in part, smooth-walled, with germ pore at one end or/and both ends, with guttules or guttules absent. Asexual morph: Hyphomycetous. Conidiophores macronematous, mononematous, gregarious, cylindrical, branched or unbranched, pale brown to brown, septate, smooth-walled. Conidia solitary, acrogenous, ellipsoidal to obclavate, 0–multi-septate, smooth-walled, hyaline to brown, with guttules or guttules absent (adapted from Fuckel 1870; Réblová 1999b; Maharachchikumbura et al. 2016).
Type genus: Helminthosphaeria Fuckel 1870
Notes: Helminthosphaeriaceae is typified by Helminthosphaeria (Fuckel 1870) and most of its species have black setose, globose ascomata and cylindrical asci with brown to hyaline, subglobose ascospores and mononematous conidiophores with oblong conidia. Initially, Echinosphaeria, Endophragmiella, Helminthosphaeria, Hilberina, Ruzenia and Synaptospora were included based on morphological and/or phylogenetic analysis (Miller and Huhndorf 2004a; Miller et al. 2014; Maharachchikumbura et al. 2015). Diplococcium and Tengiomyces were added to this family (Lumbsch and Huhndorf 2007; Wijayawardene et al. 2018). Tengiomyces was accepted in this family as its ascomata are similar to Helminthosphaeria (Réblová 1999a), whereas its ascospores are similar to Chaetosphaerella and Crassochaeta (Chaetosphaerellaceae, Coronophorales) (Réblová 1999a; Hyde et al. 2020; Wijayawardene et al. 2020). Thus, we suggest placing Tengiomyces in Coronophorales genera incertae sedis (see notes for Tengiomyces). The hyphomycetous Diplococcium and Endophragmiella asexual morphs have associated species of Helminthosphaeria and Echinosphaeria on the natural substratum (Sutton 1973; Samuels et al. 1997; Goh and Hyde 1998; Goh et al. 1998; Réblová 1999b; Huhndorf et al. 2004b). Diplococcium has phylogenetic affinities with Helotiales (Leotiomycetes) and Pleosporales (Dothideomycetes) (Shenoy et al. 2007, 2010); and Endophragmiella species are closely related to Helminthosphaeriaceae and Tubeufiales (Dothideomycetes) (Hernández-Restrepo et al. 2017; Vu et al. 2019). However, Diplococcium was transferred to Vibrisseaceae (Ekanayaka et al. 2019) and we recommend to place Endophragmiella in Ascomycota genera incertae sedis (see notes for Endophragmiella). In this study, Synaptospora is transferred to Sordariales genera incertae sedis (see notes for Synaptospora). Thus, four genera, Echinosphaeria, Helminthosphaeria, Hilberina and Ruzenia are presently accepted in Helminthosphaeriaceae.
Echinosphaeria A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 29 (2004)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary or scattered, superficial, subglobose to ovoid, membranaceous, brown to black, with setae, ostiolate, with papilla, periphysate. Setae hyaline to brown, straight, rigid, with acute apex. Peridium composed of loose arrangement, of hyaline to brown cells of textura angularis to textura prismatica. Asci 8-spored, unitunicate, cylindrical to clavate, apex blunt, with J- apical ring, pedicellate. Ascospores cylindrical to allantoid, 0–multi-septate, hyaline to brown, smooth-walled or verrucose, ends rounded, slightly curved, guttulate. Asexual morph: Undetermined (adapted from Persoon 1796; Karsten 1873; Miller and Huhndorf 2004a).
Notes: Miller and Huhndorf (2004a) established Echinosphaeria for Lasiosphaeria canescens based on LSU sequence analysis. This genus clustered with Helminthosphaeria in Helminthosphaeriaceae (Hongsanan et al. 2017; Wijayawardene et al. 2018; Hyde et al. 2020). In this study, Echinosphaeria nests in Helminthosphaeriaceae based on the LSU-TUB sequence analysis (Fig. 21).
Type species: Echinosphaeria canescens (Pers.) A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Basionym: Sphaeria canescens Pers., Observ. mycol. (Lipsiae) 1: 67 (1796)
Facesoffungi number: *****??; Fig. 22 a–r
Saprobic on wood. Sexual morph: Ascomata 365–660 × 340–590 µm (x̄ = 470 × 450 µm, n = 10), perithecial, solitary or scattered, superficial, globose to ovoid, membranaceous, brown to black, with brown, densely setae, ostiolate, with papilla, periphysate. Setae 5.5–12 µm wide at base, rigid, hyaline to pale brown, with acute apex. Peridium 25–50 µm (x̄ = 37 µm, n = 30) wide, comprising two layers, outer layer composed of loose arrangement, of brown to pale brown cells of textura angularis; inner layer composed of pale brown to hyaline cells of textura prismatica. Paraphyses 2–4 µm wide, numerous, septate, unbranched, filiform. Asci 105–115(–125) × 9–15.5 µm (x̄ = 110 × 12.5 µm, n = 30), 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex blunt, with J- apical ring. Ascospores (22.5–)24–28(–31) × 4–7.5 µm (x̄ = 26 × 6 µm, n = 50), bi-seriate, subglobose to allantoid, 0–1-septate, hyaline to pale brown, smooth-walled, ends rounded, slightly curved, guttulate. Asexual morph: Undetermined.
Material examined: USA, Michigan, Marquette, Huron Mountain Club, between Conway Lake and Lake Superior on road to Todd property, 46º00’ N/87º0’ W, on driftwood half buried in sand, 15 July 1996, S.M. Huhndorf et al. (F-SMH 2627). Sweden, Uppsala, Uppland, on dead wood, 26 August 1884, L. Romell (S-F 134887). Finland, on dead wood, 16 October 1861, P.K. Karsten (S-F 134901).
Known hosts and distribution: On dead, decorticated wood of Betula, Pinus and Quercus in Finland (type locality) (Persoon 1796; Karsten 1873); on dead wood in Denmark, Sweden and USA (Miller and Huhndorf 2004a).
Notes: Echinosphaeria canescens is the type species of Echinosphaeria (Miller and Huhndorf 2004a). This species has been reported with hyphomycetous Endophragmiella and selenosporella-like. Cultural characteristics for Ech. canescens are lacking (Hughes 1979; Sivanesan 1983; Miller and Huhndorf 2004a; Hernández-Restrepo et al. 2017; Vu et al. 2019). Strains of Ech. canescens clustered with Helminthosphaeria species in Helminthosphaeriaceae (Miller et al. 2014; Miller and Huhndorf 2004a; Hongsanan et al. 2017; this study, Fig. 21). The molecular data of two Endophragmiella species are available, End. taxi is closely related to Tubeufiales (Dothideomycetes) (Vu et al. 2019); and End. dimorphospora is nested in Helminthosphaeriaceae, but is phylogenetically distant from Ech. canescens (Hernández-Restrepo et al. 2017; this study, Fig. 21). Selenosporella strains are more closely related to Diatrypaceae (Xylariales) (Vu et al. 2019). In this study, we are unable to obtain the type material. We re-examined authentic samples collected by Karsten and Hundorf.
Helminthosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 166 (1870) [1869-70]
Saprobic or parasitic on wood or decorticated branch in terrestrial habitats, sometimes fungicolous. Sexual morph: Ascomata perithecial, solitary, scattered or gregarious, superficial or immersed, globose to subglobose, dark brown to black, carbonaceous or membranaceous, glabrous or with setae, with or without ostiole, with periphyses. Peridium carbonaceous or membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, evanescent, septate, unbranched or branched, filiform to cylindrical. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex truncate, with J- apical ring. Ascospores bi-seriate, ellipsoidal to clavate, 0–3-septate, hyaline to brown or grey, smooth-walled, with germ pore at one end or/and both ends, guttulate. Asexual morph (associated): Hyphomycetous. Conidiophores macronematous, mononematous, gregarious, cylindrical, branched, pale brown to brown, septate, smooth-walled. Conidia solitary, ellipsoidal to obclavate, 0–multi-septate, smooth-walled, hyaline to brown, guttulate (adapted from Fuckel 1870; Samuels et al. 1997; Miller et al. 2014).
Notes: Desmazières (1837) introduced a species growing on Clavaria fuliginea in northern France, and Fuckel (1870) later accepted it as the generic type of Helminthosphaeria, named as H. clavariarum. This genus was considered as member of Sordariales based on the ontogeny of the ascomata and 1-septate ascospores with germ pores are similar to Sordaria (Parguey-Leduc 1960; Lundqvist 1972; Eriksson and Hawksworth 1993). Subsequently, Samuels et al. (1997) monographed this genus, established Helminthosphaeriaceae and concluded that most species in this family were found on members of Clavariaceae or/and coriaceous basidiomycetes. They also found that Helminthosphaeria species were accompanied with hyphomycetous Diplococcium or/and heteroconium-like asexual morphs on the substratum (Samuels et al. 1997). Afterward, Miller et al. (2014) introduced several lignicolous Helminthosphaeria species based on phylogenetic analysis and reported that the polyphyletic Helminthosphaeria has Diplococcium and Selenosporella asexual morphs in nature, but not confirmed by cultural characteristics or phylogenetic analysis. Helminthosphaeria is characterized by a fungicolous and lignicolous habit, superficial or immersed ascomata, cylindrical asci with hyaline to brown ascospores and associated asexual morphs with smooth-walled or verrucose, septate conidia (Samuels et al. 1997; Goh and Hyde 1998; Réblová 1999a; Hernández-Restrepo et al. 2017). In this study, the polyphyletic Helminthosphaeria is related to Endophragmiella dimorphospora and Hilberina species, and scattered in Helminthosphaeriaceae (Fig. 21). However, we could not confirm the asexual morph of Helminthosphaeria due to lack of cultural characteristics and molecular data.
Type species: Helminthosphaeria clavariarum (Desm.) Fuckel, Jb. nassau. Ver. Naturk. 23-24: 166 (1870) [1869-70]
Basionym: Peziza clavariarum Desm., Annls Sci. Nat., Bot., sér. 2 8: 8 (1837)
Facesoffungi number: *****??; Fig. 23
Saprobic or parasitic on Clavulina. Sexual morph: Ascomata 295–390 × 195–275 µm (x̄ = 335 × 240 µm, n = 10), perithecial, solitary, superficial, globose, black, rough, surrounded by setae, ostiolate, with papilla, periphysate. Setae 9.5–15 µm wide. brown to pale brown, tapering, with a septate lumen. Ostioles inconspicuous, a loose arrangement of cylindrical to oblong cells. Peridium 25–50 µm wide, membranaceous, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 3–7 µm wide, numerous, evanescent, septate, filiform. Ascogenous hyphae discrete, smooth-walled, branched, with hyaline, ovoid to obpyriform cells, 9.5–12 × 3.5–6.5 µm (x̄ = 11 × 5.5 µm, n = 10), growing in sympodial succession, in dense clusters, each giving rise to an ascus. Asci (70–)80–85(–110) × 8–13.5 µm (x̄ = 82 × 10 µm, n = 30), 8-spored, unitunicate, cylindrical, apex rounded, pedicellate. Ascospores (9.5–)11–13(–16) × 5.5–7 µm (x̄ = 12.5 × 6.5 µm, n = 50), uni-seriate, ellipsoidal to broad-fusiform or triangular, aseptate, hyaline to pale brown, smooth-walled, with guttules and an apical germ pore. Asexual morph (associated): Hyphomycetous. Conidiophores (32.5–)42–55(–95) × 3.5–14 µm (x̄ = 53 × 7 µm, n = 10), arising from the substrate, macronematous, mononematous, cylindrical, sparingly branched, pale brown to brown, septate, smooth-walled. Conidiogenous cells (10.5–)14–20(–22) × 3–7.5 µm (x̄ = 17.5 × 5.5 µm, n = 10), terminal or intercalary, polytretic, brown, tapering, apex rounded, conidia formed through minute pores. Conidia (10–)17–20(–25) × 6–8 µm (x̄ = 18 × 7 µm, n = 30), solitary, ellipsoidal to subglobose, 0–1-septate, sometimes constricted at septum, slightly curved, smooth-walled, hyaline to brown, with guttules.
Material examined: Sweden, Uppland, Fresta, on Clavulina cinerea, 18 September 1984, von H. Eichwald (S-F 135699); Sweden, Södermanland, Hölö sn. Tullgarnsomradet, SSO, Tullgarns slot, on Clavulina sp., 9 October 2008, K. Jaederfeldt (S-F 121831).
Known hosts and distribution: On old or living Clavaria fuliginea in Franch (type locality) (Desmazières 1837); on Clavaria in Franch, Germany, Sweden, UK and USA (Samuels et al. 1997).
Notes: Helminthosphaeria clavariarum is fungicolous and has been reported on Clavariaceae members in Europe and North America associated with diplococcium-like asexual morphs (Dennis 1978; Samuels et al. 1997). Huhndorf et al. (2004b) analyzed LSU sequence data for H. clavariarum (SMH4609). Miller et al. (2014) re-examined the material of SMH4609 and analyzed TUB sequence data for H. clavariarum (ANM Acc.17). In this study, we re-examined two authentic specimens (F 135699 and F 121831) determined by Lundqvist (mentioned in the label of material).
New combination:
Helminthosphaeria plumbea (Huhndorf, F.A. Fernández & Cand.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Synaptospora plumbea Huhndorf, F.A. Fernández & Cand., Sydowia 51(2): 177 (1999)
Index Fungorum number: IF558251
Notes: See notes for Synaptospora.
Hilberina Huhndorf & A.N. Mill., Mycol. Res. 108(1): 31 (2004)
Saprobic on wood or leaves. Sexual morph: Ascomata perithecial, solitary or scattered, superficial, membranaceous, ovoid to obpyriform, brown to dark brown, tuberculate, covered with setae, ostiolate. Asci 8-spored, unitunicate, cylindrical. Ascospores biseriate, cylindrical, geniculate, one or both end(s) tapering to a distinct point or rounded ends, hyaline when young, becoming pale brown, smooth-walled, with guttules, 0–3-septate, sometimes with sheath. Asexual morph: Hyphomycetous. Conidiophores macronematous, mononematous, cylindrical, single or in group up to two, erect, arising directly from the host, multi-septate, cells expanding sometimes, brown to dark brown, apical cell swollen. Conidiogenous cells phialides, with cylindrical branches, hyaline to pale brown, discrete, smooth-walled; secondary branches, hyaline, 2–3 per branch, cylindrical, tapering at the apex. Conidia hyaline, aseptate, oblong to obovoid, smooth-walled (adapted from Fuckel 1870; Miller and Huhndorf 2004a).
Notes: Fuckel (1870) introduced Leptospora caudata has black ascomata and oblong ascospores and was accompanied with a hyphomycete with hyaline, oval conidia, on decayed wood of Betula alba in nature. This species was introduced as the generic type of Hilberina (Miller and Huhndorf 2004a). Subsequently, several taxa with bristly ascomata and ascospores distinctly curved or bent near the attenuate, lanceolate or acerose base were accepted as members of Hiberina (Declercq 2009, 2013; Miller et al. 2014). Miller et al. (2014) sequenced five Hilberina species and found that they are polyphyletic in Helminthosphaeriaceae and their asexual morphs unknown. In this study, Hilberina species are closely related to members of Helminthosphaeria and scattered in Helminthosphaeriaceae (Fig. 21). Its sexual-asexual link needs confirming with cultural characteristics and more phylogenetic data.
Type species: Hilberina caudata (Fuckel) Huhndorf & A.N. Mill., in Miller & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Basionym: Leptospora caudata Fuckel, Jb. nassau. Ver. Naturk. 23-24: 144 (1870)
Facesoffungi number: *****??; Fig. 24
Saprobic on decorticated wood. Sexual morph: Ascomata 350–600 × 375–565 µm (x̄ = 475 µm × 475 µm, n = 10), perithecial, solitary or scattered, superficial, membranaceous, ovoid to subglobose, dark brown to black, covered with setae, ostiolate, with papilla, periphysate. Setae 15–30 µm wide at base, brown to pale brown, tapering, with a septate lumen and apex rounded. Peridium 35–80 µm (x̄ = 55 µm, n = 30) wide, comprising two layers, outer layer composed of pale brown to brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci 125–180 × 9.5–13.5 µm, 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- inconspicuous apical ring. Ascospores (38–)43–50 × 3–8 µm (x̄ = 45.5 × 5.5 µm, n = 50), biseriate, cylindrical, geniculate, one end tapering to a distinct point, hyaline when young, becoming pale brown, smooth-walled, with guttules, 0–3-septate, without gelatinous appendages. Asexual morph (associated): Hyphomycetous. Conidiophores 3–8.5 µm, cylindrical, single or in group up to two, erect, arising directly from the host, macronematous, mononematous, multi-septate, not constricted, cells expanding sometimes, brown to dark brown, apical cell swollen. Conidiogenous cells phialides (3–)5–8 × 1–2 µm (x̄ = 6 × 1.5 µm, n = 30), with cylindrical branches, hyaline to pale brown, discrete, smooth-walled, aseptate; secondary branches (3.5–)4.5–5.5(–7.5) × 1–1.5 µm (x̄ = 5 × 1.2 µm, n = 30), hyaline, aseptate, 2–3 per branch, cylindrical, tapering at the apex. Conidia 2.5–3.5 × 1–2 µm (x̄ = 3 × 1.5 µm, n = 50), hyaline, aseptate, oblong to obovoid, smooth-walled (adapted from Fuckel 1870).
Material examined: Puerto Rico, Bisley Watershed 3, down slope on Quebrada 2 to Quebrada 3, Luquillo Mountains, 18º 19.00’ N / 65°0.07’ W, on dead wood, 28 January 1997, F.A. Fernández (F-SMH 3156); Germany, Hessen, 9º 0’ 19’’ E / 49° 46’ 28’’ N, on dead wood of Betula sp., K.W.G. Fuckel (G-G00266357 syntype).
Known hosts and distribution: On decorticated wood of Betula alba in Germany (type locality) (Fuckel 1870); on dead wood in Puerto Rico.
Notes: In this study, we re-examined a typical material and an authentic specimen, which were well preserved. The sexual morph of Hilberina caudata associated with haplographium-like asexual morph were found on the type materials. Hilberina caudata is sister to Hilberina punctata and nested in Helminthosphaeriaceae based on multi-gene analysis (Miller et al. 2014; this study, 96%ML/1.00BY, Fig. 21). However, cultural characteristics and phylogenetic data of H. caudata are required to gain better insights into the relationship with Haplographium and its taxonomic placement.
Ruzenia O. Hilber, The Genus Lasiosphaeria and Allied Taxa (Kelheim): 7 (2002)
Saprobic on wood. Sexual morph: Ascomata perithecial, scattered to gregarious, superficial, subglobose to obovoid, dark brown to black, rough, ostiolate, with papilla, periphysate. Peridium membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- apical ring. Ascospores allantoid, 0–1-septate, hyaline, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Hilber and Hilber 2002).
Notes: The monotypic genus Ruzenia was introduced as similar to Lasiosphaeria (Hilber and Hilber 2002). Subsequently, its generic type, Ruzenia spermoides, was clarified as member of Helminthosphaeriaceae based on phylogenetic analyses (Miller and Huhndorf 2004a; Miller et al. 2014). In this study, Ruzenia is similar to Hilberina, but differs in having less ascomata setae, lacking basal spines on the ascospores and being hyaline. It is basal to Helminthosphaeria plumbea and Hilberina munkii in Helminthosphaeriaceae (90%ML/0.93BY, Fig. 21).
Type species: Ruzenia spermoides (Hoffm.: Fr.) O. Hilber ex A. N. Mill. & Huhndorf, Mycological Research 108 (1): 31 (2004)
Basionym: Sphaeria spermoides Hoffm., Veg. Crypt. 2: 12 (1790)
Facesoffungi number: *****??; Fig. 25
Saprobic on wood. Sexual morph: Ascomata 385–620 × 360–530 µm (x̄ = 485 × 465 µm, n = 10), perithecial, scattered to gregarious, superficial, subglobose to obovoid, dark brown to black, rough, ostiolate, with papilla, periphysate. Peridium 75–105 µm (x̄ = 90 µm, n = 30) wide, membranaceous, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 3–7.5 µm wide, septate, constricted at the septum, unbranched, cylindrical, tapering towards the rounded apex. Asci (100–)110–130(–150) × 8–12 µm (x̄ = 125 × 10 µm, n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with J- apical ring. Ascospores 20–26 × 3–5.5 µm (x̄ = 23 × 4 µm, n = 50), bi-seriate, allantoid, 0–1-septate, hyaline, smooth-walled, guttulate. Asexual morph: Undetermined.
Material examined: Sweden, Uppland, Bladaker; on dead, decorticated wood of Tilia cordata, 20 September 1992, N. Lundqvist (S-F 242341); Sweden, Uppland, Österaker-Östra Ryd, on dead, decorticated wood, 17 May 1983, N. Lundqvist (S-F 242342).
Known hosts and distribution: On dead decorticated wood in Czech Republic, Denmark, Germany (type locality), Sweden and USA (Hoffmann 1790; Fries 1823a; Miller and Huhndorf 2004a).
Notes: Ruzenia spermoides strains clustered in Helminthosphaeriaceae with 100%ML/1.00BY support and this clade is basal to Helminthosphaeria plumbea and Hilberina munkii (90%ML/0.93BY, Fig. 21) in the phylogenetic analysis (Miller and Huhndorf 2004a; Miller et al. 2014). In this study, we were unable to obtain the type material. Therefore, we re-examined two authentic samples collected by Lundqvist in Sweden.
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 182 (1986)
Sordariales comprises Bombardiaceae, Chaetomiaceae, Diplogelasinosporaceae, Lasiosphaeriaceae, Lasiosphaeridaceae, Naviculisporaceae, Podosporaceae, Neoschizotheciaceae, Strattoniaceae, Sordariaceae and Zygospermellaceae as clarified by phylogenetic analyses (Hawksworth and Eriksson 1986; Huhndorf et al. 2004b; Kruys et al. 2014; Maharachchikumbura et al. 2016; Hongsanan et al. 2017; Hyde et al. 2017, 2020; Marin-Felix et al. 2020; this study). Most taxa of Sordariales occur in terrestrial or/and aquatic habitats as saprobes and have ascospores with various appendages or different wall patterns (Maharachchikumbura et al. 2015, 2016; Hyde et al. 2020). This order is basal to Phyllachorales and Boliniales in Sordariomycetidae (59%ML, Fig. 1).
Bombardiaceae S.K. Huang, Maharachch. & K.D. Hyde, fam. nov.
Index Fungorum number: IF558344
Etymology: Named after Bombardia, the oldest genus in this family.
Coprophilous or saprobic on soil or wood. Sexual morph: Ascomata perithecial or cleistothecial, solitary to scattered or gregarious, immersed to semi-immersed to superficial, globose to subglobose, glabrous or hairy, yellow to dark brown to black, sometimes semitransparent, membranaceous or coriaceous. Paraphyses numerous, filiform to cylindrical, septate, evanescent. Asci 4–8-spored, unitunicate, cylindrical to clavate, pedicellate, usually with J- apical ring, sometimes with apical globule. Ascospores uni-seriate or bi-seriate, ellipsoidal or cylindrical or maraca-shaped, 0–multi-septate, hyaline to brown to dark brown, verrucose or smooth-walled, sometimes with gelatinous sheath or irregular appendage(s). Asexual morph: Hyphomycetous. Conidiophores arising from aerial hyphae, erect, inflated at the base. Phialides monophialidic, smooth-walled, hyaline, cylindrical, unbranched or branched. Conidia small, hyaline to pale yellow, ellipsoidal to oval, aseptate, smooth-walled, forming globose masses on the apex of the phialides (adapted from Malloch and Cain 1971; Lundqvist 1964, 1972; Udagawa and Furuya 1974; Khan and Krug 1991; Krug and Scott 1994; Bell 2005; Davison et al. 2008).
Type genus: Bombardia (Fr.) P. Karst. 1873
Notes: Marin-Felix et al. (2020) mentioned that Apodospora, Bombardia, Bombardioidea, Fimetariela and several species of Cercophora, Podospora and Zopfiella formed a low support clade. In this study, these taxa clustered (85%ML) and is sister to Sordariaceae (83%ML, Fig. 26).
The species of this cluster have diverse morphological characteristics. Most of them are coprophilous fungi, except for Zopfiella attenuata collected from soil and Bombardia bombarda found on hardwood (Lundqvist 1972; Udagawa and Furuya 1974). They have hairy or glabrous, membranaceous (sometimes with carbonaceous neck), or coriaceous ascomata (Malloch and Cain 1971; Lundqvist 1972; Udagawa and Furuya 1974; Khan and Krug 1991; Krug and Scott 1994; Bell 2005; Davison et al. 2008). Zopfiella attenuata and Z. pleuropora have cleistothecial ascomata and ascospores comprise fusiform to ellipsoidal, brown upper cell(s) and a conical hyaline lower cell, without appendages or sheath (Malloch and Cain 1971; Udagawa and Furuya 1974); the other members have perithecial ascomata and ascospores have appendages or surround the gelatinous sheath (Lundqvist 1964, 1972; Khan and Krug 1991; Krug and Scott 1994; Krug 1995; Bell 2005; Davison et al. 2008). Apodospora peruviana, A. simulans, A. gotlandica, Bombardioidea anartia and Fimeteriella rabenhorstii have broadly fusiform to ellipsoidal ascospores surrounded by gelatinous sheath (Lundqvist 1964, 1972; Krug and Scott 1994; Krug 1995; Davison et al. 2008). All Apodospora species have the gelatinous sheath with an apical invagination (Lundqvist 1972; Krug and Scott 1994; Krug 1995; Davison et al. 2008). Fimeteriella rabenhorstii is the only species in the cluster with 4-spored asci (Lundqvist 1964; Krug 1995). Bombardioidea anartia and Podospora appendiculata have a special coriaceous peridium similar to Bombardia bombarda (Lundqvist 1972; Krug and Scott 1994). Whereas, Bombardia bombardia, Cercophora scortea, Podospora fabiformis, P. fibrinocaudata and P. appendiculata have ascospores comprise an ellipsoidal, brown upper cell and a cylindrical, hyaline pedicel, with terminal appendage(s) at each or one end (Lundqvist 1972; Khan and Krug 1991; Bell 2005). Podospora fabiformis has ascospores with apical and basal appendages, sometimes two lateral appendages arising from the top of the pedicel (Bell 2005). Podospora fibrinocaudata has ascospores with gelatinous sheath surrounding the pedicel’s basal (Khan and Krug 1991). Bombardia bombarda, Cercophora scortea and Podospora appendiculata have cylindrical to clavate asci with a subapical or apical globule and ascospores with appendages at each end (Lundqvist 1972).
Apodospora Cain & J.H. Mirza, Can. J. Bot. 48(5): 891 (1970)
Coprophilous or saprobic on wood. Sexual morph: Ascomata perithecial, gregarious to scattered, immersed to semi-immersed, globose to subglobose, dark brown to black, glabrous or hairy. Peridium membranaceous, composed of hyaline to brown cells of textura angularis to textura prismatica. Paraphyses numerous, filiform, septate, evanescent. Asci 8-spored, unitunicate, ellipsoidal, pedicellate, blunt at the apex. Ascospores oval to ellipsoidal, aseptate, brown to dark brown, verrucose, surrounded by gelatinous sheath, usually with an apical invagination. Asexual morph: Hyphomycetous. Mycelium grey-brown, with numerous phialides scattered on hyphae. Conidia small, ovate masses on the apex of the phialides (adapted from Cain and Mirza 1969).
Notes: Cain and Mirza (1969) introduced Apodospora typified by A. simulans, which has oblong to ellipsoidal ascospores surrounded by a gelatinous sheath and oval to globose small conidia masses on the phialides. This genus resembles Sordaria in having ascospores surrounded by gelatinous sheath (Cain and Mirza 1969). Six species are accommodated in this genus (Cain and Mirza 1969; Lundqvist 1972; Fakirova 1973; Muroi et al. 1987), and A. gotlandica, A. peruviana, and A. simulans have molecular data (Kruys et al. 2014). In this study, Apodospora species clustered in Bombardiaceae in the phylogenetic analysis (100%ML/1.00BY, Fig. 26).
Type species: Apodospora simulans Cain & J.H. Mirza, Can. J. Bot. 48(5): 891 (1970)
Facesoffungi number: *****??; Fig. 27
Coprophilous. Sexual morph: Ascomata 380–530 × 250–400 µm (x̄ = 410 × 275 µm, n = 5), perithecial, gregarious to scattered, immersed to semi-immersed, globose to subglobose, dark brown to black, ostiolate, with papilla. Peridium 25–50 µm (x̄ = 38 µm, n = 30) wide, membranaceous, comprising two layers, outer layer composed of pale brown to brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 3–4 µm wide, filiform, septate, evanescent. Asci (200–)220–280(–350) × 12–20 µm (x̄ = 250 × 16 µm, n = 30), 8-spored, unitunicate, ellipsoid, pedicellate, apex blunt, with J- apical ring distinct. Ascospores (15–)18–21(–25) × 8–10.5 µm (x̄ = 20 × 9 µm, n = 50), uni-seriate, oblong to oval, aseptate, brown to dark brown, verrucose, surrounded by gelatinous sheath 2.5–6 µm diam. Asexual morph: Hyphomycetous. Mycelium grey-brown, with numerous phialides scattered on the hyphae. Phialides pale brown, with flaring collarette. Conidia small, ovate forming globose masses on the apex of the phialides (adapted from Cain and Mirza 1969).
Material examined: Sweden, Uppland, Danmark par., 500m E. of Grynbacken, in coniferous forest, on old moose dung, 18 May 1959, N. Lundqvist (S-F122911).
Known hosts and distribution: On dung of moose (Alces americanus) in Canada (type locality) (Cain and Mirza 1969); on moose dung in Canada, Norway and Sweden (Lundqvist 1972; Kruys et al. 2014).
Notes: We re-examined authentic material (S-F122911) of Apodospora simulans which was determined by Cain (mentioned in the label of material).
Bombardia (Fr.) P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 20 (1873)
Saprobic on wood. Sexual morph: Ascomata perithecial, immersed to semi-immersed, solitary or gregarious, oval to subglobose, coriaceous to membranaceous, dark brown to black, glabrous or surrounded by branched hairs, ostiolate, with papilla. Paraphyses filiform, branched. Asci 8-spored, unitunicate, ellipsoidal to cylindrical, pedicellate, rounded or truncate apex, evanescent. Ascospores hyaline and cylindrical when young, upper cell becoming swollen to fusiform or oval, brown; lower cell cylindrical or subglobose, hyaline, smooth-walled, with guttules. Asexual morph: Undetermined (adapted from Fries 1849).
Notes: Fries (1849) introduced Sphaeria subgenus, Bombardia, based on ‘Sphaeria (Bombardia) fasciculata’, and Karsten (1873) raised this species to genus level and established Bombardia. Saccardo (1882) clarified ‘Sphaeria (Bombardia) fasciculata’ (≡ Sphaeria fasciculata Fr.) and S. bombarda Batsch. as conspecific, and named Bombardia fasciculata which is a homonym of B. fasciculata Rab. (Batsch 1786; Rabenhorst 1851). However, the illegitimate B. fasciculata is still used (Maharachchikumbura et al. 2016; Index Fungorum 2020) and caused some confusion. Bombardia bombarda (= Sphaeria bombarda Batsch.) was recommended as the generic type instead of B. fasciculata (Schröter 1894; Lundqvist 1972), and its molecular data was sequenced later (Miller and Huhndorf 2005; Spatafora et al. 2006). In this study, Bombardia is sister to Bombardioidea in Bombardiaceae (100%ML/1.00BY, Fig. 26).
Type species: Bombardia bombarda (Batsch) J. Schröt., in Cohn, Krypt.-Fl. Schlesien (Breslau) 3.2(1–2): 302 (1893)
Basionym: Sphaeria bombarda Batsch, Elench. fung. (Halle): 271 (1786)
Facesoffungi number: *****??; Fig. 28
Saprobic on wood. Sexual morph: Ascomata 1–1.5 × 0.4–0.8 mm (x̄ = 1.2 × 0.6 mm, n = 5), perithecial, immersed to semi-immersed, gregarious, oval to subglobose, coriaceous, dark brown to black, glabrous, ostiolate papilla, with 2.5 µm wide periphyses. Peridium 75–120 µm (x̄ = 95 µm, n = 30) wide, coriaceous, comprising four layers, outer layer composed of amorphous brown to black material; middle layer composed of hyaline to brown cells of textura intricata, and dark brown, strongly tight cells of textura angularis; inner layer composed of pale brown to hyaline cells of textura prismatica. Paraphyses 2–3 µm wide, filiform, septate, evanescent. Asci (180–)200–230(–270) × 10–15 µm (x̄ = 230 × 12 µm, n = 30), 8-spored, unitunicate, ellipsoidal to cylindrical, pedicellate, apex rounded, with apical globule, evanescent. Ascospores bi-seriate, hyaline and cylindrical when young, upper cell becoming swollen to fusiform, brown, (10–)12–15 × 5–10 µm (x̄ = 13.5 × 8 µm, n = 50), with an apical germ pore, sometimes with a tiny, hyaline, unconspicuous decoration on the apex; lower cell cylindrical, hyaline, (20–)24–30 × 3.5–6 µm (x̄ = 26 × 4.5 µm, n = 50), smooth-walled, guttulate, with a terminal conical appendage at the base, evanescent. Asexual morph: Undetermined.
Material examined: Italy, Veneto, Treviso, Cansiglio, on dead trunks of Fagus, October 1874, P.A. Saccardo (S-F750); Ukraine, Lvivska oblast, Stryi, riverbank near Wierzany, on stems of Alnus, 28 October 1916, F. Petrak (S-F753).
Known hosts and distribution: On dead, decorticated wood in Germany (type locality) (Saccardo 1882); on dead wood in Belgium, Denmark, England, Finland, France, Germany, Hungary, Italy, New Zealand, Poland, Russia, Sweden, UK, Ukraine and USA (Lundqvist 1972; Miller and Huhndorf 2005).
Notes: We re-examined authentic materials of Bombardia bombarda, F750 and F753, collected by Saccardo and Petrak respectively. The molecular data of B. bombarda were sequenced (Miller and Huhndorf 2005; Spatafora et al. 2006), and in this study, B. bombarda (SMH4821) is sister to Bombardioidea anartia (HHB99-1) (100%ML/1.00BY, Fig. 26).
Bombardioidea C. Moreau ex N. Lundq., Symb. bot. upsal. 20(no. 1): 274 (1972)
Coprophilous. Sexual morph: Ascomata perithecial, superficial or semi-immersed, gregarious or scattered, ovoid to subglobose, coriaceous to membranaceous, dark brown to black, verrucose, ostiolate, with papilla. Peridium coriaceous, outer layer composed of amorphous brown to black material; middle layer composed of hyaline to brown cells of textura intricata, and dark brown, strongly tight cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses filiform, septate. Asci 4–8-spored, unitunicate, ellipsoidal to cylindrical, pedicellate, usually with apical ring distinct, evanescent. Ascospores uni- or bi-seriate, ellipsoidal, ovoid or oblong, hyaline when young, becoming brown when mature, aseptate, verrucose, with a germ pore at each end, surrounded by gelatinous sheath, guttulate. Asexual morph: Hyphomycetous, phialophora-like. Conidiophores brown to dark brown, septate, branched, clusters of several phialides, producing conidia in chains or forming globose masses on the apex. Conidia numerous, aseptate, hyaline to pale brown, globose to subglobose (adapted from Niessl 1872; Moreau 1953; Lundqvist 1972).
Notes: Sordaria bombardioides was transferred to Bombardioidea as its generic type (Niessl 1872; Moreau 1953; Lundqvist 1972). Species of this genus are coprophilous usually reported on leporid dung (Lundqvist 1972; Krug and Scott 1994). Huhndorf et al. (2004b) sequenced molecular data for Bombardioidea anartia and in this study, it is sister to Bombardia bombarda (100%/ML/1.00BY) in Bombardiaceae (Fig. 26).
Type species: Bombardioidea bombardioides (Auersw.) in Lundqvist, Symb. bot. upsal. 20(no. 1): 277 (1972)
Basionym: Sordaria bombardioides Auersw., in Niessl, Verh. nat. Ver. Brünn 10: 187 (1872)
Facesoffungi number: *****??; Fig. 29
Coprophilous. Sexual morph: Ascomata 0.5–1.5× 0.5–0.8 mm (x̄ = 1.2 × 0.6 mm, n = 5), perithecial, superficial, gregarious, oval to subglobose, coriaceous, dark brown to black, verrucose, ostiolate, with papilla. Peridium 120–160 µm (x̄ = 135 µm, n = 30) wide, coriaceous, comprising four layers, outer layer composed of amorphous brown to black material; middle layer composed of hyaline to brown cells of textura intricata, and dark brown, strongly tight cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses filiform, septate. Asci (140–)145–160 × 15–25 µm (x̄ = 150 × 20 µm, n = 30) ascospore-bearing part, 8-spored, unitunicate, ellipsoidal to cylindrical, pedicellate, about 80 µm long, apex rounded, evanescent. Ascospores (20–)22–27(–30) × 10–17 µm (x̄ = 25 × 12 µm, n = 50), uni-seriate, ellipsoidal, hyaline when young, becoming brown when mature, aseptate, verrucose, with a germ pore at each end, surrounded by gelatinous sheath 1–6 µm wide, guttulate. Asexual morph: Hyphomycetous. Conidiophores brown to dark brown, becoming hyaline towards the apex, septate, branched, with clusters of 2–7 phialides. Phialides subhyaline to dark brown, subglobose to ampulliform. Conidia numerous, aseptate, hyaline to pale brown, globose to subglobose (adapted from Krug and Scott 1994).
Material examined: Czech Republic, Jihomoravsky Kraj, Brünn, on hare dung, 9 August 1883, G. Niessl von (S-F726); Poland, Silesia, Karlsbrunn, on hare dung, August 1902, G. Niess von (S-F729).
Known hosts and distribution: On hare dung in Germany (type locality) (Niessl 1872); on hare and rabbit dung, occasionally on cow, roe-deer, horse, mose, porcupine and sheep dung in Austria, Bulgaria, Canada, Croatia, Czech Republic, Denmark, England, Japan, Israel, Germany, Hungary, Netherlands, Poland, Sweden and USA (Lundqvist 1972; Krug and Scott 1994).
Notes: We re-examined authentic specimens of Bombardioidea bombardioides, F726 and F729, collected by Niessl and determined by Lundqvist (mentioned in the label of material). In this study, we provided hand-drawings of the asexual morph (Fig. 31 r) from Krug and Scott (1994).
Fimetariella N. Lundq., Bot. Notiser 117: 239 (1964)
Coprophilous. Sexual morph: Ascomata perithecial, solitary to scattered, immersed to semi-immersed, pyriform to subglobose, brown to black, ostiolate. Peridium comprising membranaceous to coriaceous, pale brown to brown cells. Paraphyses numerous, filiform, evanescent. Asci 4–8-spored, unitunicate, cylindrical, pedicellate, apex rounded. Ascospores uni-seriate, ellipsoidal to oval, aseptate, hyaline to brown to dark brown, with a germ pore at one or each end, surrounded by hyaline, gelatinous sheath. Asexual morph: Hyphomycetous. Phialides abundant, hyaline, reduced to collarettes producing conidia. Conidia ovoid, hyaline, aseptate (adapted from Lundqvist 1964b; Kruys et al. 2014).
Notes: Lundqvist (1964b) transferred Sordaria rabenhrstii to Fimetariella as its type species. It is characterized by pyriform ascomata and aseptate ascospores surrounded by a hyaline, gelatinous sheath. Kruys et al. (2014) sequenced F. rabenhorstii (20410), which was isolated from moose dung in Sweden and collected by Lundqvist. This strain is basal to Zopfiella attenuata and Z. pleuropora in Bombardiaceae (83%ML, Fig. 26).
Type species: Fimetariella rabenhorstii (Niessl) N. Lundq., Bot. Notiser 117: 239 (1964)
Basionym: Sordaria rabenhorstii Niessl, in Rabenhorst, Fungi europ. exsicc.: no. 1528 (1873)
Facesoffungi number: *****??; Fig. 30
Coprophilous. Sexual morph: Ascomata 400–500 × 350–400 µm (x̄ = 430 × 380 µm, n = 10), perithecial, solitary, immersed to semi-immersed, ovoid to subglobose, membranaceous, black, ostiolate, with papilla. Peridium 24–40 µm (x̄ = 30 µm, n = 30) wide, outer layer composed of pale brown to brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, filiform, evanescent, broader towards the base, without constrictions, septate. Asci (95–)110–128(–132) × 9.5–13 µm (x̄ = 120 × 12 µm, n = 20), 4-spored, unitunicate, cylindrical, pedicellate, apex rounded with thickened wall. Ascospores (10–)13–15(–18) × 7.5–10 µm (x̄ = 14 × 8.5 µm, n = 40), uni-seriate, ellipsoidal to oval, aseptate, hyaline when young, becoming brown to dark brown, surrounded by hyaline, gelatinous sheath 1–2.5 µm wide. Asexual morph: Undetermined.
Material examined: Sweden, Uppland, Älvkarleby par., SE of the mire Gustavmyrarna (4 km SW of Skutskär), in coniferous forest, on elk dung (Alces alces) in moist chamber, 8 June 1996, N. Lundqvist (S-F66877); Czech Republic, Jihomoravsky kraj, Brünn, on dung of Caprearum, G. Niessl (S-F66507, holotype); Czech Republic, Jihomoravsky kraj, Brünn, on dung of Caprearum, G. Niessl (S-F66508, isotype).
Known hosts and distribution: On dung of roe and hare in the Czech Republic (type locality) (Saccardo 1882); on dung of deer, elk, goat, moose, rabbit, wapiti in Belgium, Canada, Denmark, Sweden and USA (Krug 1995).
Notes: We re-examined the type materials of Sordaria rabenhorstii, which were determined as Fimetariella rabenhrstii by Lundqvist (mentioned in the label of material). It is in good condition.
Lasiosphaeriaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 50 (1932)
Coprophilous or saprobic on soil or wood. Sexual morph: Ascomata perithecial or cleistothecial, solitary to scattered or gregarious, immersed to semi-immersed to superficial, globose to subglobose, dark brown to black. Paraphyses numerous, filiform to cylindrical, septate, evanescent. Asci 4–8-spored, unitunicate, cylindrical to clavate, pedicellate. Ascospores uni-seriate or bi-seriate, ellipsoidal to oval or irregular, 0–multi-septate, hyaline to brown to dark brown, verrucose or smooth-walled, with gelatinous sheath or irregular appendage(s). Asexual morph: Hyphomycetous. Conidiophores arising from aerial hyphae, erect, inflated at the base. Conidia small, oval to globose, ovate forming globose masses on the apex of the phialides (adapted from Nannfeldt 1932; Lundqvist 1972).
Type genus: Lasiosphaeria Ces. & De Not. 1863
Notes: Lasiosphaeriaceae is typified by Lasiosphaeria and has black ascomata, cylindrical to clavate asci and varied ascospores (Cesati and de Notaris 1863; Nannfeldt 1932; Lundqvist 1972; Wang et al. 2019; Marin-Felix et al. 2020). Species in Lasiosphaeriaceae were reported to be similar to taxa in Nitschkiaceae, Sordariaceae and Tripterosporaceae based on their ascospores with various ornamentations and appendages, such as Bombardia, Lasiosphaeria (Munk 1957; Carroll and Munk 1964; Lundqvist 1972; Barr 1990; Kruys et al. 2014). The traditional classification of Lasiosphaeriaceae is based on morphology; obviously, it is inconsistent with molecular phylogeny (Miller and Huhndorf 2004; 2005; Wang et al. 2019; Marin-Felix et al. 2020). Huhndorf et al. (2004b) confirmed that Lasiosphaeriaceae is polyphyletic based on LSU sequence data. Subsequently, many polyphyletic genera in this family were repositioned based on the sequence data (Cai et al. 2006a; Chang et al. 2010; Kruys et al. 2014; Wang et al. 2019; Marin-Felix et al. 2020). Wang et al. (2019) redefined three genera, Cladorrhinum, Podospora and Triangularia, and established Podosporaceae based on phylogenetic analysis. Wijayawardene et al. (2020) listed 32 genera in Lasiosphaeriaceae. Marin-Felix et al. (2020) re-classified Cercophora, Diplogelasinospora, Immersiella, Jugulospora, Rinaldiella, Schizothecium, Zygopleurage and some species of Apiosordaria and Podospora to establish Diplogelasinosporaceae, Naviculisporaceae and Schizotheciaceae in Sordariales based on morphology and phylogenetic analyses.
In this study, based on phylogenetic analyses, we divide the remaining Lasiosphaeriaceae taxa into five clades (Fig. 26): i) Bombardiaceae clade: This is a sister clade of Sordariaceae (83%ML). Apodospora, Bombardia, Bombardioidea, Fimetariella and some species of Cercophora, Podospora and Zopfiella clustered in this clade (85%ML); ii) Lasiosphaeriaceae clade: This clade is regarded as Lasiosphaeriaceae sensu stricto (Marin-Felix et al. 2020) because several typical Lasiosphaeria species with molecular data are included in this clade; however, the ex-type strain of L. ovina has not been sequenced. This clade is sister to Naviculisporaceae (82%ML). It clusters Anopodium, Bellojisia, Corylomyces, Lasiosphaeria, Mammaria, Zopfiella and some species of Cercophora, Podospora and Ramophialophora in 96%ML support; iii) Lasiosphaeridaceae clade: This is an orphan clade in Sordariales, which clusters Lasiosphaeris species only (100%ML/1.00BY); iv) Strattoniaceae clade: This is an unstable clade which includes Strattonia species and is sister to Diplogelasinosporaceae with poor support; v) Zygospermellaceae clade: This clade contains Episternus onthosphagi, Zygospermella insignis and Ramophialophora petraea (71%ML/0.99BY), and is an orphan clade in Sordariales (88%ML/1.00BY).
In addition, the generic type of Apodus is nested in Neoschizotheciaceae in phylogenetic analysis (Fig. 26). The polyphyletic Arnium and Ramophialophora, which have no molecular data of their type species, are placed in Sordariales genera incertae sedis. Given the diversity of the morphology of lasiosphaeriaceous taxa and locating each genus more accurately, we put the genera (Biconiosporella, Camptosphaeria, Diffractella, Emblemospora, Eosphaeria, Periamphispora, Tripterosporella) without molecular data in Sordariales genera incertae sedis pending further studies.
Thaxteria is typified by T. didyma, which was synonymized under Bertia didyma (Saccardo 1891; Mugambi and Huhndorf 2010). Apiosordaria is the synonym of Triangularia based on its available molecular data of type species (Wang et al. 2019). Therefore, they will not be discussed in this study.
Marin-Felix et al. (2020) redefined Lasiosphaeriaceae and proposed that Lasiosphaeriaceae sensu stricto may divided into two clades; Lasiosphaeria clade and the clade includes the generic type of Anopodium, Bellojisia, Corylomyces, Zopfiella and several other species Cercophora sulphurella, C. sparsa, Podospora didyma and Ramophialophora humicola (clade I, Fig. 26). In this study, we added the molecular data which was derived from Vu et al. (2019) of Mammaria and found that this genus is basal to Lasiosphaeria clade and clade I in Lasiosphaeriaceae (96%ML, Fig. 26). Therefore, six genera are accommodated in Lasiosphaeriaceae based on phylogenetic analysis.
This family has high morphological variability. Ramophialophora humicola is a phialophora-like hyphomycetes (Madrid et al. 2010). All sexual species have perithecial ascomata, except for Zopfiella tabulata, which has cleistothecial ascomata; Anopodium ampullaceum, Cercophora sulphurella, C. sparsa, Mammaria echinnobotryoides, Podospora didyma and Zopfiella tabulata have cylindrical ascospores with swollen apical cell and terminal appendage(s) present or absent, whereas Bellojisia rhynchostoma and Corylomyces seleonsporus have allantoid to reniform ascospores (Saccardo 1878, 1906; Lundqvist 1964, 1972; Guarro et al. 1991; Stchigel et al. 2006; Réblová 2008). However, more fresh collections and phylogenetic data are requested to determine the affinities and taxonomic placement for the members in this clade.
Anopodium N. Lundq., Bot. Notiser 117: 356 (1964)
Coprophilous. Sexual morph: Ascomata perithecial, scattered, semi-immersed, or erumpent through the bark of host, or superficial, globose, pale brown to dark brown, ostiolate, with black papilla, surrounded by setae. Peridium membranaceous, composed of hyaline to brown cells of textura angularis to textura prismatica. Asci 8-spored, unitunicate, ellipsoidal, pedicellate, blunt at the apex. Ascospores subglobose to oval, collapsing when dry, aseptate, hyaline when young, becoming dark brown, smooth-walled, with an apical germ pore. Asexual morph: Undetermined (adapted from Lundqvist 1964a).
Notes: Anopodium was introduced by Lundqvist (1964a) with two species A. ampullaceum (type) and A. epile. This coprophilous genus is characterized by pale brown ascomata with black necks and ascospores with a pedicel at the end (Lundqvist 1964a). Anopodium epile differs from A. ampullaceum in having glabrous ascomata (Lundqvist 1964a). Kruys et al. (2014) sequenced A. ampullaceum. In this study, Fig. 26 shows Anopodium nests in Lasiosphaeriaceae clade I.
Type species: Anopodium ampullaceum N. Lundq., Bot. Notiser 117: 356 (1964)
Facesoffungi number: *****??; Fig. 31
Coprophilous. Sexual morph: Ascomata 400–550 × 275–400 µm (x̄ = 480 × 335 µm, n = 10), perithecial, scattered, semi-immersed, or erumpent through bark of host, or superficial, globose, pale brown to dark brown, ostiolate, with black papilla, surrounded by setae 1.5–4.5 wide. Necks 180–200 × 100–135 µm (x̄ = 185 × 120 µm, n = 5), membranaceous, central, dark brown to black, smooth-walled, with hyaline periphyses. Peridium 20–55 µm (x̄ = 40 µm, n = 20) wide, membranaceous, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 4–5 µm wide, numerous, broad, evanescent. Asci (135–)150–170(–200) × 35–45 µm (x̄ = 160 × 40 µm, n = 30), 8-spored, unitunicate, ellipsoidal, pedicellate, apex blunt, apical ring absent, evanescent. Ascospores (25–)28–32(–35) × 15–25 µm (x̄ = 30 × 20 µm, n = 50) for the mature swollen cells, bi-seriate, subglobose to oval, collapsing when dry, aseptate, hyaline, swollen cell towards the apex of ascus when young, becoming dark brown, partly ascospores of their swollen cell toward the apex; partly toward the base when mature, smooth-walled, with an apical germ pore and an evanescent pedicel (10–)18–25 × 1.5–3.5 µm (x̄ = 20 × 2.5 µm, n = 20) at the other end. Asexual morph: Undetermined.
Material examined: Sweden, Jämtland, Mårdsundsbodarna, Hallen par., in subalpine birch forest, on hare dung (Lepus timidus) in moist chamber, Stockholm, 11 July 1988, N. Lundqvist (S-F47944); UK, Midlothian, Edinburgh, Hillend, on rabbit dung (Oryctolagus cuniculus) in moist chamber, 26 December 1996, M.J. Richardson (S-F66862).
Known hosts and distribution: On fresh hare dung in Sweden (type locality) (Lundqvist 1964a); on rabbit dung in Belgium, Norway, Sweden and UK (Lundqvist 1972; Kruys et al. 2014).
Notes: Lundqvist (1964a) discovered Anopodium ampullaceum on hare dung in Sweden, and it was reported in the UK (Kruys et al. 2014). In this study, we re-examined authentic material collected by Lundqvist in Sweden and a sample collected in the UK. We found that most of the swollen ascospores cells in the same ascus from S-F47944 were towards the ascus apex when they were young, becoming partly toward the ascus apex and partly toward the base. Kruys et al. (2014) sequenced A. ampullaceum (MJR 40/07 and E00218015) and found that these strains are related to Bellojisia, Corylomyces and Lasiosphaeria. In this study, A. ampullaceum is sister to Cercophora sulphurella in Lasiosphaeriaceae clade I (87%ML, Fig. 26).
Bellojisia Réblová, Mycologia 100(6): 897 (2008)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary to gregarious, semi-immersed to superficial, carbonaceous, pyriform to subglobose, tomentose, brown to black, ostiolate, with necks, periphysate. Peridium composed of brown to hyaline cells of textura prismatica to textura intricata to textura angularis. Paraphyses septate. Asci 8-spored, unitunicate, cylindrical, pedicellate, with J- apical ring. Ascospores 0–1-septate, navicular to reniform to falciform, dark brown, with a large guttule in each cell and an apical germ pore. Asexual morph: Undetermined (adapted from Réblová 2008).
Notes: The monotypic Bellojisia was established based on Jobellisia rhynchostoma in phylogenetic analysis and its unique reniform ascospores different from Jobellisia (Réblová 2008). In this study, the strain of B. rhynchostoma is basal to Podospora didyma and Zopfiella tabulata in Lasiosphaeriaceae clade I (89%ML, Fig. 26).
Type species: Bellojisia rhynchostoma (Höhn.) Réblová, Mycologia 100(6): 897 (2008)
Fig. 34 i–j
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary to gregarious, semi-immersed to superficial, carbonaceous, pyriform to subglobose, tomentose, brown to black, ostiolate, with necks, periphysate. Peridium comprising two layers, outer layer composed of brown cells of textura prismatica to textura intricata, inner layer composed of hyaline to brown cells of textura angularis. Paraphyses septate. Asci 8-spored, unitunicate, cylindrical, pedicellate, with J- apical ring. Ascospores 0–1-septate, navicular to reniform to falciform, dark brown, with a large guttule in each cell and an apical germ pore. Asexual morph: Undetermined (adapted from Réblová 2008).
Known hosts and distribution: In pericarps of Juglans regia in the Austria (type locality); on decorticated wood of Robinia pseudoacacia in the France (Réblová 2008).
Notes: The reniform ascospores of Bellojisia rhynchostoma are similar to Corylomyces selenosporus and they nested in Lasiosphaeriaceae clade I in phylogenetic analysis (Fig. 26).
Corylomyces Stchigel, M. Calduch & Guarro, Mycol. Res. 110(11): 1362 (2006)
Saprobic on wood. Sexual morph: Ascomata perithecial, gregarious, superficial, pyriform to subglobose, olivaceous-yellow to olive, tomentose, ostiolate, with long necks, surrounded by hairs. Asci 8-spored, unitunicate, cylindrical, pedicellate, evanescent. Ascospores uni-seriate, reniform, 0–1-septate, hyaline to dark brown, with an apical, umbonate germ pore. Asexual morph: Undetermined (adapted from Stchigel et al. 2006).
Notes: The monotypic Corylomyces was reported on hazelnut and is placed in Lasiosphaeriaceae based on the LSU sequence data (Stchigel et al. 2006). In this study, the ex-type of C. selenosporus (CBS 113930) is sister to Cercophora sparsa in Lasiosphaeriaceae clade I (85%ML, Fig. 26).
Type species: Corylomyces selenosporus Stchigel, M. Calduch & Guarro [as 'selenospora'], in Stchigel, Cano, Miller, Calduch & Guarro, Mycol. Res. 110(11): 1362 (2006)
Fig. 42 k
Saprobic on wood. Sexual morph: Ascomata perithecial, gregarious, superficial, pyriform to subglobose, olivaceous-yellow to olive, tomentose, ostiolate, with long necks, surrounded by hairs. Asci 8-spored, unitunicate, cylindrical, pedicellate, evanescent. Ascospores uni-seriate, reniform to lunate, 0–1-septate, hyaline to dark brown, with an apical, umbonate germ pore. Asexual morph: Undetermined (adapted from Stchigel et al. 2006).
Known hosts and distribution: On hazelnut (Corylus avellana) decomposing on soil in France (type locality) (Stchigel et al. 2006).
Notes: Bellojisia rhynchostoma and Corylomyces selenosporus have long necks, tomentose ascomata and 0–1-septate ascospores; whereas the ascospores of the former are lunate and the latter are navicular (Stchigel et al. 2006; Réblová 2008). Cercophora sparsa has cylindical ascospores, which is different from Cor. selenosporus has navicular ascospores (Hilber and Hilber 1979; Stchigel et al. 2006).
Lasiosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 229 (1863)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary or gregarious, superficial to semi-immersed, globose to subglobose, brown, surrounded by white furfuraceous cover, ostiolate, with distinct black papilla. Peridium comprising membranaceous to coriaceous, brown to hyaline cells. Paraphyses filiform. Asci 8-spored, unitunicate, cylindrical, pedicellate. Ascospores cylindrical to ellipsoidal, sometimes slight curved, hyaline to brown, 0–multi-septate, guttulate, smooth-walled. Asexual morph: Hyphomycetous, phialophora-like. Conidiogenous cells phialides, monophialidic, cylindrical to oblong, hyaline, with collarette or collarette absent. Conidia oval to pyriform, hyaline, truncate at the base (adapted from Cesati and de Notaris 1863; Huhndorf et al. 2004b; Miller and Huhndorf 2004a; Miller et al. 2014).
Notes: Lasiosphaeria as the type genus of Lasiosphaeriaceae is characterized by globose ascomata, cylindrical asci and cylindrical to ellipsoidal ascospores (Cesati and de Notaris 1863; Huhndorf et al. 2004b; Miller and Huhndorf 2004a; Miller et al. 2014). Strains of Lasiosphaeria species are close to Anopodium, Bellojisia, Corylomyces and Zopfiella in phylogenetic analysis (Maharachchikumbura et al. 2016; Hyde et al. 2020, Marin-Felix et al. 2020). In this study, the Lasiosphaeria strains is sister to clade I comprises Anopodium, Bellojisia, Corylomyces and Zopfiella (85%ML), and is related to Mammaria (Fig. 26).
Type species: Lasiosphaeria ovina (Pers.) Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 229 (1863)
Basionym: Sphaeria ovina Pers., Syn. meth. fung. (Göttingen) 1: 71 (1801)
Facesoffungi number: *****??; Fig. 32
Saprobic on wood. Sexual morph: Ascomata 360–450 × 340–370 µm (x̄ = 400 × 345 µm, n = 10), perithecial, solitary or gregarious, superficial to semi-immersed, globose to subglobose, brown, membranaceous, surrounded by white furfuraceous cover, ostiolate, with distinct black papilla. Peridium comprising three layers, outer layer composed of white cells of textura intricata, 23–48 µm wide; middle layer coriaceous, composed of brown cells of textura angularis, 23–35 µm wide; inner layer membranaceous, composed of hyaline cells of textura prismatica, 6–16 µm wide. Paraphyses 1–2 µm wide, numerous, filiform. Asci (80–)95–110(–120) × 8–12 µm (x̄ = 100 × 10 µm, n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded with subapical globule, apical ring distinct. Ascospores (25–)28–34(–45) × 2–5 µm (x̄ = 30 × 3 µm, n = 50), bi-seriate, slight curved, cylindrical to geniculate, hyaline and aseptate when young, becoming pale brown and multi-septate when mature, with a large guttule in each cell, smooth-walled. Asexual morph: Undetermined.
Material examined: Sweden, Småland, Högsby par., Massemåla, on decayed branch of apple tree in old pasture, 3 October 1992, N. Lundqvist (S-F123316); Sweden, Uppland, Uppsala, “Predikstolen” N of Lurbo (5 km SSW of Uppsala), on decayed wood in deciduous forest, 7 September 1994, N. Lundqvist (S-F123318); Sweden, Västergötland, Säter par., Ăngen (2 km WNW of Stöpen), on rotten birch log in deciduous forest, 13 September 2001, N. Lundqvist (S-F123375).
Known hosts and distribution: On rotten trunk in Denmark, England, Germany (type locality), Sweden, Turkey and USA (Persoon 1801; Miller and Huhndorf 2004b; Dülger and Akata 2016; Vu et al. 2019).
Notes: Several Lasiosphaeria ovina were sequenced, but the ex-type strain is lacking (Miller and Huhndorf 2004b; Fernández et al. 2006; Raja et al. 2011; Vu et al. 2019). In this study, strains of L. ovina (CBS 126299 and SMH 1538) are nested in Lasiosphaeria clade (Fig. 26). We could not obtain the type material. Therefore, we re-examined authentic specimens collected by Lundqvist.
Lasiosphaeria rehmiana Henn., Verh. bot. Ver. Prov. Brandenb. 40: 135 (1898) [1899]
Facesoffungi number: *****??; Fig. 33
Saprobic on wood. Sexual morph: Ascomata 220–290 × 240–270 µm (x̄ = 240 × 255 µm, n = 10), perithecial, gregarious, superficial, globose to subglobose, brown, membranaceous, surrounded by white furfuraceous cover, ostiolate, with distinct black papilla. Setae 3–10 µm wide, brown to dark brown, aseptate, swollen at the base, tapering, with a swollen, hyaline apex. Peridium comprising three layers, outer layer composed of white cells of textura intricata, 10–24 µm wide; middle layer coriaceous, composed of brown cells of textura angularis, 10–24 µm wide; inner layer membranaceous, composed of hyaline cells of textura prismatica, 6–14 µm wide. Paraphyses 2–4 µm wide, numerous, septate, cylindrical. Asci (100–)115–120(–150) × 10–15 µm (x̄ = 118 × 13 µm, n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded. Ascospores (20–)24–30(–35) × (3–)4–6(–7) µm (x̄ = 27 × 5 µm, n = 50), bi-seriate, slight curved, ellipsoidal, 3–5-septate, hyaline when young, becoming brown and swollen at the middle cells when mature, slightly constricted at septum, guttulate, smooth-walled. Asexual morph: Undetermined.
Material examined: Germany, Berlin, Hoard. Bot. Berol., Warmhaus, on an imported wood from Cameroon, on decayed wood, May 1891, P. Hennings (S-F6328, holotype).
Known hosts and distribution: On dead wood in Germany (type locality) (Hennings 1898).
Notes: Lasiosphaeria rehmiana was introduced as a species closely related to Chaetosphaeria in having ellipsoidal, septate ascospores (Hennings 1898). Réblová and Huhndorf subsequently re-examined the type material, Lasiosphaeria rehmiana (F6328) and considered it as Melanochaeta hemipsila (Chaetosphaeriaceae) (mentioned in the label of material).
Mammaria Ces. ex Rabenh., Bot. Ztg. 12: 190 (1854)
Saprobic on wood or leaves. Sexual morph: Ascomata perithecial, gregarious, superficial, globose to subglobose, black, ostiolate. Asci 8-spored, unitunicate, cylindrical. Ascospores bi-seriate, cylindrical to geniculate, hyaline to brown, with a germ pore at each end. Asexual morph: Hyphomycetous. Mycelium hyaline to pale brown, branched. Conidiophores mononematous, erect, hyaline to pale brown, branched. Conidia globose, hyaline to pale brown, smooth-walled (adapted from Cesati 1854; del Valle Catania et al. 2011).
Notes: Mammaria was introduced as a hyphomycetous genus and is typified by M. echinobotryoides (Cesati 1854). Saccardo (1886) transferred this species to Trichosporum (Dothideomycetes). Vu et al. (2019) sequenced M. echinobotryoides, and in this study, M. echinobotryoides is branched out of Lasiosphaeria and clade I in Lasiosphaeriaceae (96%ML, Fig. 26). Pseudocercophora ingoldii and Cercophora solaris were reported as the sexual morph of M. echinobotryoides (Subramanian and Sekar 1986; del Valle Catania et al. 2011), and Réblová et al. (2016) recommended using the old Mammaria over Pseudocercophora. In this study, Mammaria echinobotryoides and C. solaris clustered (100%ML/1.00BY) in Lasiosphaeriaceae (Fig. 26).
Type species: Mammaria echinobotryoides Ces., in Rabenhorst, Klotzschii Herb. Viv. Mycol. 19: no. 1859 (1854)
Synonym: Cercophora solaris (Cooke & Ellis) R. Hilber & O. Hilber, Z. Mykol. 45(2): 221 (1979)
Fig. 34 k–m
Saprobic on wood. Sexual morph: Ascomata perithecial, gregarious, superficial, globose to subglobose, black, ostiolate, with papilla. Asci 8-spored, unitunicate, cylindrical. Ascospores bi-seriate, aseptate, cylindrical to geniculate, hyaline, becoming 0–1-septate, upper cell triangular, brown; lower cell cylindrical, hyaline, with a germ pore at each end. Asexual morph: Hyphomycetous. Mycelium hyaline to pale brown, branched. Conidiophores mononematous, erect, hyaline to pale brown, branched. Chlamydospores oval to ellipsoid, pale brown to brown, slightly apiculate, flattened basal, with a longitudinal germ slit. Conidia globose, hyaline to pale brown, smooth-walled (adapted from Hennebert 1968; del Valle Catania et al. 2011)
Known hosts and distribution: On hollow trunk in Belgium, Canada, Italy (type locality), Singapore and USA (Saccardo 1886; Subramanian and Sekar 1986; Vu et al. 2019).
Notes: Mammaria echinobotryoides was found as a hyphomycetous species (Cesati 1854). Later Cercophora solaris was introduced as the sexual morph of M. echinobotryoides based on similar cultural characteristics (Subramanian and Sekar 1986, del Valle Catania et al. 2011). Cercophora solaris has cylindrical to geniculate ascospores similar to Lasiosphaeria; phylogenetically, the strains of Cercophora solaris and M. echinobotryoides clustered (100%ML/1.00BY, Fig. 26). Thus, we recommend to transfer C. solaris as M. echinobotryoides.
Zopfiella G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 56 (1884)
Saprobic on wood or coprophilous. Sexual morph: Ascomata cleistothecial, scattered, superficial, globose to subglobose, brown, surrounded by septate, olive to yellow-green hairs. Peridium membranaceous, comprising brown to olivaceous brown cells of textura angularis. Asci 4–8-spored, unitunicate, cylindrical to clavate, evanescent. Ascospores bi-seriate, upper cell(s) ellipsoidal to fusiform, aseptate to multi-septate, pale brown to dark brown, with an apical germ pore and a truncate base; lower cell hyaline, cylindrical, gelatinous, evanescent. Asexual morph: Hyphomycetous, humicola-like. Conidia globose, hyaline to pale brown (adapted from Winter 1884; Guarro et al. 1991).
Notes: Zopfiella is characterized by cleistothecial ascomata and ascospores comprise ellipsoidal, brown upper cell(s) and a cylindrical, hyaline pedicel (Winter 1884; Guarro et al. 1991). Subsequently, Zopfiella species scattered in Naviculisporaceae, Podosporaceae and Neoschizotheciaceae based on phylogenetic analysis (Wang et al. 2019; Marin-Felix et al. 2020; this study, Fig. 26). However, the generic type, Z. tabulate is sister to Podospora didyma in Lasiosphaeriaceae clade I (100%ML/0.95BY, Fig. 26).
Type species: Zopfiella tabulata (Zopf) G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 56 (1884)
Basionym: Cephalotheca tabulata Zopf, Sber. Gesellschaft naturf. Freunde Berlin 2: 34 (1880)
Fig. 42 m
Coprophilous. Sexual morph: Ascomata cleistothecial, scattered, superficial, globose to subglobose, brown, surrounded by septate, olive to yellow-green hairs. Peridium membranaceous, comprising brown to olivaceous brown cells of textura angularis. Asci 4-spored, unitunicate, cylindrical to clavate, evanescent. Ascospores bi-seriate, ellipsoidal, hyaline when young, becoming upper cell(s) ellipsoidal to fusiform, 0–1-septate, pale brown to dark brown, with an apical germ pore and a truncate base; lower cell hyaline, cylindrical, gelatinous, evanescent. Asexual morph: Undetermined (adapted from Winter 1884; Guarro et al. 1991).
Known hosts and distribution: On dung of hare, porcupine, rabbit and sheep in Germany (type locality) (Winter 1884; Cai et al. 2005).
Notes: Zopfiella tabulata was isolated from dung of hare, rabbit and sheep (Winter 1884; Guarro et al. 1991). Subsequently, Cai et al. (2005) sequenced Zopfiella tabulata (CBS 230.78) collected from porcupine dung. This strain is sister to Podospora didyma (CBS 232.78) in phylogenetic analysis, whereas the latter has perithecial ascomata and ascospores with terminal appendages (Mirza and Cain 1969).
Lasiosphaeridaceae S.K. Huang, Maharachch. & K.D. Hyde, fam. nov.
Index Fungorum number: IF558345
Etymology: Named for Lasiosphaeris, the type genus of this family.
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, subglobose to obpyriform, black, ostiolate, tuberculate or/and surrounded by brown, septate setae or hairs. Asci 8-spored, unitunicate, cylindrical, with J- apical ring and a subapical globule. Ascospores cylindrical to geniculate or sigmoid, 0–multi-septate, ends rounded, hyaline to pale brown, becoming brown to dark brown, ellipsoidal to broadly fusiform at apical cell(s), sometimes with terminal, short, conical gelatinous appendage at each end. Asexual morph: Hyphomycetous, phialophora-like. Conidia oval to globose, hyaline (adapted from Tode 1791; Hilber and Hilber 1979; Miller and Huhndorf 2004a).
Type genus: Lasiosphaeris Clem. 1909
Notes: Lasiosphaeris is similar to Lasiosphaeria and Immersiella in having geniculate or sigmoid ascospores, but Lasiosphaeria has white furfuraceous tissue covering on the surface of the ascomata and Immersiella has immersed ascomata, whereas Lasiosphaeris has superficial, hairy ascomata (Tode 1791; Clements 1909; Miller and Huhndorf 2004a; Kruys et al. 2014). However, these three genera were subsequently found to be far apart in phylogenetic analysis (Miller and Huhndorf 2004a; Kruys et al. 2014; Marin-Felix et al. 2020, this study). Miller and Huhndorf (2004a) regarded Lasiosphaeris species as an independent clade in Lasiosphaeriaceae based on LSU sequence data. Marin-Felix et al. (2020) showed that Lasiosphaeris is sister to Zygospermella based on multi-gene analysis. In this study, Cercophora arenicola, Lasiosphaeris hirsuta and L. hispida form a clade (100%ML/1.00BY) which is nested in Sordariales as an orphan clade (74%ML, Fig. 26). Furthermore, Cercophora arenicola is similar to Lasiosphaeris which has hairy ascomata, cylindrical to clavate asci with a subapical globule. Still, ascospores have swollen apical cells with a terminal appendage at each end (Hilber and Hilber 1979). Therefore, we accepted C. arenicola as Lasiosphaeris arenicola (see the combination list below) and delimited the new family Lasiosphaeridaceae based on Lasiosphaeris.
Lasiosphaeris Clem., Gen. fung. (Minneapolis): 35 (1909)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, subglobose to obpyriform, black, ostiolate, tuberculate or/and surrounded by brown, septate setae or hairs. Asci 8-spored, unitunicate, cylindrical, with J- apical ring and a subapical globule. Ascospores cylindrical to geniculate or sigmoid, 0–multi-septate, ends rounded, hyaline to pale brown, becoming brown to dark brown, ellipsoidal to broadly fusiform at apical cell(s), sometimes with terminal, short, conical gelatinous appendage at each end. Asexual morph: Hyphomycetous, phialophora-like. Conidia oval to globose, hyaline (adapted from Tode 1791; Miller and Huhndorf 2004a).
Notes: Lasiosphaeris was established based on Sphaeria hispida (Clements 1909) and Réblová et al. (2011) introduced Lasiadelphia lasiosphaeriae for the asexual morph of Lasiosphaeris hispida. However, Réblová et al. (2016) recommended to use the old Lasiosphaeris over Lasiadelphia. Lasiosphaeris is characterized by hairy ascomata, cylindrical to clavate asci with apical globule and geniculate or sigmoid ascospores (Tode 1791; Hilber and Hilber 1979), which is related to Immersiella and Lasiosphaeria in morphology, but distinct in phylogenetic analysis. Lasiosphaeris species clustered as an independent clade in Sordariales based on single/multi-gene analysis (Miller and Huhndorf 2004a; Kruys et al. 2014; Marin-Felix et al. 2020). This resulted in Lasiosphaeris being placed in Sordariales genera incertae sedis (Miller and Huhndorf 2004a; Wijayawardene et al. 2020). Since a large number of lasiosphaeriaceous taxa have been repositioned according to the phylogenetic analysis (Marin-Felix et al. 2020), we raised Lasiosphaeris as a novel family, Lasiosphaeridaceae, which includes all Lasiosphaeris species and is branched out of other families in Sordariales (74%ML) in our phylogenetic analysis (Fig. 26).
Type species: Lasiosphaeris hispida (Tode) Clem., Gen. fung. (Minneapolis): [173] (1909)
Basionym: Sphaeria hispida Tode, Fung. mecklenb. sel. (Lüneburg) 2: 17 (1791)
Facesoffungi number: *****??; Fig. 35
Saprobic on wood. Sexual morph: Ascomata 440 × 600 µm (x̄ = 520 µm, n = 10), perithecial, solitary or gregarious, superficial to semi-immersed, globose to subglobose, black, membranaceous, ostiolate, with septate, brown, tapering hairs, 3.5–5.5 wide. Peridium 45–75 µm (x̄ = 62 µm, n = 30) wide, membranaceous, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 2.5–5 µm wide, numerous, filiform. Asci (185–)210–230(–250) × 14–20 µm (x̄ = 220 × 17 µm, n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with apical globule, apical ring distinct. Ascospores 60–70(–75) × 4.5–7.5 µm (x̄ = 65 × 6 µm, n = 50), bi-seriate, cylindrical to geniculate, slight curved near the base, hyaline and aseptate when young, becoming pale brown and multi-septate when mature, with a large guttule in each cell, smooth-walled. Asexual morph: Undetermined.
Material examined: USA, Michigan, Marquette, Huron Mountain Club, around Ives Lake, 45 º0.00’N, 87 º0.00’W, on decayed wood, 17 August 1997, S.M. Huhndorf and M.H. Huhndorf (F-SMH3336).
Known hosts and distribution: On dead wood in Germany (type locality) and USA (Fuckel 1870; Miller and Huhndorf 2005).
Notes: The molecular data of Lasiosphaeris hispida were sequenced (Miller and Huhndorf 2005; Vu et al. 2019). In this study, L. hispida is basal to L. arenicola and L. hirsute in Lasiosphaeridaceae (100%ML/1.00BY, Fig. 26). We could not obtain the type material. Therefore, we re-examined an authentic specimen collected by Huhndorf.
New combination:
Lasiosphaeris arenicola (R. Hilber) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cercophora arenicola R. Hilber, in Hilber & Hilber, Z. Mykol. 45(2): 214 (1979)
Index Fungorum number: IF558252
Podosporaceae X. Wei Wang & Houbraken, in Wang et al., Stud. Mycol. 93: 223 (2019)
Coprophilous or saprobic on soil or wood. Sexual morph: Ascomata perithecial or cleistothecial, solitary to scattered or gregarious, immersed to semi-immersed or superficial, obpyriform or ovoid, membranaceous to carbonaceous, brown to black, glabrous or surrounded by hairs. Paraphyses cylindrical, septate. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, evanescent. Ascospores uni- or bi-seriate, ovoid, ellipsoidal or triangular, 0–1-septate, hyaline to dark brown, with an apical germ pore. Asexual morph: Hyphomycetous. Conidia globose to oval, hyaline, smooth-walled, ovate masses on the apex of the phialides (adapted from Wang et al. 2019).
Type genus: Podospora (Corda) Ces. 1856
Notes: Three redefined genera, Cladorrhinum, Podospora and Triangularia were classified in Podosporaceae based on phylogenetic analyses (Wang et al. 2019). Apiosordaria was synonymized under Triangularia because its generic type, A. verruculosa, was nested in Triangularia clade (Wang et al. 2019). In this study, species of Cladorrhinum, Podospora and Triangularia (= Apiosordaria) are scattered in Sordariales, especially Podospora (Fig. 26). These three genera are polyphyletic with different characteristics; Cladorrhinum has ornamented ascospores, Podospora has ascospores with unique gelatinous appendages, while Triangularia has polygonal ascospores (Krug et al. 1983; Guarro and Cano 1988; Bell and Mahoney 1997). Herein we review each genus of Podosporaceae based on phylogenetic analyses and characteristics of the generic types.
Cladorrhinum Sacc. & Marchal, in Marchal, Bull. Soc. R. Bot. Belg. 24(1): 64 (1885)
Saprobic on wood or soil, coprophilous. Sexual morph: Ascomata perithecial, solitary to scattered, immersed to superficial, globose to subglobose, semi-transparent, brown to black, surrounded by hyaline, septate hyphae. Peridium membranaceous, comprising pale brown to reddish brown cells of textura intricata to textura epidermoidea. Asci 8-spored, unitunicate, cylindrical to clavate to obovoid, pedicellate, apex rounded, evanescent. Ascospores ovoid to ellipsoidal, aseptate, hyaline to dark brown, with an apical germ pore and a gelatinous appendage at one or each end. Asexual morph: Hyphomycetous. Conidiophores micronematous. Conidiogenous cells intercalary or terminal, with a flared collarette, producing blastic conidia. Conidia globose to oval, hyaline, aseptate, smooth-walled, with a truncate base (adapted from Mouchacca and Gams 1993; Wang et al. 2019)
Notes: The hyphomycete Cladorrhinum was reported as an asexual morph of Apiosordaria and Cercophora (Mouchacca and Gams 1993), and it is characterized by clustered dematiaceous hyphae, with a lateral phialide producing globose, aseptate, hyaline conidia. Cladorrhinum was transferred to Podosporaceae based on multi-gene analysis of the generic type (Wang et al. 2019). Subsequently, Marin-Felix et al. (2020) modified Cladorrhinum and transferred Arnium tomentosum and Cercophora coprophila as Cla. tomentosum and Cla. coprophilum respectively based on phylogenetic analysis. In this study, four species are transferred to Cladorrhinum based on their molecular data (see the combination list below, Fig. 26).
Type species: Cladorrhinum foecundissimum Sacc. & Marchal, in Marchal, Bull. Soc. R. Bot. Belg. 24(1): 64 (1885)
Fig. 34 p
Saprobic in soil or coprophilous. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Mycelium yellowish-grey, velvety, dense. Hyphae filiform, septate, hyaline, branched. Conidiophores micronematous. Conidiogenous cells intercalary, with a flared collarette, producing blastic conidia. Conidia globose, hyaline, smooth-walled, with a truncate base, masses on the apex of the phialides (adapted from Marchal 1885; Wang et al. 2019).
Known hosts and distribution: On wild boar dung in Belgium (type locality) (Marchal 1885); on soil in the Netherlands (Wang et al. 2019).
Notes: Hyphomycetous Cladorrhinum foecundissimum was first isolated from wild boar dung in Belgium (Marchal 1885). Wang et al. (2019) provided an ex-neotype strain for Cla. foecundissimum (CBS 180.66) which was isolated from soil in the Netherlands since the holotype was lost. In this study, we found that this strain is sister to Cla. grandiusculum with 63%ML support (Fig. 26).
New combinations:
Cladorrhinum grandiusculum (A.E. Bell & Mahoney) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cercophora grandiuscula A.E. Bell & Mahoney, in Bell, CBS Diversity Ser. (Utrecht) 3: 59 (2005)
Index Fungorum number: IF558255
Cladorrhinum leucotrichum (Speg.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria leucotricha Speg., Michelia 1(no. 2): 225 (1878)
Synonym: Zopfiella leucotricha (Speg.) Malloch & Cain, Can. J. Bot. 49(6): 876 (1971)
Index Fungorum number: IF558256
Cladorrhinum terricola (S. Ueda) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cercophora terricola S. Ueda, Mycoscience 35(3): 287 (1994)
Index Fungorum number: IF558257
Cladorrhinum olerum (Fr.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sphaeria olerum Fr., Elench. fung. (Greifswald) 2: 98 (1828)
Synonym: Arnium olerum (Fr.) N. Lundq. & J.C. Krug, Symb. bot. upsal. 20(no. 1): 212 (1972)
Facesoffungi number: *****??; Index Fungorum number: IF558330; Fig. 36
Coprophilous, saprobic on herbaceous plant. Sexual morph: Ascomata perithecial, scattered, semi-immersed, broadly obpyriform to subglobose, membranaceous, pale brown to dark brown, semi-transparent, ostiolate, with a black papilla, periphysate. Peridium composed of yellowish cells of textura intricata. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, evanescent. Ascospores (40–)47–58(–65) × 20–40 µm (x̄ = 50 × 30 µm, n = 20), uni-seriate, obovoid to ellipsoidal, aseptate, hyaline when young, becoming dark brown when mature, with ornamented wall and a gelatinous appendage at each end. Asexual morph: Undetermined.
Material examined: Norway, Nord-Trøndelag, Grong, Godejorden, on old cow dung after 1–2 months in moist chamber in Uppsala, 17 June 1962, N. Lundqvist (UPS-UPS:BOT:F-125924); Sweden, Gotland, Visby par., Skansudd, NE of Visby, on old cow dung in moist chamber in Uppsala, 30 July 1961, N. Lundqvist (UPS- UPS:BOT:F-125925).
Known hosts and distribution: On stems of Brassica oleracea in Franch (type locality) (Fries 1828); usually on cabbage stalk in Austria, Belgium, Czech Republic, Denmark, England, Finland, France, Germany, Morocco, Norway, Portugal, Sweden (Lundqvist 1972; Jeng and Krug 1977; Kruys et al. 2014).
Notes: We re-examined two specimens of Cladorrhinum olerum which was collected and determined by Lundqvist (1972), and the hand-drawing of ascus and ascospores are provided (Fig. 42 e–f) based on Kruys et al. (2014). Phylogenetically, Arnium olerum nested in Lasiosphaeriaceae (Kruys et al. 2014) and was transferred to Cladorrhinum (Marin-Felix et al. 2020). In this study, Cla. olerum is sister to Cla. grandiusculum in Podosporaceae (63%ML, Fig. 26).
Podospora Ces., Hedwigia 1(15): 103 (1856)
Synonym: Schizothecium Corda, Icon. fung. (Prague) 2: 29 (1838)
Saprobic on wood or soil. Sexual morph: Ascomata perithecial, solitary to scattered, immersed to semi-immersed, obpyriform to ovoid, brown to black, ostiolate. Peridium membranaceous to coriaceous, composed of pale brown to reddish brown-walled cells of textura porrecta to textura angularis to textura prismatica. Paraphyses cylindrical, septate. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded, evanescent. Ascospores ovoid to ellipsoid, aseptate, hyaline when young, becoming a swollen, dark brown upper cell and a hyaline lower cell when mature, with a gelatinous appendage at one or each end. Asexual morph: Hyphomycetous, cladorrhinum-like. Conidiophores micronematous. Conidiogenous cells intercalary or terminal, with a flaring collarette producing blastic conidia. Conidia globose to oval, aseptate, hyaline, smooth-walled, with a truncate base (adapted from Cesati 1856; Lundqvist 1972).
Notes: Podospora is typified by P. fimicola which has coriaceous ascomata covered with simple short hairs and ascospores with gelatinous appendages (Cesati 1856). Lundqvist (1972) clarified Podospora species and proposed that Podospora fimiseda is a name change for Podospora fimicola. Thus, these species are conspecific (Niessl 1883; Lundqvist 1972; Bell and Mahoney 1997; Doveri 2008). However, following priority, Wang et al. (2019) accepted P. fimicola as its generic type and redefined Podospora, the type genus of Podosporaceae. Phylogenetically, we transfer seven species clustered in Podospora clade, to Podospora to avoid confusion (see the combination list below, Fig. 26). Species of Podospora (s. lat.) are also scattered in Bombardiaceae, Lasiosphaeriaceae, Naviculisporaceae and Neoschizotheciaceae (Fig. 26).
Type species: Podospora fimicola (Corda) Ces., Hedwigia 1(15): 103 (1856)
Basionym: Schizothecium fimicola Corda [as 'fimicolum'], Icon. fung. (Prague) 2: 29 (1838)
Fig. 34 a–d
Coprophilous. Sexual morph: Ascomata perithecial, solitary to scattered, immersed to semi-immersed, obpyriform to ovoid, membranaceous, brown to black, ostiolate. Peridium comprising three layers, outer layer composed of pale brown to reddish brown-walled cells of textura porrecta to textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses cylindrical, septate. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded, evanescent. Ascospores bi-seriate, ovoid to ellipsoid, aseptate, hyaline when young, becoming dark brown when mature, with ornamented wall and a gelatinous appendage at each end, with longitudinally striate. Asexual morph: Hyphomycetous (Cladorrhinum on CMA). Conidia globose to oval, hyaline, smooth-walled (adapted from Bell and Mahoney 1997).
Material examined: Italy, Piemonte, Biella, Bugellensibus Mountain, on cow dung, 1855, V. Cesati (S-F26041, lectotype).
Known hosts and distribution: On dry cattle dung on dry pastures in Czech Republic (type locality) (Corda 1838); on cow, horse dung in Austria, Belgium, Brazil, Canada, Chile, China, Denmark, Finland, France, Germany, Hungary, Italy, Liberia, Norway, Poland, Puerto Rico, Romania, Russia, Scotland, Spain, Sweden, USA, and Venezuela (Lundqvist 1972).
Notes: Lundqvist (1972) reviewed samples of related Podospora fimicola and found that this species is generally found in the faeces of herbivores. Vu et al. (2019) sequenced molecular data for the epitype strain of Schizothecium fimicola (= Podospora fimicola), CBS 482.64, and this strain is sister to P. costaricensis in Podosporaceae (Fig. 26). In this study, we observed the lectotype material of P. fimicola however only the dried ascomata could be seen. We drew the ascus and ascospores of P. fimicola concerning Bell and Mahoney (1997) and Doveri (2008).
New combinations:
Podospora brunnescens (W. Gams) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cladorrhinum brunnescens W. Gams, Mycotaxon 48: 435 (1993)
Index Fungorum number: IF558259
Podospora dacryoidea S.K. Huang & K.D. Hyde, nom. nov.
Synonym: Cladorrhinum australe Gasoni, in Carmarán, Berretta, Martínez, Barrera, Munaut & Gasoni, Mycol. Progr. 14(no. 94): 9 (2015)
Index Fungorum number: IF558260
Etymology: ‘dacryoidea’ refers to Carmarán et al. (2015) mentioned that this fungus has dacryoid conidia.
Podospora flexuosa (Madrid et al.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cladorrhinum flexuosum Madrid, Cano, Gené & Guarro, Mycologia 103(4): 801 (2011)
Index Fungorum number: IF558261
Podospora jamaicensis (B.M. Robison) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Echinopodospora jamaicensis B.M. Robison, Trans. Br. mycol. Soc. 54(2): 318 (1970)
Synonym: Apiosordaria jamaicensis (B.M. Robison) J.C. Krug, Udagawa & Jeng, Mycotaxon 17: 546 (1983)
Index Fungorum number: IF558298
Podospora hamata (Wu et al.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Apiosordaria hamata B. Wu, K.D. Hyde, Jing Z. Sun & Xing Z. Liu, in Wu, Tian, Wang, Liu, Hyde & Sun, Mycotaxon 131(4): 852 (2016) [2017]
Index Fungorum number: IF558299
Podospora macrospora (Guarro & Calvo) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Zopfiella macrospora Guarro & Calvo, Nova Hedwigia 37(4): 663 (1983)
Index Fungorum number: IF558300
Podospora spinosa (Cailleux) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Tripterospora spinosa Cailleux, Cahiers de La Maboké 8(1): 16 (1970)
Synonym: Apiosordaria spinosa (Cailleux) J.C. Krug, Udagawa & Jeng, Mycotaxon 17: 546 (1983)
Index Fungorum number: IF558301
Triangularia Boedijn, Annls mycol. 32(3/4): 302 (1934)
Synonym: Apiosordaria Arx & W. Gams, Nova Hedwigia 13: 201 (1967)
Saprobic in soil. Sexual morph: Ascomata cleistothecial or perithecial, gregarious to scattered, superficial to immersed, membranaceous or carbonaceous or coriaceous, obpyriform to pyriform, black, opaque or semi-transparent, glabrous or surrounded by hairs or setae, ostiolate, with papilla. Peridium comprising membranaceous to coriaceous, pale brown to reddish brown cells. Paraphyses filiform, septate. Asci 4–8–multi-spored, unitunicate, clavate to cylindrical, pedicellate, evanescent. Ascospores ellipsoidal to triangular, aseptate, hyaline when young, becoming 1-septate, upper cell larger, brown, ellipsoidal to polygonal, smooth-walled or with ornamented wall, with numerous broad or narrow pits, with a sub-apical or apical germ pore; lower cell smaller, hyaline, sometimes collapsing, with gelatinous appendages at one or each end, smooth-walled or sometimes with ornamented on surface, guttulate. Asexual morph: Hyphomycetous, cladorrhinum-like. Conidiophores micronematous. Conidiogenous cells intercalary or terminal, with a flaring collarette producing blastic conidia. Conidia globose to oval, aseptate, hyaline, smooth-walled, with a truncate base (adapted from Boedijn 1934; Guarro and Cano 1988; Wang et al. 2019).
Notes: Triangularia was transferred from Trigonia, and is typified by T. bambusae in having glabrous ascomata or those covered with hairs and ellipsoidal to triangular ascospores (Boedijn 1934; Guarro and Cano 1988). This genus was previously accepted in Lasiosphaeriaceae (Wijayawardene et al. 2018), and Wang et al. (2019) redefined Triangularia as a member of Podosporaceae based on multi-gene analysis. Apiosordaria was synonymized under Triangularia, because the strain of its generic type A. verruculosa (Fig. 34 e–f) nested in Triangularia (Wang et al. 2019). In this study, seven species clustered in Triangularia were transferred to Triangularia to avoid confusion (see the combination list below, Fig. 26). Species of Apiosordaria and Triangularia are also scattered in Naviculisporaceae and Neoschizotheciaceae in the phylogenetic analysis (Fig. 26).
Type species: Triangularia bambusae (J.F.H. Beyma) Boedijn, Annls mycol. 32(3/4): 302 (1934)
Basionym: Trigonia bambusae J.F.H. Beyma, Centbl. Bakt. ParasitKde, Abt. II 89: 236 (1933)
Fig. 34 q
Saprobic in soil. Sexual morph: Ascomata perithecial, gregarious to scattered, superficial to immersed, ovoid to pyriform, black, glabrous or surrounded by hairs. Peridium comprising membranaceous, pale brown to reddish brown cells of textura angularis. Asci 8-spored, unitunicate, clavate to cylindrical, pedicellate, evanescent. Ascospores uni- or bi-seriate, ellipsoidal to triangular, aseptate and hyaline when young, becoming conical with an oblique septum near one end, upper cell triangular, brown, with a subapical germ pore; lower cell irregular or hemispherical, hyaline, collapsing. Asexual morph: Undetermined (adapted from Guarro and Cano 1988).
Known hosts and distribution: On culms of Bambusaceae in Netherlands (type locality) (Van Beyma Thoe Kingma 1933).
Notes: Triangularia bambusae is sister to T. batistae in Podosporaceae based on phylogenetic anaylsis (Vu et al. 2019; Wang et al. 2019; this study, 100%ML/1.00BY, Fig. 26).
New combinations:
Triangularia microsclerotigena (Madrid et al.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cladorrhinum microsclerotigenum Madrid, Cano, Gené & Guarro, Mycologia 103(4): 804 (2011)
Index Fungorum number: IF558302
Triangularia nannopodalis (Cain) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora nannopodalis Cain, Can. J. Bot. 40(3): 455 (1962)
Index Fungorum number: IF558303
Triangularia praecox (Cailleux) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora praecox Cailleux, Cahiers de La Maboké 7: 102 (1969)
Index Fungorum number: IF558304
Triangularia samala (Udagawa & T. Muroi) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cercophora samala Udagawa & T. Muroi, Trans. Mycol. Soc. Japan 20(4): 454 (1979)
Index Fungorum number: IF558305
Triangularia tarvisina (Sacc.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Philocopra setosa subsp. tarvisina Sacc., Syll. fung. (Abellini) 1: 250 (1882)
Synonym: Podospora tarvisina (Sacc.) J.H. Mirza & Cain, Can. J. Bot. 47(12): 2041 (1970) [1969]
Index Fungorum number: IF558306
Triangularia unicaudata (C. Moreau & M. Moreau ex G. Sm.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Pleurage unicaudata C. Moreau & M. Moreau ex G. Sm., Trans. Br. mycol. Soc. 40(4): 488 (1957)
Synonym: Podospora unicaudata (C. Moreau & M. Moreau ex G. Sm.) Cain, Can. J. Bot. 40: 460 (1962)
Index Fungorum number: IF558321
Triangularia yaeyamensis (Morinaga et al.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Apiosordaria yaeyamensis Morinaga, Utatsu & Minoura, Trans. Mycol. Soc. Japan 20(4): 493 (1979)
Index Fungorum number: IF558322
Sordariaceae G. Winter [as 'Sordarieae'], Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 162 (1885)
Saprobic or endophytic on herbs or woody plants, isolated from soil or/and dung, in terrestrial and/or aquatic habitats. Sexual morph: Ascomata perithecial or cleistothecial, solitary, scattered or gregarious, superficial, erumpent or immersed, globose, subglobose, pyriform to ovoid, carbonaceous, coriaceous or membranaceous, yellow, olivaceous brown, brown to black, tuberculate or glabrous, with setae or not, ostiolate or not, with papilla or indistinct or papilla absent, with hyaline periphyses or periphyses absent. Peridium comprising hyaline to brown-walled cells, carbonaceous, coriaceous or membranaceous. Paraphyses numerous, branched or unbranched, septate, filiform or cylindrical, evanescent, or absent. Asci 2–8-spored, unitunicate, cylindrical to clavate, pedicellate, with J- apical ring or ring indistinct or absent. Ascospores uni–bi-seriate, hyaline, becoming yellowish or olivaceous brown or brown or black when mature, globose to ellipsoidal, aseptate, with 0–multi germ pore(s), smooth-walled or with angular to circular reticulate patterns or pits or longitudinal ribs on surface, with sheath or sheath absent, guttulate. Asexual morph: Hyphomycetous. Hyphae flexuous, hyaline to dark brown, septate, branched. Conidiophores hyaline to brown, branched or unbranched, straight or flexuous, smooth-walled, septate. Conidia hyaline, yellow, orange or dark brown, oval to ellipsoidal, smooth-walled, with sheath or sheath absent (adapted from Lundqvist 1972; García et al. 2004; Cai et al. 2006b; Maharachchikumbura et al. 2016).
Type genus: Sordaria Ces. & De Not. 1863
Notes: Sordariaceae was established by Winter (1885), and traditionally eight genera, Copromyces, Effetia, Gelasinospora, Guilliermondia, Neurospora, Pseudoneurospora, Sordaria and Stellatospora have been accommodated (Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2018). Most of the species in this family grow on dung, soil or decaying substrata in terrestrial and/or aquatic habitats (Frederick et al. 1969; Lundqvist 1972; Guarro and von Arx 1987; Krug and Scott 1994; García et al. 2004; Cai et al. 2006b), and they are closely related to Lasiosphaeriaceae, based on phylogenetic analysis and similarities in perithecial ascomata (Lundqvist 1972; Huhndorf et al. 2004b; Maharachchikumbura et al. 2015, 2016; Figs. 1, 26). Due to the typical morphology of ascospores, the genera in Sordariaceae have undergone various taxonomic changes. The monotypic genera, Effetia and Guilliermondia were included in Sordariaceae because of their characteristics are similar to Sordaria (Boudier 1904; Lundqvist 1967; Bartoli et al. 1984; Maharachchikumbura et al. 2015). Copromyces lacked reliable molecular data and was suspected of resembling Rechingeriella (Dothideomycetes) (Petrak 1967). Thus, we recommend placing this genus in Ascomycota genera incertae sedis pending future work. Pseudoneurospora was independent of Neurospora because it has unique ascospores with irregular ornamentation on the surface and it also exists as a separate cluster in molecular analysis (García et al. 2004). The monotypic Boothiella was accepted in Sordariaceae based on its type strain (CBS 334.67) nested in Sordariaceae (Wang et al. 2019; this study). Wang et al. (2019) removed Stellatospora from Sordariaceae as its ex-type strain, Stellatospora terricola (CBS 811.95), clusters in Chaetomiaceae. Gelasinospora was synonymized under Neurospora based on their similar morphology and close phylogenetic relationship (García et al. 2004). The traditional genera Neurospora and Sordaria were previously defined by morphology: Neurospora, which means ‘nerve spore’ refers to the ascospores with axon-like ornamentation on the surface (Shear and Dodge 1927; García et al. 2004). Sordaria is a genus mainly found in the faeces and has ascospores with sheaths (Cesati and de Notaris 1863; Maharachchikumbura et al. 2016). These two genera are polyphyletic in Sordariales (Spatafora and Blackwell 1994; Dettman et al. 2001; García et al. 2004; Cai et al. 2006b; this study). In this study, we found that all Neurospora and Sordaria species clustered in Sordariaceae. Their respective positions are usually unstable in multi-gene (Fig. 37) or single-gene analyses (not shown). Hence, Neurospora and Sordaria can be mainly distinguished by morphology. Neurospora is characterized by globose to pyriform, brown to black ascomata, cylindrical to broadly clavate asci and aseptate ascospores with a variety of ribs or pits, whereas, Sordaria has subglobose to ovoid, brown to black ascomata, cylindrical to clavate asci and ellipsoidal to ovoid ascospores with smooth-walls and may or may not have sheath. However, in this study, we include five genera in Sordariaceae based on morphology and phylogenetic analyses.
Boothiella Lodhi & Mirza, Mycologia 54(2): 217 (1962)
Saprobic in soil. Sexual morph: Ascomata cleistothecial, solitary or gregarious, superficial to immersed in the medium, globose to subglobose, membranaceous, brown to black, glabrous. Peridium composed of membranaceous, hyaline to subhayline cells of textura angularis. Asci 4-spored, unitunicate, clavate to cylindrical, short pedicellate, apex rounded, evanescent. Ascospores hyaline, becoming dark brown, oval to citriform,1–2-apiculate, aseptate, with a germ pore at one or each end. Asexual morph: Undetermined (adapted from Lodhi and Mirza 1962; Wang et al. 2019).
Notes: The monotypic genus Boothiella was introduced as a member of Eurotiales and its ascospores are similar to Thielavia (Lodhi and Mirza 1962). Kirk et al. (2008) transferred Boothiella to Sordariaceae and Maharachchikumbura et al. (2015, 2016) placed this genus in Chaetomiaceae. Vu et al. (2019) and Wang et al. (2019) analyzed sequence data from isolates CBS 334.67 (ex-type) and CBS 887.97, and showed that B. tetraspora nests in Sordariaceae. In the present study, Boothiella strains form a clade in Sordariaceae (100%ML/1.00BY, Fig 26, 37).
Type species: Boothiella tetraspora Lodhi & J.H. Mirza, Mycologia 54(2): 217 (1962)
Fig. 66 r–s
Saprobic in soil. Sexual morph: Ascomata cleistothecial, solitary or gregarious, superficial to immersed in the medium, globose to subglobose, membranaceous, brown to black, glabrous. Peridium composed of membranaceous, hyaline to subhayline cells of textura angularis. Asci 4-spored, unitunicate, clavate to cylindrical, short pedicellate, apex rounded, evanescent. Ascospores hyaline, becoming dark brown, oval to citriform,1–2-apiculate, aseptate, with a germ pore at one or each end. Asexual morph: Undetermined (adapted from Lodhi and Mirza 1962; Wang et al. 2019).
Known hosts and distribution: On soil in Pakistan (type locality) Lodhi and Mirza 1962); on sand in Spain (Wang et al. 2019).
Notes: Boothiella tetraspora is similar to Thielavia, except for its translucent ascomata (Lodhi and Mirza 1962). Wang et al. (2019) proposed that this species resembles Hyalosphaerella fragilis and Pseudothielavia subhyaloderma (Chaetomiaceae), but the former has 4-spored and the latter two have 8-spored asci. However, B. tetraspora nests in Sordariaceae based on phylogenetic analyses (Wang et al. 2019; this study, Fig. 26, 37).
Guilliermondia Boud., Bull. Soc. mycol. Fr. 20: 19 (1904)
Coprophilous on horse dung. Sexual morph: Ascomata cleistothecial, solitary or gregarious, superficial, dark brown to black, tuberculate. Peridium outer layer composed of carbonaceous, dark tissues; inner layer composed of membranaceous, hyaline to dark brown cells of textura angularis. Paraphyses numerous, evanescent, filamentous, septate, branched. Asci 8-spored, unitunicate, spherical, without an apical ring, with long and evanescent pedicel. Ascospores overlapping, ellipsoidal, brown, aseptate, smooth-walled, surrounded by hyaline or brown gelatinous sheath. Asexual morph: Undetermined (adapted from Boudier 1904).
Notes: The monotypic Guilliermondia is typified by G. saccoboloides has black ascomata and ellipsoidal ascospores surrounded by a brown sheath (Boudier 1904). Initially, Boudier (1904) proposed that this genus is related to Phillipsiella (Saccardiaceae), and Guilliermondia was later considered similar to Sordaria and transferred to Sordariaceae (Kirk et al. 2008; Lumbsch and Huhndorf 2010; Hyde et al. 2020). In this study, Guilliermondia is placed in Sordariaceae, because the generic type G. saccoboloides is similar to Sordaria fimicola. Both taxa are isolated from the faeces of herbivores and have dark ascomata and ellipsoidal, brown ascospores with gelatinous sheaths.
Type species: Guilliermondia saccoboloides Boud., Bull. Soc. mycol. Fr. 20: 20 (1904)
Facesoffungi number: ??; Fig 38
Coprophilous on horse dung. Sexual morph: Ascomata 295–460 × 170–275 µm (x̄ = 350 × 230 µm, n = 10), cleistothecial, solitary or gregarious, superficial, dark brown to black, tuberculate. Peridium outer layer composed of carbonaceous, dark tissues; inner layer composed of membranaceous, hyaline to dark brown cells of textura angularis. Paraphyses 1–2 µm wide, numerous, evanescent, filamentous, septate, branched. Asci 20–30 µm (x̄ = 25 µm, n = 30) diam. at ascospore-bearing part, 8-spored, unitunicate, spherical, without an apical ring, with long and evanescent pedicel. Ascospores 10–16 × 6–9 µm (x̄ = 14 × 7 µm, n = 50), overlapping, ellipsoidal, brown, aseptate, smooth-walled, surrounded by hyaline or brown gelatinous sheath 1–2.5 µm thick. Asexual morph: Undetermined (adapted from Boudier 1904).
Material examined: France, Lyon, 69 Rhôneon, on horse dung, April 1903, M.A.A. Guilliermond (PC-MNHN-PC-PC0167691, holotype).
Known hosts and distribution: On horse dung in France (type locality) (Boudier 1904).
Notes: We re-examined the holotype, MNHN-PC-PC0167691, and found that most of the ascomata had collapsed and the pedicel of the asci were basically inconspicuous.
Neurospora Shear & B.O. Dodge, J. Agric. Res., Washington 34: 1025 (1927)
Coprophilous or isolated from soil, burned or unburned vegetation in terrestrial habitats, heterothallic or pseudohomothallic. Sexual morph: Ascomata perithecial or cleistothecial, gregarious or scattered, solitary, semi-immersed to superficial, globose, subglobose to ovoid, brown to black, ostiole with hyaline periphyses. Peridium comprising brown to hyaline cells of textura angularis, textura globulosa, textura prismatica, textura intrincata or textura epidermoidea, carbonaceous to membranaceous. Paraphyses numerous, filiform or cylindrical or lacking. Asci 4- or 8-spored, unitunicate, cylindrical to clavate, pedicellate, with J- apical ring. Ascospores hyaline, olivaceous brown, grey or black, oval to ellipsoidal, aseptate, with longitudinal ribs or irregular to circular, deep or shallow pits on surface and germ pore(s) at one/each end, with or without guttules. Asexual morph: Hyphomycetous. Mycelium hyaline to light brown, septate. Conidiophores reduced to conidiogenous cells. Conidiogenous cells phialidic, oval to clavate, hyaline, smooth-walled. Conidia ellipsoidal to oval, hyaline, yellow, orange or dark brown, smooth-walled (adapted from Shear and Dodge 1927; Tai 1935; Perkins et al. 1976; Perkins and Turner 1988; Turner et al. 2001).
Notes: Neurospora is characterized by dark brown ascospores with ornamented surface and its asexual morph is usually found in nature and/or culture and is chrysonilia-like (Shear and Dodge 1927; von Arx 1981a, b; Jacobson et al. 2004). This genus is widespread and usually found in soil and burned or unburned vegetation in terrestrial habitats (Perkins et al. 1976; Perkins and Raju 1986; Perkins and Turner 1988; Turner et al. 2001; García et al. 2004). Three different mating strategies have been shown in Neurospora: homothallism, pseudohomothallism and heterothallism (Shear and Dodge 1927; Tai 1935; Mahoney et al. 1969; Krug and Khan 1991; Krug et al. 1994). Initially, Gelasinospora was distinct from Neurospora as the latter has longitudinal orientated structures on the ascospores, while the former has foveolate ascospores (Dowding 1933). Phylogenetic analysis demonstrated that Gelasinospora and Anixiella species scattered in Neurospora clades (García et al. 2004). Hence, Gelasinospora and Anixiella were synonymized under Neurospora (García et al. 2004; Kirk et al. 2008; Index Fungorum 2020). Pseudoneurospora was independent from Neurospora because it has an irregular pattern on the ascospores surface (García et al. 2004). Species of Neurospora were traditionally distinguished based on ascospore wall ornamentation (Dowding 1933; García et al. 2004; Cai et al. 2006b). Subsequently, García et al. (2004) and Cai et al. (2006b) showed that the traditional classification of Neurospora results in species being a polyphyletic in the Sordariales in phylogenetic analysis. García et al. (2004) considered the ornamentation of the epispore and the mating strategy are the most important characters to consistent the phylogenetic relatedness in Neurospora (Whitehouse 1949; Taylor and Natvig 1989; Grognet and Silar 2015). In this study, we were unable to find characteristics consistent with polyphyletic Neurospora in phylogenetic analysis, either ascospore morphology or mating strategies. Therefore, we propose to delimit Neurospora as species nested in Sordariaceae in phylogenetic analysis and has aseptate ascospores with various ribs and/or pitted on surface. In this study, Neurospora is divided in four clades based on multi-gene analysis. We have sorted out their mainly characteristics based on mating type, form of ascomata, ascospore appearance and types of epispore, and listed in Fig. 37.
Type species: Neurospora sitophila Shear & B.O. Dodge, J. Agric. Res., Washington 34: 1026 (1927)
Facesoffungi number: *****??; Fig. 39 a–j
Saprobic on wood and burnt vegetation, heterothallic. Sexual morph: Ascomata 280–380 µm (x̄ = 315 µm, n = 10) diam., perithecial, superficial, solitary, globose, dark brown to black, with an ambiguous ostiole, covered by white hyphae. Peridium 70–120 µm (x̄ = 95 µm, n = 30) wide, comprising four layers, outer layer composed of compact, dark brown cells of textura angularis; middle layer closely aligned from external to internal with hyaline to pale brown cells of textura globulosa to textura angularis; and composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci (150–)180–190(–200) × 13–16 µm (x̄ = 185 × 14.5 µm, n = 30), 8-spored, unitunicate, cylindrical, slightly curved, short pedicellate, thin-walled and evanescent at maturity. Ascospores (17–)19–23(–28) × (10–)12–14(–15) µm (x̄ = 22 × 13 µm, n = 50), uni-seriate, subglobose to ellipsoidal or oval, thick-walled, light brown when young, becoming dark brown or olivaceous at maturity, aseptate, epispore smooth, with a germ pore at each end. Ribs longitudinal or reticulation, frequently branched, pigment deposited externally, nested straight, discrete, elongate pits. Asexual morph: Hyphomycetous. Conidia orange, globose to subglobose, aseptate, smooth-walled (adapted from García et al. 2004).
Material examined: Canada, Ottawa, on drying veneer (birch and poplar) of fruit basket, culture 5574, 5 February 1927, M. Hantsport (BPI-630404, from ex-type).
Known hosts and distribution: On corn-meal agar culture of Monilia sitophila in Germany (type locality) (Shear and Dodge 1927); in the air, on garbage, leaves, wood in Australia, Canada, France, New Zealand and USA (Shaw 1998; Francuz et al. 2010; Vojtkova et al. 2020); on burnt vegetation and non-burnt substrata distributed worldwide (García et al. 2004).
Notes: Shear and Dodge (1927) found Neurospora sitophila on a culture of Monilia sitophila and established Neurospora. Subsequently, the hyphomycetous Chrysonilia sitophila and M. sitophila were synonymized under N. sitophila (Perkins et al. 1976; Perkins and Turner 1988; Turner et al. 2001; García et al. 2004), and this species was found in the air and human waste, such as food or coffee residues, and was identified as one of the causes of allergies and asthma (Shaw 1998; Francuz et al. 2010; Vojtkova et al. 2020). In this study, N. sitophila clustered in Neurospora clade II accommodates heterothallic species (Fig. 37). Gelasinospora autosteira was considered as N. calospora based on the similar morphology (García et al. 2004); however, the ex-type strain of Gelasinospora autosteira was sequenced by Vu et al. (2019), and this strain is sister to N. nigeriensis in Neurospora clade III, whereas has far distance with N. calospora. Therefore, we recommend to transfer G. autosteira as Neurospora autosteira based on phylogenetic analysis (see the combination list below, Fig. 37).
Neurospora tetraspora Dania García et al., in García et al., Mycol. Res. 108(10): 1134 (2004)
Basionym: Gelasinospora tetrasperma Dowding, Canadian Journal of Research, Section C 9: 294 (1933)
Facesoffungi number: *****??; Fig. 40
Isolated from soil, dung or burnt vegetation, pseudohomothallic. Sexual morph (On culture): Mycelium composed of hyaline to light brown, branched, septate hyphae 3–5 µm wide (x̄ = 4 µm, n = 30). Ascomata perithecial, solitary or scattered, semi-immersed to superficial, slightly coriaceous, pyriform, ostiolate, brown to black. Asci 4-spored, unitunicate, with J- apical ring. Ascospores oval to ellipsoidal, hyaline becoming dark brown, slightly flattened on one side, aseptate, with a germ pore at each end, with circular pits. Asexual morph: Undetermined (adapted from Dowding 1933; Matsushima 1975; von Arx 1982; Fort et al. 1990; García et al. 2004).
Material examined: Culture from CBS, Netherlands (IMI-38600, from ex-type).
Known hosts and distribution: On ptarmigan and rabbit dung in Canada (type locality) (Dowding 1933); on dung and soil in Canada, England, Finland, Norway, Russia, Spain, Sweden and USA (Lundqvist 1972; García et al. 2004).
Notes: Dowding (1933) cultivated the spores which was found on ptarmigan and rabbit dung in Manitobba and Ontario and introduced a homothallic Gelasinospora tetrasperma as the generic type of Gelasinospora. Dettman et al. (2001) confirmed that G. tetrasperma is a pseudohomothallic species. Subsequently, Gelasinospora was placed as a synonym of Neurospora, and G. tetrasperma was replaced by Neurospora tetraspora (García et al. 2004). Sequences data from the ex-type strain (CBS:178.33) is available (Cai et al. 2006b; Spatafora et al. 2006), and in this study, this strain is distinct with N. tetraspora and nested in Neurospora clade I which mainly comprises homothallic species with an ornate epispore (Fig. 37). We re-examined the ex-type culture (38600) of G. tetrasperma, and found that were no ascomata, but a large number of conidia (22–28 × 12–15 µm) with a pitted surface in clumps on pale brown to brown, septate, shrunken hyphae (Fig. 40).
New combination:
Neurospora autosteira (Alexop. & S.H. Sung) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Gelasinospora autosteira Alexop. & S.H. Sung, Mycologia 42(6): 723 (1951) [1950]
Index Fungorum number: IF558325
Pseudoneurospora Dania García et al., Mycol. Res. 108(10): 1139 (2004)
Isolated from soil. Sexual morph: Mycelium composed of hyaline to brown, branched hyphae. Ascomata perithecial or cleistothecial, gregarious or scattered, superficial to immersed, yellowish to brown or black, sometimes translucent, pyriform to subglobose, ostiolate, surrounded by hyaline, septate hyphae or sinuose, brown, septate setae. Peridium membranaceous, comprising brown to hyaline cells of textura angularis or textura globulosa. Asci 8-spored, unitunicate, cylindrical to clavate, short pedicellate, apex rounded, with J- apical ring, evanescent. Ascospores uni- or bi- seriate, hyaline when young becoming dark brown or black, oval to ellipsoidal, aseptate, verrucose to tuberculate, with a germ pore at each end. Asexual morph: Undetermined (adapted from Udagawa 1980; García et al. 2004).
Notes: Pseudoneurospora amorphoporcata was initially described as Gelasinospora amorphoporcata (Udagawa 1980), and García et al. (2004) separated it from Gelasinospora and established Pseudoneurospora based on phylogenetic analysis and because its ascospores had a strongly verrucose to tuberculate irregular pattern (Fig. 45k) on the surface. Subsequently, a second species, P. canariensis, was introduced by Crous et al. (2014) having similar ascospores and molecular data. In this study, an invalid name ‘Copromyces octosporus’ (CBS 386.78) is sister to P. canariensis (see notes for Copromyces). Pseudoneurospora species clustered with 92%ML/1.00BY support and basal to Sordaria clade II in Sordariaceae (97%ML/1.00BY, Fig 37).
Type species: Pseudoneurospora amorphoporcata (Udagawa) Dania García et al., Mycological Research 108 (10): 1139 (2004)
Basionym: Gelasinospora amorphoporcata Udagawa, Transactions of the Mycological Society of Japan 21 (1): 19 (1980)
Fig. 39 k
Isolated from soil. Sexual morph: Ascomata perithecial, brown to black, gregarious or scattered, superficial to immersed, pyriform to subglobose, membranaceous, ostiolate, with papilla. Peridium comprising brown to hyaline cells of textura angularis or textura globulosa, membranaceous. Asci 8-spored, unitunicate, cylindrical, short pedicellate, apex rounded, with J- apical ring. Ascospores hyaline when young becoming dark brown or black, oval to ellipsoidal, aseptate, with a germ pore at each end, with irregular reticulate patterns. Asexual morph: Undetermined (adapted from Udagawa 1980; García et al. 2004).
Known hosts and distribution: On soil in Philippines and Thailand (type locality) (Udagawa 1980; García et al. 2004).
Notes: Pseudoneurospora amorphoporcata was considered similar to Copromyces bisporus and Emblemospora monotrema in having ornamented ascospores (García et al. 2004). However, Pseudoneurospora can be distinguished from C. bisporus because the latter lacks an apical ring in the asci and has globose ascospores with one germ pore. It also can be differentiated from E. monotrema which has ascospores with plate-like wrinkles (Lundqvist 1972; Jeng and Krug 1976). In this study, P. amorphoporcata is basal to P. canariensis and ‘Copromyces octosporus’ (92%ML/1.00BY, Fig. 37).
Sordaria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 225 (1863)
Isolated from dung and soil or saprobic on wood. Sexual morph: Ascomata perithecial, gregarious, immersed to superficial, brown, translucent, subglobose to pyriform, glabrous or with rare hairs, ostiolate, with papilla, periphysate. Peridium membranaceous, composed of dark brown to hyaline cells of textura angularis to textura prismatica. Asci 8-spored, unitunicate, cylindrical, short pedicellate, apex rounded, with J- apical ring, with or lacking paraphyses. Ascospores uni-seriate, subglobose to ellipsoidal, hyaline, becoming dark brown, aseptate, smooth-walled, germ pore at one end, with or without a gelatinous sheath. Asexual morph: Undetermined (adapted from De Notaris 1867; Winter 1884; Potebnia 1907; Cain 1957a; Lundqvist 1972; Khan and Krug 1989; Watanabe 1989).
Notes: Sordaria was first mentioned in Cesati and de Notaris (1863), and typified by Sphearia sordaria Fr., which later was transferred as Coniochaeta sordaria (Petrak 1953). Moreau (1953) introduced Sordaria fimicola as its generic type, which has been widely accepted (von Arx and Müller 1954; Munk 1957; Stafleu and Voss 1969; Lundqvist 1972). Lundqvist (1972) clarified Sordaria species as having dark perithecial ascomata, cylindrical asci and brown, smooth-walled ascospores with or without a gelatinous sheath. Traditionally, species of Sordaria are coprophilous and usually found in faeces and soil (Kirk et al. 2008; Maharachchikumbura et al. 2016). von Arx et al. (1987) re-defined species with a gelatinous sheath as Sordaria, while these lacking a gelatinous sheath were placed in Asordaria. Subsequently, Cai et al. (2006b) proposed that the presence or absence of the ascospore sheath is not a phylogenetically significant for the separation of Asordaria from Sordaria. We placed Asordaria under Sordaria, and Sordaria is divided in three clades (Sordaria clade I, II, III) in Sordariaceae based on phylogenetic analysis (Fig. 37).
Type species: Sordaria fimicola (Roberge ex Desm.) Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 226 (1863)
Basionym: Sphaeria fimicola Roberge ex Desm., Annls Sci. Nat., Bot., sér. 3 11(2): 339 (1849)
Facesoffungi number: *****??; Fig. 41
Saprobic on dead wood, homothallic. Sexual morph: Ascomata 250–350 × 190–300 µm (x̄ = 270 × 240 µm, n = 10), perithecial, solitary or gregarious, immersed, erumpent through bark of the host at maturity, globose to subglobose, membranaceous, dark brown to black, ostiolate, with papilla. Peridium 15–30 µm wide, comprising two layers, outer layer composed of brown to dark brown cells of textura angularis; inner layer composed of pale brown to hyaline cells of textura prismatica. Paraphyses evanescent. Asci (155–)165–180(–220) × 12–20 µm (x̄ = 175 × 15 µm, n = 50), 8-spored, unitunicate, cylindrical to subclavate, short pedicellate, apex truncate to rounded, with J- inconspicuous apical ring. Ascospores (17–)19–23 × (9–)10–13 µm (x̄ = 20 × 11.5 µm, n = 50), uni-seriate, subglobose to oval or ellipsoidal, smooth-walled, pale brown to dark brown, aseptate, germ pore at one end, with gelatinous sheath. Asexual morph: Undetermined.
Material examined: Germany, Berlin, Brandenburg, Berlin Bot. Garden, on rotten stems of Phragmites sp. with Claviceps sp., May 1889, P. Hennings (S-F6838, lectotype of Sordaria fimicola var. phragmiticola Henn.); Germany, Hessen, Nassau, Östrich, on horse dung, 1802, L. Fuckel (S-F6841, lectotype of Sphaeria equina Fuckel); Italy, on decaying Vitis sp., 22 March 1930, Shear (BPI-580866).
Known hosts and distribution: On horse dung in France (type locality) (Desmazières 1849); on hare, horse, guinea-pig, camel and rabbit dung, heather soil, grass culms, filter paper, ergot-infested panicles of Phragmites communis in Argentina, Austria, Belgium, Bulgaria, Canada, Czech Republic, Denmark, England, Egypt, Finland, France, Germany, Hungary, Italy, Netherlands, Norway, Poland, South Africa, Spain and Sweden (Lundqvist 1972).
Notes: We re-examined three specimens, the lectotype of Sordaria fimicola var. phragmiticola Henn. (F6838), lectotype of Sphaeria equina Fuckel (F6841) and Sordaria fimicola (BPI-580866). These authentic materials were determined as Sordaria fimicola by Lundqvist (1972). Phylogenetically, the strains of Sordaria fimicola, which collected from Canada and Netherlands (Vu et al. 2019), clustered in Sordaria clade II (62%ML, Fig. 37).
Strattoniaceae S.K. Huang, Maharachch. & K.D. Hyde, fam. nov.
Index Fungorum number: IF558346
Etymology: Named for Strattonia, the type genus of this family.
Coprophilous. Sexual morph: Ascomata perithecial, solitary, scattered or gregarious, superficial to immersed, coriaceous to membranaceous, pyriform to subglobose, brown to black, hairy, ostiolate, with a black papilla, periphysate. Paraphyses filiform-ventricose, septate. Asci 4–8-spored, unitunicate, cylindrical, apex truncate or rounded, with J- apical ring. Ascospores uni-seriate, fusiform to obovoid, hyaline to subhyaline, 1-septate near the base, sometimes finely guttulae; upper cell becoming brown to black, swollen, ellipsoidal, with an apical germ pore; pedicel short, conical or cylindrical, hyaline, collapsing when dry, surrounded by gelatinous sheath. Gelatinous sheath rounded or irregular, swelling in water, invaginated at the pedicel, with indentations. Asexual morph: Hyphomycetous, phialophora-like. Conidiophores arising from aerial hyphae, erect, inflated at the base. Phialides monophialidic, smooth-walled, hyaline, cylindrical, unbranched or branched. Conidia hyaline to pale yellow, ellipsoidal to oval, aseptate, smooth-walled (adapted from Lundqvist 1972; Bell 2000, 2005).
Type genus: Strattonia Cif. 1954
Notes: The coprophilous Strattonia was introduced based on Fimetaria tetraspora and was initially characterized by superficial to immersed, coriaceous ascomata, 4-spored asci and brown ascospores (Ciferri 1954). Subsequently, Lundqvist (1972) delimited this genus as having ostiolate ascomata, non-amyloid asci and two-celled ascospores (a dark brown fusiform or ellipsoidal upper cell and a hyaline pedicel), sometimes with gelatinous sheath, which is similar to Podospora, Sordaria and Triangularia. Since authentic material of S. tetraspora is lacking, Lundqvist (1972) described this type species based on previous literature and found a special feature, that is, if gelatinous sheath present, it surrounds the ascospores but collapses at the pedicel. Eleven species accepted in this genus based on morphology (Ciferri 1954; Lundqvist 1972; Abdullah 1983; Bell 2000, 2005).
Vu et al. (2019) sequenced Podospora petrogale and Strattonia oblecythiformis. These two strains clustered in 100%ML/1.00BY support as an unstable sister clade to Diplogelasinospora (Fig. 26). Morphologically, these two species are coprophilous and have similar hairy ascomata, non-amyloid, cylindrical asci and ascospores comprise a fusiform to ellipsoidal, brown to dark brown upper cell and a hyaline pedicel, while surrounded by gelatinous sheath, except pedicel, in addition, both of them have phialophora-like asexual morph (Bell 2000, 2005). These characters are also similar to S. tetraspora. The difference between P. petrogale and S. oblecythiformis is that the former has a cylindrical pedicel in ascospores, whereas the latter has oblecythiform pedicel (Bell 2000, 2005). Therefore, we placed Podospora petrogale in Strattonia (see the combination list below).
Strattonia carbonaria, S. karachiensis and S. minor with no gelatinous sheath were placed in Jugulospora and Lundqvistomyces (Neoschizotheciaceae) in phylogenetic analysis (Marin-Felix et al. 2020; this study). Thus, we believe that the gelatinous sheath surrounds the ascospores except pedicel should be accepted as a feature for the identifying Strattonia species.
We also raised Strattonia at a family level, Strattoniaceae. It is characterized by two-celled ascospores comprise a brown upper cell and a hyaline pedicel and surrounded by gelatinous sheath except for pedicel, which is different from other lasiosphaeriaceous taxa. Phylogenetically, Strattonia species form a sister clade to Diplogelasinosporaceae (Fig. 26).
Strattonia Cif., Sydowia 8(1-6): 245 (1954)
Coprophilous. Sexual morph: Ascomata perithecial, solitary, scattered or gregarious, superficial to immersed, coriaceous to membranaceous, pyriform to subglobose, brown to black, hairy, ostiolate, with a black papilla, periphysate. Paraphyses filiform-ventricose, septate. Asci 4–8-spored, unitunicate, cylindrical, apex truncate or rounded, with J- apical ring. Ascospores uni-seriate, fusiform to obovoid, hyaline to subhyaline, 1-septate near the base, sometimes finely guttulae; upper cell becoming brown to black, swollen, ellipsoidal, with an apical germ pore; pedicel short, conical or cylindrical, hyaline, collapsing when dry, surrounded by gelatinous sheath. Gelatinous sheath rounded or irregular, swelling in water, invaginated at the pedicel, with indentations. Asexual morph: Hyphomycetous, phialophora-like. Conidiophores arising from aerial hyphae, erect, inflated at the base. Phialides monophialidic, smooth-walled, hyaline, cylindrical, unbranched or branched. Conidia hyaline to pale yellow, ellipsoidal to oval, aseptate, smooth-walled (adapted from Lundqvist 1972; Bell 2000, 2005).
Notes: Strattonia is typified by S. tetraspora, which has hairy ascomata, cylindrical asci and ascospores comprise a brown upper cell and a hyaline pedicel and surrounded by gelatinous sheath except pedicel (Ciferri 1954; Lundqvist 1972). Lundqvist (1972) proposed that Strattonia is connected with Podospora, Sordaria and Triangularia and tried to sort out the affinities of Strattonia species by morphology. Subsequently, the no gelatinous sheath Strattonia carbonaria, S. karachiensis and S. minor were placed in Jugulospora and Lundqvistomyces respectively, based on phylogenetic analysis (Miller and Huhndorf 2005; Vu et al. 2019, Marin-Felix et al. 2020; this study, see notes for Jugulospora minor). We divide the remaining Strattonia species that do not have receivable molecular data into three groups according to the characteristics of their gelatinous sheath: i) Strattonia grandis, S. tetraspora and S. zopfii are similar to S. petrogale and S. oblecythiformis have gelatinous sheath surrounds ascospores except pedicel (Lundqvist 1972; Bell 2005); ii) Strattonia borealis and S. mesopotamica have terminal gelatinous appendages surrounding apical germ pore and pedicel (Lundqvist 1972; Abdullah 1983), which are similar to Podospora fibrinocaudata in Bombardiaceae; iii) Strattonia dissimilis and S. insignis have ascospores surrounded by gelatinous sheath except apical germ pore and pedicel (Lundqvist 1972; Barrasa et al. 1985), which are similar to Apodospora gotlandica in Bombardiaceae. However, more fresh collections and molecular information of Strattonia are expected to define its taxonomic placement more accurately.
Type species: Strattonia tetraspora (R. Stratton) Cif., Sydowia 8(1-6): 245 (1954)
Basionym: Fimetaria tetraspora R. Stratton, Ohio St. Univ. Bull. 26: 84 (1921)
On rotten paper. Sexual morph Ascomata perithecial, solitary to gregarious, superficial to immersed, coriaceous, pyriform to subglobose, black, ostiolate. Asci 4-spored, unitunicate, cylindrical. Ascospores uni-seriate, fusiform to obovoid, composed of a brown upper cell, with an apical germ pore, and a hyaline pedicel, sometimes surrounded by gelatinous sheath, but pedicel never. Asexual morph Undetermined (adapted from Ciferri 1954; Lundqvist 1972).
Known hosts and distribution: On rotten paper in Dominica (type locality) (Ciferri 1954).
Notes: Ciferri (1954) introduced Strattonia tetraspora and noted that this species was found on rotten paper from Moca, Santiago, Dominican Republic, in April 1928. Lundqvist (1972) mentioned that the authentic sample of S. tetraspora is lacking. In this study, we also could not obtain any authentic specimens or illustrations of S. tetraspora. This makes it impossible for us to identify its substrate, nor can we provide more information about this species. However, we provided the hand-drawing of Strattonia zopfii (Fig. 42 a), which is similar to S. tetraspora has ascospores surrounded by gelatinous sheath but invaginated at the pedicel, refer to Lundqvist (1972).
New combination:
Strattonia petrogale (A.E. Bell) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora petrogale A.E. Bell, Muelleria 12(2): 236 (2000) [1999]
Index Fungorum number: IF558343
Zygospermellaceae S.K. Huang, Maharachch. & K.D. Hyde, fam. nov.
Index Fungorum number: IF558347
Etymology: Named for Zygospermella, the type genus of this family.
Coprophilous, entomogenous or on the rock. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, obpyriform to subglobose, dark brown to black, verrucose, ostiolate. Peridium comprising carbonaceous to membranaceous, brown to black cells of textura angularis to textura prismatica. Paraphyses unbranched, broad, cylindrical, ventricose, septate. Asci 8-spored, unitunicate, cylindrical to subclavate, pedicellate, apex rounded, with J- apical ring. Ascospores uni- or bi-seriate, hyaline, aseptate, fusiform to ellipsoidal when young, becoming elongate-subfusiform, ends truncate, 1–3-septate, constricted at the middle septum, dark brown when mature, with an appendage and germ pore at each end. Appendages terminal, hyaline, cylindrical, tapering, smooth-walled, ends rounded, with longitudinally striate. Asexual morph: Hyphomycetous. Conidiophores arising from aerial hyphae, erect, inflated at the base. Conidia small, globose to oval, forming globose masses on the apex of the phialides (adapted from Cain 1934,1935; Lundqvist 1972; Zhang et al. 2017).
Type genus: Zygospermella Cain 1935
Notes: Zygospermella was initially introduced as Delitschia (Pleosporales) because of the similar hourglass-shaped ascospores, and then redefined as Zygospermella and placed in Lasiosphaeriaceae (Mouton 1897; Cain 1935; Lundqvist 1969). Phylogenetically, Zygospermella and Lasiosphaeris form a clade sister to Neoschizotheciaceae (Kruys et al. 2014; Marin-Felix et al. 2020). In this study, Lasiosphaeris, Zygospermella, Episternus and Ramophialophora petraea clustered as a sister clade of Neoschizotheciaceae in LSU sequence data (not shown). However, in multi-gene analysis, Episternus onthophagi, Ramophialophora petraea and Zygospermella insignis form an independent cluster and nested in Sordariales (71%ML/0.99BY) as an orphan clade (Fig. 26). In addition, the phialophora-like hyphomycetous E. onthophagi is epizoic fungus and R. petraea was collected on the rock in Karst cave (Zhang et al. 2017; Górz and Boroń 2018). Zygospermella has special hourglass-shaped ascospores attached to terminal appendages with longitudinally striate, which is different from the other members in Sordariales.
Episternus Górz & Boroń, Phytotaxa 376(1): 49 (2018)
Type species: Episternus onthophagi Górz & Boroń, Phytotaxa 376(1): 49 (2018)
Known hosts and distribution: On legs of dung beetle (Onthophagus verticicornis) in Poland (type locality) (Górz and Boroń 2018).
Notes: Górz and Boroń (2018) found the hyphae arising from the surface of a dung beetle and has 1-septate, ovoid conidia. The monotypic Episternus was established as an epizoic fungus in Lasiosphaeriaceae based on LSU-SSU-ITS sequence data (Górz and Boroń 2018). In this study, the ex-type strain of E. onthophagi (KRAM F 58223) is basal to Zygospermella insignis and Ramophialophora petraea in Zygospermellaceae (71%ML/0.99BY, Fig. 26).
Zygospermella Cain, Mycologia 27(2): 227 (1935)
Coprophilous. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, obpyriform to subglobose, dark brown to black, verrucose, ostiolate. Peridium comprising carbonaceous to membranaceous, brown to black cells of textura angularis to textura prismatica. Paraphyses unbranched, broad, cylindrical, ventricose, septate. Asci 8-spored, unitunicate, cylindrical to subclavate, pedicellate, apex rounded, with J- apical ring. Ascospores uni- or bi-seriate, hyaline, aseptate, fusiform to ellipsoidal when young, becoming hourglass-shaped to elongate-subfusiform, ends truncate, usually 1-septate, constricted at the middle septum, dark brown when mature, with a germ pore and appendage at each end. Appendages hyaline, cylindrical, tapering, smooth-walled, ends rounded, with longitudinally striate. Asexual morph: Undetermined (adapted from Cain 1934,1935; Lundqvist 1972).
Notes: Zygospermella was invalidly introduced as Zygospermum (Cain 1934). Cain (1935) proposed that Zygospermum should be corrected to Zygospermella and accepted Zygospermella insignis and Z. setosa. Lundqvist (1969, 1972) clarified that these species are conspecific, as Z. insignis, and introduced another species, Zygospermella striata. Therefore, two species are presently accepted for Zygospermella, Z. insignis (type) and Z. striata. Subsequently, Kruys et al. (2014) sequenced Z. insignis, which was collected on muskox dung in the USA, cattle dung in France and cow dung in Sweden, and found these strains clustered in Lasiosphaeriaceae (s. lat.). In this study, Z. insignis is sister to Ramophialophora petraea in Zygospermellaceae (100%ML/1.00BY, Fig. 26).
Type species: Zygospermella insignis (Mouton) Cain, Mycologia 27(2): 227 (1935)
Facesoffungi number: *****??; Fig. 43
Coprophilous. Sexual morph: Ascomata 650–900 × 250–500 µm, perithecial, solitary, superficial to semi-immersed, carbonaceous to membranaceous, pyriform to subglobose, dark brown to black, verrucose, surrounded by setae, ostiolate, with black, carbonaceous necks 180–250 × 100–150 µm, inner cells with hyaline periphyses. Setae 2.5–4.5 µm wide, brown, aseptate, tapering towards the acute apex. Peridium 25–40 µm wide, outer layer composed of carbonaceous, brown to black cells of textura intricata to textura angularis; inner layer composed of membranaceous, hyaline cells of textura angularis to textura prismatica. Paraphyses 2.5–3.5 µm wide (x̄ = 2.5 µm, n = 30), unbranched, cylindrical, septate, tapering, slightly constricted at the septa. Asci (150–)200–240(–270) × 15–25 µm (x̄ = 220 × 20 µm, n = 20), 8-spored, unitunicate, cylindrical to subclavate, pedicellate, apex rounded and thickened. Ascospores uni- or bi-seriate, hyaline, aseptate, fusiform to ellipsoidal when young, becoming hourglass-shaped to elongate-subfusiform, ends truncate, 1-septate, sometimes 3-septae, constricted at the middle septum 3–7 µm wide, dark brown when mature, (26–)30–40(–45) × 7–10 µm (x̄ = 35 × 8 µm, n = 20), with a germ pore and appendage at each end. Appendages 4.5–8 µm wide, hyaline, cylindrical, tapering, ends rounded, smooth-walled, with longitudinally striate. Asexual morph: Undetermined.
Material examined: Canada, Ontario. Bear Island, Lake Temagami, on cow dung, 21 November 1933, R.F. Cain (BPI-618905).
Known hosts and distribution: On dung of cow, sheep and horse in Belgium (type locality), Canada, England, Norway, Scotland, Spain and Sweden (Mouton 1897; Lundqvist 1969).
Notes: We re-examined authentic material collected by Cain and determined by Lundqvist (mentioned in the label of material) since the type specimen was unavailable.
Other genera in Sordariales
Chaetomiaceae G. Winter
Stellatospora Tad. Ito & Nakagiri, Mycoscience 35 (4): 413 (1994)
Isolated from rice field soil. Sexual morph: Mycelium white, superficial, effuse. Ascomata cleistothecial, solitary or gregarious, globose to subglobose, dark brown to black, covered in hyaline hairs. Peridium composed of brown to hyaline cells of textura angularis. Asci 3–8-spored, unitunicate, obpyriform to oval, evanescent. Ascospores multi-seriate, stellate to irregular with up to 9 short protuberances when mature, aseptate, pale greenish brown to brown, with a germ pore on one protuberance, guttulate. Asexual morph: Undetermined (adapted from Ito and Nakagiri 1994).
Notes: The monotypic Stellatospora typified by S. terricola has stellate ascospores and was initially placed in Sordariaceae (Ito and Nakagiri 1994). Subsequently, the ex-type strain (CBS 811.95) is available (Wang et al. 2019; Vu et al. 2019), and nests in Chaetomiaceae (Wang et al. 2019; this study, 100ML/1.00BY, Fig. 26). Therefore, this genus belongs in Chaetomiaceae based on phylogenetic analysis.
Type species: Stellatospora terricola Tad. Ito & Nakagiri, Mycoscience 35(4): 413 (1994)
Facesoffungi number: *****??; Fig. 44
Isolated from rice field soil, colonies on oatmeal agar (OA). Sexual morph: Mycelium white, superficial, effuse. Ascomata 75–120 µm diam., cleistothecial, solitary or gregarious, globose to subglobose, dark brown to black, covered in hyaline hairs. Peridium 3–5 µm wide, composed of pale brown to hyaline cells of textura angularis. Asci 3–8-spored, unitunicate, obpyriform to oval, evanescent. Ascospores (9–)11–15(–17) µm (x̄ = 12 µm, n = 50) diam., multi-seriate, stellate to irregular with up to 9 short protuberances when mature, aseptate, pale greenish brown to brown, with a germ pore on one protuberance, guttulate. Asexual morph: Undetermined (adapted from Ito and Nakagiri 1994).
Material examined: Japan, Ikeda, Osaka, rice field soil, 22 May 1990, T. Ito (NBRC-IFO-H-12166, holotype).
Known hosts and distribution: Rice field soil, Japan (type locality) (Ito and Nakagiri 1994).
Notes: We re-examined the type material and found that there are star-like ascospores filling the ascomata, but no complete asci. Stellatospora terricola (CBS 811.95) is basal to Ovatospora and Trichocladium in Chaetomiaceae (100ML/1.00BY, Fig. 26).
Neoschizotheciaceae S.K. Huang & K.D. Hyde, fam. nov.
Facesoffungi number: *****??; Index Fungorum number: IF558385
Etymology: Named for Neoschizothecium, the type genus of this family.
Coprophilous or saprobic on soil or wood. Sexual morph: Ascomata perithecial, solitary to scattered or gregarious, immersed to semi-immersed or superficial, obpyriform or ovoid, membranaceous to coriaceous, brown to black, sometimes semi-transparent, ostiolate, with long or short necks, glabrous or surrounded by hairs, periphysate. Peridium comprising yellowish to pale brown or dark brown cells of textura angularis, textura globulosa or textura epidermoidea. Paraphyses absent, or cylindrical, septate. Asci 4- to multi-spored, unitunicate, cylindrical to clavate, pedicellate, evanescent. Ascospores uni- or bi-seriate, ellipsoidal or triangular, usually 0–1-septate, hyaline to dark brown, with germ pore at one or each end, usually with gelatinous appendages at one or each end, rarely surrounded by gelatinous sheath. Asexual morph: Hyphomycetous. Conidia globose to cylindrical, hyaline to pale brown, smooth-walled, ovate masses on the apex of the phialides (adapted from Marin-Felix et al. 2020).
Type genus: Neoschizothecium S.K. Huang & K.D. Hyde
Notes: The Neoschizotheciaceae clade accommodates the genera Apodus, Cercophora, Echria, Immersiella, Jugulospora, Lundqvistomyces, Neoschizothecium, Pseudoechria, Pseudoschizothecium, Rinaldiella and Zygopleurage based on the molecular data of their generic types, and also several Arnium, Apiosordaria, Podospora, Ramophialophora and Zopfiella species (88%ML, Fig. 26). These taxa are similar to Bombardiaceae, Podosporaceae and Lasiosphaeriaceae in having ellipsoidal ascospores, sometimes with appendages, but are phylogenetically distinct from these three families (Fig. 26). Marin-Felix et al. (2020) introduced this clade as Schizotheciaceae based on Schizothecium. However, Schizothecium is a synonym of Podospora (Podosporaceae) based on the type species S. fimicola, which nests in Podospora in the phylogenetic analysis (Wang et al. 2019; Index Fungorum 2020; this study, Fig. 26). We introduce Neoschizotheciaceae as a new family in Sordariales with 74%ML support (Fig. 26), based on Neoschizothecium. This family includes 11 genera and is characterized by ostiolate ascomata, cylindrical to clavate asci and ellipsoidal ascospores, sometimes with long or short cylindrical or lash-like gelatinous appendages. Marin-Felix et al. (2020) introduced three genera Lundqvistomyces, Pseudoechria and Pseudoschizothecium which we do not discuss in our study.
Apodus Malloch & Cain, Can. J. Bot. 49(6): 872 (1971)
Saprobic on Oryza sp. or coprophilous. Sexual morph: Mycelium hyaline to brown, septate, branched hyphae. Ascomata cleistothecial, solitary, superficial, globose to subglobose, brown to black, surrounded by hairs. Peridium composed of brown to hyaline cells of textura angularis. Asci 8-spored, unitunicate, clavate to cylindrical, blunt at the apex, evanescent. Ascospores bi-seriate or irregular, subglobose to ellipsoidal, 0–1-septate, pale brown to dark brown, with a germ pore at the apex, sometimes with hyaline sheath. Asexual morph: Undetermined (adapted from Malloch and Cain 1971).
Notes: Apodus is typified by A. deciduus in having ascomata with long hairs and ellipsoidal ascospores with a germ pore at the apex (Malloch and Cain 1971). Subsequently, A. oryzae was collected from a sheath of Oryza sativa in Italy, characterized by lacking an ostiole, but with dark brown ascomata covered with light brown, flexuous, septate hairs, clavate asci and ellipsoidal to broadly fusiform, 0–1-septate ascospores with an apical germ pore and sometimes covered with a hyaline gelatinous sheath (von Arx 1975). Apodus deciduus and A. oryzae were recognized as congeneric because of their similar morphology, but they do not cluster together in a phylogenetic analysis (Cai et al. 2006b). In this study, strains of A. deciduus and A. oryzae nested in Neoschizotheciaceae and Naviculisporaceae, respectively (Fig. 26). Therefore, we place Apodus in Neoschizotheciaceae based on phylogenetic placement of the generic type, A. deciduus. Apodus oryzae belongs to Naviculisporaceae but requires further data.
Type species: Apodus deciduus Malloch & Cain, Can. J. Bot. 49(6): 872 (1971)
Facesoffungi number: *****??; Fig. 45
Coprophilous. Sexual morph: Mycelium hyaline to brown, septate, branched hyphae 2–4.5 µm wide. Ascomata 195–500 µm (x̄ = 300 µm, n = 5) diam., cleistothecial, solitary, superficial, globose to subglobose, brown to black, surrounded by long, septate, brown, flexuous, tapering hairs. Peridium 15–30 µm wide, composed of brown to hyaline cells of textura angularis. Asci (45–)52–58(–60) × (12–)14–17 µm (x̄ = 55 × 15 µm, n = 20), 8-spored, unitunicate, clavate to cylindrical, with short pedicel, apex blunt, with an indistinct apical ring, evanescent. Ascospores 12–16 × 8–11.5 µm (x̄ = 14 × 9.5 µm, n = 20), bi-seriate or irregular, subglobose to ellipsoid, usually aseptate, often producing a single septum in culture, pale brown to dark brown, sometimes collapsing at the middle, with a germ pore at the apex. Asexual morph: Undetermined (adapted from Malloch and Cain 1971).
Material examined: USA, California, San Mateo Co., Crystal Springs Reservoir, on dung of dusky-footed wood rat in nest, 30 December 1969, D. Malloch (TRTC-045704, holotype).
Known hosts and distribution: On rat dung in the USA (type locality) (Malloch and Cain 1971).
Notes: We requested the holotype but the sample was too dry to find any clear structure except ascomata. The features of ascospores were found from a permanent mount. Cai et al. (2006b) sequenced the ex-type strain Apodus deciduus (CBS 506.70) which is basal to Cercophora newfieldiana and Zopfiella tardifaciens in Neoschizotheciaceae (100ML/0.96BY, Fig. 26).
Cercophora Fuckel, Jb. nassau. Ver. Naturk. 23-24: 244 (1870) [1869-70]
Saprobic on woody or herbaceous plants or coprophilous. Sexual morph: Ascomata perithecial, scattered to gregarious, superficial to semi-immersed, globose to subglobose, brown to black, glabrous or hairy, ostiolate, with papilla. Peridium composed of pale brown to brown cells of textura angularis. Asci 8-spored, unitunicate, cylindrical, pedicellate, evanescent. Ascospores bi-seriate, straight or slight curved, ellipsoidal to cylindrical, aseptate, hyaline, with guttules and bipolar appendages when young, becoming 2–multi-celled, upper cell(s) swelling to become fusiform, dark brown when mature; lower cell(s) hyaline when mature, smooth-walled, with guttules, appendages disappear with age. Asexual morph: Hyphomycetous. Conidiophores micronematous, reduced to conidiogenous cells. Conidia asepate, hyaline to brown, smooth-walled, globose to subglobose or usually with a truncate base and rounded apex (adapted from Fuckel 1870; del Valle Catania et al. 2011).
Notes: Cercophora is characterized by membranaceous to carbonaceous ascomata and ascospores with a swollen head and hyaline appendage(s) (Fuckel 1870). Chrysoporium, Cladorrhinum and Phialophora species have been published as the asexual morphs of Cercophora (Udagawa and Muroi 1979; von Arx 1981a; Ueda 1994; del Valle Catania et al. 2011). In this study, we found that Cercophora species are scattered in Lasiosphaeriaceae, Naviculisporaceae, Podosporaceae and Neoschizotheciaceae in Sordariales (Fig. 26). Molecular data for the generic type, Cercophora mirabilis, has been sequenced and found that this species grouped with hyphomycetous Ramophialophora globispora in Neoschizotheciaceae (Miller and Huhndorf 2005; Marin-Felix et al. 2020; this study, 100%ML/1.00BY, Fig. 26). Therefore, we place this genus in Neoschizotheciaceae.
Type species: Cercophora mirabilis Fuckel, Jb. nassau. Ver. Naturk. 23-24: 245 (1870) [1869-70]
Facesoffungi number: *****??; Fig. 46
Saprobic on woody or herbaceous plants or coprophilous. Sexual morph: Ascomata perithecial, scattered, semi-immersed, globose to subglobose, brown to black, surrounded by hairs, ostiolate, with papilla. Peridium composed of pale brown cells of textura angularis. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with subapical globule, evanescent. Ascospores bi-seriate, slight curved, ellipsoid, aseptate, hyaline, with guttules and bipolar appendages when young, becoming 2-celled, upper cell smaller than lower cell: the upper cell swelling to become fusiform, truncate at the base, dark brown when mature, (15–)18–25 × 10–15 µm (x̄ = 20 × 12 µm, n = 20); lower cell cylindrical, hyaline when mature, 32–40 × 4–5 µm (x̄ = 35 × 4.5 µm, n = 20), smooth-walled, with guttules, appendages disappear with age. Asexual morph: Hyphomycetous. Conidiophores micronematous, reduced to conidiogenous cells. Conidia asepate, hyaline to brown, globose to subglobose, smooth-walled (adapted from Udagawa and Muroi 1979; del Valle Catania et al. 2011).
Material examined: Sri Lanka, Hambantota Distr., southernmost part of Yala National Park near the coast, open dry jungle, on buffalo (Bubalus bubalis) dung in moist chamber, 11 February 1974, N. Lundqvist (UPS-UPS:BOT:F-698102).
Known hosts and distribution: On rotten stalks of cabbage in Germany (type locality) (Fuckel 1870); on dead woody or herbaceous plants or dung in Brazil, Canada, Corsiea, Denmark, Finland, Germany, Italy, Morocco, Norway, Poland, Scotland, Sri Lanka, Sweden (Lundqvist 1972).
Notes: Cercophora is typified by C. mirabilis (Fuckel 1870), which has vermiform ascospores with a swollen head, hyaline appendage(s) and a phialophora-like species was introduced as its asexual morph (Udagawa and Muroi 1979). Strain of C. mirabilis which nested in Neoschizotheciaceae were sequenced (Miller and Huhndorf 2005; Marin-Felix et al. 2020). We re-examined an authentic specimen determined by Lundqvist (mentioned in the label of material). The type strain Cercophora mirabilis (CBS 120402) is sister to Ramophialophora globispora (LC6218) in Neoschizotheciaceae (100%ML/1.00BY, Fig. 26).
Echria (N. Lundq.) Kruys, Huhndorf & A.N. Mill., Fungal Diversity 70: 106 (2014) [2015]
Coprophilous or saprobic on wood. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, globose to subglobose, with rigid hairs, ostiolate, with papilla. Asci 8-spored, unitunicate, cylindrical, pedicellate. Ascospores uni-seriate, ellipsoidal to fusiform, aseptate, smooth-walled or roughened wall, with a large central guttule and surrounded by gelatinous sheath. Asexual morph: Undetermined (adapted from Lundqvist 1972; Kruys et al. 2014).
Notes: Echria initially was introduced as a subgenus of Arnium and typified by Arnium macrotheca (Lundqvist 1972), and Kruys et al. (2014) raised Echria to generic level and transferred two Arnium species, A. gigantospora and A. macrotheca, to Echria based on LSU and TUB sequences data. Subsequently, Marin-Felix et al. (2020) showed that these two Echria species clustered and, in this study, two Echria species clustered (100%ML/1.00BY) and is sister to Rinaldiella pentagonospora in Neoschizotheciaceae (100%ML/0.96BY, Fig. 26). Echria different from Rinaldiella in having subglobose ascomata with rigid hairs and ellipsoidal ascospores surrounded by gelatinous sheaths, whereas Rinaldiella has subglobose ascomata covered with hairs and clavate ascospores (Lundqvist 1972; Crous et al. 2014).
Type species: Echria macrotheca (P. Crouan & H. Crouan) Kruys, Huhndorf & A.N. Mill., Fungal Diversity 70: 106 (2014) [2015]
Basionym: Sphaeria macrotheca P. Crouan & H. Crouan, Florule Finistère (Paris): 24 (1867)
Coprophilous or saprobic on wood. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, globose to subglobose, ostiolate, with papilla, surrounded by brown setae. Asci 8-spored, unitunicate, cylindrical, pedicellate. Ascospores uni-seriate, ellipsoidal to subglobose, aseptate, with a large central guttule and surrounded by gelatinous sheath. Asexual morph: Undetermined (adapted from Lundqvist 1972; Kruys et al. 2014).
Known hosts and distribution: On old cow dung among the Splachnum ampullaceum in France (type locality) (Crouan and Crouan 1867); on dung of cow, hare, horse, mule, rabbit, rat and sheep in Bulgaria, Canada, Denmark, England, France, Morocco, Puerto Rico, Sweden and USA (Lundqvist 1972).
Notes: Marin-Felix et al. (2020) established Schizotheciaceae and placed E. macrotheca in this family based on phylogenetic analysis. In this study, Echria macrotheca is sister to E. gigantospora in Neoschizotheciaceae (100%ML/1.00BY, Fig. 26).
Immersiella A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary to scattered, immersed to semi-immersed, ovoid to obpyriform, brown, verrucose, ostiolate, with papilla, covered with brown hairs. Peridium comprising membranaceous, hyaline to brown cells of textura angularis or textura prismatica. Asci 8-spored, unitunicate, cylindrical, pedicellate, with J- apical ring distinct, evanescent. Ascospores bi-seriate, cylindrical to geniculate, hyaline to pale brown, 0–multi-septate, guttulate, smooth-walled, with gelatinous appendages or appendage absent. Asexual morph: Undetermined (Miller and Huhndorf 2004a; Kruys et al. 2014).
Notes: Miller and Huhndorf (2004a) established Immersiella with I. caudata and I. immersa (type) in Sordariales based on phylogenetic analysis of LSU sequence data. Subsequently, Kruys et al. (2014) found that the clade comprising I. caudata and I. immersa was closely related to Arnium hirtum, which was isolated from dung in Scotland. In our study, A. hirtum, I. caudata and I. immersa also form a strongly supported clade in Neoschizotheciaceae (100%ML/0.97BY, Fig. 26). Therefore, we recommend to treat A. hirtum as Immersiella hirta.
Type species: Immersiella immersa (P. Karst.) A.N. Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Basionym: Lasiosphaeria immersa P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 163 (1873)
Facesoffungi number: *****??; Fig. 47
Saprobic on wood. Sexual morph: Ascomata 270–450 × 360–460 µm (x̄ = 325 × 370 µm, n = 10), perithecial, solitary to scattered, superficial to semi-immersed, sitting on subiculum, ovoid to obpyriform, brown, membranaceous, verrucose, ostiolate, with papilla. Subiculum composed of brown, septate, hyphae,1.5–3.5 µm wide. Peridium 55–95 µm (x̄ = 75 µm, n = 30) wide, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, branched, septate, filiform. Asci (200–)220–250(–270) × 14–17 µm (x̄ = 235 × 15 µm, n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with subapical globule, distinct J- apical ring, evanescent. Ascospores 35–45(–50) × 4.5–6.5 µm (x̄ = 42 × 5 µm, n = 50), bi-seriate, slight curved, cylindrical to geniculate, hyaline and aseptate when young, becoming pale brown and multi-septate, swelling at end when mature, with a large guttule in each cell, smooth-walled, with a germ pore at each end. Asexual morph: Undetermined.
Material examined: USA, Wisconsin, 45º0.00’ N / 90°0.00’ W, on dead wood, 9 June 1999, S.M. Huhndorf (F-SMH 4104).
Known hosts and distribution: On dead wood of Betula in Finland (type locality) (Karsten 1873); on dead wood in USA (Miller and Huhndorf 2004a).
Notes: Immersiella immersa is characterized by pyriform ascomata with cylindrical to geniculate ascospores (Miller and Huhndorf 2004a). We could not obtain the type material. Therefore, we re-examined an authentic specimen collected by Huhndorf.
New combination:
Immersiella hirta (E.C. Hansen) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria hirta E.C. Hansen, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 59: 22 (1877)
Synonym: Arnium hirtum (E.C. Hansen) N. Lundq. & J.C. Krug, Symb. bot. upsal. 20(no. 1): 218 (1972)
Index Fungorum number: IF558326
Jugulospora N. Lundq., Symb. bot. upsal. 20(no. 1): 256 (1972)
Coprophilous. Sexual morph: Ascomata perithecial, gregarious, superficial to semi-immersed, ovoid to subglobose, brown to black, membranaceous to carbonaceous, ostiolate, with central papilla, periphysate. Peridium composed of brown cells of textura angularis. Paraphyses numerous, filiform, flexuous, septate. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex truncate, with apical ring, evanescent. Ascospores uni-seriate, broadly ellipsoidal to ovoid, with an apical germ pore, hyaline when young, becoming upper cell aseptate, brown to dark brown, with plate-like wrinkles on surface when mature and truncate at the base; lower cell tiny, hyaline conical. Asexual morph: Undetermined (adapted from Lundqvist 1972).
Notes: Jugulospora is typified by J. rotula and characterized by ovoid ascomata and ovoid ascospores with a gelatinous cap-like cell at the base and a plate-like wrinkles (Lundqvist 1972). Marin-Felix et al. (2020) transferred this genus to Schizotheciaceae and introduced three more species, J. antarctrica, J. carbonaria and J. vestita based on morphology and phylogenetic analysis. This study also confirms that Jugulospora strain nest in Neoschizotheciaceae (50%ML/0.96BY, Fig. 26).
Type species: Jugulospora rotula (Cooke) N. Lundq., Symb. bot. upsal. 20(no. 1): 260 (1972)
Basionym: Sphaeria rotula Cooke, Handb. Brit. Fungi 2: 868 (1871)
Facesoffungi number: *****??; Fig. 48
Coprophilous. Sexual morph: Ascomata 230–250 × 150–230 µm (x̄ = 240 × 170 µm, n = 10), perithecial, gregarious, superficial to semi-immersed, ovoid to subglobose, brown to black, membranaceous to carbonaceous, ostiolate, with central, papilla, periphysate. Peridium 15–45 µm wide, composed of brown cells of textura angularis. Paraphyses numerous, filiform, flexuous, septate. Asci (80–)100–120(–150) × 9–15 µm (x̄ = 110 × 10 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex truncate, with apical ring, evanescent. Ascospores (10–)14–15(–16) × 8–10 µm (x̄ = 14.5 × 9 µm, n = 50), uni-seriate, broadly ellipsoidal to ovoid, with an apical germ pore, hyaline when young; becoming upper cell aseptate, brown to dark brown when mature, with plate-like wrinkles on surface and truncate at the base connected a tiny, hyaline conical lower cell. Asexual morph: Undetermined.
Material examined: UK, Surrey, Abinger, on burnt ground, October 1816, M.C. Cooke (NY-01050505, Isolectotype).
Known hosts and distribution: On burnt ground in Belgium, Canada, Denmark and England (type locality) (Cooke 1871; Lundqvist 1972).
Notes: The sequence data of Jugulospora rotula were sequenced (Miller and Huhndorf 2005; Marin-Felix et al. 2020) and in this study, these strains form a sister clade to J. minor (100%ML, Fig. 26) in Neochizotheciaceae. We re-examined a type of Jugulospora rotula which was collected by Cooke (mentioned in the label of material).
New combination:
Jugulospora minor (N. Lundq.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Strattonia minor N. Lundq., Symb. bot. upsal. 20(no. 1): 271 (1972)
Facesoffungi number: *****??; Index Fungorum number: IF558327; Fig. 42 b–j
Saprobic in soil. Sexual morph: Ascomata 300–400 × 320–340 µm (x̄ = 350 × 330 µm, n = 5), perithecial, solitary to gregarious, superficial to semi-immersed, membranaceous, pyriform to subglobose, brown to black, verrucose, ostiolate, with iso-radiating cells at surface, inner cells with hyaline periphyses. Peridium 15–60 µm wide, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 6–10 µm wide, filiform. Asci (120–)160–170(–230) × 8–12 µm (x̄ = 165 × 10 µm, n = 20), 8-spored, unitunicate, cylindrical. Ascospores 20–22(–25) × 6–10 µm (x̄ = 21 × 8 µm, n = 50), uni-seriate, oval to ellipsoidal, with a germ pore at the apex, hyaline when young, becoming upper cell aseptate, brown to dark brown when mature, verrucose; lower cell small, hyaline, caudal, triangular 2–3.5 × 2–3 µm. Asexual morph: Undetermined.
Material examined: Sweden, Uppland, Uppsala, Stadsskogen Forest, on burnt ground among Anthracobiae and small mosses, N. Lundqvist (UPS-UPS:BOT:F-117927, holotype).
Known hosts and distribution: On burnt soil in Belgium, Denmark, France and Sweden (type locality) (Lundqvist 1972).
Notes: Strattonia is characterized by ascospores with a large, brown cell and a short, hyaline cell, surrounded by gelatinous sheath (Lundqvist 1972). Miller and Huhndorf (2005) and Vu et al. (2019) sequenced Strattonia carbonaria and S. minor, and these two strains clustered with Jugulospora in Neoschizotheciaceae (100%ML/1.00BY, Fig. 26). These two species, have ascospores comprise a brown and a hyaline cell, but gelatinous sheath absent (Lundqvist 1972), are more similar to Jugulospora. Strattonia carbonaria has been transferred as Jugulospora carbonaria (Marin-Felix et al. 2020). Therefore, we transfer S. minor as Jugulospora minor based on morphology and phylogenetic analyses.
Neoschizothecium S.K. Huang & K.D. Hyde, gen. nov.
Etymology: Name refers to genus similar to, but different from Schizothecium
Facesoffungi number: *****??; Index Fungorum number: IF558329
Saprobic on wood, coprophilous. Sexual morph: Ascomata perithecial, solitary, scattered or gregarious, superficial to semi-immersed, pyriform, olivaceous-brown to dark brown, semi-transparent, surrounded by pale brown hyphae, ostiolate, with papilla. Peridium comprising membranaceous, pale brown cells of textura angularis. Paraphyses cylindrical, septate. Asci 8–multi-spored, unitunicate, cylindrical, evanescent. Ascospores uni-seriate to irregularly arranged, ellipsoidal to broadly fusiform, aseptate, hyaline to dark brown, truncate at the base with a cylindrical, slender, hyaline pedicel, umbonate apex, verrucose, with an apical germ pore. Asexual morph: Undetermined (adapted from Corda 1838; Lundqvist 1972; Munngai et al. 2012; Wang et al. 2019).
Notes: Schizothecium was introduced by Corda (1838) and is typified by S. fimicola and has membranaceous ascomata and ascospores with lash-like caudae (Corda 1838; Lundqvist 1972; Hu et al. 2006; Mungai et al. 2012). The generic type, Schizothecium fimicola, was synonymized as Podospora fimicola based on the epitype strain S. fimicola (CBS 482.64), which nested in Podospora (Podosporaceae) (Wang et al. 2019; Index Fungorum 2020). Marin-Felix et al. (2020) established Schizotheciaceae and raised Schizothecium as its type genus; however, the generic type of Schizothecium has not been clarified. Currently, Schizothecium is a synonym of Podospora under Podosporaceae, whereas several Schizothecium species clustered in Schizotheciaceae (Marin-Felix et al. 2020; this study, 100ML/1.00BY, Fig. 26). In this study, a novel genus Neoschizothecium is therefore proposed to accommodate species in the Neoschizotheciaceae clade (Fig. 26) and is typified by Neo. curvisporum (see the combination list below). This genus nests in Neoschizotheciaceae (95%ML) and is distinct from Podospora in multi-gene analysis (Fig. 26).
Type species: Neoschizothecium curvisporum (Cain) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria curvispora Cain, Canadian Journal of Research, Section C 26: 492 (1948)
Synonym: Schizothecium curvisporum (Cain) N. Lundq., Symb. bot. upsal. 20(no. 1): 334 (1972)
Index Fungorum number: IF558333; Fig. 49 q–r
Saprobic on seeds. Sexual morph: Ascomata perithecial, gregarious, superficial, pyriform, olivaceous brown to dark brown, semi-transparent, surrounded by pale brown hyphae, ostiolate, with papilla, periphysate. Peridium composed of membranaceous, pale brown cells of textura angularis. Asci 8-spored, unitunicate, cylindrical, apex rounded, pedicellate. Ascospores uni- or bi-seriate, reniform, aseptate, brown to dark brown, truncate at the base with a clavate, hyaline pedicel, with an apical germ pore. Asexual morph: Undetermined.
Known hosts and distribution: On seeds of Apium graveolens and Daucus carota in the USA (type locality) (Cain 1948).
Notes: Schizothecium curvisporum was established based on Sordaria curvispora which was isolated in culture from seed of Daucus carota L. var. sativa DC. in the USA (DAOM 20504) (Cain 1948; Lundqvist 1972). Subsequently, Vu et al. (2019) sequenced the ex-type strain S. curvisporum (CBS 506.50), and this strain nests in Neoschizothecium (100%ML/1.00BY, Fig. 26). In this study, we were unable to obtain authentic materials. Ascomata of N. curvisporum is similar to N. conicum and the hand-drawing is provided for the asci and ascospores (Fig. 49 q–r) based on Cain (1948).
New combinations:
Neoschizothecium aloides (Fuckel) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria aloides Fuckel, Fungi rhenani exsic., suppl., fasc.: no. 2549 (1868)
Synonym: Schizothecium aloides (Fuckel) N. Lundq., Symb. bot. upsal. 20(no. 1): 253 (1972)
Index Fungorum number: IF558331
Neoschizothecium carpinicola (Mouch.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora carpinicola Mouch., Persoonia 13(1): 197 (1986)
Synonym: Schizothecium carpinicola (Mouch.) L. Cai, Fungal Diversity 19: 14 (2005)
Index Fungorum number: IF558332
Neoschizothecium conicum (Fuckel) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cercophora conica Fuckel, Jb. nassau. Ver. Naturk. 23-24: 245 (1870)
Synonym: Schizothecium conicum (Fuckel) N. Lundq., Symb. bot. upsal. 20(no. 1): 253 (1972)
Facesoffungi number: *****??; Index Fungorum number: IF558328; Fig. 49 a–p
Coprophilous. Sexual morph: Ascomata 450–600 × 200–400 µm (x̄ = 500 × 320 µm, n = 5), perithecial, solitary or gregarious, superficial to semi-immersed, pyriform, brown to dark brown, membranaceous, ostiole, central, papillate with iso-radiating cells, surrounded by pale brown hyphae 2–4 µm wide, with hyaline, septate periphyses. Peridium 20–40 µm (x̄ = 30 µm, n = 30) wide, comprising two layers, outer layer composed of pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses tapering, cylindrical, septate. Asci (150–)165–175(–195) × 20–35 µm (x̄ = 170 × 20 µm, n = 30), 8-spored, unitunicate, cylindrical, apex rounded, with thickened wall, pedicellate, evanescent. Ascospores (16–)18–23(–25) × 10–16 µm (x̄ = 20 × 13 µm, n = 50), uni-seriate, ellipsoidal to broadly fusiform, aseptate, hyaline to dark brown, truncate at the base with a conical, hyaline pedicel, umbonate apex with a cylindrical, evanescent gelatinous appendage, verrucose, with an apical germ pore, collapsing when dry. Asexual morph: Undetermined.
Material examined: Sweden, Öland, Ås par., Ottenby lund., on fallen deer dung (Dama dama) in moist chamber in Stockholm, 16 May 1984, N. Lundqvist (S-F124742); Sweden, Skåne, Östra Vemmerlöv par., W of Hörte., in seashore meadow, on horse dung in moist chamber in Storvreta, 19 September 1996, N. Lundqvist (S-F124745).
Known hosts and distribution: On various kinds of dung, particularly cow and horse dung in all over the Nordic area, like Germany (type locality) and Sweden (Fuckel 1870; Lundqvist 1972); on roots in Canada (Vu et al. 2019).
Notes: Schizothecium conicum was found with Cercophora mirabilis on rotten cow dung under an oak tree (Fuckel 1870). Subsequently, Lundqvist (1972) clarified the relevant materials to Sch. conicum and proposed Sordaria curvula (= Podospora curvula) is a synonym. Vu et al. (2019) sequenced P. curvula (= Sch. conicum) (CBS 434.50), and this strain clustered with other Schizothecium strains with 100%ML/1.00BY support in Neoschizotheciaceae (Marin-Felix et al. 2020; this study, Fig. 26). In this study, we placed Sch. conicum in Neoschizothecium and re-examined an authentic specimen collected by Lundqvist.
Neoschizothecium fimbriatum (A. Bayer) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria fimbriata A. Bayer, Práce Mor. Přirodověd. Společn. Brno 1(3): 83 (1924)
Synonym: Schizothecium fimbriatum (A. Bayer) Barrasa & Soláns, Revta Ibér. Micol. 6(1): 3 (1989)
Index Fungorum number: IF558334
Neoschizothecium glutinans (Cain) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria glutinans Cain, Univ. Toronto Stud., Biol. Ser. 38: 40 (1934)
Synonym: Schizothecium glutinans (Cain) N. Lundq., Symb. bot. upsal. 20(no. 1): 254 (1972)
Index Fungorum number: IF558338
Neoschizothecium inaequale (Cain) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria inaequalis Cain, Canadian Journal of Research, Section C 26: 489 (1948)
Synonym: Schizothecium inaequale (Cain) N. Lundq., Symb. bot. upsal. 20(no. 1): 334 (1972)
Index Fungorum number: IF558339
Neoschizothecium minicaudum (Faurel & Locq.-Lin.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora minicauda Faurel & Locq.-Lin., Revue Mycol., Paris 42: 344 (1978)
Index Fungorum number: IF558340
Neoschizothecium selenosporum (Stchigel, Guarro & M. Calduch) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora selenospora Stchigel, Guarro & M. Calduch, Mycologia 94(3): 554 (2002)
Synonym: Schizothecium selenosporum (Stchigel, Guarro & M. Calduch) Y. Marín & Stchigel, in Marin-Felix et al., Microorganisms 8(9, no. 1430): 34 (2020)
Index Fungorum number: IF558341
Neoschizothecium tetrasporum (G. Winter) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria tetraspora G. Winter, Hedwigia 11: 161 (1872)
Synonym: Schizothecium tetrasporum (G. Winter) N. Lundq., Symb. bot. upsal. 20(no. 1): 256 (1972)
Index Fungorum number: IF558342
Rinaldiella Deanna A. Sutton, Y. Marín, Guarro & E.H. Thomps., in Crous et al., Persoonia 32: 301 (2014)
Human pathogen. Sexual morph: Ascomata perithecial, immersed, scattered, pyriform to subglobose, membranaceous, dark brown to black, surrounded by filiform hairs, ostiolate, with papilla. Peridium membranaceous, translucent, composed of brown to yellowish brown cells of textura epidermoidea. Paraphyses filiform, septate. Asci 8-spored, unitunicate, clavate to cylindrical, pedicellate, evanescent. Ascospores uni- to bi-seriate, hyaline, clavate and aseptate when young, upper cell becoming swollen to polygonal, brown, with truncate base and slightly acuminate apex; lower cell conical, subhyaline, slightly warted. Asexual morph: Undetermined (adapted from Crous et al. 2014).
Notes: The monotypic genus Rinaldiella was isolated from a contaminated human lesion and is characterized by pyriform ascomata with filiform brown hairs and clavate, 1-septate ascospores with a swollen, verrucose upper cell (Crous et al. 2014). It is different from Echria has subglobose ascomata with rigid hairs and ellipsoidal ascospores surrounded by gelatinous sheath (Lundqvist 1972). In this study, we place this species to Neoschizotheciaceae, based on its ex-type strain, Rinaldiella pentagonospora (CBS 132344) which is sister to Echria in Neoschizotheciaceae (100%ML/1.00BY, Fig. 26).
Type species: Rinaldiella pentagonospora Deanna A. Sutton, Y. Marín, Guarro & E.H. Thomps., in Crous et al., Persoonia 32: 301 (2014)
Fig. 34 n–o
Human pathogen. Sexual morph: Ascomata perithecial, immersed, scattered, pyriform to subglobose, membranaceous, dark brown to black, surrounded by filiform hairs, ostiolate, with papilla. Peridium membranaceous, translucent, composed of brown to yellowish brown cells of textura epidermoidea. Paraphyses filiform, septate. Asci 8-spored, unitunicate, clavate to cylindrical, pedicellate, evanescent. Ascospores uni- to bi-seriate, hyaline, clavate and aseptate when young, upper cell becoming swollen to polygonal, brown, with truncate base and slightly acuminate apex; lower cell conical, subhyaline, slightly warted. Asexual morph: Undetermined (adapted from Crous et al. 2014).
Known hosts and distribution: On human lesion in the USA (type locality) (Crous et al. 2014).
Notes: Rinaldiella is sister to Echria (100%ML/1.00BY, Fig. 26). The former is characterized by ascospores comprise a brown, polygonal upper cell and a subhyaline, conical lower cell, but different from the latter in having ellipsoidal ascospores surrounded by gelatinous sheath (Crous et al. 2014; Kruys et al. 2014).
Zygopleurage Boedijn, Persoonia 2(3): 316 (1962)
Coprophilous. Sexual morph: Ascomata perithecial, solitary to scattered, superficial to semi-immersed, semi-transparent, pyriform to subglobose, olivaceous-brown to black, verrucose, ostiolate, with short necks. Peridium comprising membranaceous, hyaline to brown cells of textura angularis. Asci 4–8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded. Ascospores irregularly arranged or spirally coiled around each other, hyaline, cylindrical to fusiform, aseptate when young, becoming elongate, the terminal cells swelling fusiform or oval and dark brown or olivaceous-brown with truncate base; the intercalary cell elongate, hyaline, sometimes slightly swelling when mature, usually 3-celled, sometimes multi-septate, with gelatinous appendages. Asexual morph: Undetermined (adapted from Boedijn 1962; Mirza and Nasir 1968; Lundqvist 1969, 1972).
Notes: All three species of Zygopleurage were collected from dung, especially cow dung, and the generic type, Zygopleurage zygospora, was first found in Italy and Zygopleurage faiyumensis and Z. multicaudata were collected in Egypt and Pakistan respectively (Boedijn 1962; Mirza and Nasir 1968; Lundqvist 1969). Lundqvist (1972) proposed that this genus is similar to Lasiosphaeria and Podospora based on ascospores comprise two brown cells connected by a long hyaline cell. Huhndorf et al. (2004b) sequenced Z. zygospora (SMH4219). In this study, this strain nested in Neoschizotheciaceae with 92%ML support (Fig. 26).
Type species: Zygopleurage zygospora (Speg.) Boedijn, Persoonia 2(3): 316 (1962)
Basionym: Sordaria zygospora Speg., Michelia 1(no. 2): 227 (1878)
Facesoffungi number: *****??; Fig. 50
Coprophilous. Sexual morph: Ascomata 0.7–1 × 0.5–0.8 mm, perithecial, solitary to scattered, superficial to semi-immersed, membranaceous, semi-transparent, pyriform to subglobose, olivaceous-brown, verrucose, ostiolate, with dark brown necks 300–400 × 150–300 µm, inner cells with hyaline periphyses. Peridium 25–60 µm wide, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci (250–)280–315(–330) × (35–)40–50(–55) µm (x̄ = 300 × 45 µm, n = 20), 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded and thick-walled. Ascospores bi-seriate, 3-celled, the two terminal cells hyaline to dark brown, ellipsoidal to fusiform with an apical germ pore and truncate at the base, (25–)27–32(–35) × (12–)14–18(–20) µm (x̄ = 30 × 15 µm, n = 50), with 4 distinct hyaline gelatinous processes at each end; connected by a long, hyaline 4–7 µm wide cell, with gelatinous ornamentation, evanescent. Asexual morph: Undetermined.
Material examined: Egypt, Faiyum Province, Al Fayyum, Libyan Desert, along the Cairo-Faiyum desert road, in the dried-up salt marsh at Kom Aushim 10 km N.E. of Lake Qarun, on cow dung in moist chamber in Uppsala, 1 March 1968, N. Lundqvist (S-F43798); Sweden, Småland, Vimmerby parish, 3 km S of Storebro, on cow dung after 3 weeks in moist chamber in Uppsala, 31 May 1959, N. Lundqvist (S-F43801).
Known hosts and distribution: On dung of cow, guinea pig, horse in Canada, Egypt, France, Italy (type locality), Liberia, Puerto Rico, Sweden and USA (Boedijn 1962; Lundqvist 1969).
Notes: We re-examined the authentic specimen collected by Lundqvist (mentioned in the label of material).
Sordariales genera incertae sedis
Arnium Nitschke ex G. Winter, Bot. Ztg. 31: 450 (1873)
Saprobic on wood, coprophilous. Sexual morph: Ascomata perithecial, scattered, superficial to semi-immersed, obpyriform to subglobose, membranaceous, pale brown to dark brown, ostiolate, with necks. Necks cylindrical to conical, black, glabrous or covered with unbranched, tapering, brown hairs, with hyaline, filiform periphyses. Paraphyses filiform to cylindrical, septate. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded, evanescent. Ascospores uni-seriate to irregularly arranged, obovoid to ellipsoidal to fusiform, aseptate, hyaline when young, becoming dark brown when mature, with ornamented wall and a gelatinous appendage at each end or surrounded by a gelatinous sheath. Asexual morph: Undetermined (adapted from Nitschke 1873; Lundqvist 1974).
Notes: Arnium is typified by A. lanuginosum and has subglobose ascomata, cylindrical to clavate asci and ascospores with a gelatinous appendage at each end (Nitschke 1873). Subsequently, A. lanuginosum was synonymized under Podospora brassicae (Ellis and Everhart 1892). This genus was rarely mentioned until it was clarified by Lundqvist (1972), and several Arnium species were then introduced (Cain and Mirza 1972; Krug and Cain 1972; Lundqvist 1974). Arnium species are phylogenetically scattered among the families of Sordariales, and several taxa have been transferred to Cladorrhinum, Podospora and Triangularia (Wang et al. 2019; Marin-Felix et al. 2020). In this study, Arnium species nested in Naviculisporaceae, Podosporaceae and Neoschizotheciaceae, but there is no molecular data for A. lanuginosum. Thus, we place the genus in Sordariales genera incertae sedis pending more data.
Type species: Arnium lanuginosum Nitschke [as 'lanuginosa'], Bot. Ztg. 31: 450 (1873)
Saprobic on wood. Sexual morph: Ascomata perithecial, superficial, globose to subglobose, tomentose. Asci 8-spored, unitunicate, cylindrical. Ascospores oval to ellipsoidal, dark brown, with gelatinous appendage at each end. Asexual morph: Undetermined (adapted from Nitschke 1873; Ellis and Everhart 1892).
Notes: Ellis and Everhart (1892) believed that Arnium lanuginosum and Podospora brassicae were conspecific, and these species were synonymized under A. olerum (Lundqvist 1972) based on similar ellipsoidal ascospores with cylindrical gelatinous appendages. In this study, A. olerum was transferred as C. olerum based on phylogenetic analysis (see notes for Cladorrhinum olerum). However, there is no molecular data for A. lanuginosum and we could not obtain any related specimens.
Biconiosporella Schaumann, Veröff. Inst. Meeresf. Bremerhaven 14(1): 24 (1972)
Saprobic on wood. Sexual morph: Ascomata perithecial, scattered, semi-immersed to superficial, globose to subglobose, collapsing when dry, membranaceous, black, ostiolate, with papilla. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, evanescent. Ascospores uni-seriate, obovoid to ellipsoidal, hyaline when young, becoming pale brown to brown when mature, smooth-walled, 3-septate, with a germ pore at each end and with tubercules at the middle. Asexual morph: Undetermined (adapted from Schaumann 1972).
Notes: Biconiosporella is characterized by obovoid to ellipsoidal ascospores with tubercules at the middle (Schaumann 1972). Jones et al. (2009) referred this genus to Lasiosphaeriaceae based on morphological observations. We found that its unique ascospores are similar to Stellatospora (Chaetomiaceae). Molecular data for B. corniculate is lacking, therefore, we place Biconiosporella in Sordariales genera incertae sedis pending fresh collection.
Type species: Biconiosporella corniculata Schaumann, Veröff. Inst. Meeresf. Bremerhaven 14(1): 24 (1972)
Facesoffungi number: *****??; Fig. 51
Saprobic on wood. Sexual morph: Ascomata 275–470 × 150–240 µm (x̄ = 370 × 180 µm, n = 5), perithecial, scattered, semi-immersed to superficial, globose to subglobose, collapsing when dry, membranaceous, black, ostiolate, with papilla. Peridium comprising dark brown to hyaline cells of textura angularis. Asci (200–)220–250(–300) × 20–35 µm (x̄ = 230 × 26 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, evanescent. Ascospores 28–35(–38) × 19–25 µm (x̄ = 33 × 22.5 µm, n = 50), uni-seriate, obovoid to ellipsoidal, hyaline when young, becoming pale brown to brown when mature, smooth-walled, with a germ pore at each end and 3–4 obvious coniform tubercules evenly distributed at middle, 3-septate, one septum near apex, two septa near the truncate base. Asexual morph: Undetermined.
Material examined: Germany, on rotten wood in sea water, 25 November 1969, K. Schaumann (NY-01388911, holotype).
Known hosts and distribution: On dead wood in Germany (type locality) (Schaumann 1972).
Notes: The monotypic Biconiosporella has unique ascospores with coniform tubercules at center. We re-examined the holotype which is well preserved, but there were no mature asci.
Camptosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 140 (1870) [1869-70]
Saprobic on stems or wood or coprophilous. Sexual morph: Ascomata perithecial, scattered, superficial to semi-immersed, erumpent through bark of host, pyriform to subglobose, black, glabrous or hairy, ostiolate, with papilla, periphysate. Peridium membranaceous. Paraphyses filiform. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, with J- apical ring. Ascospores broadly clavate to obpyriform, aseptate, hyaline to brown, smooth-walled or verrucose, with conical gelatinous appendage at the apex. Asexual morph: Undetermined (adapted from Fuckel 1870; Lundqvist 1972).
Notes: Fuckel (1870) established Camptosphaeria which is typified by Cam. sulphurea. Lundqvist (1972) reduced this genus as a subgenus of Cercophora because the cylindrical to broadly fusiform ascospores are similar to Cercophora and Podospora. Krug and Jeng (1977) re-defined Camptosphaeria as a genus which has pod-like ascospores with conical gelatinous appendages and suggested that its ascospores are similar to Bombardia and Lasiosphaeria.
Type species: Camptosphaeria sulphurea Fuckel, Jb. nassau. Ver. Naturk. 23-24: 140 (1870) [1869-70]
Facesoffungi number: *****??; Fig. 52
Saprobic on stems. Sexual morph: Ascomata 520–650 × 400–450 µm (x̄ = 560 × 420 µm, n = 5), perithecial, scattered, semi-immersed, erumpent through bark of host, pyriform to subglobose, black, ostiolate, with papilla, periphysate. Peridium 20–50 µm (x̄ = 30 µm, n = 30) wide, membranaceous, comprising two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses filiform. Asci (130–)140–170(–180) × 12–25 µm (x̄ = 150 × 18 µm, n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with apical globule. Ascospores (25–)28–30(–35) × 10–15 µm (x̄ = 30 × 13 µm, n = 30), bi-seriate, cylindrical to broadly clavate to obpyriform, aseptate, hyaline to pale brown, smooth-walled, with conical gelatinous appendage at the apex. Asexual morph: Undetermined (adapted from Fuckel 1870; Lundqvist 1972).
Material examined: Germany, Hessen, Nessen, Östrich, on stems of Peucedanum officinale, 1894, L. Fuckel (G-318793/1, holotype).
Known hosts and distribution: On stems of Peucedanum in Germany (type locality) (Fuckel 1870).
Notes: Fuckel (1870) introduced Camptosphaeria sulphurea with pod-like ascospores having conical gelatinous appendages. We re-examined the holotype (318793/1) which was determined by Lundqvist. Complete asci and ascospores were not observed, so we drew them with reference to Fuckel (1870) and Lundqvist (1972).
Diffractella Guarro, P.F. Cannon & Aa, Syst. Ascom. 10: 107 (1991)
Saprobic on wood. Sexual morph: Ascomata cleistothecial, solitary, superficial, carbonaceous, globose, with alveolate-reticulate ornamentation, brown to black, surrounded by long setae. Peridium composed of brown to reddish brown, radial, septate, cylindrical cells. Asci 8-spored, unitunicate, globose to oval. Ascospores multi-seriate, fusiform to falciform, upper cell(s) brown to dark brown, aseptate or with 1-septate near the base, guttulate; lower cell cap-like, hyaline. Asexual morph: Undetermined (adapted from Fuckel 1872; Guarro et al. 1991).
Notes: Guarro et al. (1991) transferred Cephalotheca curvata (Fuckel 1872) to Diffractella as the generic type, which is characterized by cleistothecial ascomata, globose asci and fusiform ascospores with cap-like basal cell. The genus was considered as a member of Lasiosphaeriaceae because it has falciform ascospores with a special cap-like cell, which is similar to Bellojisia and Zopfiella (Guarro et al. 1991; Réblová 2008).
Type species: Diffractella curvata (Fuckel) Guarro, P.F. Cannon & Aa, Syst. Ascom., Reprint of Volumes 1-4 (1982-1985) 10(2): 108 (1991)
Basionym: Cephalotheca curvata Fuckel, Jb. nassau. Ver. Naturk. 25-26: 298 (1871)
Facesoffungi number: *****??; Fig. 53
Saprobic on wood. Sexual morph: Ascomata 245–400 µm (x̄ = 300 µm, n = 5) diam., cleistothecial, solitary, superficial, carbonaceous, globose, with alveolate-reticulate ornamentation, brown to black, surrounded by long setae. Setae 4–6 µm wide, filiform, septate, brown and tapering, lighter at the rounded apex. Peridium composed of brown to reddish brown, radial-shaped, septate, cylindrical cells. Asci 15–22 µm (x̄ = 18 µm, n = 20) diam., 8-spored, unitunicate, globose to oval. Ascospores 15–18(–21) × 4.5–7.5 µm (x̄ = 17 × 6 µm, n = 20), multi-seriate, fusiform to falciform, upper cell(s) brown to dark brown, aseptate or with 1-septate near the base, guttulate; lower cell cap-like, hyaline, sometimes collapsing when dry. Asexual morph: Undetermined.
Material examined: Germany, Baden-Wurttemberg, Ca. Eberbach, 9 º0’11’’E/48 º35’33’’N, on rotten oak wood, 1894, L. Fuckel (G-352276/1, holotype).
Known hosts and distribution: On rotten hollow trunks of oak in Germany (type locality) (Fuckel 1872).
Notes: We re-examined the holotype of Diffractella curvata (G-352276/1).
Emblemospora Jeng & J.C. Krug, Can. J. Bot. 54(16): 1971 (1976)
Coprophilous. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, pyriform to subglobose, brown to black, ostiolate, with black necks, surrounded by setae. Peridium membranaceous, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci 8-spored, unitunicate, subglobose to clavate, pedicellate, apex rounded or truncate, with J- apical ring, evanescent. Ascospores bi-seriate, ellipsoidal to subglobose, brown to dark brown, aseptate, with plate-like wrinkled wall, germ pore at one or both end(s), guttulate. Asexual morph: Undetermined (adapted from Jeng and Krug 1976).
Notes: Jeng and Krug (1976) established Emblemospora, which includes two species, E. ditrema and E. monotreme (type), in Sordariaceae. Species are similar to Diplogelasinospora, Jugulospora and Neurospora in having pyriform ascomata and clavate asci with one-celled ascospores with ornamentation (Jeng and Krug 1976). Molecular data is unavailable for this genus.
Type species: Emblemospora monotrema Jeng & J.C. Krug, Can. J. Bot. 54(16): 1972 (1976)
Facesoffungi number: *****??; Fig. 54
Coprophilous. Sexual morph: Ascomata 450–600 × 230–380 µm (x̄ = 500 × 300 µm, n = 5), perithecial, solitary, superficial to semi-immersed, pyriform to subglobose, brown to black, membranaceous, ostiolate, with black necks (65–100 × 100–120 µm), surrounded by setae, with hyaline, septate periphyses. Setae 1.5–3.5 µm wide, numerous, cylindrical, erect, septate, apex rounded, brown and tapering lighter at the apex. Peridium 18–35 µm (x̄ = 27 µm, n = 20) wide, two layers, outer layer composed of brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Asci 160–225 × 25–55 µm (x̄ = 190 × 40 µm, n = 30), 8-spored, unitunicate, subglobose to clavate, pedicellate, apex rounded, with J- apical ring, evanescent. Ascospores 27–35 × 15–25 µm (x̄ = 30 × 20 µm, n = 50), bi-seriate, ellipsoidal to subglobose, brown to dark brown, aseptate, with plate-like wrinkled wall and an apical germ pore, guttulate. Asexual morph: Undetermined.
Material examined: Venezuela, Edo. Scure, NW of Irapa, trail between Manacal and Los Pocitos, on dung of burro, 13 July 1972, K.P. Dumont, R.F. Cain, G.J. Samuels, G. Morillo and J. Farfan (TRTC-175786, holotype).
Known hosts and distribution: On dung of burro in Venezuela (type locality) (Jeng and Krug 1976).
Notes: Emblemospora monotrema is similar to Neurospora species and is characterized by pyriform ascomata and clavate asci with one-celled ornamented ascospores (Jeng and Krug 1976).
Eosphaeria Höhn., Annls mycol. 15(5): 362 (1917)
Saprobic in soil. Sexual morph: Ascomata perithecial, gregarious, superficial, globose to subglobose, brown to black, carbonaceous, verrucose, ostiolate, papillate, periphysate. Peridium carbonaceous to membranaceous, composed of black to pale brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, filiform, evanescent. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with subapical globule, evanescent. Ascospores bi-seriate, slightly curved, aseptate, multi-guttulate, hyaline and ellipsoidal when young, becoming multi-septate, geniculate, 2/3 upper cell swelling, truncate at the base, dark brown when mature; 1/3 lower cell hyaline when mature, smooth-walled, with bipolar short appendages, disappearing with age. Asexual morph: Hyphomycetous, phialophora-like. Conidia small, oval to globose (adapted from von Höhnel 1917).
Notes: The monotypic genus Eosphaeria is typified by E. uliginosa and is characterized by subglobose ascomata and L-shaped ascospores with short appendage at each end (von Höhnel 1917). The genus was considered similar to Lasiosphaeria, and was thus initially accommodated in Lasiosphaeriaceae (Barr 1990; Kirk et al. 2001; Huhndorf et al. 2004b). Except for appendages at each end in ascospores, Eosphaeria is similar to Immersiella in having black ascomata, cylindrical asci with an apical globule, and L-shaped ascospores. Molecular data is unavailable for Eosphaeria.
Type species: Eosphaeria uliginosa (Fr.) Höhn., Annls mycol. 15(5): 362 (1917)
Basionym: Sphaeria uliginosa Fr., in Kunze & Schmidt, Mykologische Hefte (Leipzig) 2: 39 (1823)
Facesoffungi number: *****??; Fig. 55
Saprobic in soil. Sexual morph: Ascomata 280–620 µm (x̄ = 480 µm, n = 10) diam., perithecial, gregarious, superficial, globose to subglobose, brown to black, carbonaceous, verrucose, ostiolate, with papilla, periphysate. Peridium 60–100 µm (x̄ = 75 µm, n = 20) wide, comprising two layers, outer layer carbonaceous, composed of black cells; inner layer membranaceous, composed of pale brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, filiform, evanescent. Asci (170–)190–250(–280) × 13–17 µm (x̄ = 225 × 16 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with subapical globule, evanescent. Ascospores bi-seriate, slightly curved, aseptate, multi-guttulate, hyaline and ellipsoidal when young, becoming multi-septate, geniculate, 2/3 upper cells swelling, truncate at the base, dark brown when mature, 30–35(–45) × 4–7 µm (x̄ = 34 × 5.5 µm, n = 50); 1/3 lower cells hyaline when mature, 12–18(–20) × 4–6 µm (x̄ = 15 × 4.5 µm, n = 50), smooth-walled, with bipolar short conical appendages (upper cauda 3.5–5.5 × 1–2.5 µm; lower cauda 6.5–10 × 1.5–3 µm), disappearing with age. Asexual morph: Hyphomycetous, phialophora-like. Conidia small, oval to globose (adapted from von Höhnel 1917).
Material examined: USA, Maine, Hancock Co. Township of 728MD, Lead Mountain (Humpback), along road near parking area, 44°51’28’’N, 68°05’52’’W, beech-maple forest, in soil, 2 September 2014, R.C. Harris (NY-01818643).
Known hosts and distribution: On soil in marshy places in Germany (type locality) (Fries 1823b; von Höhnel 1917); on clay soil in USA (Ellis and Everhart 1893).
Notes: Lasiosphaeria dichroospora, which was collected on clay soil in USA, was synonymized under Eosphaeria uliginosa (Ellis and Everhart 1893; von Höhnel 1917). We could not obtain type material of E. uliginosa and, therefore, re-examined an authentic specimen collected from USA by Harris with features similar to those described in Ellis and Everhart (1893) and von Höhnel (1917).
Isia D. Hawksw. & Manohar., Trans. Br. mycol. Soc. 71(2): 332 (1978)
Saprobic on leaves or soil. Sexual morph: Ascomata perithecial, solitary, immersed, membranaceous, globose, dark brown to black, ostiolate, with papilla. Peridium composed of membranaceous, brown to black cells of textura angularis. Asci 8-spored, unitunicate, cylindrical, evanescent. Ascospores uni-seriate, subglobose to ellipsoid, 0–1-septate, hyaline to pale brown, with dense irregular verrucose ornamentation on surface. Asexual morph: Undetermined (adapted from Hawksworth and Manoharachary 1978; Udagawa and Sugiy 1982).
Notes: Hawksworth and Manoharachary (1978) introduced Isia with I. neocaledoniensis as the type. It was transferred from Thielavia, and is similar to Copromyces and Jugulospora in having ascospores with verrucose ornamentation on the surface, although Copromyces has globose ascospores and Jugulospora has a conical pedicel on ascospores (Hawksworth and Manoharachary 1978). Therefore, this genus is accommodated in Sordariales genera incertae sedis (Hawksworth and Manoharachary 1978; Maharachchikumbura et al. 2016; Hyde et al. 2020).
Type species: Isia neocaledoniensis (C. Moreau) D. Hawksw. & Manohar., Trans. Br. mycol. Soc. 71(2): 334 (1978)
Basionym: Thielavia neocaledoniensis C. Moreau, Bull. trimest. Soc. mycol. Fr. 81(4): 699 (1966)
Facesoffungi number: *****??; Fig. 56
Saprobic on leaves. Sexual morph: Ascomata perithecial, solitary, immersed, membranaceous, globose, dark brown to black, ostiolate, with papilla, periphysate. Peridium composed of membranaceous, brown to black cells of textura angularis. Asci (118–)120–135(–150) × 13–18(–22) µm (x̄ = 130 × 16 µm, n = 20) ascospore-bearing part, 8-spored, unitunicate, cylindrical, apex rounded or truncate, with J- apical ring. evanescent. Ascospores 18–23(–25) × (7.5–)9–12(–14) µm (x̄ = 20 × 10 µm, n = 50), uni-seriate, subglobose to ellipsoid, 0–1-septate, hyaline when young, becoming pale brown at maturity, apex cuspidal and base blunt, wall with densely verrucose ornamentation. Asexual morph: Undetermined.
Material examined: India, Bihar, on leaves of Pandanus sp., 23 April 1977, M.A. Rizwi (IMI-213076, holotype).
Known hosts and distribution: On leaves of Pandanus in New Caledonia and India (type locality) (Hawksworth and Manoharachary 1978).
Notes: Thielavia neocaledoniensis was transferred to Isia which was considered as a member of Sordariaceae (Hawksworth and Manoharachary 1978). We re-examined the type (213076) that contains a permanent mount but we could find only a few ascospores in its ascomata.
Lockerbia K.D. Hyde, Sydowia 46(1): 23 (1994)
Saprobic on palm fronds or wood. Sexual morph: Ascomata cleistothecial, solitary, superficial, membranaceous, globose, dark brown to black. Peridium composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses numerous, filamentous, branched, embedded in a gelatinous matrix. Asci 8-spored, unitunicate, cylindrical to subclavate, pedicellate, apex rounded and thick-walled. Ascospores uni-seriate, limoniform to oval, hyaline to brown to dark brown when mature, wall minutely verrucose, aseptate, guttulate, surrounded by inconspicuous hyaline mucilaginous sheath. Asexual morph: Undetermined (adapted from Hyde 1994; Raja and Shearer 2008).
Notes: Lockerbia palmicola was introduced as generic type, and is similar to Diplogelasinospora, Neurospora and Sordaria species in having ellipsoidal ascospores with slightly irregular verrucose wall and surrounded by a hyaline mucilaginous sheath (Hyde 1994; Raja and Shearer 2008). Hence, Lockerbia was considered as a member in Sordariales genera incertae sedis (Maharachchikumbura et al. 2016; Hyde et al. 2020).
Type species: Lockerbia palmicola K.D. Hyde, Sydowia 46(1): 24 (1994)
Facesoffungi number: *****??; Fig. 57
Saprobic on palm fronds. Sexual morph: Ascomata 220–390 µm (x̄ = 280 µm, n = 5) diam., cleistothecial, solitary, superficial, membranaceous, globose, dark brown to black. Peridium 15–23 µm (x̄ = 20 µm, n = 20) wide, comprising two layers, outer layer composed of brown to black cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 1–2 µm wide, numerous, filamentous, branched, embedded in a gelatinous matrix. Asci (120–)145–160(–170) × (13–)16–23(–25) µm (x̄ = 150 × 20 µm, n = 20), 8-spored, unitunicate, cylindrical to subclavate, pedicellate, apex rounded and thick-walled. Ascospores (16–)18–21(–25) × 10–16(–18) µm (x̄ = 20 × 15 µm, n = 50), uni-seriate, limoniform to oval, hyaline when young, brown to dark brown when mature, aseptate, guttulate, wall minutely verrucose, surrounded by inconspicuous hyaline mucilaginous sheath. Asexual morph: Undetermined.
Material examined: Australia, north Queensland, Bamaga, ‘Lockerbie’ rainforest, on dead palm rachides (Archontophoenix sp.) on forest floor, February 1992, K.D. Hyde (BRIP-21334, holotype).
Known hosts and distribution: On dead rachides of palm in Australia (type locality) (Hyde 1994).
Notes: Hyde (1994) found Lockerbia palmicola is characterized by cleistothecial ascomata and ellipsoidal to oval ascospores. We re-examined the type (BRIP-21334).
Periamphispora J.C. Krug, Mycologia 81(3): 476 (1989)
Coprophilous. Sexual morph: Ascomata perithecial, solitary to scattered, superficial to semi-immersed, globose to subglobose, brown, membranaceous to coriaceous, tuberculate, surrounded by hairs, ostiolate, with papilla. Peridium composed of pale brown to hyaline cells of textura porrecta to textura angularis to textura prismatica. Paraphyses numerous, filiform, septate, evanescent. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded. Ascospores bi-seriate, ellipsoidal to oval, aseptate, ends rounded, with a germ pore at the apex, hyaline when young, becoming brown to reddish brown, rough-walled, surrounded by hyaline, gelatinous sheath. Asexual morph: Undetermined (adapted from Krug 1989).
Notes: The monotypic genus Periamphispora is characterized by globose to subglobose ascomata with hairs and ellipsoidal ascospores irregularly ribbed on the surface, which is similar to Fimetariella, Podospora and Sordaria (Krug 1989). This genus was previously considered a member in Lasiosphaeriaceae (Krug 1989; Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2020). This genus is also similar to Podospora and Sordaria in having hairy ascomata, clavate to cylindrical asci and ellipsoidal ascospores with sheath, however molecular data are unavailable.
Type species: Periamphispora phacelodes J.C. Krug, Mycologia 81(3): 476 (1989)
Facesoffungi number: *****??; Fig. 58
Coprophilous. Sexual morph: Ascomata 650–700 × 360–380 µm (x̄ = 670 × 370 µm, n = 5), perithecial, solitary to scattered, superficial to semi-immersed, globose to subglobose, brown, membranaceous to coriaceous, tuberculate, ostiolate, with necks. Necks brown to dark brown, surrounded by numerous, pale brown, filiform, septate, hairs 1.5–3.5 µm wide. Peridium 33–55 µm (x̄ = 43 µm, n = 30) wide, outer layer composed of pale brown to reddish brown cells of textura porrecta to textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, filiform, septate, evanescent. Asci 200–300 × 40–65 µm (x̄ = 250 × 55 µm, n = 30) ascospore-bearing part, 8-spored, unitunicate, cylindrical to clavate, pedicellate, about 100 µm long, apex rounded. Ascospores (30–)40–45(–50) × 20–25(–30) µm (x̄ = 42 × 23 µm, n = 50), bi-seriate, ellipsoidal to oval, aseptate, ends rounded, with a germ pore at the apex, hyaline when young, becoming brown to reddish brown, rough-walled, surrounded by hyaline, gelatinous sheath, 2–5 µm wide. Asexual morph: Undetermined.
Material examined: Spain, Málaga, about 20km W of Málaga, Benalmádena (36°36’N, 4°34’W), on horse dung, 30 April 1971, P. Blaser (TRTC-55023, holotype).
Known hosts and distribution: On horse dung in Spain (type locality) (Krug 1989).
Notes: We re-examined the type, Periamphispora phacelodes 55023, which was grown on dung extract agar medium culture (Weitzman and Silva-Hutner 1967), and it is well preserved.
Ramophialophora M. Calduch, Stchigel, Gené & Guarro, Stud. Mycol. 50(1): 84 (2004)
Saprobic on soil and wood. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Mycelium superficial to immersed. Hyphae pale brown, septate, branched. Conidiophores macronematous, mononematous, erect, cylindrical, septate, branched, brown to dark brown. Conidiogenous cells monophialidic or polyphialidic, terminal or/and lateral. Conidia brown, globose to oval, aseptate, with a protuberant basal hilum (adapted from Calduch et al. 2004).
Notes: Ramophialophora is typified by R. vesiculosa and has macronematous, mononematous conidiophores and aseptate, brown conidia in slimy masses (Calduch et al. 2004). The genus was accepted as a member of Lasiosphaeriaceae based on phylogenetic analysis (Hyde et al. 2020; Wijayawardene et al. 2020). In this study, the strains of Ramophialophora are scattered in Lasiosphaeriaceae, Neoschizotheciaceae and Zygospermellaceae in Sordariales (Fig. 26), but there is no molecular data for R. vesiculosa. Therefore, we recommend transferring this genus to Sordariales genera incertae sedis until more evidence is available.
Type species: Ramophialophora vesiculosa M. Calduch, Stchigel, Gené & Guarro, in Calduch, Gené, Cano, Stchigel, Cano & Guarro, Stud. Mycol. 50(1): 84 (2004)
Facesoffungi number: *****??; Fig. 59
Saprobic on soil and wood. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on sterilized wood, hairy, brown. Mycelium superficial to immersed on substrate. Hyphae 1.5–2.5 µm wide, pale brown, septate, branched. Conidiophores 2–6.5 µm wide, macronematous, mononematous, erect, cylindrical, septate, branched, pale brown to dark brown, becoming paler towards the apex. Conidiogenous cells 5.5–9 × 2.5–4 µm (x̄ = 7 × 3.5 µm, n = 30), monophialidic or polyphialidic, terminal and lateral, lageniform, smooth-walled, with conspicuous collarettes, 1.5–2.5 µm wide. Conidia 2–3 µm (x̄ = 2.5 µm, n = 30) diam., aggregated, pale brown to brown, globose to ellipsoidal, aseptate, smooth-walled, with a truncate and protuberant base.
Material examined: Spain, Asturias, Muniellos Integral Biological Reserve, in soil (isolated on sterilized wood), 26 June 1999, M. Calduch and A.M. Stchigel (IMI-389151, holotype).
Known hosts and distribution: On soil in Spain (type locality) (Calduch et al. 2004).
Notes: In this study, we re-examined the type, R. vesiculosa (IMI-389151), but molecular data is lacking.
Reconditella Matzer & Hafellner, Biblthca Lichenol. 37: 46 (1990)
Saprobic on lichen. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, membranaceous, pyriform to subglobose, dark brown to black, verrucose, surrounded by brown, septate setae, ostiolate, with papilla, periphysate. Peridium composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses unbranched, filiform. Asci 4–8-spored, unitunicate, cylindrical, pedicellate, apex rounded. Ascospores uni- or bi-seriate, ellipsoidal to fusiform, slightly curved, 0–1-septate, hyaline to pale brown, striate and/or verrucose, guttulate. Asexual morph: Undetermined (adapted from Matzer and Hafellner 1990).
Notes: The monotypic genus Reconditella is typified by R. physconiarum which was initially considered as a member of Sordariales (Matzer and Hafellner 1990). We re-examined the type material and found that this genus is characterized by ellipsoidal to broadly fusiform, 0–1-septate ascospores with fine warts forming ridges, which is similar to Podospora and Neurospora. Therefore, we place this genus in Sordariales genera incertae sedis.
Type species: Reconditella physconiarum Hafellner & Matzer, in Matzer & Hafellner, Biblthca Lichenol. 37: 47 (1990)
Facesoffungi number: *****??; Fig. 60
Saprobic on lichen. Sexual morph: Ascomata 270–350 × 200–275 µm (x̄ = 320 × 255 µm, n = 5), perithecial, solitary, superficial to semi-immersed, membranaceous, pyriform to subglobose, dark brown to black, verrucose, surrounded by brown, septate setae 3.5–6.5 µm wide, ostiolate, with papilla, periphysate. Peridium 40–60 µm (x̄ = 45 µm, n = 20) wide, comprising two layers, outer layer composed of brown to black cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 5–6 µm wide, unbranched, filiform. Asci (70–)95–110(–120) × 10–17(–20) µm (x̄ = 100 × 15 µm, n = 20), 4–8-spored, unitunicate, cylindrical, pedicellate, apex rounded. Ascospores 15–20(–25) × 6–8(–9.5) µm (x̄ = 18 × 7.5 µm, n = 50), uni- or bi-seriate, ellipsoidal to fusiform, slightly curved, 0–1-septate, hyaline to pale brown, striate and/or verrucose, guttulate. Asexual morph: Undetermined.
Material examined: Austria, Steiermark, Gesäuse area, Johnsbach, c. 0.5 km E of Gasthof Kölbl. Alt. c. 875 m. MTB 8453/4, on Fraxinus excelsior, on thallus of Physconia distorta, 20 May 1988, S.J. Hafellner and E. Schreiner (GZU-287606, holotype); ibid. (S-F20696, isotype).
Known hosts and distribution: On Physconia distorta in Austria (type locality) (Matzer and Hafellner 1990).
Notes: We re-examined the type specimens of Reconditella physconiarum. Specimen 287606 has fragile ascomata and has good quality permanent mount; F20696 is well preserved. There is no molecular data available for R. physconiarum.
Roselliniopsis Matzer & Hafellner, Biblthca Lichenol. 37: 97 (1990)
Saprobic on wood or lichens. Sexual morph: Ascomata perithecial, solitary, superficial to semi-immersed, membranaceous, pyriform to subglobose, black, verrucose, ostiolate, with papilla. Peridium composed of pale brown to hyaline cells of textura angularis to textura prismatica. Paraphyses evanescent. Asci 8-spored, unitunicate, cylindrical to ellipsoidal, pedicellate, apex rounded. Ascospores uni-seriate, ellipsoidal to fusiform, aseptate, pale brown to dark brown, smooth-walled, with guttules. Asexual morph: Undetermined (adapted from Matzer and Hafellner 1990; Matzer 1993; Alstrup et al. 1994; Kondratyuk et al. 2013).
Notes: Roselliniopsis was established as a lichenicolous genus in Sordariales and is typified by R. groedensis (Matzer and Hafellner 1990; Maharachchikumbura et al. 2016; Hyde et al. 2020; Wijayawardene et al. 2020). It is placed in Sordariales genera incertae sedis and characterized by superficial ascomata with brown setae and pale brown ascospores (Matzer and Hafellner 1990), but no molecular data is available.
Type species: Roselliniopsis groedensis (Zopf) Matzer & Hafellner, Biblthca Lichenol. 37: 99 (1990)
Basionym: Rosellinia groedensis Zopf, Hedwigia 35(6): 350 (1896)
Facesoffungi number: *****??; Fig. 61
Lichenicolous. Sexual morph: Ascomata 300–370 × 300–400 µm (x̄ = 330 × 350 µm, n = 5), perithecial, solitary, superficial to semi-immersed, membranaceous, pyriform to subglobose, black, verrucose, ostiolate, with papilla. Setae 5–10 µm wide, brown, septate, apex rounded, cells swollen near the septum. Peridium 50–120 µm (x̄ = 75 µm, n = 20) wide, comprising two layers, outer layer composed of pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses evanescent. Asci (100–)110–125(–132) × (10–)14–17(–22) µm (x̄ = 120 × 15 µm, n = 20), 8-spored, unitunicate, cylindrical to ellipsoidal, pedicellate, apex rounded. Ascospores 14–20(–22) × (8–)10–12(–14) µm (x̄ = 18 × 11 µm, n = 50), uni-seriate, oval to fusiform, aseptate, pale brown to dark brown, smooth-walled, with germ pore at each end, guttulate. Asexual morph: Undetermined.
Material examined: Italy, Trentino-Alto Adige, Bolzano, St. Ulrich in Gröden (Südtirol), on Pertusaria sulphurella Köerb. var. variolosa, W. Zopf (S-F47825).
Known hosts and distribution: On lichen in Italy (type locality) (Zopf 1896; Matzer and Hafellner 1990).
Notes: Zopf (1896) introduced lichenicolous Rosellinia groedensis and Matzer and Hafellner (1990) re-examined its isotype from Italy and designated it as the type species of Roselliniopsis. We re-examined an authentic specimen collected by Zopf and found it to be similar to Podospora and Sordaria in having membranaceous, black ascomata and cylindrical asci with smooth-walled, ellipsoidal to fusiform ascospores, but it lacks molecular data.
Synaptospora Cain, Beih. Sydowia 1: 4 (1957) [1956]
Saprobic on wood. Sexual morph: Ascomata perithecial, superficial, gregarious to scattered, globose to subglobose, carbonaceous, with ostiole, black, verrucose, surrounded by setae. Peridium carbonaceous to membranaceous, composed of hyaline to brown cells of textura angularis. Paraphyses septate, cylindrical, evanescent. Asci 2–8-spored, unitunicate, cylindrical to clavate, apex broadly rounded or truncate. Ascospores uni- or bi-seriate, globose to subglobose, aseptate, hyaline when young, becoming pale brown to dark brown, fused in groups. Asexual morph: Undetermined (adapted from Cain 1957b; Huhndorf et al. 1999b; Réblová 2002).
Notes: Cain (1957b) established Synaptospora and introduced two distinctive taxa, S. petrakii (type) and S. tartaricola. The genus has mature ascospores which become fused in groups and Cain (1957b) inferred that this genus is closely related to Bombardia, Podospora and Sordaria, due to its carbonaceous ascomata. Subsequently, S. olandica, S. plumbea and S. setosa were introduced, all of which have fused ascospores, except for S. plumbea with unfused, ellipsoidal ascospores (Huhndorf et al. 1999b; Réblová 2002). Miller et al. (2014) sequenced S. plumbea (ANM963 and SMH3962) and found that it clustered with Helminthosphaeria species. Thus, Synaptospora was transferred to Helminthosphaeriaceae based on phylogenetic analyses (Miller et al. 2014; Hyde et al. 2020; Wijayawardene et al. 2020). We found that S. plumbea is similar to Helminthosphaeria in having globose ascomata, cylindrical asci and brown aseptate, ellipsoidal, unfused ascospores, differing from other Synaptospora species with fused ascospores. Phylogenetically, S. plumbea nested in Helminthosphaeriaceae (61%ML, Fig. 21). Therefore, we recommend transferring S. plumbea to Helminthosphaeria. The remaining four Synaptospora species have globose ascomata, cylindrical asci and globose, fused ascospores, but no molecular information is available. We place Synaptospora in Sordariales genera incertae sedis pending future work.
Type species: Synaptospora petrakii Cain, Beih. Sydowia 1: 5 (1957) [1956]
Facesoffungi number: ??; Fig. 62
Saprobic on decorticated wood. Sexual morph: Ascomata 200–350 × 250–450 µm (x̄ = 300 × 350 µm, n = 5), perithecial, superficial, gregarious to scattered, globose to subglobose, carbonaceous, with ostiole, black, verrucose, surrounded by sparse, brown, septate setae, 3.5–7 µm wide, with a rounded apex. Subiculum ascomata seated on sparse, brown, septate, branched, friable hyphae, 5.5–7.5 µm wide. Peridium 30–75 µm (x̄ = 55 µm, n = 20) wide, comprising two layers, outer layer carbonaceous, composed of brown cells of textura angularis; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses 2.5–8.5 µm wide, septate, slender to cylindrical, evanescent. Asci (70–)80–95(–100) × 7.5–10 µm (x̄ = 85 × 8 µm, n = 30), normally 2-spored, unitunicate, cylindrical, broadly rounded and thickened at the apex, long pedicellate. Ascospores (20–)22–24(–26) × 4–8 µm (x̄ = 23 × 6 µm, n = 50) for four cells, uni-seriate, subglobose to ellipsoidal, hyaline and aseptate when young, gradually becoming cylindrical to oblong, dark brown, usually 2–4-septate, distinctly constricted at the septa, the cells swollen, ends broadly rounded, straight to slightly curved, smooth-walled, with a large guttule in each cell, breaking into part spores. Part spores 6–9 × 4.5–7 µm (x̄ = 7.5 × 5.5 µm, n = 50), hyaline to dark brown, globose to ellipsoidal, aseptate, smooth-walled, with a large guttule, discharging through constricting septa. Asexual morph: Undetermined (adapted from Cain 1957b).
Material examined: Canada, Ontario, north of Mississauga, in willow grove by Credit River on Creditview Road, on decaying logs, 11 November 1987, L.A. Novak (TRTC-51203); Canada, Ontario, Muskoka district, Haliburton, 11 km south of Dorset, on decorticated wood, October 1975, D. Tighe (TRTC-51205).
Known hosts and distribution: On dead decorticated log of Betula papyrifera in Canada (type locality) (Cain 1957b).
Notes: The holotype material (TRTC-32168) was collected by Cain (1957b) in Canada. We were unable to obtain the type material and, therefore, we re-examined two authentic samples, TRTC-51203 and TRTC-51205, collected in Canada and determined by M. Matzer (mentioned in the label of material). Ascospores of Synaptospora was described as ellipsoidal to globose and fused in groups when mature (Cain 1957b; Barr 1990; Huhndorf et al. 1999b). We could not find mature ascospores fused in groups in TRTC-51203 and TRTC-51205. However, we found that the ascospores were initially hyaline, ellipsoidal to oblong or cylindrical and gradually formed distinct septa, and finally the septa split the brown ascospores into individual globose to subglobose part spores (Fig. 62 m–r). Fresh collections of Synaptospora species are required to determine the nature of ascospores and placement of the genus.
Tripterosporella Subram. & Lodha, Curr. Sci. 37: 246 (1968)
Coprophilous Sexual morph: Ascomata cleistothecial, scattered, superficial, globose to subglobose, semi-translucent to opaque, dark brown to black, surrounded by brown, septate hairs. Peridium membranaceous, composed of brown cells of textura angularis to textura prismatica. Asci 8-spored, unitunicate, clavate to cylindrical, pedicellate, evanescent, apex rounded or truncate, with J- apical ring. Paraphyses filiform or absent. Ascospores bi-seriate or irregularly arranged, cylindrical, aseptate and hyaline when young, upper cell(s) becoming swollen and fusiform, 0–1-septate, brown to olivaceous brown, guttulate, with truncate base and an apical germ pore, collapsing when dry; lower cell(s) tail-like, hyaline, 0–multi-septate, smooth-walled, slightly curved near the base. Asexual morph: Undetermined (adapted from Subramanian and Lodha 1968; Abdullah and Rattan 1978).
Notes: Tripterosporella was isolated from herbivores dung (horses and cows), and is similar to lasiosphaeriaceous taxa in having ascospores with brown upper cell(s) and a long hyaline pedicel (Subramanian and Lodha 1968; Doveri 2010; Wijayawardene et al. 2020). We place Tripterosporella in Sordariales genera incertae sedis but phylogenetic data is required to confirm its placement.
Type species: Tripterosporella coprophila Subram. & Lodha, Curr. Sci. 37: 246 (1968)
Fig. 34 r
Coprophilous Sexual morph: Ascomata cleistothecial, scattered, superficial, globose, black, membranaceous, surrounded by brown, septate hairs. Peridium membranaceous, composed of brown to hyaline cells of textura angularis to textura prismatica. Asci 8-spored, unitunicate, clavate to cylindrical, pedicellate, apex rounded, with J- apical ring. Ascospores bi-seriate, cylindrical, aseptate and hyaline when young, upper cell(s) becoming swollen and ellipsoidal to subglobose, 0–1-septate, brown, with truncate base and an apical germ pore, collapsing when dry; lower cell(s) tail-like, hyaline, 0–mulit-septate, smooth-walled, slightly curved near the base. Asexual morph: Undetermined (adapted from Abdullah and Rattan 1978).
Known hosts and distribution: On dung of Bos taurus in India (type locality) and Iraq (Subramanian and Lodha 1968; Abdullah and Rattan 1978).
Notes: Tripterosporella coprophila has brown ascospores with ellipsoidal upper cell(s) and a long, cylindrical, hyaline, multi-septate pedicel (Subramanian and Lodha 1968; Abdullah and Rattan 1978). However, the septate pedicel was rarely mentioned in subsequent researches (Abdullah and Rattan 1978; Doveri 2010). Tripterosporella coprophila is similar to Podospora, Lasiosphaeria and Zopfiella in having cleistothecial ascomata and multi-septate ascospores comprising two types of cells. We were unable to obtain the authentic material of this species and therefore, we refer to Subramanian and Lodha (1968) for hand-drawing (Fig. 34 r).
Other genera studied
Subclass Xylariomycetidae O.E. Erikss. & Winka
Xylariales Nannf.
Diatrypaceae Nitschke
Monosporascus Pollack & Uecker, Mycologia 66(2): 348 (1974)
Saprobic on wood. Sexual morph: Colonies white and hyaline. Ascomata perithecial, solitary to gregarious, semi-immersed, membranaceous, ellipsoidal to globose, dark brown to black, ostiolate, with papilla. Peridium membranaceous, comprising brown to olivaceous-brown cells of textura angularis to textura epidermoidea. Paraphyses numerous, filiform, cylindrical, flexuous, septate. Asci 1, 2, 3 or 4-spored, unitunicate, clavate to cylindrical, pedicellate. Ascospores globose, aseptate, hyaline to bright brown to brown to black, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Pollack and Uecker 1974; Negreiros et al. 2019).
Notes: Pollack and Uecker (1974) introduced Monosporascus typified by M. cannonballus. It has a Xylaria-type centrum and 1- or 2-spored asci with aseptate, globose ascospores with guttules. This genus was shown to belong to Diatrypaceae (Xylariales) based on phylogenetic analyses (Schoch et al. 2014; Maharachchikumbura et al. 2016; Negreiros et al. 2019; Hyde et al. 2020; this study, Fig. 1).
Type species: Monosporascus cannonballus Pollack & Uecker, Mycologia 66(2): 348 (1974)
Facesoffungi number: *****??; Fig. 63
Culture on malt yeast agar from Cucumis melo. Sexual morph: Colonies white and hyaline. Ascomata 310–500 × 270–420 µm (x̄ = 380 × 330 µm, n = 10), perithecial, solitary to gregarious, semi-immersed, membranaceous, ellipsoidal to globose, dark brown to black, ostiolate, with papilla. Peridium 35–80 µm (x̄ = 55 µm, n = 20) wide, comprising 5-layers, outer layer composed of hyaline, plectenchymatous tissues; next layer composed of olive-brown cells of textura epidermoidea; third layer composed of hyaline cells of textura angularis cells; fourth layer composed of olive-brown to brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 3–6 µm wide, numerous, filiform, cylindrical, flexuous, septate. Asci (40–)45–53(–60) × (18–)20–30(–35) µm (x̄ = 50 × 24 µm, n = 20), 1-spored, unitunicate, clavate to pyriform, thick-walled, pedicellate, persistent. Ascospores 1-celled, globose, hyaline when young, becoming bright brown to brown (15–25 µm diam.) to black (35–45 µm diam.) at maturity, smooth-walled, guttulate. Asexual morph: Undetermined.
Material examined: USA, Arizona, Yuma, from Cucumis melo L., on culture of malt yeast agar-1/4 dilution, 21 June 1972, F.G. Pollack and F.A. Uecker (BPI-579840, holotype).
Known hosts and distribution: On Cucumis melo in Brazil, Spain and USA (type locality) (Pollack and Uecker 1974; Schoch et al. 2014).
Notes: We re-examined the type of Monosporascus cannonballus (BPI-579840). The strain of M. cannonballus is sister to M. mossoroensis and M. nordestinus (Diatrypaceae) (Fig. 1).
Sordariomycetes genera incertae sedis
Conidiotheca Réblová & L. Mostert, Mycol. Res. 111(3): 305 (2007)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary to scattered, immersed to semi-immersed, dark brown to black, subglobose to broadly conical, ostiolate, with papilla, periphysate. Peridium membranaceous, comprising two layers, outer layer composed of black to pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses numerous, septate, tapering, longer than the asci. Asci 8-spored, cylindrical to clavate, short pedicellate, apex obtuse, with J- apical ring. Ascospores 1–2-seriate, hyaline, ellipsoidal to subglobose, 0–1-septate, sometimes slightly curved when young, becoming fusiform, with transverse and longitudinal septa, constricted at septa, producing several lateral appendices as ascoconidia. Ascoconidia hyaline, allantoid to ellipsoidal, aseptate, smooth-walled, guttulate, filling the whole ascus and discharged through constricted apical ring. Asexual morph: Undetermined (adapted from Réblová and Mostert 2007).
Notes: Barr (1985) described Romellia tympanoides (≡ Conidiotheca tympanoides) with clavate asci and ascospores producing ascoconidia similar to Tympanis (Leotiomycetidae). Barr et al. (1993) transferred it to Jattaea based on the spicate clusters of clavate asci. Réblová and Mostert (2007) transferred J. tympanoides to Conidiotheca which was placed in Ascomycota genera incertae sedis. Maharachchikumbura et al. (2015, 2016) and Wijayawardene et al. (2018) placed this genus in Calosphaeriales and Togniniales, respectively. We re-examined the type and found that Conidiotheca is similar to Jattaea (Calosphaeriaceae, Diaporthomycetidae) and Pleonectria (Nectriaceae, Hypocreomycetidae), which have clavate asci and septate ascospores with numerous ascoconidia. However, Conidiotheca has brown to black ascomata whereas those of Pleonectria are yellow to red (Réblová and Mostert 2007; Hirooka et al. 2012). Conidiotheca tympanoides is also similar to Jattaea taediosa, as they both have black ascomata, clavate asci and septate ascospores with numerous ascoconidia when mature, however, the former does not have the ascogenous hyphae unique to the latter (Réblová et al. 2015). Unfortunately, there is no molecular data available for C. tympanoides. We place this genus in Sordariomycetes genera incertae sedis pending more data.
Type species: Conidiotheca tympanoides (M.E. Barr) Réblová & L. Mostert, Mycol. Res. 111(3): 305 (2007)
Basionym: Romellia tympanoides M.E. Barr, Mycologia 77(4): 561 (1985)
Facesoffungi number: ??; Fig. 64
Saprobic on wood. Sexual morph: Ascomata 250–455 × 350–550 µm (x̄ = 345× 425 µm, n = 10), perithecial, solitary to scattered, papillate, immersed, erumpent through the bark of the host, dark brown to black, subglobose to broadly conical, ostiolate, with papilla, periphysate. Peridium 25–50 µm (x̄ = 35 µm, n = 30) wide, comprising two layers, outer layer composed of black to pale brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses 2–3.5 µm wide (x̄ = 2.8 µm, n = 30), numerous, septate, tapering, longer than the asci. Asci (120–)150–180(–200) × 15–18(–25) µm (x̄ = 170 × 16 µm, n = 50), 8-spored, cylindrical to clavate, short pedicellate, apex obtuse, with J- apical ring. Ascospores (10–)13–20(–25) × 3.5–6.5 µm (x̄ = 18 × 5 µm, n = 50), 1–2-seriate, hyaline, ellipsoidal to subglobose, 0–1-septate, sometimes slightly curved when young, becoming fusiform, with transverse and longitudinal septa, constricted at septa, producing several lateral appendices as ascoconidia. Ascoconidia (2.5–)3–5(–6) × 1–2.5 µm (x̄ = 4 × 1.5 µm, n = 50), hyaline, allantoid to ellipsoidal, aseptate, smooth-walled, guttulate, filling the whole ascus and discharged through constricted apical ring. Asexual morph: Undetermined.
Material examined: USA, Massachusetts, Baptist Hill, Conway, 21 April 1969, M.E. Barr, on twigs of Prunus pennsylvanica (NY-00912076, holotype).
Known hosts and distribution: On decayed wood of Prunus pennsylvanica in the USA (type locality) (Réblová and Mostert 2007).
Notes: The monotypic Conidiotheca is typified by C. tympanoides (Réblová and Mostert 2007). We found the holotype NY00912076 is well preserved, but there are few complete asci, and production of ascoconidia is unclear. Therefore, the hand-drawing is provided for the asci (Fig. 63 l–n) and the process of producing ascoconidia (Fig. 63 s–v) based on Réblová and Mostert (2007).
Ascomycota genera incertae sedis
Copromyces N. Lundq., Ark. Bot. 6: 327 (1967)
Coprophilous. Sexual morph: Ascomata cleistothecial, solitary, superficial, globose to subglobose, membranaceous, dark brown, with hyaline surface hairs. Peridium membranaceous, composed of dark brown to hyaline cells of textura angularis. Asci 2-spored, ovoid, apex rounded, short pedicel. Ascospores globose to subglobose, bright yellow, becoming olivaceous, brown to black at maturity, aseptate, guttulate, with a germ pore and irregular plate-like wrinkles on the surface. Asexual morph: Undetermined (adapted from Lundqvist 1967; Avila et al. 2002).
Notes: The monotypic genus Copromyces is typified by C. bisporus and has cylindrical to clavate, 2-spored asci and globose ascospores (Lundqvist 1967; Avila et al. 2002). The genus was initially placed in Sordariaceae (Lundqvist 1967). Petrak (1967) transferred C. bisporus to Rechingeriella (Zopfiaceae, Dothideomycetes), because it has 2-spored asci and tubercular ascospores similar to those of R. insignis. Subsequently, Lundqvist (1972) suggested that Rechingeriella and Copromyces are unrelated, because R. insignis has black, carbonaceous ascomata, distinctly bitunicate asci and ascospores with no germ pore, whereas, C. bisporus has membranaceous ascomata and ascospores with germ pore. Later, Avila et al. (2002) recorded that C. bisporus has cleistothecial ascomata and unitunicate asci with globose ascospores. Unfortunately, molecular data for C. bisporus is unavailable.
Copromyces sp. (CBS 386.78) was reported in the taxa sequence list of Huhndorf et al. (2004b) and noted as strain TRTC 51747 (& CBS 386.78, named as C. octosporus Jeng & Krug). This species, however, has not been formally described and is invalid. ‘Copromyces octosporus’ (CBS 386.78) was listed as the second species in Copromyces, and placed in Sordariaceae based on LSU and ITS sequence data (Huhndorf et al. 2004b; Maharachchikumbura et al. 2015, 2016; Vu et al. 2019; Hyde et al. 2020). However, in this study, the strain CBS 386.78 is sister to Pseudoneurospora canariensis (91%ML, Fig. 37). As the species has not been formally described and there is no apparent holotype it needs recollecting and introducing.
We re-examined the isotype (749020) of Copromyces bisporus, which is well preserved, and found that it has inconspicuously evanescent asci and dissolved hyaline filaments between the asci. We place Copromyces in Ascomycota genera incertae sedis pending molecular data.
Type species: Copromyces bisporus N. Lundq., Arch. für Botanik, Ser. 2 6: 328 (1967)
Facesoffungi number: *****??; Fig. 65
Coprophilous on rabbit dung. Sexual morph: Ascomata 200–250 µm (x̄ = 235 µm, n = 10) diam., cleistothecial, solitary, uniloculate, superficial, globose to subglobose, membranaceous, dark brown, with hyaline surface hairs. Peridium 30–50 µm wide, membranaceous, composed of dark brown to hyaline cells of textura angularis. Asci (24–)26–32(–35) × 12–15(–17) µm (x̄ = 30 × 13 µm, n = 50), 2-spored, ovoid, apex rounded, short pedicellate. Ascospores (11–)12–16(–20) µm (x̄ = 15 µm, n = 50) diam., globose to subglobose, bright yellow, becoming olivaceous, brown to black at maturity, aseptate, guttulate, with a germ pore and irregular plate-like wrinkles on the surface. Asexual morph: Undetermined.
Material examined: Sweden, Skane, Ivetofta parish, Aby, in pasture, on rabbit dung, 8 June 1962, N. Lundqvist (BPI-749020, isotype).
Known hosts and distribution: On dung of rabbit (Oryctolagus cuniculus) in Sweden (type locality) (Lundqvist 1967); on rabbit and fox dung in Venezuela (Avila et al. 2002).
Effetia Bartoli, Maggi & Persiani, Mycotaxon 19: 517 (1984)
Saprobic in forest soil. Sexual morph: Ascomata perithecial, solitary, brown to dark brown, globose to subglobose, ostiolate, with short necks. Peridium membranaceous. Asci 8-spored, unitunicate, cylindrical, apex rounded. Ascospores hyaline to brown, ellipsoidal, aseptate, with an apical germ pore. Asexual morph: Hyphomycetous. Mycelium dark brown, superficial, forming a thick subiculum. Hyphae flexuous, brown, septate, branched, becoming brittle. Conidiophores brown, unbranched, straight or flexuous, smooth-walled, septate, narrowed towards apex. Conidiogenous cells polyblastic, terminal ones give rise to new ones becoming intercalary, sympodial, swollen at the apex. Conidia acropleurogenous, oval to ellipsoidal, aseptate, light brown to dark brown, smooth-walled, with sheath. Chlamydospores single or like a string of beads, irregular, brown, thick-walled (adapted from Bartoli et al. 1984).
Notes: Bartoli et al. (1984) introduced the monotypic genus Effetia and proposed that its sexual morph is similar to Apodus and Fimetariella based on its black ascomata, cylindrical asci and ellipsoidal, hyaline to light brown ascospores, while its asexual morph (in culture) was described as virgariella-like, which is more similar to Xylariaceae. However, Bartoli et al. (1984) proposed to place Effetia in Sordariaceae based on its sexual morph, but no molecular data is available. We recommend transferring Effetia to Ascomycota genera incertae sedis pending future work.
Type species: Effetia craspedoconidica Bartoli, Maggi & Persiani, Mycotaxon 19: 517 (1984)
Facesoffungi number: ??; Fig. 66 a–q
Saprobic in forest soil. Sexual morph: Ascomata perithecial, solitary, brown to dark brown, globose to subglobose, ostiolate, with short necks. Peridium membranaceous. Asci 8-spored, unitunicate, cylindrical, apex rounded. Ascospores hyaline to brown, ellipsoidal, aseptate, with a germ pore. Asexual morph: Hyphomycetous. Mycelium dark brown, superficial, forming a thick subiculum. Hyphae 1.5–3.5 µm (x̄ = 2.5 µm, n = 30) wide, flexuous, brown, septate, branched, becoming brittle. Conidiophores brown, unbranched, straight or flexuous, smooth-walled, septate, narrowed towards apex. Conidiogenous cells polyblastic, terminal ones give rise to new ones becoming intercalary, sympodial, swollen at the apex. Conidia 7.5–11 × 3–5 µm (x̄ = 9.5 × 4 µm, n = 50), acropleurogenous, oval to ellipsoidal, aseptate, light brown to dark brown, smooth-walled, with sheath 1–1.5 µm wide. Chlamydospores (5–)7–10(–12) µm (x̄ = 9 µm, n = 20) diam., single or like a string of beads, irregular, brown, thick-walled (adapted from Bartoli et al. 1984).
Material examined: Ivory Coast, Tai forest National Park, on rain forest litter, December 1976, B. Rambelli (IMI-223521, holotype).
Known hosts and distribution: On woodland soil in Ivory Coast (type locality) (Bartoli et al. 1984).
Notes: Bartoli et al. (1984) noted that the sexual morph of Effetia craspedoconidica was produced only in the initial isolates, and then it lost viability. We re-examined the type (223521) and only found the asexual morph in culture. We draw the sexual morph with reference to Bartoli et al. (1984).
Endophragmiella B. Sutton, Mycol. Pap. 132: 58 (1973)
Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Mycelium immersed to superficial, composed of pale brown to brown, septate, branched, smooth-walled hyphae. Conidiophores macronematous, mononematous, erect, simple or branched, straight or flexuous, pale brown, smooth-walled. Conidiogenous cells monoblastic, integrated, terminal, tapering towards the truncate apex. Conidia solitary, acrogenous, simple, ellipsoidal, pale brown to brown, smooth-walled, 1–multi-septate (adapted from Sutton 1973).
Notes: The hyphomycete genus Endophragmiella is typified by End. pallescens which was found associated with stromata of Cytospora chrysosperma (Valsaceae) on wood of Populus tremuloides (Sutton 1973). Hughes (1979) suggested that Endophragmiella maybe the sexual morph of Echinosphaeria and the synanamorph of Selenosporella. Hughes (1979) also described two specimens of Echinosphaeria canescens, one of which was accompanied by End. biseptata; and the other was related to Selenosporella (Hughes 1979). These associations are not confirmed through culture or phylogenetic analysis. Molecular data is available for End. dimorphospora (CBS 438.74) and End. taxi (CBS 614.84), but not for End. pallescens (Hernández-Restrepo et al. 2017;Vu et al. 2019). End. dimorphospora nests in Helminthosphaeriaceae (Fig. 21), but End. taxi is closely related to members of Tubeufiales (Dothideomycetes) (not shown).
Type species: Endophragmiella pallescens B. Sutton, Mycol. Pap. 132: 62 (1973)
Fig. 22 s
Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies effuse, reddish brown, hairy, originating around effete stromata of immersed microfungi and spreading over the bark. Mycelium about 4 µm wide, immersed to superficial, composed of brown, septate, branched, smooth-walled hyphae. Conidiophores macronematous, mononematous, erect, simple or branched, flexuous, infrequently rigid, septate, light brown, becoming pale brown towards the apex, smooth-walled. Conidiogenous cells monoblastic, integrated, cylindrical, terminal, apex with 1–3 unflared annellations, pale brown. Conidia solitary, acrogenous, simple, ellipsoidal, pale brown, smooth-walled, 1–2-septate, not constricted at the septa, with a basal marginal frill (adapted from Sutton 1973).
Known hosts and distribution: On dead wood of Populus tremuloides in Canada (type locality) (Sutton 1973).
Notes: The mycelium of Endophragmiella pallescens associated with stromata of Cytospora chrysosperma on natural substratum in Manitoba, Canada (Sutton 1973). The hand-drawing is provided for the type species (Fig. 22 s) based on Sutton (1973).
Tulipispora Révay & Gönczöl, in Révay, Gönczöl & Descals, Nova Hedwigia 88(1-2): 42 (2009)
Saprobic on wood, rarely on leaves. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Mycelium semi-immersed in wood. Conidiophores hyaline, semi-macronematous, mononematous, simple, septate. Conidiogenous cells integrated, terminal, proliferations sympodial. Conidia solitary, acrogenous to subacrogenous, multi-septate, composed of elongate main axis and branches, branches mostly subverticillate, sometimes grouped near the base of the axis, curved (adapted from Révay et al. 2009).
Notes: The monotypic Tulipispora resembles Triscelophorus and Triramulispora in having multi-septate conidia with an elongate main axis and branches (Révay et al. 2009). It has been accommodated in Calosphaeriaceae, Sordariomycetes and Ascomycota genera incertae sedis (Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2018, 2020). However, as an asexual genus, Tulipispora is dissimilar to the known asexual morph of other members in Calosphaeriaceae. Tulipispora species have multi-septate and elongate conidia, whereas species of Calospaheriaceae usually have asepate, allantoid conidia. We place Tulipispora in Ascomycota genera incertae sedis.
Type species: Tulipispora ingoldii Révay & Gönczöl, Nova Hedwigia 88(1-2): 42 (2009)
Fig. 67
Saprobic on wood, rarely on leaves. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Mycelium semi-immersed in wood. Conidiophores hyaline, semi-macronematous, mononematous, simple, septate. Conidiogenous cells integrated, terminal, proliferations sympodial. Conidia solitary, acrogenous to subacrogenous, multi-septate, composed of elongate main axis and branches, branches mostly subverticillate, sometimes grouped near the base of the axis, curved (adapted from Révay et al. 2009).
Known hosts and distribution: On decayed wood and leaves in Hungary (type locality) (Révay et al. 2009).