Table of contents and contributors

Abrothallales

Lichenoconiaceae (S Pérez-Ortega, A Suija, S Hongsanan, R Jeewon)


Ascospermales

Ascospermaceae (D Pem, D Bundhun, S Hongsanan, KD Hyde)


Asterinales

Asterinaceae (XY Zeng, S Hongsanan, KD Hyde, R Jeewon)

Asterotexaceae (S Hongsanan, KD Hyde, EHC McKenzie)

Hemigraphaceae (DQ Dai, S Hongsanan, EHC McKenzie)

Lembosiaceae (D Sandamali, KD Hyde, S Hongsanan, EHC McKenzie)

Melaspileellaceae (S Hongsanan, Q Tian, KD Hyde, R Jeewon)

Morenoinaceae (S Hongsanan, KD Hyde, EHC McKenzie)

Neobuelliellaceae (S Hongsanan, KD Hyde, EHC McKenzie)

Stictographaceae (DQ Dai, S Hongsanan, R Jeewon)


Botryosphaeriales

Aplosporellaceae (AH Ekanayaka, AJL Phillips, KD Hyde)

Botryosphaeriaceae (IS Manawasinghe, AJ Dissanaya, A Phillips)

Melanopsaceae (S Hongsanan, AJL Phillips, KD Hyde)

Phyllostictaceae (AJ Dissanaya, P Abeywickrama, AJL Phillips, S Hongsanan, KD Hyde)

Planistromellaceae (J Monkai, AJL Phillips, KD Hyde)

Saccharataceae (AJ Dissanaya, AJL Phillips, S Hongsanan KD Hyde)


Catinellales

Catinellaceae (AH Ekanayaka, HA Ariyawansa, JS Zheng, KD Hyde)


Cladoriellales

Cladoriellaceae (S Hongsanan, DJ Bhat, EHC McKenzie, KD Hyde)


Collemopsidiales

Xanthopyreniaceae (S Pérez-Ortega, A Suija, S Hongsanan, EHC McKenzie)


Dyfrolomycetales

Pleurotremataceae (JF Zhang, VV Sarma, S Hongsanan, EBG Jones, KD Hyde)


Eremithallales

Melaspileaceae (V Thiyagaraja, D Ertz, S Hongsanan, KD Hyde)


Eremomycetales

Eremomycetaceae (D Pem, S Boonmee, KD Hyde, S Hongsanan, EHC McKenzie)


Jahnulales

Aliquandostipitaceae (ZL Luo, DF Bao, EBG Jones, KD Hyde)

Manglicolaceae (S Suetrong, S Preedanon, EBG Jones, KD Hyde)


Kirschsteiniotheliales

Kirschsteiniotheliaceae (ZL Luo, DF Bao, R Jeewon, KD Hyde)


Lembosinales

Lembosiaceae (S Hongsanan, KD Hyde, EHC McKenzie)


Lichenotheliales

Lichenotheliaceae (V Thiyagaraja, S Hongsanan, R Jeewon, KD Hyde)


Microthyriales

Microthyriaceae (XY Zeng, S Hongsanan, DJ Bhat, KD Hyde, EHC McKenzie)


Minutisphaerales

Agrogenosporaceae (J Yang, DJ Bhat, EHC McKenzie, KD Hyde)

Minutisphaeraceae (KM Thambugala, HA Raja, DJ Bhat, KD Hyde)


Monoblastiales

Monoblastiaceae (R Lücking, A Aptroot, H Kashiwadani, KH Moon, C Gueidan, F Schumm, R Phookamsak)


Murramarangomycetales

Murramarangomycetaceae (S Hongsanan, DJ Bhat, KD Hyde, EHC McKenzie)


Muyocopronales

Muyocopronaceae (N Chaiwan, S Hongsanan, C Senwanna, KD Hyde)


Natipusillales

Natipusillaceae (S Hongsanan, ZL Luo, KD Hyde, EHC McKenzie)


Parmulariales

Parmulariaceae (DQ Dai, S Hongsanan, R Jeewon, KD Hyde)


Patellariales

Patellariaceae (AH Ekanayaka, S Hongsanan, R Jeewon, KD Hyde)


Phaeotrichales

Phaeotrichaceae (D Sandaruwan, JS Zheng, KD Hyde, S Hongsanan, EHC McKenzie)


Stigmatodiscales

Stigmatodiscaceae (S Hongsanan, A Rathnayaka, EHC McKenzie, KD Hyde)


Strigulales

Strigulaceae (SH Jiang, R Lücking, E Sérusiaux)

Tenuitholiascaceae (SH Jiang, R Lücking)


Superstratomycetales

Superstratomycetaceae (P Abeywickrama, S Hongsanan)


Trypetheliales

Polycoccaceae (S Hongsanan, D Ertz, EHC McKenzie)

Trypetheliaceae (R Lücking, SN Zhang, R Miranda-González, A Aptroot)


Tubeufiales

Bezerromycetaceae (YZ Lu, J Bezerra, EHC McKenzie, KD Hyde)

Tubeufiaceae (YZ Lu, J Bezerra, EHC McKenzie, KD Hyde)

Wiesneriomycetaceae (C Senwanna, J Bezerra, EHC McKenzie)


Valsariales

Valsariaceae (C Senwanna, D Pem, S Hongsanan, EHC McKenzie, KD Hyde)


Venturiales

Sympoventuriaceae (D Sandamali, KD Hyde, R Jeewon)

Venturiaceae (D Sandamali, KD Hyde, S Hongsanan, R Jeewon)


Zeloasperisporiales

Zeloasperisporiaceae (S Hongsanan, DJ Bhat R Jeewon, KD Hyde)


Dothideomycetes, families incertae sedis

Alinaceae (S Hongsanan, S Boonmee, EHC McKenzie)

Argynnaceae (S Boonmee, S Hongsanan, S Konta)

Ascoporiaceae (S Boonmee, EHC McKenzie)

Aulographaceae (D Sandamali, S Hongsanan)

Balladynaceae (S Hongsanan, D Pem, S Boonmee)

Cleistosphaeraceae (S Hongsanan, D Pem, S Boonmee)

Coccoideaceae (S Boonmee, EHC McKenzie, KD Hyde)

Cookellaceae (S Boonmee, EHC McKenzie, KD Hyde)

Dimeriaceae (S Boonmee, KD Hyde, EHC McKenzie, S Konta)

Dubujianaceae (D Pem, S Hongsanan, KD Hyde)

Dysrhynchisceae (S Boonmee, KD Hyde, S Konta)

Endosporiaceae (D Pem, KD Hyde)

Englerulaceae (HX Wu, S Hongsanan, EHC McKenzie, KD Hyde)

Homortomycetaceae (KM Thambugala, EHC McKenzie, KD Hyde)

Hyalomeliolinaceae (S Boonmee, S Konta)

Leptopeltidaceae (R Phookamsak, EHC McKenzie, KD Hyde)

Macrovalsariaceae (D Pem, KD Hyde)

Meliolinaceae (XY Zeng, S Hongsanan, EHC McKenzie

Mesnieraceae (P Chomnunti, S Hongsanan, S Boonmee)

Naetrocymbaceae (S Hongsanan, DQ Dai, KD Hyde)

Nematotheciaceae (S Hongsanan, S Boonmee, KD Hyde)

Neoparodiaceae (S Hongsanan, S Boonmee, KD Hyde)

Palawaniaceae (A Mapook, S Hongsanan, EHC McKenzie)

Paranectriellaceae (S Boonmee, EHC McKenzie, KD Hyde)

Parodiellaceae (S Hongsanan, P Chomnunti, KD Hyde)

Perisporiopsidaceae (S Boonmee, EHC McKenzie, KD Hyde)

Phaeodimeriellaceae (S Hongsanan, S Boonmee, KD Hyde)

Pododimeriaceae (S Hongsanan, S Boonmee KD Hyde)

Pododimeriellaceae (S Hongsanan, D Sandamali, S Boonmee, KD Hyde)

Polyclypeolinaceae (S Hongsanan, S Boonmee, KD Hyde)

Polystomellaceae (D Pem, S Boonmee, KD Hyde)

Protoscyphaceae (S Boonmee, EHC McKenzie, KD Hyde)

Pseudoperisporiaceae (S Boonmee, EHC McKenzie, KD Hyde)

Pseudorobillardaceae (S Tibpromma, WJ Li, S Hongsanan, EHC McKenzie)

Pyrenidiaceae (S Hongsanan, N Huanraluek, D Ertz, KD Hyde)

Seynesiopeltidaceae (D Pem, S Boonmee, EHC McKenzie, KD Hyde)

Stomatogeneceae (S Hongsanan, S Boonmee, EHC McKenzie, KD Hyde)

Toroaceae (S Hongsanan, S Boonmee, KD Hyde, EHC McKenzie)

Thyrinulaceae (XY Zeng, S Hongsanan, KD Hyde, EHC McKenzie)

Trichopeltinaceae (D Sandamali, S Hongsanan, KD Hyde)

Trichothyriaceae (XY Zeng, HX Wu, KD Hyde)

Vizellaceae (HX Wu, S Hongsanan, S Boonmee, KD Hyde)

Introduction

Dothideomycetes were previously known as Loculoascomycetes, and is the largest class of ascomycetes (Nannfeldt 1932; Luttrell 1955; Janex-Favre 1971; Barr 1979a, b; Eriksson 1981; Reynolds 1971; Tehler 1990, 1995; Letrouit-Galinou et al. 1994; Barr and Huhndorf 2001; Liu and Hall 2004; Hyde et al. 2013; Hongsanan et al. 2020). Members of this class mostly have bitunicate, fissitunicate asci (Nannfeldt 1932; Luttrell 1955; Eriksson 1981; Barr and Huhndorf 2001; Hyde et al. 2013; Hongsanan et al. 2020). Hyde et al. (2013) provided descriptions, notes and discussed taxonomic placement of families in Dothideomycetes. They also confirmed that Dothideomycetes comprises two subclasses viz. Dothideomycetidae and Pleosporomycetidae, while other uncertain orders and families were treated as incertae sedis in Dothideomycetes. Based on morphology and phylogeny, Dothideomycetidae comprised Capnodiales, Dothideales, and Myrangiales, and Pleosporomycetidae comprised Gloniales, Hysteriales, Mytilinidiales, and Pleosporales (Hyde et al. 2013, Hongsanan et al. 2020). Hongsanan et al. (2020) provided a monograph of families in Dothideomycetidae and Pleosporomycetidae based on ‘Families of Dothideomycetes’ (Hyde et al. 2013) which was the latest update and the most complete monograph of families in Dothideomycetes, included in Wijayawardene et al. (2017a) who provided ‘Notes for genera: Ascomycota’ and Wijayawardene et al. (2020) who provided ‘Outline of Fungi and fungus-like taxa’. Hongsanan et al. (2020) provided an updated description, notes, including figures to represent the morphology, list of accepted genera, and economic and ecological significance of families in Dothideomycetidae and Pleosporomycetidae. They accepted three orders with 25 families in Dothideomycetidae and four orders with 94 families in Pleosporomycetidae (Hongsanan et al. 2020). Liu et al. (2017) established guidelines of using divergence estimates in the classification of Dothideomycetes, and this was followed by Hongsanan et al. (2020) who provided an updated divergence estimation of families in Dothideoycetes using two representative strains from each family. In our paper, we used the divergence time estimation from Hongsanan et al. (2020) to discuss orderal and familial status of incertae sedis groups.

The aim of this paper is to bring together data of orders and families which were not assigned to Dothideomycetidae nor Pleosporomycetidae due to their uncertain phylogenetic placement or lack of sequence data. We included all published data that we could locate in 2019 and have also included some data that we were aware of up to the date of publication. This work refined account of the families of Dothideomycetes and will provide a baseline for research on Dothideomycetes over the next decade. As an online database, dothideomycetes.org (Pem et al. 2019b) and fungalgenera.org (Monkai et al. 2019), the contents will be be continually refined and updated and stay abreast of classification changes as new data and additional evidence are published.

Materials and methods

Layout of the paper


Each family which is currently not assigned to Dothideomycetidae and Pleosporomycetidae is introduced with descriptions and important notes. Accepted genera including the type species, and notes are provided with basionyms (other synonyms can be found in Index Fungorum 2020). Faces of Fungi and Index Fungorum numbers are provided as in Jayasiri et al. (2015) and Index Fungorum (2020). Estimated numbers of species for each genus is provided mainly based on data in Species Fungorum (2020), with species transferred to other genera or species not being included. The number of species with sequence data available in GenBank is provided. The morphology of each family is illustrated by representative photographic plates from type herbarium specimens, or new species/new collections with sequence data. We provide drawing for families which we were unable to obtain herbarium specimens or fresh collections. Notes on the ecological and economic significance are also provided.


Molecular phylogeny

Phylogenetic trees of each order are provided, except for orders where too little sequence data is available. Trees for each order include representatives from all genera that have sequence data (Supplemental material 1). The methods below were applied for most of the phylogenetic trees shown in this paper. Other additional methods are mentioned in the legend of each analysis.

Gene regions used in each analysis were selected based on the most recent study or the availability of sequence data. Datasets for each partition were aligned using MAFFT (Katoh et al. 2019), and adjusted manually using Bioedit (Hall 2004). All missing genes were coded as missing data. Phylogenetic trees based on individual gene partitions (data not shown) were congruent with the combined data sets. The phylogenetic analyses of the combined dataset were performed using maximum likelihood algorithms in RAxML and Bayesian analyses.

Maximum likelihood analysis (ML) was carried out by using raxmlGUIv.0.9b2 (Silvestro and Michalak 2012). The search strategy was set to bootstrapping and the analysis performed using the GTRGAMMAI model. The number of replicates was inferred using the stopping criterion (Pattengale et al. 2009). The bootstrap values expressed from 1,000 repetitions by RAxML analysis which are equal or greater than 70% are given on each node. The best fit model of evolution was selected by MrModeltest 2.2 (Nylander et al. 2008). Bayesian phylogenetic analyses, posterior probabilities (PP) were set by MCMC sampling in MrBayes v3.1.2 (Huelsenbeck and Ronquist 2001; Zhaxybayeva and Gogarten 2002), following the details in Cai et al. (2008). The first 20% were discarded, and the remaining trees were used for calculating posterior probabilities. Posterior probabilities values (PP) from Bayesian analysis which are equal or greater than 0.90 are given on each node. Phylogenetic tree was viewed in FigTree v.1.4.0 (Rambaut 2014).


Order of Dothideomycetes

For the subclasses Dothideomycetidae and Pleosporomycetidae see Hongsanan et al. (2020) and Dothideomycetes genera incertae sedis see Wijayawardene et al. (2020).


Abrothallales Pérez-Ort. & Suija

Lichenoconiaceae Diederich & Lawrey (= Abrothallaceae Pérez-Ort. & Suija) (57)

Abrothallus De Not.

Lichenoconium Petr. & Syd.


Acrospermales Minter et al.

Acrospermaceae Fuckel

Acrospermum Tode

Gonatophragmium Deighton

Oomyces Berk. & Broome

Pseudovirgaria H.D. Shin et al.


Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss. (= Asterotexales Firmino et al.)

Asterinaceae Hansf.

Asterina Lév.

Asterinella Theiss.

Asterolibertia G. Arnaud

Asterostomella Speg.

Batistinula Arx

Cirsosia G. Arnaud

Dothidasteromella Höhn.

Echidnodella Theiss. & Syd.

Halbania Racib.

Meliolaster Höhn.

Parasterinopsis Bat.

Platypeltella Petr.

Prillieuxina G. Arnaud

Pycnocarpon Theiss.

Schenckiella Henn.

Trichasterina G. Arnaud

Trichopeltospora Bat. & Cif.

Uleothyrium Petr.

Vizellopsis Bat. et al.


Asterotexaceae Firmino et al.

Asterotexis Arx


Hemigraphaceae D.Q. Dai & K.D. Hyde

Hemigrapha (Müll. Arg.) D. Hawksw.


Lembosiaceae Luttrell ex P.M. Kirk et al.

Lembosia Lév.


Melaspileellaceae D.Q. Dai & K.D. Hyde

Melaspileella (P. Karst.) Vain.


Morenoinaceae Hongsanan & K.D. Hyde

Morenoina Theiss.


Neobuelliellaceae Hongsanan & K.D. Hyde

Neobuelliella Hongsanan & K.D. Hyde


Stictographaceae D.Q. Dai & K.D. Hyde

Buelliella Fink

Karschia Körb.

Labrocarpon Etayo & Pérez-Ort.

Melaspileopsis (Müll. Arg.) Ertz & Diederich

Stictographa Mudd


Asterinales genera incertae sedis

Andamanomyces Hosag.

Caribaeomyces Cif.

Caudella Syd. & P. Syd.

Discopycnothyrium Hongsanan & K.D. Hyde

Hazslinszkya Körb.

Inocyclus Theiss. & Syd.

Melanographa Müll. Arg.

Pirozynskiella S. Hughes

Vishnumyces Hosag.


Botryosphaeriales C.L. Schoch et al.

Aplosporellaceae Slippers et al.

Alanomyces Sharma

Aplosporella Speg.


Botryosphaeriaceae Theiss. & H. Syd.

Alanphillipsia Crous & M.J. Wingf.

Barriopsis A.J.L. Phillips et al.

Botryobambusa Phook. et al.

Botryosphaeria Ces. & De Not.

Cophinforma Doilom et al.

Diplodia Fr.

Dothiorella Sacc.

Endomelanconiopsis Rojas & Samuels

Eutiarosporella Crous

Lasiodiplodia Ellis & Everh.

Macrophomina Petr.

Marasasiomyces Crous

Mucoharknessia Crous et al.

Neodeightonia C. Booth

Neofusicoccum Crous et al.

Neoscytalidium Crous & Slippers

Oblongocollomyces Tao Yang & Crous

Phaeobotryon Theiss. & Syd.

Sakireeta Subram. & K. Ramakr.

Sardiniella Linaldeddu et al.

Sphaeropsis Sacc.

Tiarosporella Höhn.


Melanopsaceae Phillips et al.

Melanops Nitschke ex Fuckel


Phyllostictaceae Fr.

Phyllosticta Pers.

Pseudofusicoccum Mohali et al.


Planistromellaceae M.E. Barr

Kellermania Ellis & Everh.

Mycosphaerellopsis Höhn.

Planistroma A.W. Ramaley

Umthunziomyces Crous & M.J. Wingf.


Saccharataceae Slippers et al.

Pileospora Tanney & Seifert

Saccharata Denman & Crous (= Neoseptorioides Crous et al.)

Septorioides Quaedvl. et al.


Botryosphaeriales genera incertae sedis

Auerswaldiella Theiss. & Syd.

Coccostromella Petr.

Leptoguignardia E. Müll.

Metameris Theiss. & Syd.

Phyllachorella Syd.

Pilgeriella Henn.

Sivanesania W.H. Hsieh & C.Y. Chen

Vestergrenia Rehm


Catinellales Ekanayaka et al.

Catinellaceae Ekanayaka et al.

Catinella Boud.


Cladoriellales Crous

Cladoriellaceae Crous

Cladoriella Crous


Collemopsidiales Perez-Ortega et al.

Xanthopyreniaceae Zahlbr.

Collemopsidium Nyl.

Didymellopsis (Sacc.) Clem. & Shear

Frigidopyrenia Grube

Rhagadodidymellopsis Fdez.-Brime et al.

Xanthopyrenia Bachm.

Zwackhiomyces Grube & Hafellner

Zwackhiomacromyces Etayo & van den Boom


Dyfrolomycetales K.L. Pang et al.

Pleurotremataceae Walt. Watson (= Dyfrolomycetaceae K.D. Hyde et al.)

Dyfrolomyces K.D. Hyde et al.

Melomastia Nitschke ex Sacc.

Pleurotrema Müll. Arg.


Eremithallales Lücking & Lumbsch

Melaspileaceae W. Watson (= Eremithallaceae Lücking & Lumbsch)

Encephalographa A. Massal.

Melaspilea Nyl.


Eremomycetales Crous et al.

Eremomycetaceae Malloch & Cain

Eremomyces Malloch & Cain

Rhexothecium Samson & Mouch.


Eremomycetales genus incertae sedis

Arthrographis G. Cochet ex Sigler & J.W. Carmich.


Jahnulales Pang et al.

Aliquandostipitaceae Inderbitzin

Aliquandostipite Inderbitzin

Brachiosphaera Nawawi

Jahnula Kirschst.

Megalohypha A. Ferrer & Shearer

Neojahnula W. Dong et al.

Pseudojahnula W. Dong et al.

Xylomyces Goos et al.


Manglicolaceae Suetrong & E.B.G. Jones

Manglicola Kohlm. & E. Kohlm.


Kirschsteiniotheliales Hern.-Restr. et al.

Kirschsteiniotheliaceae Boonmee & K.D. Hyde

Kirschsteiniothelia D. Hawksw.


Kirschsteiniotheliales genus incertae sedis

Brachysporiella Bat. (Brachysporiella sensu lato)

Taeniolella S. Hughes sensu lato


Lembosinales Crous

Lembosinaceae Crous

Lembosina Theiss.


Lichenotheliales K. Knudsen et al.

Lichenotheliaceae Henssen

Lichenothelia D. Hawksw.

Endococcus Nyl.


Microthyriales G. Arnaud

Microthyriaceae Sacc.

Arnaudiella Petr.

Calothyriopsis Höhn.

Chaetothyriothecium Hongsanan & K.D. Hyde

Hamatispora L.T.H. Yen et al.

Microthyrium Desm.

Neoanungitea Crous

Paramicrothyrium H.X. Wu & K.D. Hyde

Pseudomicrothyrium X.Y. Zeng et al.

Pseudopenidiella Crous & Koukol

Seynesiella G. Arnaud

Tumidispora Hongsanan & K.D. Hyde


Microthyriales genera incertae sedis

Heliocephala V. Rao et al.

Mitopeltis Speg.

Neoscolecobasidium Crous

Parazalerion Madrid et al.

Thyriodictyella Cif.

Tothia Bat.


Minutisphaerales Raja et al.

Acrogenosporaceae Jayasiri & K.D. Hyde.

Acrogenospora M.B. Ellis


Minutisphaeraceae Raja et al

Minutisphaera Shearer et al.


Monoblastiales Lücking et al.

Monoblastiaceae Walt. Watson (= Eriomycetaceae Huanraluek & K.D. Hyde)

Acrocordia A. Massal.

Anisomeridium (Müll. Arg.) M. Choisy

Caprettia Bat. & H. Maia

Eriomyces Huanraluek, Thambugala & K.D. Hyde

Funbolia Crous & Seifert

Heleiosa Kohlm. et al.

Megalotremis Aptroot

Monoblastia Riddle

Phellinocrescentia Crous & Decock

Pseudopassalora Crous

Trypetheliopsis Asahina


Murramarangomycetales Crous

Murramarangomycetaceae Crous

Phaeothyriolum Syd.


Muyocopronales Mapook et al.

Muyocopronaceae K.D. Hyde

Arxiella Papendorf

Leptodiscella Papendorf

Muyocopron Speg.

Mycoleptodiscus Ostaz.

Neocochlearomyces Pinruan et al.

Neomycoleptodiscus Hern.-Restr. et al.

Paramycoleptodiscus Crous & M.J. Wingf.

Pseudopalawania Mapook & K.D. Hyde

Setoapiospora Mapook & K.D. Hyde


Natipusillales Raja et al.

Natipusillaceae Raja et al.

Natipusilla A. Ferrer


Parmulariales D.Q. Dai & K.D. Hyde

Parmulariaceae E. Müll. & Arx ex M.E. Barr

Aldona Racib.

Aldonata Sivan. & A.R.P. Sinha

Antoniomyces Inácio

Aulacostroma Syd. & P. Syd.

Campoa Speg.

Cirsosiopsis Butin & Speer

Cocconia Sacc.

Cycloschizon P. Henn.

Cyclostomella Pat.

Dothidasteroma Höhn.

Ferrarisia Sacc.

Hysterostomella Speg.

Kiehlia Viégas

Mintera Inácio & P.F. Cannon

Pachypatella Theiss. & Syd.

Palawaniella Doidge

Parmularia Lév.

Parmulariopsella Sivan.

Parmulariopsis Petr.

Parmulina Theiss. & Syd.

Placoasterella Sacc. ex Theiss. & Syd

Placosoma Syd.

Placostromella Petr.

Pleiostomellina Bat. et al.

Polycyclina Theiss. & Syd.

Polycyclus Höhn.

Protothyrium G. Arnaud

Pseudolembosia Theiss.

Rhagadolobiopsis Guatim. & R.W. Barreto

Rhagadolobium P. Henn. & Lindau

Rhipidocarpon (Theiss.) Theiss. & Syd.

Symphaeophyma Speg.

Syrropeltis Bat. et al.

Thallomyces H.J. Swart

Viegasiella Inácio & P.F. Cannon


Patellariales D. Hawksw. & O.E. Erikss.

Patellariaceae Corda

Baggea Auersw.

Banhegyia L. Zeller & Tóth

Colensoniella Hafellner

Endotryblidium Petr.

Glyphium Nitschke ex F. Lehm.

Haematomyxa Sacc

Holmiella Petrini et al.

Hysteropatella Rehm

Hysteropeltella Petr.

Lahmiomyces Cif. & Tomas.

Lecanidiella Sherwood

Lirellodisca Aptroot

Murangium Seaver

Patellaria Fr.

Poetschia Körb.

Pseudoparodia Theiss. & Syd.

Rhizodiscina Hafellner

Rimula Velen.

Schrakia Hafellner

Stratisporella Hafellner

Tryblidaria (Sacc.) Sacc.


Phaeotrichales Ariyaw. et al.

Phaeotrichaceae Cain

Echinoascotheca Matsush.

Phaeotrichum Cain & M.E. Barr

Trichodelitschia Munk


Stigmatodiscales Voglmayr & Jaklitsch

Stigmatodiscaceae Voglmayr & Jaklitsch

Stigmatodiscus Voglmayr & Jaklitsch


Strigulales Lücking et al.

Strigulaceae Zahlbr. (= Phyllobatheliaceae Bitter & F. Schill.)

Dichoporis Clem.

Flagellostrigula Lücking et al.

Flavobathelium Lücking et al.

Phyllobathelium (Müll. Arg.) Müll. Arg

Phyllocharis Fée,

Phyllocraterina Sérus. & Aptroot

Phylloporis Clem.

Puiggariella Speg.

Raciborskiella Höhnel

Racoplaca Fée

Serusiauxiella S.H. Jiang et al.

Strigula Fr.

Swinscowia S.H. Jiang et al.


Tenuitholiascaceae S.H. Jiang et al.

Tenuitholiascus S.H. Jiang et al.


Superstratomycetales van Nieuwenhuijzen et al.

Superstratomycetaceae van Nieuwenhuijzen et al.

Superstratomyces van Nieuwenhuijzen et al.


Trypetheliales Lücking et al.

Polycoccaceae Ertz et al.

Clypeococcum D. Hawksw.

Polycoccum Saut. ex Körb.


Trypetheliaceae Zenker

Alloarthopyrenia Phukhams. et al.

Aptrootia Lücking & Sipman

Architrypethelium Aptroot

Astrothelium Eschw.

Bathelium Ach.

Bogoriella Zahlbr.

Constrictolumina Lücking et al.

Dictyomeridium Aptroot et al.

Macroconstrictolumina Lücking et al.

Marcelaria Aptroot

Nigrovothelium Lücking et al.

Polymeridium (Müll. Arg.) R.C. Harris

Polypyrenula D. Hawksw.

Pseudobogoriella Lücking et al.

Pseudopyrenula Müll. Arg.

Schummia Lücking et al.

Trypethelium Sprengel

Viridothelium Lücking et al.


Tubeufiales Boonmee & K.D. Hyde

Bezerromycetaceae J.D.P. Bezerra et al.

Bezerromyces J.D.P. Bezerra et al.

Neorhamphoria Boonmee et al.

Xiliomyces J.D.P. Bezerra et al.


Tubeufiaceae M.E. Barr

Acanthohelicospora Boonmee & K.D. Hyde

Acanthophiobolus Berl.

Acanthostigma De Not.

Acanthostigmina Höhn.

Acanthotubeufia Y.Z. Lu & K.D. Hyde

Aquaphila Goh et al.

Artocarpomyces Subram.

Berkleasmium Zobel

Bifrontia Norman

Boerlagiomyces Butzin

Camporesiomyces D.P. Wei & K.D. Hyde

Chaetosphaerulina I. Hino

Chlamydotubeufia Boonmee & K.D. Hyde

Dematiohelicoma Y.Z. Lu et al.

Dematiohelicomyces Y.Z. Lu et al.

Dematiohelicosporum Y.Z. Lu et al.

Dematiotubeufia Y.Z. Lu et al.

Dictyospora Brahamanage et al.

Discotubeufia Jayasir et al.

Helicangiospora Boonmee et al.

Helicoarctatus Y.Z. Lu et al.

Helicodochium J.S. Monteiro et al.

Helicohyalinum Y.Z. Lu et al.

Helicoma Corda

Helicomyces Link

Helicosporium Nees

Helicotruncatum Y.Z. Lu et al.

Helicotubeufia Y.Z. Lu & J.K. Liu

Kamalomyces R.K. Verma et al.

Kevinhydea N.G. Liu et al.

Manoharachariella Bagyan. et al.

Muripulchra Z.L. Luo et al.

Neoacanthostigma Boonmee et al.

Neochlamydotubeufia Y.Z. Lu et al.

Neohelicoma Y.Z. Lu et al.

Neohelicomyces Z.L. Luo et al.

Neohelicosporium Y.Z. Lu et al.

Neotubeufia Chaiwan et al.

Pleurohelicosporium Y.Z. Lu et al.

Podonectria Petch

Pseudohelicomyces Y.Z. Lu et al.

Pseudohelicoon Y.Z. Lu & K.D. Hyde

Tamhinispora Rajeshkumar & Rahul Sharma

Thaxteriella Petr.

Thaxteriellopsis Sivan. et al.

Tubeufia Penz. & Sacc.


Wiesneriomycetaceae Suetrong et al.

Parawiesneriomyces Crous & M.J. Wingf.

Phalangispora Nawawi & J. Webster

Pseudogliophragma Phadke & V.G. Rao

Setosynnema D.E. Shaw & B. Sutton

Speiropsis Tubaki

Wiesneriomyces Koord.


Valsariales Jaklitsch et al.

Valsariaceae Jaklitsch et al.

Bambusaria Jaklitsch et al.

Myrmaecium Nitschke ex Fuckel

Valsaria Ces. & De Not.


Venturiales Y. Zhang ter et al.

Sympoventuriaceae Y. Zhang ter et al.

Acroconidiellina M.B. Ellis

Clavatispora Boonmee & K.D. Hyde

Echinocatena R. Campb. & B. Sutton

Fusicladium Bonord.

Matsushimaea Subram.

Mycosisymbrium Carris

Ochroconis de Hoog & Arx

Scolecobasidiella M.B. Ellis

Scolecobasidium E.V. Abbott

Sympoventuria Crous & Seifert

Veronaeopsis Arzanlou & Crous

Verruconis Samerp. et al.

Yunnanomyces Tibpromma & K.D. Hyde


Venturiaceae E. Müll. & Arx ex M.E. Barr

Apiosporina Höhn.

Atopospora Petr.

Caproventuria U. Braun

Coleroa (Fr.) Rabenh.

Cylindrosympodium W.B. Kendr. & R.F. Castañeda

Dimeriella Speg.

Dimerosporiopsis Henn.

Magnohelicospora R.F. Castañeda et al.

Metacoleroa Petr.

Neocoleroa Petr.

Protoventuria Berl. & Sacc.

Pseudoanungitea Crous

Pseudoparodiella F. Stevens

Sympodiella W.B. Kendr.

Tyrannosorus Unter. & Malloch

Venturia Sacc.


Venturiales genus incertae sedis

Cylindrosympodioides Crous & M.J. Wingf.

Lasiobotrys Kunze


Zeloasperisporiales Hongsanan & K.D. Hyde

Zeloasperisporiaceae Crous

Zeloasperisporium R.F. Castañeda


Dothideomycetes families incertae sedis

Alinaceae Boonmee & K.D. Hyde

Alina Racib.


Argynnaceae Shearer & J.L. Crane

Argynna Morgan

Lepidopterella Shearer & J.L. Crane


Ascoporiaceae Kutorga & D. Hawksw.

Ascoporia Samuels & A.I. Romero


Aulographaceae Luttr. ex P.M. Kirk et al.

Aulographum Lib.

Echidnodes Theiss. & Syd.

Lembosiella Sacc.

Thyriopsis Theiss. & Syd.


Balladynaceae Boonmee & K.D. Hyde

Balladyna Racib.

Balladynocallia Bat.

Balladynopsis Theiss. & Syd.


Cleistosphaeraceae Boonmee & K.D. Hyde

Cleistosphaera Syd. & P. Syd.


Coccoideaceae P. Henn. ex Sacc. & D. Sacc.

Coccoidea P. Henn.

Coccoidella Höhn.

Englerodothis Theiss. & Syd.


Cookellaceae Höhn. ex Saccardo & Trotter

Cookella Sacc.

Pycnoderma Syd. & P. Syd.


Dimeriaceae E. Müll. & Arx ex Arx & E. Müll.

Dimerium (Sacc. & P. Syd.) McAlpine


Dubujianaceae D. Pem et al.

Dubujiana D.R. Reynolds & G.S. Gilbert


Dysrhynchisceae Boonmee & K.D. Hyde

Dysrhynchis Clem.


Endosporiaceae D. Pem

Endosporium Tsuneda


Englerulaceae P. Henn.

Allosoma Syd.

Digitosarcinella S. Hughes

Englerula P. Henn.

Goosia B. Song

Parenglerula Höhn.

Rhytidenglerula Höhn.

Schiffnerula Höhn.

Thrauste Theiss.


Homortomycetaceae Thambug. et al.

Homortomyces Crous & M.J. Wingf.


Hyalomeliolinaceae Boonmee & K.D. Hyde

Hyalomeliolina F. Stevens


Leptopeltidaceae Höhn. ex Trotter

Dothiopeltis E. Müll.

Leptopeltis Höhn.

Ronnigeria Petr.

Staibia Bat. & Peres


Macrovalsariaceae D. Pem et al.

Macrovalsaria Petr.


Meliolinaceae S. Hughes

Briania D.R. Reynolds

Meliolina Syd. & P. Syd.


Mesnieraceae Arx & E. Müll.

Bondiella Piroz.

Mesniera Sacc. & P. Syd.

Stegasphaeria Syd. & P. Syd.


Naetrocymbaceae Höhn. ex R.C. Harris

Bonaria Bat.

Jarxia D. Hawksw.

Leptorhaphis Körb.

Naetrocymbe Körb.

Tomasellia A. Massal.


Nematotheciaceae Boonmee & K.D. Hyde

Nematothecium Syd. & P. Syd.

Nematostigma Syd. & P. Syd.

Ophioparodia Petr. & Cif.


Neoparodiaceae Boonmee & K.D. Hyde

Neoparodia Petr. & Cif.


Palawaniaceae Mapook & K.D. Hyde

Palawania (Niessl) Syd. & P. Syd.


Paranectriellaceae Boonmee & K.D. Hyde

Paranectriella (Henn. ex Sacc. & D. Sacc.) Magnus. (= Araneomyces Höhn.)

Puttemansia Henn.


Parodiellaceae Theiss. & H. Syd. ex M.E. Barr

Parodiella Speg.


Perisporiopsidaceae E. Müll. & Arx ex R. Kirschner & T.A. Hofm. (= Parodiopsidaceae Toro)

Asteronia (Sacc.) Henn.

Byssocallis Syd.

Chevalieropsis G. Arnaud

Parodiellina Henn. ex G. Arnaud

Perisporiopsis Henn.


Phaeodimeriellaceae Boonmee et al.

Phaeodimeriella Speg.


Pododimeriaceae Boonmee & K.D. Hyde

Chaetoscutula E. Müll.

Pododimeria E. Müll.


Polyclypeolinaceae Boonmee & K.D. Hyde

Polyclypeolina Bat. & I.H. Lima


Polystomellaceae Theiss. & H. Syd.

Dermatodothella Viégas

Dothidella Speg.

Munkiella Speg.

Parastigmatea Doidge


Protoscyphaceae Kutorga & D. Hawksw.

Protoscypha Syd.


Pseudoperisporiaceae Toro

Bryomyces Döbbeler

Eudimeriolum Speg.

Lasiostemma Theiss.

Nematostoma Syd. & P. Syd.


Pseudorobillardaceae Crous

Pseudorobillarda M. Morelet


Pyrenidiaceae Zahlbr.

Pyrenidium Nyl.


Seynesiopeltidaceae K.D. Hyde

Seynesiopeltis F. Stevens & R.W. Ryan


Stomatogeneceae Boonmee & K.D. Hyde

Stomatogene Theiss.


Thyrinulaceae X.Y. Zeng et al.

Blastacervulus H.J. Swart

Paraopeba V.P. Abreu et al.

Thyrinula Petr. & Syd.


Toroaceae Boonmee & K.D. Hyde

Toroa Syd.


Trichopeltinaceae Bat. et al.

Acrogenotheca Cif. & Bat.

Brefeldiella Speg.

Saccardinula Speg.

Trichopeltella Höhn.

Trichopeltheca Bat.

Trichopeltina Theiss.

Trichothyrinula Petr.


Trichothyriaceae Theiss.

Lichenopeltella Höhn.

Macrographa Etayo

Pachythyrium G. Arnaud ex Spooner & P.M. Kirk

Trichothyrium Speg.


Vizellaceae H.J. Swart

Acarella Syd.

Blasdalea Sacc. & P. Syd.

Vizella Sacc.


Abrothallales Pérez-Ort. & Suija.

= Lichenoconiales Diederich, Lawrey & K.D. Hyde.

Index Fungorum number: IF 805297; Facesoffungi number: FoF 08045.

Abrothallales was established by Pérez-Ortega and Suija (in Pérez-Ortega et al. 2014) based on its monophyletic placement and morphological distinctness within the class Dothideomycetes. Diederich et al. (in Hyde et al. 2013) had introduced Lichenoconiales, which Liu et al. (2017) recovered as a sister clade of Abrothallales, raising the question of whether both orders should be maintained. Diederich et al. (2018) synonymized Lichenoconiales with Abrothallales and tentatively Lichenoconiaceae with Abrothallaceae. Pérez-Ortega and Suija agreed with Diederich et al. (2018) to use the name Abrothallales since Abrothallus is the most outstanding, easily recognizable and better-known genus of the group. Lichenoconiaceae is the older name, therefore, we retain Lichenoconiaceae over Abrothallaceae. Abrothallales contains species which are lichenicolous, parasymbiontic or parasitic on macrolichens from a single family Abrothallaceae (Pérez-Ortega et al. 2014; Wijayawardene et al. 2017a). The divergence time for Abrothallales is estimated as 204 MYA (stem age, Hongsanan et al. 2020) (Fig. 1).

Fig. 1
figure 1

Phylogram generated from maximum likelihood analysis (RAxML) of Abrothallales based on combined ITS, LSU and tef1 sequence data. Maximum likelihood bootstrap values equal or above 70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/PP) are given at the nodes. An original isolate number is noted after the species name. The tree is rooted to Patellaria quercus (CPC 27232 and BHI F828a). The ex-type strains are indicated in bold. Hyphen (-) represents support values below 70% MLBS and 0.90 PP

Accepted families: Lichenoconiaceae.


Lichenoconiaceae Diederich & Lawrey, in Hyde et al., Fungal Diversity 63: 131 (2013).

= Abrothallaceae Pérez-Ort. & Suija, in Pérez-Ortega, Suija, Crespo & Ríos, Fungal Diversity 64(1): 303 (2014).

Index Fungorum number: IF 803667; Facesoffungi number: FoF 08046, 58 species.

Parasitic or parasymbiotic on lichen thalli. Vegetative hyphae immersed in lichen thallus, usually hyaline, I+ violet in some species. Sexual morph: Ascomata apothecioid, solitary or in groups, superficial, rounded, convex to almost globose, rarely flattened, black or dark brown, some species with greenish pruina, at least in young stages. Proper exciple thin, usually disappearing when mature, composed of radiating hyphae. Hymenium hyaline or greenish in upper part, covered with light to dark brown or reddish granules dissolving in KOH. Hypothecium light to dark brown, composed of isodiametric cells (textura angularis type). Hamathecium comprising septate, anastomosed and irregularly dichotomously branching interascal filaments. Asci 4–8-spored, thick-walled, bitunicate, functionally fissitunicate, ovate to clavate, apedicellate, with a distinct ocular chamber, I-. Ascospores 2-seriate, irregularly or 2-seriately arranged within the ascus, ellipsoid to broadly ellipsoid, light to dark brown, 1- to 3-septate, asymmetric and soleiform, splitting into part-spores within the ascus in some species, wall finely ornamented to verrucose. Asexual morph: Coelomycetous. Vouauxiomyces-type, pycnidium immersed, semi-immersed or almost superficial, black, with a small ostiole. Pycnidial wall thick-walled, comprises isodiametric cells (textura angularis-type). Conidiophores reduced. Conidiogenous cells percurrently proliferating, ampulliform to lageniform, lining the cavity of the pycnidium, hyaline, smooth-walled. Conidia holoblastic, clavate to obpyriform or almost roundish, hyaline to dark brown, smooth to slightly echinulate, usually embedded in mucilage.

Type: Lichenoconium Petr. & Syd.

Notes: Lawrey et al. (2011) studied the phylogenetic affinities of Lichenoconium, a genus of lichenicolous fungi known only in its asexual morph. They found that Lichenoconium is also a member of Dothideomycetes, and subsequently Hyde et al. (2013) introduced a new order Lichenoconiales to accommodate it. Pérez-Ortega et al. (2014) introduced Abrothallaceae and the new order Abrothallales for the lichenicolous genus Abrothallus. The classification of this genus had long been debated. This is one of the few Dothideomycetes genera having apothecioid ascomata, but the bitunicate asci and the shape of interascal filaments clearly segregates it from other similar groups (Pérez-Ortega et al. 2014). Pérez-Ortega et al. (2011) established the connection between the sexual morph Abrothallus and the asexual morph Vouauxiomyces. Vouauxiomyces and Lichenoconium share many features, such as the mode of conidiogenesis and shape of conidia (detailed descriptions in Hawksworth 1981b), although the conidia are hyaline in Abrothallus (except in Abrothallus kamchatica) and dark brown in Lichenoconium. Recent phylogenetic analyses recovered Abrothallus and Lichenoconium as sister genera, with a relatively young split (9 MYA) raising the question of the need to keep two separate orders (Liu et al. 2017). Diederich et al. (2018) in their review of lichenicolous fungi treated Lichenoconiales as a synonym of Abrothallales, and Lichenoconiaceae as a possible synonym of Abrothallaceae.


Lichenoconium Petr. & Syd., Beih. Reprium nov. Spec. Regni veg. 42(1): 432 (1927) [1926].

Index Fungorum number: IF 8772; Facesoffungi number: FoF 08047; 16 morphological species (Species Fungorum 2020), 4 species with molecular data.

Type species: Lichenoconium lichenicola (P. Karst.) Petr. & Syd. [as ‘lichenicolum’], Beih. Reprium nov. Spec. Regni veg. 42(1): 432 (1927) [1926].

Dactylium dendroides subsp. lichenicola P. Karst., Meddn Soc. Fauna Flora fenn. 14: 107 (1887).

Notes: Lichenoconium species are distributed worldwide as lichenicolous taxa on a variety of lichen hosts (Lawrey et al. 2011; Hyde et al. 2013; Wijayawardene et al. 2017a; Diederich et al. 2018). Many are host-specific, while others occur on disparate genera (Lawrey et al. 2011). The asexual morph is characterized by coelomycetous, immersed to sessile, dark brown pycnidia, an undifferentiated pycnidial ostiole, simple conidia, globose to ellipsoid, basally often truncate, thick-walled, brown and commonly verrucose and conidiogenous cells that are hyaline or poorly pigmented, holoblastic, elongate and annellidic (Lawrey et al. 2011). The sexual morph has not been recorded.


Other genera included

Abrothallus De Not. Giorn. Bot. Ital. 1: 194 (1846).

= Vouauxiomyces Dyko & D. Hawksw., Lichenologist 11(1): 57 (1979).

Index Fungorum number: IF 4; Facesoffungi number: FoF 08048; – 42 morphological species (Species Fungorum 2020), 20 species with molecular data.

Type species: Abrothallus bertianus De Not. Giorn. Bot. Ital. 1: 194 (1846).

Notes: De Notaris described Abrothallus as a lichenized taxon (De Notaris 1846, 1849). Tulasne (1852) and Lindsay (1857) unequivocally established its lichenicolous habit. Suija et al. (2018) fixed the nomenclatural problems concerning the exact date of publication, confirming A. bertianus as the type species. They also reviewed the material described by Giuseppe De Notaris, Søren Christian Sommerfelt, and Ignaz Kotte lectotypifying Abrothallus species described by these authors (Suija et al. 2018). These authors also introduced the combination A. santessonii (≡ Vouauxiomyces santessonii), providing an updated description for this taxon. Pérez-Ortega et al. (2011) established the connection between the sexual and asexual morphs. The number of species is tentative since there are numerous single collections occurring on unusual hosts that may represent new species and there are only a few collections from certain regions, e.g. from Africa and Asia. Recent studies focused on the taxonomy of species growing on Peltigerales (Suija et al. 2011, 2015). However, species delimitation, especially those on specimens growing on Parmeliaceae, is difficult and in need of a thorough revision (Suija et al. 2018). For morphology of the type species, see Suija et al. (2018) (Fig. 2).

Fig. 2
figure 2

Morphological and anatomical features of Abrothallaceae. a Habitus Abrothallus welwitschii, mature apothecia. b Habitus A. welwitschii, young apothecia with greenish pruina. c Cross-section of A. parmotrematis on Parmotrema sp. d Interascal elements from A. bertianus. e One-septate ascospore of A. welwitschii, with surface showing ornamentation. f Ascus of A. boomii. g Three-septate ascospores of A. suecicus. h Scanning electron micrograph of A. welwitschii ascospores. i Ascospore of A. buellianus showing occasional splitting at the septum. j A. caerulescens asexual stage (pycnidium, Vouauxiomyces-type). k Conidia from A. boomii. l Lichenoconium sp. on A. santessonii ascoma. Scale bars: a–b = 0.5 mm, c = 50 µm, d–h, k = 10 µm; i = 5 µm, j = 30 µm, l = 20 µm


Economic and ecological significance

Species in Abrothallaceae are not harmful to plants or animals. However, their lichenicolous, parasymbiontic or parasitic lifestyles on macrolichens are interesting for biodiversity and ecological research.


Acrospermales Minter, Peredo & A.T. Watson.

Index Fungorum number: IF 90786; Facesoffungi number: FoF 06407.

Acrospermales was introduced by Minter (2007) to accommodate Acrospermaceae. The ordinal position of Acrospermaceae was previously unresolved and was referred to various orders by many authors. First, it was placed in Hysteriales or Dothideales in the 1st and 2nd editions of Ainsworth & Bisby’s Dictionary of the Fungi (Kirk et al. 2001). In the 3rd to 5th editions it was transferred to the Dothideales and in the 6th and 7th it was placed in the Ostropales and Clavicipitales, respectively. The family was also placed, rather hesitantly, in Pyrenulales by Eriksson (1982). Barr (1990) argued its ordinal position in more detail, and transferred it to Xylariales. Minter et al. (2007) introduced a new species in Acrospermaceae and discussed the ordinal position of Acrospermaceae with the introduction of Acrospermales. The divergence time for Acrospermales is estimated as 156 MYA (stem age, Hongsanan et al. 2020) (Fig. 3).

Fig. 3
figure 3

Phylogram generated from maximum likelihood analysis based on combined LSU and SSU sequence data representing Acrospermaceae. Strigula nemathora (MPN72) (Strigulaceae, Pleosporales) is used as the outgroup taxon. Bootstrap values for maximum likelihood (ML) equal to or greater than 70% and clade credibility values greater than 0.90 (the rounding of values to 2 decimal proportions) from Bayesian-inference analysis are labeled on the nodes. Ex-type strains are in bold and black, the new isolate is indicated in bold and blue

Accepted families: Acrospermaceae.


Acrospermaceae Fuckel, Jb. nassau. Ver. Naturk. 23–24: 92 (1870) [1869–70].

Index Fungorum number: IF 80430; Facesoffungi number: FoF 06380, 52 species.

Saprobic, epiphytic or symbiotic on herbaceous plants. Sexual morph: Ascomata solitary or in groups, superficial or immersed in stromata, erect, elongate, with smooth or sometimes rough surface, dark brown to black, flattened, club-shaped to conoid, with a short stipe, swelling when moist, ostiolate. Peridium comprising two or three layers, an outer layer composed of dark brown cells of textura angularis, a central layer, composed of hyaline, sometimes pale brown tissue of elongated cells intertwined, and an inner layer composed of dense tissue of small, light brown cells. Hamathecium comprising narrow, long, hyaline, filiform pseudoparaphyses. Asci typically 8-spored, bitunicate, long, narrowly cylindrical, pedicellate, apically rounded with an ocular chamber. Ascospores fasciculate, filiform, hyaline, multi-septate, almost as long as the asci, smooth-walled, not fragmenting, without sheath, typically intertwined in a fascicle within the ascus. Asexual morph: Hyphomycetous. Conidiophores micronematous, pale brown, septate, branched or unbranched. Conidiogenous cells holoblastic, sympodial with denticles, pale brown, smooth-walled. Conidia cylindrical, long ellipsoid, pale yellow, 1–3-septate, smooth-walled.

Type: Acrospermum Tode.

Notes: Acrospermaceae was introduced with a single genus Acrospermum by Fuckel (1870) and since then its higher taxonomic placement has undergone various changes (Saccardo 1883; Rehm 1887; Ellis and Everhart 1892; Ainsworth et al. 1973; Barr 1990). The family was previously placed in the class Dothideomycetes family incertae sedis, due to its uncertain position (Hyde et al. 2013). Two genera, Acrospermum and Oomyces are currently accepted (Lumbsch and Huhndorf 2010; Wijayawardene et al. 2018). Acrospermum is characterised by erect, elongate, usually brown, superficial ascomata that are more or less club-shaped and are solitary or in small groups, with long paraphyses which resemble ascospores. Oomyces is considered to be clavicipitalean (Diehl 1950) as it has conoid, yellowish white, multi-locular stromata, bitunicate asci and lacks pseudoparaphyses (Eriksson 1981). No monograph of the genus is available. Riddle (1920) reviewed Acrospermum and introduced A. maxoni and A. graminum var. foliicolum based on asexual morphs. Presently, the asexual morphs of Acrospermaceae comprise members of Dactylaria and Gonatophragmium (Wijayawardene et al. 2018). Dactylaria was shown to be polyphyletic by Bussaban et al. (2005) and is heterogenous (Seifert et al. (2011). Gonatophragmium was also found to be an asexual morph of Acrospermum by Kirk et al. (2008), but Seifert et al. (2011) did not assign it to any taxonomic rank. A Blast search of an LSU sequence of Gonatophragmium triuniae showed closed hits to Acrospermum adeanum (Crous et al. 2014). We therefore, agree with Wijayawardene et al. (2018) and include Gonatophragmium in Acrospermaceae until further data becomes available. The hyphomycetous genus Pseudovirgaria was introduced by Shin et al. in Arzanlou et al. (2007), with P. hyperparasitica as type species. Pseudovirgaria is assigned to Capnodiales, genera incertae sedis in Index Fungorum (2020), while it was mentioned as Dothideomycetes genera incertae sedis in Wijayawardene et al. (2018). In our phylogenetic analysis, two species of Pseudovirgaria clustered in Acrospermales. Pseudovirgaria is a hyphomycetous genus and morphologically resembles Gonatophragmium in having cylindric-clavate, thin-walled conidia. Therefore, we include Pseudovirgaria in Acrospermaceae based on phylogenetic evidence, and morphological resemblance to asexual genera of Acrospermaceae. Descriptions and illustrations of the asexual morphs of Acrospermaceae can be seen in previous studies (i.e. Crous et al. 2014; Berger et al. 2015; Shamsi et al. 2017)


Acrospermum Tode, Fung. mecklenb. sel. (Lüneburg) 1: 8 (1790).

Index Fungorum number: IF 54; Facesoffungi number: FoF 06381; 28 morphological species (Species Fungorum 2020), 8 species with molecular data.

Type species: Acrospermum compressum Tode, Fung. mecklenb. sel. (Lüneburg) 1: 8 (1790).

Notes: Tode (1790) introduced Acrospermum with A. compressum as the type species based on fruiting body and ostiole type. Acrospermum is characterised by superficial, club-like ascomata, bitunicate asci and fasciculate, filiform, hyaline, multi-septate ascospores (Riddle 1920). Species of Acrospermum are mostly saprobic and are distributed worldwide. The specimens of A. compressum were found on dry stems of Heracleum sphondylium in Germany. Acrospermum compressum can also be observed on dead stems of Urtica dioica, and A. graminum on grass culms. Acrospermum adeanum is a necrotrophic parasite and has been observed on 32 different moss species from 22 different genera, most of which belong to the pleurocarpous superorder Hypnanae (Döbbeler 1979; Bell and Newton 2004). The asexual morph of Acrospermum is hyphomycetous (Wijayawardene et al. 2018).


Acrospermum urticae D. Pem, Camporesi & K.D. Hyde, sp. nov.

Index Fungorum number: IF 556687; Facesoffungi number: FoF 06382; Fig. 4

Fig. 4
figure 4

Acrospermum urticae (IT 3999, holotype). a–d Ascomata on host surface. e Ascoma in vertical section f Peridium. g–i Narrowly cylindrical asci. j–k Filiform ascospores Scale bars: a = 2000 µm, b, e = 500 µm, c, d = 300 µm, f, g = 100 µm, h = 25 µm, i = 30 µm, j = 50 µm, k = 40 µm

Etymology: Name reflects the host from which the fungus is isolated.

Holotype: MFLU 18-1666.

Saprobic on dead stem of Urtica dioica. Sexual morph: Ascomata 940–1057 high × 301–345 µm diam. (\( \bar{x} \) = 1021 × 318 µm), solitary or in groups, superficial, club-shaped to conoid, erect, uni-locular, brown to blackish when dry, with a short stipe or sessile, flattened when dry, swelling when moist, ostiole large, apex rounded. Peridium 11–12 µm in vertical section comprising three layers, an outer layer comprising dark brown cells of textura angularis, a central thick layer, comprising pale brown to hyaline tissue of gelatinized hyphae with elongated cells, and an inner layer comprising dense tissue of small, hyaline cells. Hamathecium comprising narrow, long, pseudoparaphyses. Asci 195–319 × 6.2–6.7 μm (\( \bar{x} \) = 252.5 × 6.4 µm), 8-spored, bitunicate, narrowly cylindrical, pedicellate, with an ocular chamber. Ascospores 122–170 × 1.1–1.2 μm (\( \bar{x} \) = 146.2 × 1.2 µm), fasciculate, filiform, hyaline, multi-septate, nearly as long as the asci, smooth-walled. Asexual morph: undetermined

Material examined: Italy, Ravenna [RA], San Cassiano di Brisighella, on dead aerial stem of Urtica dioica (Urticaceae), 13 August 2018, Erio Camporesi (IT 3999, holotype; MFLU 18-1666, isotype).

GenBank numbers: LSU: MN597994, SSU: MN597996.

Notes: Acrospermum urticae differs from Acrospermum longisporium by its smaller ascomata (940–1057 high × 301–345 µm diam. v.s. 1500–2000 high × 400–500 µm diam.) and wider ascospores (122.1–175.3 × 1.1–1.2 μm v.s. 150–170 × 0.5–1 μm). Phylogenetic analyses of a combined LSU, SSU sequence dataset show that A. urticae forms a distinct lineage in Acrospermaceae with strong ML and BYPP support (80% ML, 1.0 BYPP; Fig. 3). Therefore, we introduce Acrospermum urticae as a new species.


Other genera included

Gonatophragmium Deighton, in Cejp & Deighton, Mycol. Pap. 117: 13 (1969).

Index Fungorum number: IF 8376; Facesoffungi number: FoF 06486; – 17 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Gonatophragmium mori (Sawada) Deighton 1969, in Cejp & Deighton, Mycol. Pap. 117: 13 (1969).

Spondylocladium mori Sawada, Spec. Bull. Agric. Exp. Station Formosa 19: 665 (1919).

Notes: Gonatophragmium was described in Cejp and Deighton (1969) with G. mori as the type species, a combination based on Spondylocladium mori. Gonatophragmium mori is a tropical-subtropical leaf-spotting species and is found on a wide range of hosts. Takahashi and Teramine (1986) considered Acrospermum viticola to be the sexual morph of this species, however this association was not proven by molecular data. Gonatophragmium is distinct in having pigmented, branched conidiophores formed as erect to decumbent threads with terminal and intercalary conidiogenous cells, which are regularly swollen around fertile portions with mostly numerous noticeable conidiogenous loci. The sexual morph of this genus is undetermined.


Oomyces Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 7: 185 (1851).

Index Fungorum number: IF 8376; Facesoffungi number: FoF 06488; – 7 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Oomyces carneoalbus (Lib.) Berk. & Broome, Ann. Mag. Nat. Hist., Ser. 2 7: 185 (1851).

Sphaeria carneoalba Lib., Pl. crypt. Arduenna, fasc. (Liège) 3(nos 201–300): no. 241 (1834).

Notes: Oomyces (= Coscinaria Ellis & Everh. 1886 fide Species Fungorum 2017) was introduced by Berk and Broome (1851), with O. carneoalbus as the type species. The genus is characterized by conoid, yellowish white, multi-locular stromata and lacks pseudoparaphyses (Eriksson 1981). Oomyces was described as similar to the egg of some insects, such as Crioceris, because its perithecia are visible as little dimples in the truncate apex of the fruiting body.

Pseudovirgaria H.D. Shin, U. Braun, Arzanlou & Crous, in Arzanlou et al., Shin & Crous, Stud. Mycol. 58: 87 (2007)

Index Fungorum number: IF504564; Facesoffungi number: FoF 06487; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Pseudovirgaria hyperparasitica H.D. Shin, U. Braun, Arzanlou & Crous, in Arzanlou et al., Stud. Mycol. 58: 87 (2007)

Notes: Phylogenetic analyses indicated that two species of Pseudovirgaria clustered within Acrospermaceae (Hudson et al. 2019, this study). Morphologically, it resembles Gonatophragmium in having cylindric-clavate, thin-walled conidia. Therefore, we agree with Hudson et al. (2019) to accept this genus in Acrospermaceae.


Economic and ecological significance

Species of Acrospermaceae are parasitic or endophytic and may play a negative role by infecting ferns. In Mexico, several species of Terpsichore (T. subtilis and T. taxifolia) are infected by Acrospermum, typically as black clavate stromata on the hosts.

Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.

Index Fungorum number: IF 90461; Facesoffungi number: FoF 07605.

Asterinales contains epifoliar fungi which have superficial mycelium forming a network on host plants, 1-celled appressoria, with a star-like opening to the thyriothecium. The order was revised by Hongsanan et al. (2014b), however, its classification is unclear due to insufficient sequence data. Sequence data of Asterinales clustered in two unrelated clades in phylogenetic trees (Ertz et al. 2016; Hyde et al. 2016b; Liu et al. 2017). The clade containing the type species Asterotexis cucurbitacearum was treated as Asterotexales (Ertz et al. 2016). However, Hyde et al. (2016b) synonymized it under Asterinales sensu stricto because most Asterinales strains cluster in this clade. The other clade, including species of Parmulariaceae were treated as Asterinales sensu lato (Liu et al. 2017). Parmulariaceae was transferred to its own order Parmulariales by Dai et al. (2018).

Although the type species Asterina melastomatis and a few other Asterinales-like taxa fell within the clade Asterinales sensu lato (Ertz et al. 2016; Hyde et al. 2016b), we did not include these sequence data in our analysis. This is because Asterinales-like taxa forming in the Asterinales sensu lato need to be rechecked since other hyphomycetous strains were added into this clade, thus, there is possibility that these sequence data are not Asterinales. Thyrinulaceae is introduced to accommodate a clade sister to Parmulariales (= Asterinales sensu lato) with Thyrinula as generic type based on the rules of nomenclatural priority. Hongsanan et al. (2014b) synonymised Lembosiaceae under Asterinaceae, however, adding more sequence data for Lembosia (Fig. 5) indicates that Lembosia should be raised to a family in Asterinales.

Fig. 5
figure 5

Phylogram generated from maximum likelihood analysis (RAxML) of Asterinales based on LSU sequence data. Maximum likelihood bootstrap values equal to or greater than 70%, Bayesian posterior probabilities equal to or greater than 0.90 (MLBS/PP) are given at the nodes. Isolate numbers are noted after each species name. The tree is rooted to Venturia inaequalis (ATCC 60070) and Venturia populina (CBS 256.38). Newly sequence data generated in this study are in blue. Ex-types are indicated in bold. Hyphen (-) represents support values less than 70% MLBS and 0.90 PP

By considering phylogenetic trees (Fig. 5), we retain Lembosiaceae in Asterinales and introduce Morenoinaceae and Neobuelliellaceae to accommodate the clades of Morenoina and Neobuelliella, respectively. Based on morphology and phylogeny, the current Asterinales comprises eight families. The classification of Asterinales is questionable and needs more morphological and molecular data to clarify its phylogenetic relationship. The divergence time for Asterinales is estimated as 221 MYA (stem age, Hongsanan et al. 2020).

Accepted families: Asterinaceae, Asterotexaceae, Hemigraphaceae, Lembisiaceae, Melaspileellaceae, Morenoinaceae, Neobuelliellaceae and Stictographaceae.

Asterinaceae Hansf., Mycol. Pap. 15: 188 (1946).

Index Fungorum: IF 80492; Facesoffungi number: FoF 06726, >1000 species.

Colonies epiphyllous or hypophyllous. Hyphae superficial, straight to substraight, dark brown, reticulate, with appressoria. Appressoria 1-celled, mostly lateral, alternate to unilateral. Sextual morph: Thyriothecia superficial, flattened, with stellate or longitudinal dehiscence. Upper walls brown, comprising radial, septate cells of textura prismatica. Asci 8-spored, bitunicate, ellipsoid, usually thickened at the apex. Hamathecium cellular pseudoparaphyses present or absent. Ascospores 2–5-seriate or fasciculate or conglobate, fusoid to ellipsoid, hyaline to dark brown, mostly 1-septate, smooth-walled or toughened. Asexual morph: Coelomycetous states with pycnidia or pycnothyria, and hyphomycetous states without conidiomata or sporodochia then gelatinous, pale. Hyphae brown, superficial, with appressoria. Conidiomata pycnothyria, flattened, dimidiate, radiate, orbicular, stellately dehisced at the centre. Conidiophores branched or unbranched, hyaline or brown. Conidiogenous cells monoblastic or percurrent, hyaline or brown. Conidia ovate, pyriform, angular, or wall straight to sinuate, brown.

Type: Asterina Lév.

Notes: Asterinaceae was established as a member of Microthyriales by Hansford (1946). Members of the family typically have upper walls comprising radiating cells with star-like or longitudinal splits and dark brown hyphae with appressoria. There are 18 genera in this family based on morphology (Hongsanan et al. 2014b; Guatimosim et al. 2015; Wijayawardene et al. 2017a; Dai et al. 2018). Although Wijayawardene et al. (2017a) included Echidnodes in Asterinaceae, we exclude it from Asterinaceae as Hongsanan et al. (2014b) transferred this genus to Aulographaceae. Phylogenetic studies have several different interpretations of this family. In this study, we reappraise the phylogenetic relationship of Asterinaceae and related families based on all available sequence data and previous studies.


Asterina Lév., Annls Sci. Nat., Bot., sér. 3 3: 59 (1845).

Index Fungorum number: IF 409; Facesoffungi number: FoF 06727; >1000 morphological species (Species Fungorum 2020), 9 species with molecular data.

Type species: Asterina melastomatis Lév., Annls Sci. Nat., Bot., sér. 3 3: 59 (1845).

Notes: Asterina was introduced as a member of Sphaeriaceae with A. azarae, A. compacta, A. pulla and the type A. melastomatis. It is the largest genus in Asterinaceae, but only nine species have sequence data available in GenBank due to its unculturable character. Members of the genus have circular thyriothecia with stellate dehiscence, lateral appressoria, globose asci, and dark brown, 1-septate ascospores.


Asterina magnoliae X.Y. Zeng, T.C. Wen & K.D. Hyde, in Hyde et al., Mycosphere 9(2): 349 (2018).

Index Fungorum number: IF 554238; Facesoffungi number: FoF 04089; Fig. 6

Fig. 6
figure 6

Asterina magnoliae (MFLU 16-0071). a Host leaves. b, c Colonies on leaf surface. d Thyriothecium. e Pycnidioma. f Young asci. g Mature asci. h Ascospores. i Pycnidia. Scales bars: b–c = 500 μm, d–e = 50 μm, f–g = 20 μm, h–i = 10 μm

Description: see Hyde et al. (2018).

Material examined: Thailand, Chiang Mai, Mae Taeng, Pa Pae, Bahn Pa Deng, Mushroom Research Centre, 128 Moo 3, on living leaves of Magnolia odora (Magnoliaceae), 8 July 2015, Xiang-Yu Zeng (MFLU 16-0071).

GenBank number: LSU: MN629745.

Notes: Our new collection of Asterina magnoliae is identified by morphological characters (Fig. 6) and phylogenetic evidence (Fig. 5). Hyde et al. (2018) provided a full description of this species.


Other genera included

Asterinella Theiss., Annls mycol. 10(2): 160 (1912).

Index Fungorum number: IF 411; Facesoffungi number: FoF 06729; – 35 morphological species (Species Fungorum 2020), molecular data available for an unnamed species in the genus.

Type species: Asterinella puiggarii (Speg.) Theiss., Brotéria, sér. bot. 10(2): 116 (1912).

≡ Asterina puiggarii Speg., Anal. Soc. cient. argent. 12(3): 99 (1881).

Notes: Asterinella was introduced as a member of Microthyriaceae. It is characterised by superficial hyphae with intercalary appressoria and thyriothecia with a stellate ostiole. Wu et al. (2014) transferred it to Asterinaceae based on morphology.


Asterolibertia G. Arnaud 1918, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 165 (1918) [1917].

Index Fungorum number: IF 421; Facesoffungi number: FoF 06731; – 35 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Asterolibertia couepiae (Henn.) G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 165 (1918) [1917].

Asterina couepiae Henn., Hedwigia 34: 104 (1895).

Notes: Asterolibertia is characterised by intercalary appressoria. Hongsanan et al. (2014b) synonymised Asterolibertia under Asterina, while Firmino et al. (2016) questioned the intercalary appressoria in Asterolibertia as not homologous to the lateral appressoria in Asterina. However, both justifications are based on morphology.


Asterostomella Speg., Anal. Soc. cient. argent. 22(4): 198 (1886).

Index Fungorum number: IF 7271; Facesoffungi number: FoF 06730; – 87 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Asterostomella paraguayensis Speg., Anal. Soc. cient. argent. 22(4): 198 (1886).

Notes: Asterostomella is a coelomycetous genus characterised by brown, ovoid, aseptate pycnidia, sometimes with a non-pigmented band in the middle. The genus is considered as a member of Asterinaceae based on its scutellate conidiomata with stellate dehiscence, which is similar to Asterina.


Batistinula Arx, Publicações Inst. Micol. Recife 287: 4 (1960).

Index Fungorum number: IF 523; Facesoffungi number: FoF 06732; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Batistinula gallesiae Arx, Publicações Inst. Micol. Recife 287: 6 (1960).

Notes: Batistinula typically has 3-septate ascospores. A specimen that shares the same morphological and biometric characteristics of the type was collected and sequenced by Guatimosim et al. (2015). However, that fresh collection was found on a different host family, which may contradict the host-specificity of Asterinaceae.


Cirsosia G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 127 (1918) [1917].

Index Fungorum number: IF 1065; Facesoffungi number: FoF 06734; – 15 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Cirsosia manaosensis (Henn.) G. Arnaud [as ‘manaoensis’], Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 127 (1918) [1917].

Lembosia manaosensis Henn. [as ‘manaoensis’], Hedwigia 43(4): 265 (1904).

Notes: Cirsosia is mostly similar to Lembosia in having a linear fissure, but has intercalary appressoria.


Dothidasteromella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 421 (1910).

Index Fungorum number: IF 1692; Facesoffungi number: FoF 06236; – 11 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Dothidasteromella sepulta (Berk. & M.A. Curtis) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 421 (1910).

Asterina sepulta Berk. & M.A. Curtis, Proc. Amer. Acad. Arts & Sci. 4: 129 (1860).

Notes: Dothidasteromella is mostly similar to Echidnodella, Halbania and Uleothyrium in lacking appressoria, but has Y-shaped dehiscence, 1-septate ascospores, and lack pseudoparaphyses.


Echidnodella Theiss. & Syd., Annls mycol. 15(6): 422 (1918) [1917].

Index Fungorum number: IF 1731; Facesoffungi number: FoF 06761; – 33 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Echidnodella linearis (Syd. & P. Syd.) Syd., Annls mycol. 15(6): 422 (1918) [1917].

Morenoella linearis Syd. & P. Syd., Annls mycol. 15(3/4): 250 (1917).

Notes: Echidnodella is similar to Lembosia and Cirsosia in having linear fissures, but lacks appressoria and has cellular pseudoparaphyses.


Halbania Racib., Crypt. Par. Java: no. 89 (1889).

Index Fungorum number: IF 2201; Facesoffungi number: FoF 06735; – 3 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Halbania cyathearum Racib., Crypt. Par. Java: no. 89 (1889).

Notes: Halbania is mostly similar to Batistinula in having 3-septate ascospores, but lacks appressoria on the hyphae. Sequence data is needed to confirm its phylogenetic placement.


Meliolaster Höhn., Ber. dt. bot. Ges. 35(10): 701 (1918).

Index Fungorum number: IF 3102; Facesoffungi number: FoF 06738; – 2 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Meliolaster clavisporus (Pat.) Höhn., Ber. dt. bot. Ges. 35(10): 701 (1918).

Meliola clavispora Pat., J. Bot., Paris 4: 61 (1890).

Notes: Thyriotheica in Meliolaster are composed of radially arranged of cells, which open by star-like fissures when mature. Meliolaster is similar to Batistinula and Halbania in having 3-septate ascospores, but differs in the presence and shape of appressoria on the hyphae.


Parasterinopsis Bat., Atas Inst. Micol. Univ. Recife 1: 327 (1960).

Index Fungorum number: IF 3722; Facesoffungi number: FoF 06739; – 3 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Parasterinopsis sersalisiae (Hansf.) Bat., Atas Inst. Micol. Univ. Recife 1: 327 (1960).

Patouillardina sersalisiae Hansf., Proc. Linn. Soc. London 156: 117 (1944) [1943–44].

Notes: Parasterinopsis is retained in Asterinaceae based on its thyriothecia with irregular fissures and the superficial hyphae with appressoria. However, it typically has cylindrical, 1–4-septate ascospores.


Platypeltella Petr., in Sydow & Petrak, Annls mycol. 27(1/2): 62 (1929).

Index Fungorum number: IF 4178; Facesoffungi number: FoF 06741; – 3 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Platypeltella smilacis Petr., Annls mycol. 27(1/2): 62 (1929).

Notes: Platypeltella was introduced as a member of Microthyriaceae. Wu et al. (2014) included it in Asterinaceae based on the superficial hyphae with intercalary capitate appressoria. Platypeltella is similar to Asterinella, but differs in having paraphyses and a round ostiole.


Prillieuxina G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4).

Index Fungorum number: IF 4365; Facesoffungi number: FoF 06742; – 62 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Prillieuxina winteriana (Pazschke) G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 162 (1918) [1917].

Asterina winteriana Pazschke, Hedwigia 31(3): 104 (1892).

Notes: Prillieuxina is typical of Asterinaceae in having stellate fissures and brown, 1-septate ascospores. However, appressoria are very rare in Prillieuxina, and the cells of the upper walls are radially arranged as in Microthyrium. Guatimosim et al. (2015) provided the only sequence for this genus.


Pycnocarpon Theiss., Abh. K.K. Zool.-Bot. Ges. Wien 7(3): 31 (1913).

Index Fungorum number: IF 4564; Facesoffungi number: FoF 07606; – 4 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Pycnocarpon magnificum (Syd., P. Syd. & E.J. Butler) Theiss., Abh. K.K. Zool.-Bot. Ges. Wien 7(3): 31 (1913).

Asterina magnifica Syd., P. Syd. & E.J. Butler, Annls mycol. 9(4): 391 (1911).

Notes: Wijayawardene et al. (2018) accepted this genus in Dothideomycetes genera incertae sedis. However, Doilom et al. (2018) included Pycnocarpon in Asterinaceae based on its superficial, web-like hypha, flattened thyriothecia, opening by radiating star-like or longitudinal splits, saccate asci, and conglobose, hyaline to brown, 1-septate ascospores strongly constricted at the septum (Doilom et al. 2018). We accept Pycnocarpon in Asterinaceae but note that the upper wall of ascomata of Pycnocarpon differs from members of Asterinaceae in having radially arranged, subglobose cells instead of cells of textura prismatica. Thus, sequence data is needed to confirm its placement.


Schenckiella Henn., Bot. Jb. 17: 523 (1893).

Index Fungorum number: IF 4885; Facesoffungi number: FoF 06743; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Schenckiella marcgraviae Henn., Bot. Jb. 17: 523 (1893).

Notes: This is a very unusual genus with a unique combination of characters, with surface hyphae lacking appressoria, Asterina-like thyriothecia, elongated clavate asci, brown, cellular pseudoparaphyses, which are rarely observed in the Dothideomycetes, and 4–5-septate brown ascospores.


Trichasterina G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 172 (1918) [1917].

Index Fungorum number: IF 5544; Facesoffungi number: FoF 06744; – 11 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Trichasterina styracis (Theiss.) G. Arnaud, Annals d’École National d’Agric. de Montpellier, Série 2 16(1–4): 172 (1918) [1917].

Asterina styracis Theiss., Abh. K.K. Zool.-Bot. Ges. Wien 7(3): 41 (1913).

Notes: Thyriothecia shares almost the same morphology as Asterina, but with setae on the hyphae. Whether such a character difference justifies separate genera should be tested using molecular data.


Trichopeltospora Bat. & Cif., in Batista, Costa & Ciferri, Publicações Inst. Micol. Recife 90: 17 (1958) [1957].

Index Fungorum number: IF 5569; Facesoffungi number: FoF 06745; – 2 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Trichopeltospora pipericola Bat., Cif. & C.A.A. Costa, in Batista, Costa & Ciferri, Publicações Inst. Micol. Recife 90: 17 (1958) [1957].

Notes: Trichopeltospora was introduced as a member of Microthyriaceae. Wu et al. (2011b) transferred Trichopeltospora to Asterinaceae based on its irregular ostiole and appressoria on the hyphae. This transfer needs to be confirmed by phylogenetic analyses.


Uleothyrium Petr., Annls mycol. 27(5/6): 388 (1929).

Index Fungorum number: IF 5661; Facesoffungi number: FoF 06762; – 2 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Uleothyrium amazonicum Petr., Annls mycol. 27(5/6): 388 (1929).

Notes: Uleothyrium is similar to Platypeltella in having rounded ostioles, which are distinct from any other genera in Asterinaceae. However, Uleothyrium lacks appressoria, while Platypeltella has intercalary appressoria.


Vizellopsis Bat., J.L. Bezerra & T.T. Barros, Publicações Inst. Micol. Recife 637: 5 (1969).

Index Fungorum number: IF 5748; Facesoffungi number: FoF 06746; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Vizellopsis grevilleae Bat., J.L. Bezerra & T.T. Barros, Publicações Inst. Micol. Recife 637: 5 (1969).

Notes: Vizellopsis was introduced as a member of Microthyriaceae, and Lumbsch and Huhndorf (2010) included this genus in Dothideomycetes incertae sedis. Dai et al. (2014a) transferred Vizellopsis to Asterinaceae based on the small, black thyriothecia, forming below the dark brown mycelium, comprising radiating cells and a concentrically ridged surface, but strongly thickened and septate hyphae which is different from other genera of Asterinaceae.


Economic and ecological significance

Asterinaceae species produce haustoria to gain nutrients from host plants without causing pathogenic damage. They may reduce photosynthesis by covering the host surface, and increase the temperature and respiration in those areas as do other black mildews.


Asterotexaceae Firmino, O.L. Pereira & Crous [as ‘Asterotexiaceae’], in Guatimosim et al., Persoonia 35: 238 (2015).

Index Fungorum number: IF 548079; Facesoffungi number: FoF 07607, 2 species.

Epiphytes, phytopathogens, forming dark colonies irregular to star-shaped, solitary to confluent. External mycelium growing through ascomatal cavity or fusing with the host epidermis cells, septate, hyaline, smooth. Appressoria formed underneath the ascomata, solitary or forming in small clusters, globose, cone-shaped or ovoid to elongate, brown, with a central hyaline penetration peg. Sexual morph: Ascomata superficial to erumpent, scutellate, dimidiate, brown to black. Scutellum of radially arranged rows of cells, poorly developed base, opening by numerous irregular fissures. Hamathecium comprising septate, anastomosing, cellular pseudoparaphyses, embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, oblong to cylindrical, with short and rounded pedicel or pedicel sometimes absent. Ascospores overlapping 2–3-seriate, ellipsoidal, hyaline to slightly yellowish, 1-septate, slightly constricted at the septum, upper cell broader than lower cell (adapted from Hongsanan et al. 2014b; Guatimosim et al. 2015). Asexual morph: Undetermined.

Type: Asterotexis Arx.

Notes: Asterotexaceae was established by Guatimosim et al. (2015), with the generic type Asterotexis. A phylogenetic tree provided by Guatimosim et al. (2015) showed that species of Asterotexis formed a distinct clade sister to the Inocyclus angularis (Incertae sedis clade). They introduced Asterotexiales to accommodate Asterotexaceae (Guatimosim et al. 2015). Ertz et al. (2016) indicated that the Asterotexiales clade contains Asterotexis species, Inocyclus angularia (Parmulariaceae) and some Asterinales species. However, the Asterotexiales clade in Ertz et al. (2016) was treated as Asterinaceae sensu stricto (Hyde et al. 2016b). Asterotexales was synonymized under Asterinales by Liu et al. (2017). In our phylogenetic analyses (Fig. 5), Asterotexis species cluster with the clade of Asterina species as an unstable clade. In another analysis, which did not include Lembosia mimusopis (data not shown), Asterotexis clustered with Inocyclus angularis (Incertae sedis clade). Thus, we retain Asterotexaceae within Asterinales and note that more sequence data are needed to clarify its phylogenetic placement.


Asterotexis Arx, Fungus, Wageningen 28: 6 (1958).

Index Fungorum number: IF 430; Facesoffungi number: FoF 06766; 2 morphological species (Index Fungorum 2020), 1 species with molecular data.

Type species: Asterotexis cucurbitacearum (Rehm) Arx.

Notes: Asterotexis is a plant-pathogen found on leaves, and was identified as a member of Asterinaceae (Inácio and Cannon 2008; Guerrero et al. 2011; Hongsanan et al. 2014b). Morphologically and phylogenetically, the genus could not be placed in any family of Asterinales (this study). Thus, Asterotexis is placed in its own family, Asterotexaceae.


Asterotexis cucurbitacearum (Rehm) Arx [as ‘cucurbitarum’], Fungus, Wageningen 28: 6 (1958).

Dothidella cucurbitacearum Rehm, Hedwigia 36(6): 376 (1897).

Index Fungorum number: IF 118911; Facesoffungi number: FoF 07608; Fig. 7

Fig. 7
figure 7

Asterotexis cucurbitacearum (a, b, d, f from S-F7565, holotype and c, e, g–n from from S-F22084). a Herbarium specimen. b Ascomata on substrate. c Ascoma when viewed in squash mount. d Drawing from von Arx and Müller (1975). e Upper wall of ascoma. f Asci. g Hamathecium. h–j Asci. k Ocular chamber strained in Melzer’s reagent. l, m 1-septate ascospores. n 2-septate ascospore. Scale bars: c = 100 μm, e–j = 20 μm, k–n = 10 μm

Description: see Hongsanan et al. (2014b).

Material examined: Brazil, Brazilia, Rio de Janeiro, on Cucurbitaceae, May 1887, E. Ule 676, Ex Herb. Sydow (SF7565, holotype); COSTA RICA, San José, Finca La Caja, on surface of leaves of Sechium edule (Cucurbitaceae), 25 March 1927, Det. J.A. Stevensen (S-F220847).


Economic and ecological significance

Species in this family are phytopathogens. The appearance of colonies on leaves can mainly reduce the photosynthesis, respiration, disrupt other plants mechanism and can make host tissues become pale.


Hemigraphaceae D.Q. Dai & K.D. Hyde, in Dai et al., MycoKeys 369(2): 67 (2018).

Index Fungorum number: IF 554062; Facesoffungi number: FoF 03910, 9 species.

Biotrophic on lichens. Sexual morph: Ascostromata solitary to gregarious, superficial, stellate, irregularly opening from the centre to margin, conical in section, coriaceous, black to dark brown. Peridium comprises two layers, black and thick-walled cells of textura angularis in outer part, thin and light brown cells of textura angularis at inner layers. Hamathecium comprising few, brown, unbranched, filamentous, septate, cellular pseudoparaphyses. Asci 8-spored, bitunicate, clavate to cylindric-clavate, subglobose, with an ocular chamber and a short pedicel. Ascospores 3-seriate to irregularly arranged, ellipsoid, brown, 1-septate, with larger upper cell and narrower lower cell, smooth-walled. Asexual morph: Undetermined.

Type: Hemigrapha (Müll. Arg.) R. Sant. ex D. Hawksw.

Notes: Dai et al. (2018) studied the syntype of Hemigrapha asteriscus (≡ Melanographa asteriscus), and concluded that it is different from the family type of Parmulariaceae. Phylogenetically (LSU), H. atlantica forms a distint lineage within Asterinales (Ertz and Diederich 2015; Dai et al. 2018; this study). Thus, Hemigraphaceae was established in Asterinales to accommodate a single genus Hemigrapha (Dai et al. 2018).


Hemigrapha (Müll. Arg.) R. Sant. ex D. Hawksw., Kew Bull. 30(1): 9 (1975).

Melanographa sect. Hemigrapha Müll. Arg., Flora, Regensburg 65(33): 519 (1882).

Index Fungorum number: IF 2282; Facesoffungi number: FoF 02311; 8 morphological species (Index Fungorum 2020), 1 species with molecular data.

Type species: Hemigrapha asteriscus (Müll. Arg.) R. Sant. ex D. Hawksw., Kew Bull. 30(1): 191 (1975).

Notes: Hemigrapha was included in Parmulariaceae by Lumbsch and Huhndorf (2010), Hyde et al. (2013) and Wijayawardene et al. (2014a). However, it is dissimilar with the family type of Parmulariaceae having dark, superficial, star-shaped ascomata forming on a thallus of lichens (Diederich and Wedin 2000; Dai et al. 2018). Ertz and Diederich (2015) included H. atlantica in their phylogenetic analysis and indicated that this genus belongs to Asterinales based on LSU sequence data, without assigning the family placement. The same result was shown in Dai et al. (2018) and in this study (Fig. 5).


Hemigrapha asteriscus (Müll. Arg.) R. Sant. ex D. Hawksw., Kew Bull. 30(1): 191 (1975).

Melanographa asteriscus Müll. Arg., Flora, Regensburg 65(33): 519 (1882).

Index Fungorum number: IF 530383; Facesoffungi number: FoF 02312; Fig. 8

Fig. 8
figure 8

Hemigrapha asteriscus (G 00292584, syntype). a, b Herbarium material. c, d Appearance of ascostromata on host surface. d Ascostromata in the water. e Section of ascostroma. f Asci with hamathecial tissues. g Ascospores. Scale bars: b = 10 mm, c, d = 500 µm, e = 50 µm, f, g = 5 µm

Description: see Dai et al. (2018).

Material examined: Australia, New South Wales, Mount Kosciuszko, on thallus of Peltigera dolichorrhiza (Lichen). Müller J. 1882 (G 00292584, syntype).


Economic and ecological significance

Members of this family are biotrophic on lichens and appear as black colonies on the host surface.


Lembosiaceae Hosag, in Hosagoudar et al., J. Mycopathol. Res. 39(1): 61 (2001).

Index Fungorum number: IF 80503; Facesoffungi numbers: FoF 07609, 160 species.

Epiphytic on living leaves. Superficial hyphae with lateral appressoria. Sexual morph: Thyriothecia solitary, scattered, superficial, oval, ellipsoidal, X- or Y-shaped, easily removed from the host surface, black, opening by a linear fissure, with basal peridium poorly developed. Upper wall comprising linear, dark cells, which are branched at the margin. Hamathecium comprising vertical asci inclined upwards, pseudoparaphyses not observed. Asci bitunicate, fissitunicate dehiscence not observed, subglobose to ovoid, apedicellate, apical region of asci usually with a thick opaque region, ocular chamber not observed, not staining blue in IKI. Ascospores overlapping, oblong to obvoid, hyaline to brown, 1-septate. Asexual morph: Undetermined.

Type: Lembosia Lév.

Notes: Hosagoudar et al. (2001) introduced Lembosiaceae using the morphological character of elongate thyriothecia with longitudinal or X- or Y-shaped slits (Hongsanan et al. 2014b). However, Hongsanan et al. (2014b) treated Lembosiaceae as a synonym of Asterinaceae based on the first phylogenetic evidence of LSU sequence data of Lembosia albersii. Here, we re-introduced Lembosiaceae using sequence data currently available in GenBank.


Lembosia Lév., Annls Sci. Nat., Bot., sér. 3 3: 58 (1845).

Index Fungorum number: IF 2724; Facesoffungi number: FoF 06736, – 160 morphological species (Species Fungorum 2020), 6 species with molecular data

Type species: Lembosia tenella Lév., Annls Sci. Nat., Bot., sér. 3 3: 58 (1845).

Notes: Species of Lembosia and Asterina are obligately biotrophic, having appressoria and similar thyriothecia (Hosagoudar et al. 2001). However, Lembosia differs from Asterina in having elongate thyriothecia which dehisce to open by a longitudinal or X- or Y-shaped slit (Hosagoudar 1991).


Lembosia xyliae X.Y. Zeng, T.C. Wen & K.D. Hyde, in Ariyawansa et al., Fungal Diversity: 75: 50 (2015).

Index Fungorum number: IF 551345; Facesoffungi number: FoF 00933, Fig. 9

Fig. 9
figure 9

Lembosia xyliae (MFLU 14-0004, holotype). a, b Colonies on leaf surface. c Squash mount of ascomata with asci. d Upper walls and hyphae with hyphopodia. e Immature ascospores. f Mature ascospores. Scale bars: a = 400 μm, b = 200 μm, c = 100 μm, d = 50 μm, e–f = 10 μm

Description: see Ariyawansa et al. (2015a).

Material examined: Thailand, Chiang Rai, Mae Fah Luang University, on leaves of Xylia sp. (Fabaceae), 18 January 2014, XY Zeng (MFLU 14-0004, holotype).


Economic and ecological significance

Species in Lembosiaceae are considered as obligatory biotrophs, however, there are no records available to show its economic significance.


Melaspileellaceae D.Q. Dai & K.D. Hyde, in Dai et al., Phytotaxa 369(2): 70 (2018).

Index Fungorum number: IF 554063; Facesoffungi number: FoF 03911, 1 species.

Saprobic on trees and shrubs. Sexual morph: Ascostromata solitary, superficial, dark to black, coriaceous, small rounded. Peridium comprises 2 layers, black and thick-walled cells at outer layers, light brown to hyaline cells of textura angularis of inner layers. Hamathecium comprising dense, hyaline, mainly unbranched, filamentous, septate, cellular pseudoparaphyses around asci, with brownish tips. Asci 8-spored, bitunicate, broadly clavate to subglobose, with a rounded apex and a short pedicel. Ascospores 2-seriate to irregularly arranged, ellipsoid, hyaline, 1-septate, with slightly larger upper cell, with slightly narrower lower cell, smooth-walled, bearing 2–3 appendages which disappear when dry. Asexual morph: Undetermined (adapted from Dai et al. 2018).

Type: Melaspileella (P. Karst.) Vain.

Notes: This family was established by Dai et al. (2018) to accommodate a single genus Melaspileella based on phylogenetic placement generated from LSU and SSU of M. proximella. The family formed a clade sister to Hemigraphaceae within Asterinales with high bootstrap support in Dai et al. (2018), while sister to Pirozynskiella laurisilvatica (FMR 13133) genus incertae sedis in Asterinales in our analyses (Fig. 5). Thus, we retain Melaspileellaceae in Asterinales.


Melaspileella (P. Karst.) Vain., Ann. Acad. Sci. fenn., Ser. A 15(no. 6): 317 (1921).

= Mycomelaspilea Reinke, in Pringsheim, Jb. wiss. Bot. 28: 136 (1895).

Index Fungorum number: IF 3095; Facesoffungi number: FoF 07733; 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Melaspileella proximella (Nyl.) Ertz & Diederich.

Notes: Melaspileella was introduced by Vainio (1921b). Ertz and Diederich (2015) lectotypified the genus with M. proximella. They also transferred the genus to Asterinales without assigning it to any family. A phylogenetic tree based on LSU and SSU sequence data in Dai et al. (2018) indicated that M. proximella should be assigned in a new family, Melaspileellaceae.


Melaspileella proximella (Nyl.) Ertz & Diederich, Fungal Diversity 71: 161 (2015).

Arthonia proximella Nyl., Lich. Scand. (Helsinki): 262 (1861).

Index Fungorum number: IF 811374; Facesoffungi number: FoF 07610; Fig. 10

Fig. 10
figure 10

Melaspileella proximella. a Fruiting body on host (redrawn from ERD-7986. Photo: Enrique Rubio). b Section through ascoma, in water (redrawn from Finland, H-NYL 4827, lectotype). c Ascus with ascospores, in water (redrawn from Finland, H-NYL 4827, lectotype). d Ascus with ascospores (Hungary, on Juniperus, Zeller & Tóth 2835, BP – paratype of Banhegyia setispora). e Ascospore with polar setulae, in water (redrawn from Belgium, Ertz 19187). f Ascospore with polar setulae (redrawn from Finland, Nylander, H – lectotype). Scale bars: b = 20 µm, c, d = 10 µm, e, f = 5 µm

Description: see Zeller and Tóth (1960).


Economic and ecological significance

Members of this family are saprobic on plants and involved in recycling organic matter. Melaspileella proximella grows on trees and shrubs including trunks of Pinus and Tilia, and twigs of Juniperus (Ertz and Diederich 2015).


Morenoinaceae Hongsanan & K.D. Hyde, fam. nov.

Index Fungorum number: IF 557813; Facesoffungi number: FoF 07611, 26 species.

Saprobic on stem or leaves, forming blackened areas. Sexual morph: Thyriothecia solitary, aggregated, or gregarious, superficial, easily removed from the host surface, black, ellipsoid, oblong, curved, opening by X- or Y-shaped or linear fissures, branched at the margin, from the center to the outer rim, lacking free hyphae and appressoria at the margin. Upper wall comprising linear, dark cells, which are branched at the margin, radiating from the center to the outer rim. Hamathecium pseudoparaphyses not observed. Asci 8-spored, bitunicate, fissitunicate dehiscence not observed, subglobose to oblong or saccate to globose, apedicellate, ocular chamber not observed, or with a distinct, thickened apical region. Ascospores overlapping 2–3-seriate, oblong to obvoid, or fusiform, hyaline, 1-septate, constricted at the septum, with or without guttules, smooth-walled (Hongsanan et al. 2014b; Tibpromma et al. 2017). Asexual morph: Coelomycetous. “Sirothyriella”, Pycnothyria circular, radially scutellate. Conidiogenous cells holoblastic, simple, hyaline. Conidia cylindrical, hyaline, 1-celled (asexual morph from Sivanesan 1984; drawing of asexual characters can be seen in Ellis 1980).

Type: Morenoina Theiss.

Notes: Morenoinaceae resembles Aulographaceae (Hongsanan et al. 2014b). In our phylogenetic analyses (Fig. 5), Morenoina calamicola is closely related to Morenoina palmicola (MFLUCC 15-0284) with 67% ML and 0.93 BYPP support as a distinct clade within Asterinales. Therefore, we introduce Morenoinaceae to accommodate the Morenoina clade.


Morenoina Theiss., Annls mycol. 11(5): 434 (1913).

= Aulographella Höhn., Ber. dt. bot. Ges. 35: 359 (1917).

Index Fungorum number: IF 3270; Facesoffungi number: FoF 07612, 26 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Morenoina antarctica (Speg.) Theiss.

Notes: Morenoina was placed in Leptopeltidaceae and Asterinaceae (von Arx and Müller 1975; Lumbsch and Huhndorf 2010). Hongsanan et al. (2014b) designated an epitype and placed this genus in Aulographaceae as Morenoina did not have free hyphae with appressoria as other members of Asterinaeceae and it has different ascus form from members of Leptopeltidaceae. Morenoina resembles Aulographum and differs only in the morphology of the scutellum which comprises inordinately arranged cells and a hypostroma of subcuticular hyphae beneath the thyriothecium, which is lacking in Morenoina (Ellis 1980). Phylogenetic placement of Aulographaceae is uncertain due to lack of sequence data. Sequence data of Aulographum hederae (type species) formed an unstable clade, while two species of Morenoina form a distinct clade within Asterinales (Fig. 5). Therefore, we place Morenoina in the new family Morenoinaceae.


Morenoina antarctica (Speg.) Theiss., Annls mycol. 11(5): 434 (1913).

Morenoella antarctica Speg., Boln Acad. nac. Cienc. Córdoba 11(2): 240 (1887) [1888].

Index Fungorum number: IF 174884; Facesoffungi number: FoF 07613; Fig. 11

Fig. 11
figure 11

Morenoina antarctica (IMI 349680, epitype). a, b Herbarium and specimen. c, d Appearance of thyriothecia on surface of plant. e Squash mount of thyriothecium. f Upper wall of thyriothecium. g–h Asci when immature. i Asci at maturity. j–l Ascospores. Scale bars: e = 100 μm, f = 50 μm, g–l = 20 μm

Material examined: Argentina, Tierra del Fuego, Ushuaia, south of Paso e Garibaldi, on the stem of Gramineae, 29 October 1989, collected by P.F. Cannon and D.W. Minter, identified by P.F. Cannon (IMI 349680, epitype).


Economic and ecological significance

Species of this family are saprobic and play a role in recycling organic matter.


Neobuelliellaceae Hongsanan & K.D. Hyde, fam. nov.

Index Fungorum number: IF 558103;

Facesoffungi number: FoF 07614, 1 species.

Lichenicolous. Vegetative hyphae intercellular. Sexual morph: Ascomata apothecioid, dispersed over the host thallus as black dots, grouped, rarely single, rounded, globose when young, pressed at the edges, oval when mature, often bleaching the host thallus, occasionally sunken. Excipulum dark brown, K+ greenish-brown. Hymenium hyaline to brownish. Disc olive-brown, I-, KI-. Hamathecium comprising branched, anastomosing, cellular pseudoparaphyses, capitate at apical and intercalary cells with pigment, thick at tips, with dark-brown pigment. Asci 4–6-spored, bitunicate, fissitunicate, cylindrical. Ascospores long, cells rounded, the upper cell sometimes tapered, hyaline, later ± brown, 1-septate, clearly constricted at the septa, thin-walled (adapted from Hafellner et al. 2008; Etayo 2010; Yazici and Etayo 2013). Asexual morph: Undetermined.

Type: Neobuelliella Hongsanan & K.D. Hyde

Notes: Neobuelliellaceae is similar to species of Buelliella which are placed in genus incertae sedis in Dothideomycetes. Buelliella minimula (type species) together with B. physciicola cluster within Stictographaceae (Asterinales) in the phylogenetic analyses of Dai et al. (2018) and this study. Two strains of Neobuelliella poetschii (≡ Buelliella poetschii) form a distinct clade separately from Stictographaceae, and are sister to Hemigraphaceae (Ertz et al. 2015; Dai et al. 2018; this study). Therefore, we introduce a new family Neobuelliellaceae to accommodate this clade.


Neobuelliella Hongsanan & K.D. Hyde, gen. nov.

Index Fungorum number: IF 557815; Facesoffungi number: FoF 07615; 1 morphological species (this study), 1 species with molecular data.

Lichenicolous. Vegetative hyphae intercellular. Sexual morph: Ascomata apothecioid, dispersed over the host thallus as black dots, grouped, rarely single, rounded, globose when young, pressed at the edges, oval when mature, often bleaching the host thallus, occasionally sunken. Hymenium hyaline to brownish. Disc olive-brown, I-, KI-. Excipulum dark brown, K+ greenish-brown. Hamathecium comprising branched, anastomosing, cellular pseudoparaphyses, capitate at apical and intercalary cells with pigment, thick at tips, with dark-brown pigment. Asci 4–6-spored, bitunicate, fissitunicate cylindrical. Ascospores long, cells rounded, the upper cell sometimes tapered, hyaline, later ± brown, 1-septate, clearly constricted at the septa, thin-walled (Hafellner et al. 2008; Etayo 2010; Yazici and Etayo 2013). Asexual morph: Undetermined.

Type species: Neobuelliella poetschii (Hafellner) Hongsanan & K.D. Hyde.

Notes: The genus is introduced to accommodate Neobuelliella poetschii which was known as Buelliella poetschii. Phylogenetic analyses from Dai et al. (2018) and this study (Fig. 5) indicate that it does not cluster with the type species of Buelliella. Thus, we introduce the new genus Neobuelliella to accommodate a single species Buelliella poetschii (current name is Neobuelliella poetschii).


Neobuelliella poetschii (Hafellner) Hongsanan & K.D. Hyde, comb. nov.

≡ Buelliella poetschii Hafellner, in Hafellner, Herzog & Mayrhofer, Mitt. naturw. Ver. Steierm. 137: 187 (2008).

Index Fungorum: IF 557814, Facesoffunginumber: FoF 07616; Fig. 12

Fig. 12: Neobuelliella poetschi
figure 12

. a Apothecioid ascomata (redrawn from Yazici and Etayo 2013). b Ascospores (redrawn from holotype in Hafellner et al. 2008). c Young ascus among hamathecium (redrawn from Ertz and Diederich 2015)

Description: see Yazici and Etayo (2013).

Notes: Phylogenetically, Buelliella poetschii formed a distinct lineage within Asterinales, and did not cluster with B. minimula (type species of Buelliella). Thus, B. poetschii is synonymized in Neobuelliella.


Economic and ecological significance

Neobuelliellaceae comprises lichenicolous taxa which play insignificant roles in ecosystems. There is only a single species in this family. Fresh collections are needed to understand the ecological significance of Neobuelliellaceae.


Stictographaceae D.Q. Dai & K.D. Hyde, in Dai et al., MycoKeys 369(2): 70 (2018).

Index Fungorum number: IF 554064; Facesoffungi number: FoF 03912, 39 species.

Lichenicolous. Sexual morph: Ascomata solitary, scattered to clustered, superficial, cymbiform to lirelliform, with slit-like disc, with a slit or star-shaped fissure, black to dark brown. Peridium thick, outer layers of black to dark brown, thick-walled cells of textura angularis, inner layers of light brown to hyaline cells of textura angularis. Hamathecium comprising dense, filamentous, septate, unbranched or occasionally branched at upper part around the asci, cellular pseudoparaphyses. Asci 8-spored, bitunicate, clavate to subglobose, with an ocular chamber, apically rounded, short pedicellate. Ascospores 2-seriate to irregularly arranged, ellipsoid, hyaline, becoming light brown to dark brown, 1-septate, with slightly larger upper cell, and narrower lower cell, smooth-walled. Asexual morph: Undetermined (Dai et al. 2018).

Type: Stictographa Mudd.

Notes: Stictographaceae was introduced to accommodate four lichenicolous or saprobic genera which have cymbiform to lirelliform, or a slit-like discs, black to dark brown ascomata, wide clavate to subglobose asci, and ellipsoid, brown, 1-septate ascospores (viz. Karschia, Labrocarpon, Melaspileopsis and Stictographa; Dai et al. 2018). In the phylogenetic tree, these four genera clustered as a distinct clade within Asterinales (Dai et al. 2018). As the type species of Buelliella clustered within the clade of Stictographaceae, we therefore include Buelliella in Stictographaceae.

Stictographa Mudd, Man. Brit. Lich.: 226 (1861).

Index Fungorum number: IF 5238; Facesoffungi number: FoF 07617; 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Stictographa lentiginosa (Lyell ex Leight.) Mudd.

Notes: Stictographa can be compared with Labrocarpon in having lichenicolous, black, simple to occasionally short, branched, lirelliform ascomata with a slit-like opening (Ertz and Diederich 2015; Diederich et al. 2017). Ertz and Diederich (2015) inferred that Stictographa and Labrocarpon might be synonyms, however, they do not phylogenetically cluster together (Ertz and Diederich 2015; Dai et al. 2018). Thus, Labrocarpon is accepted as a distinct genus. More collections and species of these genera are needed to clarify their relationship.

Stictographa lentiginosa (Lyell ex Leight.) Mudd, Man. Brit. Lich.: 226 (1861).

≡ Opegrapha lentiginosa Lyell ex Leight., Ann. Mag. nat. Hist., Ser. 2 13: 211 (1854).

Index Fungorum number: IF 406525; Facesoffungi number: FoF 07618; Fig. 13

Fig. 13: Stictographa lentiginos
figure 13

(redrawn Ertz and Diederich 2015). a Ascomata growing on lichen. b Clavate to subglobose asci and pseudoparaphyses. c Ascospores. Scale bars: a = 200 µm, b, c = 10 µm

Description: see Redinger (1938), Sanderson et al. (2009), Ertz and Diederich (2015).


Other genera included

Buelliella Fink, Lich. Fl. U.S.: 372 (1935).

Index Fungorum number: IF 676; Facesoffungi number: FoF 03625; 12 morphological species (Species Fungorum 2020, this study), 2 species with molecular data.

Type species: Buelliella minimula (Tuck.) Fink, Lich. Fl. U.S.: 372 (1935).

Buellia minimula Tuck., Syn. N. Amer. Lich. (Boston) 2: 106 (1888).

Notes: The original diagnosis of this genus was provided by Fink (1935), and valided by Hafellner (1979). Buelliella is characterised by a non-lichenized, lichenicolous habit, initially subsphaerical and closed ascomata, with the upper ascomatal breaking gradually away to expose the hymenium when mature, becoming apothecioid, with bitunicate, broadly clavate to subcylindrical asci, and a distinct ocular chamber, and ellipsoid, 1-septate, pale to medium brown ascospores. Ertz and Diederich (2015) noted that the genus is not monophyletic due to B. physciicola and B. poetschii being distantly related, and sequence data of the type species (B. minimula) was unavailable. Thus, it was placed as incertae sedis in Dothideomycetes (Kirk et al. 2008; Ertz and Diederich 2015; Wijayawardene et al. 2017a). We included sequence data of B. minimula in our analyses and found that it forms a lineage within Sticgraphaceae. Buelliella poetschii is transferred to a new genus, Neobuelliella in this paper.


Karschia Körb., Parerga lichenol. (Breslau) 5: 459 (1865).

Index Fungorum number: IF 2545; Facesoffungi number: FoF 07619; – 24 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Karschia talcophila (Ach.) Körb., Parerga lichenol. (Breslau) 5: 460 (1865).

Lecidea talcophila Ach., Lich. univ.: 183 (1810).

Notes: More than 100 species of Karschia are listed in Index Fungorum (2020), however, most taxa have been transferred to other genera, such as Buelliella, Cycloschizon, Dothidea and others by Hafellner (1979). Two species were accepted by Hawksworth et al. (1995), and later four species were included (Kirk et al. 2008). Karschia is characterized by a lichenicolous life-style, immersed to erumpent, rounded, black ascomata with a large, irregular opening, producing pseudoparaphyses with a slightly enlarged and brown apex, bitunicate, clavate to subcylindrical asci, ellipsoid, brown, smooth ascospores with one septum (Ertz and Diederich 2015).


Labrocarpon Etayo & Pérez-Ort., in Pérez-Ortega & Etayo, Lichenologist 42(3): 271 (2010).

Index Fungorum number: IF 515228; Facesoffungi number: FoF 07620; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Labrocarpon canariense (D. Hawksw.) Etayo & Pérez-Ort. [as ‘canariensis’], in Pérez-Ortega & Etayo, Lichenologist 42(3): 272 (2010).

Melaspilea canariensis D. Hawksw., Lichenologist 14(1): 84 (1982).

Notes: Labrocarpon is a monotypic genus which was introduced by Pérez-Ortega and Etayo (2010) based on morphology of ascomata with periphyses (Ertz and Diederich 2015). Labrocarpon is more or less similar to Stictographa (Pérez-Ortega and Etayo 2010; Ertz and Diederich 2015). However, it formed a separate clade from Stictographa (Ertz and Diederich 2015; Dai et al. 2018). Labrocarpon is characterized by a lichenicolous life-style, superficial, rounded to slightly elliptic, black, disk-like ascomata with a slit-like opening and a well-developed exciple, producing pseudoparaphyses with a swollen and brown apex, bitunicate, clavate, thick-walled asci with an ocular chamber, and two-celled ascospores (Ertz and Diederich 2015).


Melaspileopsis (Müll. Arg.) Ertz & Diederich, Fungal Diversity 71: 161 (2015).

Index Fungorum number: IF 811387; Facesoffungi number: FoF 07621; – 1 morphological species (Dai et al. 2018), 1 species with molecular data.

Type species: Melaspileopsis diplasiospora (Nyl.) Ertz & Diederich, Fungal Diversity 71: 161 (2015).

Opegrapha diplasiospora Nyl., Acta Soc. Sci. fenn. 7(2): 476 (1863).

Notes: The genus was established by Ertz and Diederich (2015), with the type species M. diplasiospora (≡ Melaspilea diplasiospora in Index Fungorum 2020). Phylogenetically, it does not belong to Melaspileaceae but is sister to Stictographa within Stictographaceae (Dai et al. 2018). This genus was placed in Melaspileaceae, order Eremithallales (Wijayawardene et al. 2018). However, we agree with Dai et al. (2018) to retain Melaspileopsis within Stictographaceae.


Economic and ecological significance

Stictographaceae accommodates several lichenicolous and one saprobic genus. Members of this family usually occur on the thallus of lichens. They are distributed in tropiccal to temperate areas. Stictographa lentiginosa is much rarer than the host (Phaeographis dendritica) and thus can act as a useful indicator of the quality of ancient woodlands in western Europe (Ertz and Diederich 2015).


Botryosphaeriales C.L. Schoch, Crous & Shoemaker.

Index Fungorum number: IF 501513; Facesoffungi number: FoF 07659.

Botryosphaeriales was introduced to accommodate a single family Botryosphaeriaceae (Schoch et al. 2006). Previously, nine families (Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae, Phyllostictaceae, Planistromellaceae, Saccharataceae, Septorioideaceae, Endomelanconiopsidaceae, Pseudofusicoccaceae) were accepted in Botryosphaeriales in the Outline of Ascomycetes (Wijayawardene et al. 2018). However, currently only six families (Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae, Phyllostictaceae, Planistromellaceae, Saccharataceae) are accepted in this order (Phillips et al. 2019), the remaining three families were synonymized with existing families. Thus, Endomelanconiopsidaceae is considered a synonym of Botryosphaeriaceae, Pseudofusicoccaceae a synonym of Phyllostictaceae and Septorioideaceae a synonym of Saccharataceae (Phillips et al. 2019). Species in the order are characterized by uni- or multilocular ascostromata with multi-layered dark brown walls, occurring individually or in clusters, frequently embedded in stromatic tissue. Asci are bitunicate, with a thick endotunica, pedicellate or sessile, clavate, with a well-developed ocular chamber, intermixed with hyaline, septate pseudoparaphyses, branched or not. Ascospores are hyaline or pigmented, septate or not, ellipsoid to ovoid, with or without mucoid appendages or mucilaginous sheath. Conidiomata are pycnidial, uni- to multilocular, frequently embedded in stromatic tissue. Conidiogenous cells are hyaline, phialidic or annelidic. Conidia are hyaline or pigmented, septate or not, thin- or thick-walled, with or without mucoid appendages or sheaths (Phillips et al. 2019). The divergence time for Botryosphaeriales is estimated as 150 MYA (stem age, Hongsanan et al. 2020) (Fig. 14).

Fig. 14
figure 14

Phylogram generated from maximum likelihood analysis (RAxML) of Botryosphaeriales based on ITS and LSU sequence data. Maximum likelihood bootstrap values equal to or greater than 70%, Bayesian posterior probabilities equal to or greater than 0.90 (MLBS/PP) are given at the nodes. Isolate numbers are noted after each species name. The tree is rooted to Patellaria quercus (CPC 27232 and BHI F828a). Newly sequence data generated in this study are in blue. Ex-type and references strains are indicated in bold. Hyphen (-) represents support values less than 70% MLBS and 0.90 PP

Accepted families: Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae, Phyllostictaceae, Planistromellaceae, Saccharataceae.


Aplosporellaceae Slippers, Boissin & Crous, Stud. Mycol. 76(1): 41 (2013).

Index Fungorum: IF 805795; Facesoffungi number: FoF 00113, 13 species.

Sexual morph: Stromata pseudothecial, multiloculate ascomata with multi-layered, dark brown walls, embedded in stromatic tissue. Hamathecium comprising hyphoid, hyaline pseudoparaphyses. Asci bitunicate, mostly with a thick endotunica, clavate, with a well-developed ocular chamber and intermixed with pseudoparaphyses. Ascospores 1–2-seriate, ellipsoid to ovoid, mostly aseptate, or septate, hyaline to pigmented, lacking mucoid appendages or sheaths (Slippers et al. 2013; Liu et al. 2012; Thambugala et al. 2014a; photoplates of sexual characters can be seen in Ekanayaka et al. 2016). Asexual morph: Coelomycetous. Conidiomata pycnidial, multilocular, embedded in stromatic tissues. Conidiophores reduced to conidiogenous cells. Conidiogenous cells hyaline, phialidic. Conidia ellipsoid to subcylindrical, initially hyaline, thin-walled and smooth, becoming pigmented, thick-walled and spinulose, aseptate (Sutton 1980).

Type: Aplosporella Speg.

Notes: Aplosporellaceae was introduced by Slippers et al. (2013) to accommodate Aplosporella and Bagnisiella. Currently two genera (Aplosporella and Alanomyces) are accepted in this family. This family is phylogenetically close to Botryosphaeriaceae. Liu et al. (2017), using evolutionary evidence, questioned whether this family was introduced unnecessarily and should be regarded as a genus. However, based on morphology, phylogeny and evolutionary divergence times, Phillips et al. (2019) regarded Aplosporellaceae as a distinct family.


Aplosporella Speg., Anal. Soc. cient. argent. 10(4): 157 (1880).

= Bagnisiella Speg., Anal. Soc. cient. argent. 10(4): 146 (1880).

Index Fungorum number: IF 7191; Facesoffungi number: FoF 01427; 12 morphological species (botryosphaeriales.org 2020), 10 species with molecular data.

Type species: Aplosporella chlorostroma Speg.

Notes: Aplosporella was introduced by Spegazzini (1880). This genus has frequently been incorrectly referred to as “Haplosporella” (Tilak and Rao 1964; Petrak and Sydow 1927; Tai 1979; Wei 1979). About 340 epithets for Aplosporella are listed in Index Fungorum (July 2019). This genus is characterised by multiloculate pycnidia, brown, aseptate, spinulose conidia, hyaline, aseptate spermatia and fusiform paraphyses (Du et al. 2017). Its sexual morph is not common (Ekanayaka et al. 2016). Slippers et al. (2013) indicated that Aplosporella should be separate from Botryosphaeriaceae as a distinct family, which they named Aplosporellaceae, and confirmed that Aplosporella is well-resolved with species described from living culture. They also found a close genetic relationship between the genera Aplosporella and Bagnisiella and considered that Bagnisiella represents the sexual morph and should be reduced to synonymy with Aplosporella (Schoch et al. 2009a; Slippers et al. 2013). Ekanayaka et al. (2016) introduced a new species Aplosporella thailandica from Thailand with morphology corresponding to Bagnisiella. Phillips et al. (2019) formally placed Bagnisiella as a synonym of Aplosporella. The most recent phylogenetic tree of species in Aplosporella is provided by Hyde et al. (2020a).


Aplosporella javeedii Jami, Gryzenh., Slippers & M.J. Wingf., Fungal Biology 118(2): 174 (2013).

Index Fungorum number: IF 803637; Facesoffungi number: FoF 07383; Fig. 15

Fig. 15
figure 15

Asexual morph of Aplosporella javeedii (HKAS 102118). a Substrate. b Conidiomata on substrate. c Cross section of multiloculate conidioma. d Conidioma wall. e, f Different stages of conidiogenesis. g, h Conidia. i Spermatogenous cells. j Spermatia. Scale bars: c = 1.5 mm, d = 100 μm, e–h, j = 10 μm, i = 25 μm

Description: see Phillips and Alves (2009).

Saprobic on dead stems. Sexual morph: Undetermined. Asexual morph: Conidiomata 1–2 mm diam., erumpent from bark surface, separate, discoid, dark-brown to black, multi-locular. Peridium composed cells of textura angularis, 4–10 layers, thick-celled, dark-brown to black cells at outer layers, thin-walled and hyaline cells at inner layers. Conidiophores reduced to conidiogenous cells. Conidiogenous cells hyaline, holoblastic, smooth, cylindrical. Conidia 18–25 × 9–13 μm (\( \bar{x} \) = 20 × 11 μm, n = 20) ellipsoidal to subcylindrical, with rounded ends, initially light brown, slightly granulated walls, becoming dark brown (black in mass), aseptate. Spermatophores reduced to Spermatogenous cells, occurring intermingled among conidiogenous cells in same conidioma, subcylindrical, hyaline, smooth. Spermatia 4–7 × 1–3 μm (\( \bar{x} \) = 6 × 2 μm, n = 20) subcylindrical, straight or slightly curved, hyaline, smooth-walled, granular.

Material examined: China, Yunnan Province, Kunming, Kunming Institute of Botany, Botanical Garden, on dead stems, 24 May 2018, A.H. Ekanayaka (HKAS 102118).

GenBank number: ITS: MW136694, LSU: MW142386, SSU: MW127178

Notes: This collection is morphologically and phylogenetically similar to the type specimen of Aplosporella javeedii from South Africa (Jami et al. 2014). Comparison of ITS and LSU sequence data of this collection with sequence data of A. javeedii (CMW38165) has 99% similarity. However, spermatia were found in HKAS 102118.


Other genus included

Alanomyces Roh. Sharma, Phytotaxa 297(2): 170 (2017).

Index Fungorum number: IF 804153; Facesoffungi number: FoF 07623; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Alanomyces indica Roh. Sharma, Phytotaxa 297(2): 172 (2017).

Notes: Alanomyces, introduced by Sharma et al. (2017), is characterised by rapidly spreading dark greenish black colonies. Pycnidia are multilocular with a long neck and cylindrical, hyaline, guttulate, single celled spermatia (Sharma et al. 2017).


Economic and ecological significance

Some taxa in Aplosporellaceae are recorded as plant pathogens. Aplosporella prunicola was recorded as a probable tree pathogen on gymnosperms (Aylward et al. 2017). Aplosporella beaumontiana was recorded as a major pathogen associated with a disease of okra (Yan et al. 2018).


Botryosphaeriaceae Theiss. & Syd. [as ‘Botryosphaeriacae’], Annls mycol. 16(1/2): 16 (1918).

Index Fungorum number: IF 80530; Facesoffungi number: FoF 00116, 279 species.

Endophytic, pathogenic and saprobic on a wide range of hosts worldwide. Sexual morph: Ascomata uniloculate, solitary or clustered, becoming fully or partially erumpent with maturity, wall multi-layered, dark brown, infrequently embedded in stromatic tissues. Asci 8-spored, bitunicate, fissitunicate, with well-developed ocular chamber, thick endotunica, short pedicellate, clavate. Pseudoparaphyses cellular, intermixed with asci, hyaline, septate, frequently constricted at septa, hyphae-like, branched or not, deliquescing with maturity. Ascospores 2–3-seriate, fusoid to ellipsoid or ovoid, hyaline or pigmented, smooth to verruculose, septate or not, mucoid sheath mostly absent (photoplates of sexual characters can be seen in Phillips et al. 2013, 2019). Asexual morph: Coelomycetous. Conidiomata uni or rarely multiloculate, pycnidial, infrequently embedded in stromatic tissue. Conidiophores mostly reduced to conidiogenous cells. Conidiogenous cells hyaline, phialidic, proliferating percurrently or internally giving rise to periclinal thickenings, or proliferating percurrently forming annelations, without collarettes. Conidia hyaline or pigmented, aseptate or one or multi-septate, sometimes muriform, smooth or striate, thin- or thick-walled, without mucoid sheaths or appendages. Spermatogonia similar to conidiomata in anatomy. Spermatogenous cells ampulliform to lageniform or subcylindrical, hyaline smooth, phialidic. Spermatia developing in conidiomata or spermatogonia, hyaline, smooth, granular, subcylindrical or dumbbell-shaped, with rounded ends. (Liu et al. 2012; Slippers et al. 2013; Phillips et al. 2013).

Type: Botryosphaeria Ces. & De Not.

Notes: Botryosphaeriaceae was introduced by Theissen and Sydow (1918) for three genera, Botryosphaeria, Phaeobotryon and Dibotryon. Over decades of taxonomic revisions and updates based on morphology, the family became increasingly complex. However, based on LSU sequence data Crous et al. (2006a) revealed ten lineages within Botryosphaeriaceae, which they considered to represent individual genera. Phillips et al. (2008) defined, introduced and reinstated a further five genera in Botryosphaeriaceae. Aplosporella was shown to reside in Botryosphaeriaceae by Damm et al. (2007). Endomelanconiopsis was assigned to this family by Rojas et al. (2008) while Phillips and Alves (2009) considered that Melanopsis is another genus in Botryosphaeriaceae. Two new genera (Botryobambusa and Cophinforma) were introduced by Liu et al. (2012). Phillips et al. (2013) resolved Botryosphaeriaceae into 17 genera and 110 species known from culture. Thereafter six new genera and 85 new species/species combinations have been introduced (Dissanayake et al. 2016). According to a recent study by Phillips et al. (2019) currently 22 genera are accepted in Botryosphaeriaceae. In earlier studies Dothiorella and Spencermartinsia were regarded as two separate genera in the family (Phillips et al. 2008, 2013). However, Yang et al. (2017) synonymized Spencermartinsia under Dothiorella. These two genera are morphologically distinguishable only in sexual morphs by the hyaline apiculi in Spencermartinsia. So far, these characters are reported only in S. viticola but no other species in this genus have been observed with asexual morphs. Furthermore, phylogenetic distance is also small between these two genera. Therefore, in this paper we accept that Spencermartinsia is a synonym of Dothiorella. Phyllostictaceae was re-instated for Phyllosticta (Wikee et al. 2013a). Another three families were introduced in Botryosphaeriales by Slippers et al. (2013), namely Saccharataceae for Saccharata, Aplosporellaceae for Aplosporella and Melanopsaceae for Melanops. Wyka and Broders (2016) introduced Septorioideaceae for Septorioides while Yang et al. (2017) raised Endomelanconiopsis and Pseudofusicoccum to familial status as Endomelanconiopsidaceae and Pseudofusicoccaceae respectively. Taking into account morphology, phylogeny and evolutionary divergence times, Phillips et al. (2019) synonymised Endomelanconiopsidaceae with Botryosphaeriaceae, Pseudofusicoccaceae with Phyllostictaceae, and Septorioideaceae with Saccharataceae. In addition to that, Pilgeriella which fit within Botryosphaeriaceae, but do not have cultures or DNA (Wijayawardene et al. 2017a). Therefore, we place this genus incertae sedis in Botryosphaeriales.


Botryosphaeria Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 211 (1863).

Index Fungorum number: IF 635; Facesoffungi number: FoF 00141; 283 morphological species (Index Fungorum January, 2020), 13 species with molecular data.

Type species: Botryosphaeria dothidea (Moug.) Ces. & De Not.

Notes: Botryosphaeria was introduced by Cesati and De Notaris (1863). Species in Botryosphaeria, based on the type species B. dothidea, are characterised by ascospores that are hyaline and aseptate, although they can become pale brown and septate with age (Shoemaker 1964; Sivanesan 1984; Denman et al. 2000; Alves et al. 2004; Phillips et al. 2005; Jayawardena et al. 2018).


Other genera included

Alanphillipsia Crous & M.J. Wingf., in Crous et al., Persoonia 31: 197 (2013).

Index Fungorum number: IF 805816; Facesoffungi number: FoF 01417; – 5 morphological species (Species Fungorum 2020), 5 species with molecular data.

Type species: Alanphillipsia aloes Crous & M.J. Wingf., in Crous et al. Persoonia 31: 197 (2013).

Notes: Alanphillipsia was introduced by Crous et al. (2013) to accommodate species that are aplosporella-like in morphology, but have conidia with a hyaline outer layer.


Barriopsis A.J.L. Phillips, A. Alves & Crous (2008).

Index Fungorum number: IF 511712; Facesoffungi number: FoF 01679; – 6 morphological species (Species Fungorum 2020), 6 species with molecular data.

Type species: Barriopsis stevensiana A.J.L. Phillips & Pennycook, in Wijayawardene et al., Fungal Diversity 86: 56 (2017).

Based on Barriopsis fusca A.J.L. Phillips, A. Alves & Crous, in Phillips, Alves, Pennycook, Johnston, Ramaley, Akulov & Crous, Persoonia 21: 39 (2008).

Notes: Barriopsis is characterised by aseptate, brown ascospores without apiculi (Phillips et al. 2008). Absence of apiculi on the ascospores differentiates this genus from Sphaeropsis.


Botryobambusa Phook., Jian K. Liu & K.D. Hyde, in Liu et al., Fungal Diversity 57(1): 166 (2012).

Index Fungorum number: IF 801313; Facesoffungi number: FoF 02408; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Botryobambusa fusicoccum Phook., Jian K. Liu & K.D. Hyde, in Liu et al., Fungal Diversity 57(1): 166 (2012).

Notes: Botryobambusa was introduced by Liu et al. (2012) with a single species with both sexual and asexual morphs. Phylogenetically this genus is clearly distinguished from Botryosphaeria. However, it is similar to Botryosphaeria except for smaller asci and ascospores that are surrounded by a mucilagenous sheath.


Cophinforma Doilom, Jian K. Liu & K.D. Hyde, in Liu et al., Fungal Diversity 57(1): 174 (2012).

Index Fungorum number: IF 801315; Facesoffungi number: FoF 07624; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Cophinforma eucalypti Doilom, Jian K. Liu & K.D. Hyde, in Liu et al., Fungal Diversity 57(1): 174 (2012).

Notes: Cophinforma was introduced by Liu et al. (2012) as a monotypic genus for C. eucalypti. Phillips et al. (2013), based on a multi-gene analysis, introduced two species to this genus that were previously included in Botryosphaeria. Cophinforma is phylogenetically distinguished from Botryosphaeria despite being similar, except for conidia that are longer than any known species in Botryosphaeria.


Diplodia Fr., in Montagne, Annls Sci. Nat., Bot., sér. 2 1: 302 (1834).

Index Fungorum number: IF 8047; Facesoffungi number: FoF 00147; – 397 morphological species (Species Fungorum 2020), 30 species with molecular data (Jayawardena et al. 2019b, c).

Type species: Diplodia mutila (Fr. : Fr.) Fr., Summa Veg. Scand. 2: 417. 1849.

Sphaeria mutila Fr. : Fr., Syst. Mycol. 2: 424. 1823.

Physalospora mutila (Fr. : Fr.) N.E. Stevens, Mycologia 28: 333. 1936.

Botryosphaeria stevensii Shoemaker, Can. J. Bot. 42: 1299. 1964.

Notes: Diplodia comprises two morphological groups that are supported by two distinct phylogenetic lineages. In one type the conidia are initially hyaline and aseptate, later becoming brown and 1-septate. Pigmentation is often delayed and in some species dark conidia are never seen. In the second type the conidia become pigmented at an early stage, even while they are enclosed within the pycnidia, they only rarely become septate.


Dothiorella Sacc., Michelia 2(no. 6): 5 (1880).

Index Fungorum number: IF 8098; Facesoffungi number: FoF 00148; – 257 morphological species (Species Fungorum 2020), 44 species with molecular data.

Type species: Dothiorella pyrenophora Berk. ex Sacc., Michelia 2(no. 6): 5 (1880)

Notes: Dothiorella has often been confused with Diplodia in morphological characterisations. The type species, Dothiorella pyrenophora differs from Diplodia by conidia that are brown and 1-septate early in their development, while they are still attached to the conidiogenous cells. Sexual morphs of Dothiorella have pigmented, septate ascospores. However, the sexual stage of the species is rarely found in nature and has never been reported in culture.


Dothiorella ostryae Manawasinghe, Camporesi & K.D. Hyde, sp. nov.

Index Fungorum number IF556874, Facesoffungi number: FoF 06578; Fig. 16

Fig. 16
figure 16

Dothiorella ostryae (MFLU 18-0177, holotype). a Material examined. b Submerged conidiomata on host. c, d Cross section of conidiomata on host. e Developing conidia while attached to the pycnidial wall on host. f, g Conidia on host. h Conidiogenous cells in culture. 1 Upper view of colony on PDA. j reverse view of 7 days old culture on PDA. Scale bars: a, b = 2 mm, c–h = 10 µm

Etymology: Name reflects the host genus of Ostrya carpinifolia.

Holotype: MFLU 18-0177.

Saprobic on dead aerial branch of Ostrya carpinifolia. Sexual morph Undetermined. Asexual morph Conidiomata pycnidial, stromatic, solitary, dark brown, semi-immersed, unilocular, globose papillate with a short neck, wall 5–7 cell layers, outer layers composed of light-brown cells of textura angularis, inner layer hyaline and thin-walled Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, hyaline, cylindrical, proliferating percurrently. Conidia 15–21 × 7–10 (\( \bar{x} \) = 19 × 9 µm, n = 30) μm, ellipsoidal initially hyaline and aseptate, becoming pale brown while attached to the conidiogenous cells, finally dark brown or sepia, 1-septate, with 2 cells of equal length, thick-walled.

Culture characteristics: Colonies on PDA reach 70 mm after 5 d in the dark at 25 °C.

Material examined: Italy, Passo delle Forche - Galeata (province of Forlì-Cesena), dead aerial branch of Ostrya carpinifolia L. (Betulaceae), 9 April 2018, Erio IT Camporesi (MFLU 18-0177, holotype), ex-type living culture, JZB3150026.

GenBank number: ITS: MN533805, tef1 MN537429.

Notes: The species identified in this study is similar to other species in Dothiorella (Phillips et al. 2013). In phylogenetic analysis (Fig. 14), Dothiorella ostryae is closely related to the clade that contains Dothiorella citricola (ICMP 16828) and D. viticola (CBS 117009). There are four Dothiorella species associated with the host genus Ostrya, namely D. iberica, D. omnivora, D. parva (Farr and Rossman) and D. ostryae (this study).


Endomelanconiopsis Rojas & Samuels, Mycologia 100: 770. (2008).

Index Fungorum number: IF 555482; Facesoffungi number: FoF 07625; – 3 morphological species (Species Fungorum 2020), 3 species with molecular data.

Type species: Endomelanconiopsis endophytica Rojas & Samuels, Mycologia 100: 770 (2008).

Notes: This genus was introduced by Rojas et al. (2008) with two species. Tibpromma et al. (2018a) introduced a third species, Endomelanconiopsis freycinetiae, bringing the total to three species. Endomelanconiopsis is characterized by eustromatic conidiomata and holoblastically produced, brown, non-apiculate, unicellular conidia, each with a longitudinal germ slit (Rojas et al. 2008).


Eutiarosporella Crous, in Crous et al., Phytotaxa 202(2): 85 (2015).

Index Fungorum number: 811248; Facesoffungi number: FoF 07365; – 7 morphological species (Species Fungorum 2020), 7 species with molecular data.

Type species: Eutiarosporella tritici (B. Sutton & Marasas) Crous, in Crous et al., Phytotaxa 202(2): 85 (2015).

Tiarosporella tritici B. Sutton & Marasas, Trans. Br. mycol. Soc. 67(1): 74 (1976)

Notes: This genus was introduced by Crous et al. (2015a) to accommodate tiarosporella-like taxa with long conidiomatal necks and holoblastic conidiogenesis. Eutiarosporella differs from Marasasiomyces by developing clustered conidiomata.


Lasiodiplodia Ellis & Everh., Bot. Gaz. 21: 92 (1896).

Index Fungorum number: IF 8708; Facesoffungi number: FoF 00151; – 64 morphological species (Species Fungorum 2020), 35 species with molecular data.

Type species: Lasiodiplodia theobromae (Pat.) Griff. & Maubl. Bull. trimest. Soc.Mycol. Fr. 25: 57 (1909).

Botryodiplodia theobromae Pat., in Patouillard & Lagerheim, Bull. Soc. mycol. Fr. 8(3): 136 (1892).

Notes: Lasiodiplodia was introduced by Ellis in 1894 with L. tubericola as the type species. Ellis did not provide a description of the genus and species, but this was provided by Clendenin (1896) who attributed both the genus and species to Ellis and Everhart. Griffon and Maublanc (1909) considered that Botryodiplodia theobromae was more suitably accommodated in Lasiodiplodia. Since the epithet theobromae (1892) is older than tubericola (1896), L. theobromae should be regarded as the type species of Lasiodiplodia. In earlier works on Lasiodiplodia and Diplodia it was considered that Lasiodiplodia could represent a possible synonym of Diplodia (Denman et al. 2000). However, in phylogenetic analyses these two genera were clearly separated in two distinct clades (Zhou and Stanosz 2001; Slippers et al. 2004a, b; Phillips et al. 2008). Lasiodiplodia is distinguished from Diplodia by striations on the conidia. Conidiomatal paraphyses distinguishes Lasiodiplodia from Neodeightonia. Although Barriopsis has striate conidia, they are unique in Botryosphaeriaceae because they are also present on immature, hyaline conidia.


Macrophomina Petr., Annls mycol. 21(3/4): 314 (1923).

Index Fungorum number: IF 8814; Facesoffungi number: FoF 07626; – 4 morphological species (Species Fungorum 2020), 3 species with molecular data.

Type species: Macrophomina philippinensis Petr., Annls mycol. 21(3/4): 314 (1923).

Other synonyms can be seen in Index Fungorum.

Notes: Macrophomina comprises two species with cultures and moleular data. This genus is similar to Tiarosporella in having apical mucoid appendages (Sutton and Marasas 1976). Apart from the phylogeny, these two species can be distinguished, since Macrophomina has percurrently proliferating conidiogenous cells (Nag-Raj 1993; Crous et al. 2006a) and conidia that become dark brown at maturity, and microsclerotia (Phillips et al. 2013).


Marasasiomyces Crous, in Crous et al., Phytotaxa 202(2): 86 (2015).

Index Fungorum number: IF 811252; Facesoffungi number: FoF 07440; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Marasasiomyces karoo (B. Sutton & Marasas) Crous, Crous et al., Phytotaxa 202(2): 85 (2015).

Tiarosporella graminis var. karoo B. Sutton & Marasas, Trans. Br. mycol. Soc. 67(1): 73 (1976).

Notes: This monotypic genus was introduced by Crous et al. (2015a) for M. karooo to accommodate tiarosporella-like taxa. Marasasiomyces is distinguished from Tiarosporella by conidiomata with long necks covered in brown setae, and holoblastic conidiogenesis. These characters are similar to Eutiarosporella, but differ on account of the non-clustered conidiomata.


Mucoharknessia Crous, R.M. Sánchez & Bianchin., in Crous et al., Phytotaxa 202(2): 86 (2015).

Index Fungorum number: IF 811254; Facesoffungi number: FoF 01651; – 2 morphological species (Species Fungorum 2020), 2 species with with molecular data.

Type species: Mucoharknessia cortaderiae Crous, R.M. Sánchez & Bianchin Crous, in Crous et al., Phytotaxa 202(2): 85 (2015).

Notes: This genus was introduced by Crous et al. (2015a) for species resembling Harknessia. The genus is characterised by conidia bearing mucoid appendages. Mucoharknessia comprises two species.


Neodeightonia C. Booth, in Punithalingam, Mycol. Pap. 119: 17 (1970) [1969].

Index Fungorum number: IF 3450; Facesoffungi number: FoF 07627; – 8 morphological species (Species Fungorum 2020), 8 species with with molecular data.

Type species: Neodeightonia subglobosa C. Booth, Mycol. Pap. (1970).

Notes: Neodeightonia was regarded as a synonym of Botryosphaeria (von Arx and Müller 1975). However, it was reinstated by Phillips et al. (2008) based on morphological characters including the dark, 1-septate ascospores, and its phylogenetic placement. Longitudinal striations on the conidial wall are an additional characteristic feature of this genus (Crous et al. 2006a; Phillips et al. 2008). Neodeightonia can be differentiated from Lasiodiplodia by the absence of conidiomatal paraphyses, and conidial striations distinguish it from Diplodia.


Neofusicoccum Crous, Slippers & A.J.L. Phillips, in Crous et al., Stud. Mycol. 55: 247 (2006).

Index Fungorum number: IF 500870; Facesoffungi number: FoF 00153; – 45 morphological species (Species Fungorum 2020), 43 species with molecular data (Jayawardena et al. 2019a, c).

Type species: Neofusicoccum parvum (Pennycook & Samuels) Crous, Slippers & A.J.L. Phillips., in Crous et al., Stud. Mycol. 55: 247 (2006).

Fusicoccum parvum Pennycook & Samuels, Mycotaxon 24: 455 (1985).

Notes: Neofusicoccum was introduced by Crous et al. (2006a) to accommodate species that are similar to Botryosphaeria, but phylogenetically distinct. Dichomera synasexual morph in Neofusicoccum has been used to differentiate it from Botryosphaeria, even though not all Neofusicoccum species form such a synasexual morph (Phillips et al. 2005; Barber et al. 2005). Development of paraphyses can be observed in all accepted Botryosphaeria species, but have not been reported in conidiomata of any Neofusicoccum species. However, this character is difficult to use as developing conidiogenous cells can resemble paraphyses. Accepted species in this genus have been separated on the basis of conidial dimensions and pigmentation, pigment production in culture media and ITS sequence data (Phillips et al. 2013). However, morphological characterisation of these species remains controversial (Abdollahzadeh et al. 2013). To resolve the species, multi-gene sequence data are essential. Species in Neofusicoccum have been reported as endophytes, pathogens and saprobes on a wide range of hosts.


Neoscytalidium Crous & Slippers, Stud. Mycol. (2006).

Index Fungorum number: IF 500868; Facesoffungi number: FoF 07628; – 3 morphological species (Species Fungorum 2020), 7 species with molecular data.

Type species: Neoscytalidium dimidiatum (Penz.) Crous & Slippers, in Crous et al., Stud. Mycol. 55: 244 (2006).

≡ Torula dimidiata Penz., in Saccardo, Michelia 2(no. 8): 466 (1887).

Notes: In earlier studies, species belonging to this genus was introduced as Hendersonula toruloidea, which have conidia that become septate with a darker central cell and formed a Scytalidium-like synanamorph (Crous et al. 2006a). Since the description, the production of both arthroconidial and pycnidial synasexual morphs has been shown and led to several controversies in the nomenclature. However, the current genus name was introduced by Crous et al. (2006a) and other relevant epitypifications and synonymies by Madrida et al. (2009). Species in this genus have been reported as pathogens on human skin and nails (Campbell and Mulder 1977) and plants. Diseases reported associated with this genus tend to be more common in tropical regions (Polizzi et al. 2009, 2011; Phillips et al. 2013).


Oblongocollomyces Tao Yang & Crous., in Yang et al., Fungal Biology 121: 339 (2016).

Index Fungorum number: IF 817640; Facesoffungi number: FoF 08049; 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Oblongocollomyces variabilis (F.J.J. Van der Walt, Slippers & G.J. Marais) Tao Yang & Crous, in Yang et al., Fungal Biology 121: 339 (2016).

Sphaeropsis variabilis F.J.J. Van der Walt, Slippers & G.J. Marais, in Slippers et al., Persoonia 33: 164 (2014).

Notes: Oblongocollomyces was introduced by Yang et al. (2017) as a monotypic genus in Botryosphaeriaceae. In earlier studies, this species was identified as Sphaeropsis variabilis and it was different from Sphaeropsis in having long conidiomatal necks on PNA, aggregated in dense clusters of conidiomata on OA (Yang et al. 2017). This species has conidia with up to 3 septa. Phylogenetically Oblongocollomyces close to Alanphillipsia (Fig. 14).


Phaeobotryon Theiss. & Syd., Annls mycol. 13(5/6): 664 (1915).

Index Fungorum number: IF 3892; Facesoffungi number: FoF 07629; – 8 morphological species (Species Fungorum 2020), 6 species with molecular data.

Type species: Phaeobotryon cercidis (Cooke) Theiss. & Syd., Annls mycol. (1915).

Dothidea cercidis Cooke, Grevillea 13(no. 67): 66 (1885).

Notes: Phaeobotryon has 2-septate, brown ascospores with an apiculus at each end.


Sakireeta Subram. & K. Ramakr., J. Indian bot. Soc. 36: 83 (1957).

Index Fungorum number: IF 9775; Facesoffungi number: FoF 07583; – 1 morphological species (Wijayawardene et al. 2017a), 1 species with molecular data.

Type species: Sakireeta madreeya Subram. & K. Ramakr., J. Indian bot. Soc. 36: 84 (1957).

Notes: Sakireeta comprises a single species, S. madreeya, which was previously known as Tiarosporella madreeya. This genus is distinguished from the closely related Tiarosporella by conidiomata that are aggregated in a stroma, and being plurilocular whereas, Tiarosporella has solitary unilocular conidiomata. In addition, Sakireeta has holoblastic conidiogenous cells (Crous et al. 2015a).


Sardiniella Linald., A. Alves & A.J.L. Phillips, Mycosphere (2016).

Index Fungorum number: IF 817511; Facesoffungi number: FoF 02405; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Sardiniella urbana Linald., A. Alves & A.J.L. Phillips, Mycosphere (2016).

Notes: Sardiniella was introduced as a monotypic genus for Sardiniella urbana. This genus is characterised by thick-walled, oblong conidia that become pigmented and 1-septate, which distinguishes it from Neofusicoccum. This genus is similar to Diplodia and Dothiorella. However, in Dothiorella, and in some species of Diplodia the conidia become pigmented while still attached to the conidiogenous cell and this character is not seen in Sardiniella. The conidial wall in species of Diplodia is thicker than in Sardiniella. Phylogenetically these two genera are clearly separated.


Sphaeropsis Sacc., Michelia 2(no. 6): 105 (1880).

Index Fungorum number: IF 562041; Facesoffungi number: FoF 01704; – 142 morphological species (Species Fungorum 2020), 5 species with molecular data.

Type species: Sphaeropsis visci (Alb. & Schwein.) Sacc., Michelia 2(no. 6): 105 (1880).

Dothidea visci Kalchbr., Hedwigia 8: 117 (1869).

Notes: Sphaeropsis was introduced by Saccardo (1880) to accommodate species of Diplodia with brown, aseptate conidia. Since then species were described based on host association, resulting in more than 600 species names. Pycnidial paraphyses distinguish Sphaeropsis from Diplodia. The striate conidia of Lasiodiplodia differentiate it from Sphaeropsis, which has smooth-walled conidia.


Tiarosporella Höhn., in Weese, Ber. dt. bot. Ges. (1919).

Index Fungorum number: IF 10233; Facesoffungi number: FoF 00333; – 8 morphological species (Species Fungorum 2020), 5 species with molecular data.

Type species: Tiarosporella paludosa (Sacc. & Fiori) Höhn., in Weese, Ber. dt. bot. Ges. 37: 159 (1919).

Neottiospora paludosa Sacc. & Fiori, in Sydow, Hedwigia 38(Beibl.): (137) (1899).

Notes: This genus is characterised by conidia formed from smooth, hyaline conidiogenous cells that lack periclinal thickenings and percurrent proliferations. The conidia are hyaline, subcylindrical to fusiform with irregular mucoid appendages. There are 22 species names for this genus in Index Fungorum.


Economic and ecological significance

Botryosphaeriaceae comprises a diverse range of taxa that are pathogens, endophytes or saprobes on a wide range of hosts including dicotyledonous monocotyledonous, and gymnosperms (Schoeneweiss 1981; Manawasinghe et al. 2016). Their distribution is vast, covering all geographic and climatic regions, except Polar Regions (Crous et al. 2007a; Rodas et al. 2009; Wunderlich et al. 2011; Manawasinghe et al. 2016). At present, their ecological role as phytopathogens has gained a great attention (Manawasinghe et al. 2016). The pathogenic taxa in Botryosphaeriaceae cause several important diseases such as leaf spots, cankers, dieback, fruit rot, gummosis and even plant death on many economically important crops (Rodas et al. 2009, Hyde et al. 2014). Some species are pathogenic on many hosts (Diplodia seriata, Lasiodiplodia spp., and Neofusicoccum parvum) in the same geographic area. Pathogenicity of some taxa (e.g. B. dothidea) varies with the country for the same host species (Chen et al. 2014; Netto et al. 2014; Linaldeddu et al. 2015). The pathogenic fungal taxa of this family are recognized as “opportunistic plant fungal pathogens” (Chethana et al. 2016). It is not clear whether changes in the environment triggers pathogenicity or if disease development is due to weakening of the host defenses (Chethana et al. 2016; Manawasinghe et al. 2016). Since phytopathogenic botryosphaeriaceous taxa also have an endophytic lifestyle they may act as latent pathogens. Some species in this family have been reported as human pathogens (Polizzi et al. 2009, 2011; Phillips et al. 2013).


Melanopsaceae A.J.L. Phillips, Slippers, Boissin & Crous, Stud. Mycol. 76(1): 43 (2013).

Index Fungorum number: IF 805796; Facesoffungi numbers: FoF 07630, 111 species.

On woody hosts. Sexual morph: Ascostromata pseudothecial, multiloculate, locules at various levels in ascoma, immersed, partially erumpent at maturity, black, subglobose, thick-walled. Ascomatal wall thick-walled, comprised of cells of textura angularis. Asci 8-spored, bitunicate, fissitunicate, pedicellate, clavate. Hamathecium comprising hyaline, thin-walled, hypha-like, septate, not constricted at septa, cellular pseudoparaphyses. Ascospores overlapping, ellipsoid to rhomboid, hyaline, aseptate, thin-walled, with a persistent mucus sheath. Asexual morph: Coelomycetous. Conidiomata often formed in the same stroma as ascostromata. Paraphyses septate, branched or not, filiform, hyaline, arising from between the conidiogenous cells. Conidiophores 1–2-septate, branched or not, hyaline, smooth, or reduced to conidiogenous cells. Conidiogenous cells subcylindrical, branched or unbranched, discrete, hyaline, formed from the inner wall of the conidioma, proliferating percurrently at apex, or with periclinal thickening. Conidia fusoid, hyaline, aseptate, with a persistent mucus sheath, rarely with minute marginal frill (adapted from Slippers et al. 2013; Phillips et al. 2019)

Type: Melanops Nitschke ex Fuckel.

Notes: Slippers et al. (2013) established Melanopsaceae in Botryosphaeriales to accommodate Melanops. Melanops has a mucilage sheath around the ascospores and in that respect is identical to Phyllostictaceae. However, Phyllostictaceae has uniloculate ascostromata, while those of Melanops are multiloculate (Slippers et al. 2013). Phylogenetic analyses supported the placement of Melanops as a distinct family in Botryosphaeriales (Wikee et al. 2013b; Slippers et al. 2013; Phillips et al. 2019). Divergence time estimates for this family also support its familial status (Slippers et al. 2013; Phillips et al. 2019). However, Jiang et al. (2018b) suggested that the divergence time estimation in the previous studies might not be accurate estimates due to the fact that sequence data from only two taxa were used in the phylogenetic placement, an opinion also voiced by Phillips et al. (2019).


Melanops Nitschke ex Fuckel, Jb. nassau. Ver. Naturk. 23–24: 225 (1870) [1869–70].

Index Fungorum number: IF 3078; Facesoffungi number: FoF 07442; 111 morphological species (Species Fungorum 2020), 3 species with molecualar data.

Type species: Melanops tulasnei Fuckel.

Notes: Over 100 species names are listed under Melanops in Index Fungorum (2020), but, only three species have sequence data (Jiang et al. 2018b). We were unable to obtain fresh collections to represent this genus. Thus, a drawing is provided. A description of the type species can be seen in Phillips and Alves (2009).


Melanops tulasnei Fuckel, Jb. nassau. Ver. Naturk. 23–24: 225 (1870) [1869–70].

Index Fungorum number: IF 150956; Facesoffungi number: FoF 07444; Fig. 17

Fig. 17
figure 17

Melanops tulasnei (redrawn from neotype and epitype specimens in Phillips and Alves 2009). a Stroma erumpent through the bark. b Stroma cut through horizontally revealing ascomata and conidiomata. c Ascus. d Ascus tip. e Ascospores. f Conidium and conidiogenous cell with mucus sheath around the conidium. g Conidium with mucous sheath. h Conidia. i Conidiogenous with conidia developing on conidiogenous cells. j Paraphyses. Scale bars: b = 250 μm, c = 20 μm, d, e, g–j = 10 μm, f = 5 μm

Description: see Phillips and Alves (2009).


Economic and ecological significance

The ecology and distribution of Melanops is poorly known. Although its appearance is similar to other Botryosphaeriales, it is not clear whether it is pathogenic or endophytic (Slippers et al. 2013).


Phyllostictaceae Fr. (as ‘‘Phyllostictei’’), Summa vegetabilium Scandinaviae 2: 420 (1849).

Index Fungorum number: IF 81162; Facesoffungi numbers: FoF 02296, >2000 species.

Foliicolous, pathogenic, endophytic or saprobic on plants. Sexual morph: Ascostromata pseudothecial, uniloculate, separate to gregarious, globose, brown to black, with a central ostiole. Asci 8-spored, bitunicate, fissitunicate, clavate to subcylindrical, fasciculate, pedicellate, with an ocular chamber. Pseudoparaphyses mostly absent at maturity, when present broad-cylindric, septate, cellular. Ascospores 2–3-seriate, ellipsoid-fusoid to limoniform, hyaline, aseptate, smooth-walled, usually with mucilaginous caps, or surrounded by a mucilaginous sheath. Asexual morph: Coelomycetous. Conidiomata pycnidial globose, dark brown, separate to aggregated, with a central ostiole; wall of 3–6 layers of brown cells of textura angularis. Conidiogenous cells lining the inner wall, hyaline, smooth, subcylindrical to ampulliform or doliiform, proliferating percurrently near apex, frequently covered in mucilaginous sheath. Conidia ellipsoid-fusoid to obovoid or ovoid, hyaline, aseptate, smooth-walled, guttulate or granular, frequently surrounded by a mucilaginous sheath, often bearing a single mucilaginous apical appendage.

Type: Phyllosticta Pers.

Notes: Phyllostictaceae (as Phyllostictei) was proposed by Fries (1849). Seaver (1922) used Phyllostictales and Phyllostictaceae for Phyllosticta. Phyllostictaceae was accepted by Hawksworth and David (1989). Schoch et al. (2006) suggested that Phyllosticta belongs to Botryosphaeriaceae and this was accepted by Crous et al. (2006a) and Liu et al. (2012) who noted that Phyllosticta is distinct from other genera in Botryosphaeriaceae. Wikee et al. (2013b) suggested Phyllostictaceae as a distinct family in Botryosphaeriales and Slippers et al. (2013) formally re-instated the family.

Ascospores in Melanopsaceae and Phyllostictaceae have a mucilage sheath. Phyllostictaceae differs from Melanopsaceae by uniloculate ascostromata. Though mucoid apical appendages on the conidia have been regarded as characteristic of Phyllosticta, this feature is not necessarily a characteristic of the family since they are absent in some species such as P. leucothoicola or P. neopyrolae (Wikee et al. 2013b).


Phyllosticta Pers., Traite´sur les Champignons Comestibles (Paris): 147 (1818).

Index Fungorum number: IF 9384; Facesoffungi number: FoF 00155; >2000 records (Species Fungorum 2020), several species with molecular data.

Type species: Phyllosticta cruenta (Kunze: Fr.) J. Kickx f.

Sphaeria cruenta Fr., Syst. mycol. (Lundae) 2(2): 581 (1823).

Notes: Phyllosticta is a geographically widespread genus of plant pathogenic fungi with a diverse host range and a Guignardia sexual morph (Fig. 18). Seaver (1922) used Phyllostictales and Phyllostictaceae for Phyllosticta. Phyllosticta was placed in Botryosphaeriales by Schoch et al. (2006), who proposed that Botryosphaeriaceae contained both Botryosphaeria and Phyllosticta, although no support was obtained for this relationship. Crous et al. (2006a) and Liu et al. (2012) also classified Phyllosticta in Botryosphaeriaceae. In both studies it was noted that Phyllosticta was distinct from other genera in Botryosphaeriaceae, and that these authors eventually expected it to be placed elsewhere. Wikee et al. (2013b) redefined Phyllosticta, and showed that it clusters sister to Botryosphaeriaceae, for which the older family name Phyllostictaceae was resurrected. In moving away from a dual nomenclature for fungi, the generic name Phyllosticta was chosen over Guignardia in previous studies. There are 3214 Phyllosticta epithets listed in Index Fungorum. In this entry we illustrate Phyllosticta hostae (Fig. 19).

Fig. 18
figure 18

Sexual morph of Phyllosticta ellipsoidea (redrawn from Wulandari et al. 2011, MFLU 10-0475). a Section of ascoma. b Asci. c Ascospores with polar mucilaginous appendage at each end. Scale bars: 25 µm


Phyllosticta hostae Y.Y. Su & L. Cai, Persoonia 28: 79 (2012).

Index Fungorum number: IF 564904; Facesoffungi number: FoF 07380; Fig. 19

Fig. 19
figure 19

Phyllosticta hostae (JZB 3290001). a Leaf spot on rose leaf. b Front view of the colony on PDA. c Rear view of the colony on PDA. d Conidiomata in culture. e–h Conidia. Scale bars: d = 100 µm, e–h = 10 µm

Leaf spots circular to somewhat irregularly rotundate, pale brown to brown. Sexual morph: Undetermined. Asexual morph: Colonies on PDA, Pycnidia black, sub epidermal, globose, 40–150 µm in diam. dark brown to black, darker around ostiole. Conidiogenous cells holoblastic, phialidic, cylindrical, subcylindrical to ampulliform, hyaline, thin-walled, smooth. Conidia 8–15 × 5–9 µm (\( \bar{x} \) = 10 × 7, n = 20), unicellular, thin- and smooth-walled, ellipsoid, subglobose to obovoid, with large central guttule, truncate at the base when young, later rounded at both ends, enclosed in a 1–3 µm thick mucilaginous sheath, and bearing a hyaline, mucoid apical appendage, 4–8 × 1–3 µm, straight to flexuous, unbranched, tapering towards an acute tip.

Material examined: China, Beijing Tongzhou-Yongle Eco Park, leaf spot on rose (Rosaceae) leaves, 18 August 2018, Jia Jing Yi, 28-10-1, living culture: JZB 3290001.

GenBank number: ITS: MW130255.

Notes: Phyllosticta is an important genus of plant pathogens and contains quarantinable pathogenic species in some countries, especially in Europe (Wulandari et al. 2013). Phyllosticta hostae was introduced by Su and Cai (2012) from Hosta plantaginia (Liliaceae). This is the first record of P. hostae causing leaf spots in rose in China.


Other genera included

Pseudofusicoccum Mohali, Slippers & M.J. Wingf., Stud. Mycol. 55: 249 (2006).

Index Fungorum number: IF 555584; Facesoffungi number: FoF 05299; – 8 morphological species (Species Fungorum 2020), 7 species with molecular data.

Type species: Pseudofusicoccum stromaticum (Mohali, Slippers & M.J. Wingf.) Mohali, Slippers & M.J. Wingf., in Crous et al., Stud. Mycol. 55: 249 (2006).

Fusicoccum stromaticum Mohali, Slippers & M.J. Wingf., Mycol. Res. 110(4): 408 (2006).

Notes: The type species of Pseudofusicoccum, P. stromaticum was introduced as Fusicoccum stromaticum and was distinguished from other species by its large conidiomata, the ability to grow at 35°C, and thick-walled conidia. Crous et al. (2006a) recognized that its conidia were covered by a mucous sheath, which is lacking in all species of Fusicoccium. Therefore, they introduced Pseudofusicoccum as a new genus to accommodate this taxon. Pseudofusicoccum adansoniae, P. ardesiacum and P. kimberleyense were proposed and distinguished from others by production of a pigment in culture and the size of conidia (Pavlic et al. 2008). Later, P. olivaceum was identified as a new species because of the discrepancy between ITS and tef1 BLAST results (Mehl et al. 2011). Pseudofusicoccum artocarpi was introduced as its conidia were clearly longer than others (Trakunyingcharoen et al. 2015). Pseudofusicoccum violaceum is distinctive because of the violet/purple pigment formed in culture and this has not been observed in any other Pseudofusicoccum species (Mehl et al. 2011). Pseudofusicoccum africanum was introduced as it is different from its neighbor P. violaceum by unique fixed alleles in ITS and tef1 (Jami et al. 2018). Yang et al. (2017) raised the genus to familial status as Pseudofusicoccaceae. The phylogenies reported in Phillips et al. (2019), as well as those of Minnis et al. (2012) and Liu et al. (2017), place Pseudofusicoccum in the same clade as Phyllosticta species. In view of this consistent relationship, together with the evolutionary divergence evidence as well as morphological data presented in Phillips et al. (2019), Pseudofusicoccum is considered to be an additional genus within Phyllostictaceae (Fig. 20).

Fig. 20
figure 20

Morphology of Pseudofusicoccum. a, b Conidiophores and conidia of P. stromaticum. c–e Conidiomatal wall, conidiophore and conidia of P. africanum. f, g Conidia becoming brown and septate when mature and young conidia of P. artocarpi. h–j Conidiogenous cells, germinating conidia and conidia of P. adansoniae Scale bars: a = 5 μm, b, c, e, f–j = 10 μm, d = 25 μm


Economic and ecological significance

Phyllosticta species have globally been recorded as endophytes, plant pathogens and saprobes from economically and ecologically important plant hosts. Some cause important diseases such as Citrus black spot and tan spot, subjected to phyto-sanitary legislation in the European Union and the USA. Further leaf- and fruit-spotting disease of Musa spp. (freckle disease), leaf spots in turmeric, cashew, ginger, orchids and black rot of grapes diseases are also caused by Phyllosticta species.


Planistromellaceae M.E. Barr, Mycotaxon 60: 437 (1996).

Index Fungorum: IF 81919; Facesoffungi number: 06689, 33 species.

Biotrophic, hemibiotrophic or saprobic on leaves and stems of various plants in terrestrial habitats. Sexual morph: Ascostromata multi- or uni-loculate, immersed to erumpent through cracking or splitting of the host tissue, solitary to gregarious, with periphysate ostioles, with or without papilla. Cells of ascostromata thick-walled, composed of several layers of dark brown cells, arranged in a textura angularis. Locules ovoid to globose, developing in the same stroma of the conidiogenous and/or spermatogenous locules, collapsed with the empty locule, which previously produced conidia or spermatia or both, ostiole periphysate. Peridium of locules composed of a few layers of hyaline to light brown flattened cells. Hamathecium lacking pseudoparaphyses, interascal cells abundant even at maturity. Asci 8-spored, bitunicate, fissitunicate, oblong, clavate to nearly cylindrical, with a pedicel and with an ocular chamber, forming in a basal layer, often interspersed with and covered by cellular remnants of interthecial tissues. Ascospores overlapping 1–3-seriate, ellipsoid to broadly obovoid, hyaline or lightly pigmented, yellowish to brownish, aseptate or 1–2-trans-septate, thin-walled, with or without gelatinous sheath, guttulate. Asexual morph: Coelomycetous. Conidiomata subepidermal, dark, immersed to erumpent, solitary to gregarious, pycnidia, locules or acervuli in a stroma or bearing conidia over stroma surface prior to locule formation, ostiolate. Conidiomata walls comprising several layers with cells of textura angularis, the outer layers composed of dark thick-walled cells, lighter towards the inner layers of hyaline cells. Conidiogenous cells short cylindric, conidiogenesis holoblastic, hyaline, smooth. Conidia oblong, ellipsoid-cylindric, hyaline to brown, aseptate or 1-multi-trans-septate, smooth-walled or verruculose, with or without one or more apical appendages. Spermatial state developing in the same or separate locules. Spermatogenous cells discrete or integrated, phialidic, cylindric to elongate-conical, determinate, hyaline, smooth. Spermatia bacillary, hyaline, aseptate, smooth-walled.

Type: Kellermania Ellis & Everh.

Notes: Planistromellaceae was introduced by Barr (1996) and belongs to Botryosphaeriales (Minnis et al. 2012; Monkai et al. 2013). Lumbsch and Huhndorf (2010) included the genera Comminutispora, Eruptio, Loratospora., Microcyclus, Mycosphaerellopsis, Planistroma and Planistromella in Planistromellaceae based on morphology. Phylogenetic analysis based on SSU, ITS and LSU sequence data by Minnis et al. (2012) showed that Kellermania, Piptarthron, Planistroma and Planistromella should be treated as a single genus, thus, they synonymized them under Kellermania (oldest name). However, Monkai et al. (2013) indicated that the type species of Kellermania, which is also the type of Planistromella, clustered separately from Planistroma based on LSU and ITS sequence data. Planistroma was accepted as a distinct genus. The family was revised to accommodate Kellermania, Planistroma Mycosphaerellopsis and Umthunziomyces based on morphology and molecular data (Monkai et al. 2013; Phillips et al. 2019).


Kellermania Ellis & Everh., J. Mycol. 1(12):153 (1885).

Index Fungorum number: IF 8668; Facesoffungi number: FoF 06690; 23 morphological species (Species Fungorum 2020), 18 species with molecular data.

Type species: Kellermania yuccifoliorum A.W. Ramaley.

Notes: Species in this genus are known from the host genera Agave and Nolina (Asparagaceae). Five species have been reported with both sexual and asexual morphs in Ramaley (1993, 1995, 1998) and Barr (1996). Many species of Kellermania were illustrated by Minnis et al. (2012) and conidia range from 0 to several trans-septa, and with or without appendages.


Kellermania yuccifoliorum A.W. Ramaley, Mycotaxon 47: 262 (1993).

Index Fungorum: IF 360150; Facesoffungi number: FoF 06691; Figs. 21 and 22

Fig. 21
figure 21

Kellermania yuccifoliorum (UC 1202973, holotype of P. yuccifoliorum). a, b Ascostromata on the host surface. c, d Section of ascostroma. e, f Ascoma. g Immature ascus. h Mature asci. i Cells of ascostroma. j Immature ascospore. k, l Mature ascospores. Scale bars: a = 1 cm, b = 1000 µm, c = 200 µm, d = 100 µm, e, f = 50 µm, g, h = 30 µm, i = 20 µm, jl = 10 µm

Fig. 22
figure 22

Kellermania yuccifoliorum (redrawn from Ramaley 1993). a Conidioma. b Microconidia. c Spermatia and spermatiogenesis. d Conidiogenesis and origin appearance of apical appendage. e Conidia. Scale bars: a = 80 µm, b, d = 27 µm, e = 40 µm

Description: see Monkai et al. (2013).

Material examined: USA, Califonia, San Bernardino County, Roadside 20 miles east of Baker (Hwy. 91/466), on leaves of Yucca brevifolia Engelmann, 14 April 1960, Isabelle Tavares No. 466 (UC 1202973, holotype of Planistromella yuccifoliorum).

Notes: The type species of Kellermania (K. yuccifoliorumPlanistromella yuccifoliorum) differs from other genera in Planistromellaceae in having 1–2-septate ascospores (Ramaley 1993; Barr 1996). The asexual morph of K. yuccifoliorum is characterized by uniloculate conidiomata, which develops in the same stroma as the ascogenous locules and 2-septate conidia with a unique apical appendage (Ramaley 1993).


Other genera included

Mycosphaerellopsis Höhn., Annal. Mycol. 16(1/2): 157 (1918).

Index Fungorum number: IF 3345; Facesoffungi number: FoF 06263; – 2 morphological species (Monkai et al. 2013), molecular data unavailable.

Type species: Mycosphaerellopsis myricariae (Fuckel) Höhn., Annal. Mycol. 16(1/2):157 (1918).

≡ Sphaeria myricariae Fuckel, Jb. nassau. Ver. Naturk. 27–28: 22 (1874) [1873–74].

Notes: Mycosphaerellopsis differs from other genera in Planistromellaceae in having uniloculate ascomata and 1-septate, broadly obovoid ascospores. We accept Mycosphaerellopsis in this family based on its morphology (Monkai et al. 2013).


Planistroma A.W. Ramaley, Mycotaxon 42: 69 (1991).

Index Fungorum number: IF 25358; Facesoffungi number: FoF 06264; – 7 morphological species (Monkai et al. 2013), 6 species with molecular data.

Type species: Planistroma yuccigenum A.W. Ramaley, Mycotaxon 42: 69 (1991), MycoBank: MB 358836.

Notes: Planistroma was established by Ramaley (1991), and is similar to Kellermania in ascostromata characters, but Planistroma differs on account of its aseptate ascospores (Barr 1996). The asexual morphs of Planistroma resemble Kellermania except in having apically appendaged conidia in the latter genus (Ramaley 1995). Minnis et al. (2012) synonymized Planistroma under Kellermania based on molecular data support. However, we follow Monkai et al. (2013) who assigned Planistroma as a distinct genus based on both morphological and molecular data.


Umthunziomyces Crous & M.J. Wingf., in Crous et al., Persoonia 37: 315 (2016).

Index Fungorum number: IF 819069; Facesoffungi number: FoF 06718; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Umthunziomyces hagahagensis Crous & M.J. Wingf., Persoonia 37: 315 (2016).

Notes: The monotypic genus Umthunziomyces was introduced by Crous et al. (2016a) and is known only from its asexual morphs. Umthunziomyces is similar to Kellermania in having septate conidia, but they are longer and narrower than in the latter genus. Phillips et al. (2019) included Umthunziomyces in Planistromellaceae with evidence from the phylogenetic analyses of ITS and LSU sequence data.


Economic and ecological significance

Members in this family usually grow on living or dead leaves or on stems of various plants, and are mostly saprobes, but some species are pathogens such as Kellermania agaves, and Mycosphaerellopsis moravica (Barr 1996; Ramaley 1993, 1995, 1998; Minnis et al. 2012; Crous et al. 2013). Most species inhabit Asparagaceae.


Saccharataceae Slippers, Boissin & Crous, Studies in Mycology 76: 41 (2013).

Index Fungorum number: IF 805794; Facesoffungi numbers: FoF 02296, 22 species.

Saprobic, endophytic, or pathogenic on plants. Sexual morph: Ascostromata pseudothecial, uniloculate, solitary or in clusters, with multi-layered, dark brown walls, infrequently embedded in stromatic tissue, with upper ascomatal layer darkened and thickened, developing under a very small epidermal clypeus, ascomatal wall continuous with clypeus. Asci 8-spored, bitunicate, fissitunicate, with a thick endotunica, pedicellate or sessile, cylindrical, with a well-developed ocular chamber. Hamathecium comprising 3–5 µm wide, filiform, septate, hypha-like, branched or not, abundant, intermixed with asci, cellular pseudoparaphyses. Ascospores 2–3-seriate, ellipsoidal, hyaline to pigmented, pale golden brown, granular, septate or not, without mucoid appendages or sheath. Asexual morph: Coelomycetous. Conidiomata unilocular pycnidial, infrequently embedded in stromatic tissue with thickened, darkened upper layer. Conidiophores sparingly branched, hyaline, subcylindrical, or reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, phialidic, proliferating via periclinal thickening or percurrent proliferation, with or without collarettes. Conidia fusoid, hyaline, aseptate, thin-walled, granular. Synasexual morph formed in separate conidiomata, or in same conidiomata with asexual morph. Synasexual morph: Conidia ellipsoid or oval, pigmented, aseptate, thick-walled, finely verruculose. Spermatogonia similar to conidiomata in anatomy. Spermatogenous cells ampulliform to lageniform or subcylindrical, hyaline smooth, phialidic. Spermatia developing in conidiomata or spermatogonia, subcylindrical or dumbbell-shaped, with rounded ends, hyaline, smooth, granular.

Type: Saccharata Denman & Crous.

Notes: The narrow, aseptate, filiform pseudoparaphyses in Saccharataceae are unique in Botryosphaeriales and the pale golden brown ascospores are also distinctive. Slippers et al. (2013) accommodated this family in Botryosphaeriales. Saccharataceae grouped separately from all other families that were basal in the phylogenetic tree, suggesting a long, separate evolutionary history. Saccharataceae has previously been known only from southern Africa, and is most diverse on the Proteaceae. Recent research has shown, however, that it has also been introduced as endophytes in other countries where South African Proteaceae are now being cultivated (Marincowitz et al. 2008).


Saccharata Denman & Crous, in Crous et al., CBS Diversity Ser. (Utrecht) 2: 104 (2004).

Index Fungorum number: IF 28918; Facesoffungi number: FoF 02297; 19 morphological species (Species Fungorum 2020), 15 species with molecular data.

Type species: Saccharata proteae (Wakef.) Denman & Crous.

Phyllachora proteae Wakef., Bull. Misc. Inf., Kew(5): 164 (1922).

Notes: Crous et al. (2004) described the first species of this genus from Proteaceae in the South Western Cape region of South Africa. Three other species were added to the genus, two from Proteaceae and one from Encephalartos (Marincowitz et al. 2008; Crous et al. 2008, 2009). Apart from its restricted distribution and host range, Saccharata is also unique in its asexual morphology, which includes a hyaline, fusicoccum-like and a pigmented diplodia-like asexual morph. Based on phylogenetic analysis, evolutionary estimates and morphological data, Phillips et al. (2019) considered Septorioideaceae a synonym of Saccharataceae. We were unable to obtain fresh collections of Saccharata or other genera of Saccharataceae, therefore, a drawing of Saccharata proteae is provided (Fig. 23).

Fig. 23
figure 23

Saccharata proteae (redrawn from Slippers et al. 2013, CBS 121406). a Immersed ascomata with clypeus-like structure. b Asci and ascospores. c–e Conidiogenous cells and paraphyses. f Conidia. Scale bars: b–f = 10 µm


Saccharata proteae (Wakef.) Denman & Crous, in Crous et al., CBS Diversity Ser. (Utrecht) 2: 104 (2004).

Phyllachora proteae Wakef., Bull. Misc. Inf., Kew(5): 164 (1922).

Index Fungorum number: IF 370531; Facesoffungi number: FoF 07631; Fig. 23

Description: see Slippers et al. (2013).


Other genera included


Pileospora J.B. Tanney & K.A. Seifert, Mycol. Prog. 18: 169 (2019).

Index Fungorum number: IF 824738; MycoBank number: MB82473; – 1 species with molecular data.

Type species: Pileospora piceae J.B. Tanney & K.A. Seifert, Mycol. Prog. 18: 169 (2019).

Notes: Pileospora is distinct from Septorioides by its aseptate, ellipsoidal-fusiform conidia with apical appendages versus subcylindrical to fusiform-ellipsoidal, 1–10-septate conidia lacking appendages. Pileospora is similar to Neofusicoccum, Saccharata, and other genera of Botryosphaeriales having fusicoccum-like asexual states with co-occurring irregularly-shaped conidia. An irregular, mucoid, apical appendage distinguishes conidia of Pileospora from above mentioned similar genera.


Septorioides Quaedvl., Verkley & Crous, Stud. Mycol. 75: 383 (2013).

Index Fungorum number: IF 804464; Facesoffungi number: FoF 06278; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Septorioides pini-thunbergii (S. Kaneko) Quaedvlieg et al. Stud. Mycol. 75: 383 (2013).

Septoria pini-thunbergii S. Kaneko, in Kaneko et al., Trans. Mycol. Soc. Japan 30(4): 463 (1989).

Notes: This genus is characterized by acervular conidiomata that open by means of an irregular rupture and paraphyses that are intermingled among conidiogenous cells (Quaedvlieg et al. 2013). Septorioides pini-thunbergii was associated with needle blight and the cause of sooty mould of Pinus thunbergii in Japan (Suto 2000). Later, this species was isolated as an endophyte from P. densiflora in South Korea (Yoo and Eom 2012). The trend of raising established genera to familial status continued when Wyka and Broders (2016) introduced Septorioideaceae within the Botryosphaeriales to accommodate Septorioides species. They introduced a novel species Septorioides strobi as the second species of this family. However, Phillips et al. (2019) placed Septorioides as a genus in Saccharataceae taking into consideration the results of phylogenetic analyses, evolutionary divergence and morphological data.


Economic and ecological significance

Saccharataceae has previously been known only from southern Africa, and is most diverse on Proteaceae. Research has shown, however, that it has also been introduced as an endophyte into other countries where South African Proteaceae are now cultivated (Marincowitz et al. 2008). This plant family, which has a high endemic richness in southern Africa, has evolved in the region for more than 100 million years (Barker et al. 2007). This date allows for the estimated 57 (28–100) MYA (based on rDNA SSU) separation of the Saccharataceae as a family and to have evolved with these endemic plant hosts in the region. The species are typically associated with leaf spots and stem cankers and they appear to be pathogens. Separate studies have also shown that they are endophytes (Swart et al. 2000, Taylor et al. 2001), similar to members of the Botryosphaeriaceae.


Catinellales Ekanayaka, K.D. Hyde & Ariyawansa, ord. nov.

Index Fungorum number: IF 557327; Facesoffungi number: FoF 07646

Saprobic on dead wood in terrestrial habitats. Sexual morph: Ascomata apothecial, discoid. Excipulum composed of angularis to globose cells. Hamathecium comprising cylindric, septate, paraphyses. Asci cylindric, clavate, inoperculate with an ocular chamber, J-. Ascospores ellopsoidal, uni-seriate. Asexual morph: Undetermined.

Type: Catinellaceae Ekanayaka, K.D. Hyde & Ariyawansa.

Notes: Catinella was established by Boudier (1907). Several authors have misidentified this fungus and placed it within Leotiomycetes (Durand 1922). However phylogenetic study of Greif et al. (2007) strongly supported its placement within Dothideomycetes. Catinella formed a distinct lineage within Dothideomycetes in Hongsanan et al. (2020), thus we introduced this new order to accommodate Catinella and allied species in the class Dothideomycetes. The divergence time for Catinellales is estimated as 163 MYA (Hongsanan et al. 2020).

Accepted families: Catinellaceae.


Catinellaceae Ekanayaka, K.D. Hyde & Ariyawansa, fam. nov.

Index Fungorum number: IF 557328; Facesoffungi number: FoF 07647, 1 species.

Saprobic on dead wood in terrestrial habitats. Sexual morph: Ascomata apothecial, discoid, cupulate, flattened disc. Excipulum composed of subangularis to globose cells, ectal cells are pigmented. Hamathecium comprising cylindric, septate, swollen apices agglutinate to form a pseudoepithecium, paraphyses. Asci 8-spored, cylindric, clavate, inoperculate, unitunicate, J-, with well-developed ocular chamber. Ascospores ellopsoidal, uni-seriate, darkly pigmented, one-celled. Asexual morph: Undetermined.

Type genus: Catinella Boud.

Notes: The new family Catinellaceae bears some morphological characteristics similar to Leotiomycetes, i.e. cupulate apothecia, unitunicate asci, aseptate ascospores. However, Dothideomycetes also show these characters, although they are not common within the class (Greif et al. 2007).


Catinella Boud., Hist. Class. Discom. Eur. (Paris): 150 (1907).

Index Fungorum number: IF 845; Facesoffungi number: FoF 07648; 3 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type: Catinella olivacea (Batsch) Boud.

≡ Peziza olivacea Batsch, Elench. fung. (Halle): 127 (1783).

Notes: Catinella was established by Boudier (1907), with C. olivacea as the type species. The genus was recognised as a member in Leotiomycetes based on its morphology. However, sequence data of C. olivacea indicated that this genus formed a distinct clade within Dothideomycetes (Fig. 24).

Fig. 24
figure 24

Catinella olivacea (a = redrawn from Gamundí 1981; b–e = redrawn from Akata et al. 2016). a Peridium. b Apothecial ascomata. c Ascospores. d Asci. e Anchor hyphae. Scale bars: a = 10 μm, b = 20 μm


Economic and ecological significance

Catinellaceae is saprobic on dead wood and plays a role in recycling of organic matter.


Cladoriellales Crous.

Index Fungorum number: IF 823435; Facesoffungi number: FoF 07649.

Cladoriellales was established by Crous et al. (2017), with a single genus Cladoriella. Molecular data (LSU) confirmed its status as a distinct order within Dothideomycetes. This order contains asexual, cladosporium-like species which are found as saprobes or pathogens on leaf surface. In Fig. 25, four species of Cladoriella (Cladoriella eucalypti, C. kinglakensis, C. rubrigena, and C. xanthorrhoeae) grouped with high support. The divergence time for Cladoriellales is estimated as 183 MYA (stem age, Hongsanan et al. 2020, Fig. 2).

Fig. 25
figure 25

Phylogram generated from maximum likelihood analysis (RAxML) of Cladoriellales based on LSU sequence data. Maximum likelihood bootstrap values equal or above 70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/PP) are given at the nodes. An original isolate number is noted after the species name. The tree is rooted to Capnodium coffeae (OSC 100414) and Leptoxyhium fumago (CBS 123.26). The ex-type strains are indicated in bold. Hyphen (-) represents support values below 70% MLBS and 0.90 PP

Accepted families: Cladoriellaceae.


Cladoriellaceae Crous, in Crous et al., Persoonia 39: 417 (2017).

Index Fungorum number: IF 823436; Facesoffungi numbers: FoF 07650, 5 species.

Saprobes or pathogens on leaf surface. Sexual morph: Undetermined. Asexual morph: Hyphae coiling, branched, septate, with swollen cells giving rise to conidiophores, with hyphopodium-like structures at the base, simple, intercalary, brown to dark brown, thick-walled, smooth to finely verruculose. Conidiophores separate, erect, subcylindrical, straight, septate, thick-walled, brown to dark brown, smooth to finely verruculose. Conidiogenous cells terminal or intercalary, mono- or poly-tretic, sympodial, with 1–2 conspicuous loci, thickened, darkened, refractive, with a minute central pore. Conidia remains in long acropetal chains, narrowly ellipsoidal to cylindrical or fusoid, brown, non or 1-septate, thick-walled, finely verruculose, with apical conidium rounded at the apex; additional conidia with 1–2 truncate, conspicuous hila; thickened, darkened, refractive, with a minute central pore. Chlamydospores absent (Crous et al. 2006b, 2017).

Type: Cladoriella Crous.

Notes: Cladoriellaceae was established by Crous et al. (2017) to accommodate a single genus Cladoriella. Sequence data for this family are available in GenBank and it also supports Cladoriellaceae as a family of Cladoriellales in Dothideomycetes.


Cladoriella Crous, Stud. Mycol. 55: 54 (2006).

Index Fungorum number: IF 500799; Facesoffungi number: FoF 07651; 5 morphological species (Species Fungorum 2020), 4 species with molecular data.

Type species: Cladoriella eucalypti Crous.

Notes: Crous et al. (2006b) introduced Cladoriella to accommodate cladosporium-like species. This genus is similar to Devriesia, but lacks chlamydospores, and produces a distinct red pigment on media. Phylogenetic analyses also support this distinctness as Cladoriella formed a clade apart from Cladosporium complex (Mycosphaerellaceae), Cladophialophora complex (Herpotrichiellaceae), or Pseudocladosporium complex (Venturiaceae) (Crous et al. 2006b) (Fig. 26).

Fig. 26
figure 26

Conidiophore and conidia of Cladoriella spp. a Cladoriella eucalypti (redrawn from Crous et al. 2006, CBS H-18043). b–d Cladoriella xanthorrhoeae (redrawn from Crous et al. 2017, CBS H-23300)


Economic and ecological significance

Species in this family are saprobic and probably are plant pathogenic (Wijayawardene et al. 2017a). They are found on leaf surfaces, however, the lifestyle of species in Cladoriellaceae is not well-studied.


Collemopsidiales Pérez-Ort.

Index Fungorum number: IF 815627; Facesoffungi number: FoF 07652.

This recently erected order includes a single family, Xanthopyreniaceae. It encompasses species of lichen-forming fungi, lichenicolous or forming loose associations with cyanobacteria. The diversity of the order has been largely underestimated and the actual number of species could exceed 200. The divergence time for Collemopsidiales is estimated as 230 MYA (stem age, Hongsanan et al. 2020) (Fig. 27).

Fig. 27
figure 27

Phylogram generated from maximum likelihood analysis (RAxML) of Collemopsidiales based on LSU and SSU sequence data. Maximum likelihood bootstrap values equal or above 70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/PP) are given at the nodes. An original isolate number is noted after the species name. The tree is rooted to and Abrothallus cladoniae (AB53) and A. parmeliarum (AB36). Hyphen (-) represents support values below 70% MLBS and 0.90 PP

Accepted families: Xanthopyreniaceae.


Xanthopyreniaceae Zahlbr., in Engler & Prantl, Nat. Pflanzenfam., Edn 2 (Leipzig) 8: 91 (1926).

Index Fungorum number: IF 81525; Facesoffungi number: FoF 07653, 53 species.

Lichen-forming fungi, lichenicolous or forming loose associations with cyanobacteria. Vegetative hyphae usually hyaline. Sexual morph: Ascomata perithecioid, solitary or in groups, superficial or immersed in the substrarum, subglobose. Peridium thin to thick, composed by elongated to roundish, more or less compresed cells of textura angularis. Involucrelum present in some species, with textura intricata. Hamathecium comprising thin, septate, highly branched and anastomosed interascal filaments. Asci 4–8-spored, thick-walled, bitunicate, fissitunicate, ovate to clavate, apedicellate, with a distinct ocular chamber. Ascopores irregularly arranged within the ascus, ellipsoid to broadly ellipsoid, hyaline, 1-septate, wall cell smooth, usually with a gelatinous shell. Asexual morph: Coelomycetous. Pycnidium, superficial or immersed in the substratum, black, with a small ostiole. Pycnidial wall thick-walled cells of textura angularis, composed of ± isodiametric or elongated cells. Conidiogenous cells ± cylindrical, phialidic. Conidiospores bacilliform to ellipsoid.

Type: Xanthopyrenia Bachm.

Notes: Xanthopyreniaceae was introduced by Zahlbruckner to accommodate the single species Xanthopyrenia tichothecioides (= Arthopyrenia tichothecioides), a species growing on periodically inundated calcareous rocks. The species was later combined into Pyrenocollema (P. tichothecioides) and subsequently many species associated with cyanobacteria, and with perithecioid ascomata, highly branched and anastomosed interascal filaments and hyaline 1-septate ascospores were transferred or described in Pyrenocollema (e.g. Santesson 1992; Coppins et al. 1992; McCarthy and Kantvilas 1999; Nordin 2002). However, Grube and Ryan (2002) stated that the type species of the genus, P. tremelloides is in fact a parasite on Nostoc and is not congeneric with X. tichothecioides, suggesting that the correct generic name for those species should be Collemopsidium, and many species of Pyrenocollema were subsequently combined into this genus (Grube and Ryan 2002; Mohr et al. 2004). Pérez-Ortega et al. (2016) studied the phylogenetic relationships of Xanthopyreniaceae using six molecular markers. Although they found Xanthopyreniaceae belongs to Dothiodeomycetes, and described the new order Collemopsidiales, their exact position remained elusive despite of using different taxa sampling, very likely due to the use of Arthoniomycetes as outgroup. Subsequent work by Liu et al. (2017) found high support for the sister relationships of Collemopsidiales with Monoblastiales. Based on uncertainties by Pérez-Ortega et al. (2016), Tedersoo et al. (2018) described the new class Collemopsidiomycetes which should be considered as a synonym of Dothiodeomycetes. Lumbsch and Huhndorf (2007) included the genera Collemopsidium, Didymellopsis, Frigidopyrenia, Pyrenocollema, and Zwackhiomyces as members of the Xanthopyreniaceae. Later, Wijayawardene et al. (2018) excluded Frigidopyrenia and Pyrenocollema from the family and placed them in Pezizomycotina genera incertae sedis. Unfortunately, no molecular data is available for Didymellopsis, Frigidopyrenia and Pyrenocollema. Pérez-Ortega et al. (2016) discussed that the number of species in the family may have been highly underestimated.


Xanthopyrenia Bachm., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 105(1): 65 (1919).

Index Fungorum number: IF 5810; Facesoffungi number: FoF 07654; – 2 morphological species (Pérez-Ortega et al. 2016), molecular data unavailable.

Type species: Xanthopyrenia tichothecioides (Arnold) Bachm.

Arthopyrenia tichothecioides Arnold, Flora, Regensburg 52: 268 (1869).

Notes: The genus was described to accommodate the single species Arthopyrenia tichothecioides, a species occurring with seawater or spray on calcareous rocks (Thüs and Schultz 2009). This species and X. heardensis, the only other species described in the genus, were subsequently transferred to Pyrenocollema. Pérez-Ortega et al. (2016) showed that Collemopsidium as currently circumscribed is paraphyletic, so Xanthopyrenia might be the correct generic name for non-marine Collemopsidium species (Fig. 28).

Fig. 28
figure 28

Morphological and anatomical features of Xanthopyreniaceae. a Habitus Collemopsidium foveolatum-morphotype. b C. halodytes-morphotype. c C. ostrearum-morphotype. d Zwackhiomyces coepulonus. e Ascus from C. angermannicum. F Interascal filaments from C. cf. foveolatum. g Ascospore from C. cf. sublitorale. h Ascospore from C. angermannicum with gelatinous sheath. Scale bars: a–c = 1 mm; d = 0.5 mm; e–h = 10 µm


Other genera included

Collemopsidium Nyl., Flora, Regensburg 64: 6 (1881).

Index Fungorum number: IF 1182; Facesoffungi number: FoF 07655; – 16 morphological species (Species Fungorum 2020), 3 species with molecular data (Pérez-Ortega et al. 2016).

Type species: Collemopsidium iocarpum (Nyl.) Nyl., Flora, Regensburg 64: 6 (1881).

Pyrenopsis iocarpa Nyl., Not. Sällsk. Fauna et Fl. Fenn. Förh., Ny Ser. 5: 26 (1861).

Notes: Grube and Ryan (2002) studied the types of Collemopsidium and Pyrenocollema and stated that the former is the correct generic name for species previously included in Pyrenocollema. Pérez-Ortega et al. (2016) showed that the genus as currently understood is paraphyletic and species occurring in marine and terrestrial and freshwater habitats belong to different lineages. Collemopsidium iocarpum was described growing close to the seashore and it is likely the correct name for the marine clade. Species concepts and taxonomy have to be thoroughly studied. Pérez-Ortega et al. (2016) showed that each of the species commonly accepted in the literature likely encompass dozens of different taxa. Cyanobacteria associated with Collemopsidium are largely undetermined, marine species likely belong to Hyella (Swinscow 1965b). Xanthocapsa may be involved in freshwater symbioses (Grube and Hafellner 1990).


Didymellopsis (Sacc.) Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 66 (1931).

= Didymella subgen. Didymellopsis Sacc. & D. Sacc., Syll. fung. (Abellini) 17: 657 (1905).

Index Fungorum number: IF 1550; Facesoffungi number: FoF 07656; – 8 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Didymellopsis latitans (Nyl.) Sacc. ex Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 265 (1931).

Obryzum latitans Nyl., Flora, Regensburg 68: 298 (1885).

Notes: All species in Didymellopsis are found on lichens with cyanobacterial photobionts, but D. perigena which grows on Catapyrenium squamulosum might be associated with cyanobacteria surrounding the host thallus. Differences between Didymyellopsis and Zwackhiomyces are subtle (Zhurbenko et al. 2015) and a thorough revision of the group, including DNA sequence analyses should be carried out.


Frigidopyrenia Grube, Phyton, Horn 45(2): 307 (2005).

Index Fungorum number: IF 521258; Facesoffungi number: FoF 07657; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Frigidopyrenia bryospila (Nyl.) Grube, Phyton, Horn 45(2): 308 (2005).

Verrucaria bryospila Nyl., Flora, Regensburg 47: 357 (1864).

Notes: Grube (2005) described the monotypic genus Frigidopyrenia for Verrucaria bryospila occurring on mosses in arctic-alpine areas, forming distinct squamules and with cyanobacteria as photobiont. Grube (2005) discussed the morphological differences of the genus with similar genera such as Collemopsidium, Magmopsis and Pyrenocollema, which have different peridial structure. It is not clear that the genus belongs in Xanthopyreniaceae.


Rhagadodidymellopsis Fdez.-Brime, Gaya, Llimona & Nav.-Ros., Plant and Fungal Systematics 65(1): 177 (2020).

Index Fungorum number: IF 835454; Facesoffungi number: FoF 09130; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Rhagadodidymellopsis endocarpi Fdez.-Brime, Gaya, Llimona & Nav.-Ros., Plant and Fungal Systematics 65(1): 177 (2020).

Notes: This recently erected genus, with the only species R. endocarpi growing on Endocarpon pusillum, was described based on the presence of ascomata occurring in stromata with uneven wall. It seems to be closely related to Didymellopsis and Zwackhiomyces (Fernández-Brime et al. 2020). It is here tentatively placed in Collemopsidiales and Xanthopyreniaceae until molecular data is available.


Zwackhiomyces Grube & Hafellner, Nova Hedwigia 51(3–4): 305 (1990).

Index Fungorum number: IF 25388; Facesoffungi number: FoF 07658; – 26 morphological species (Species Fungorum 2020), 1 species with molecular data.

Type species: Zwackhiomyces coepulonus (Norman) Grube & R. Sant., in Grube & Hafellner, Nova Hedwigia 51(3–4): 310 (1990).

Arthopyrenia coepulona Norman, Bot. Notiser: 192 (1868) [1867].

Notes: Zwackhiomyces is one of the most diverse genera of lichenicolous fungi with 35 known species (Diederich et al. 2018). Species mostly grow on crustose lichens with green photobionts, although five species grow on hosts with cyanobacteria as photobiont. Pérez-Ortega et al. (2016) sequenced the type species and confirmed the genus is a member of Xanthopyreniaceae.


Zwackhiomacromyces Etayo & van den Boom, Opuscula Philolichenum 13: 70 (2014).

Index Fungorum number: IF 803963; Facesoffungi number: FoF 08929; – 2 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Zwackhiomacromyces constrictocarpus Etayo & van den Boom, Opuscula Philolichenum 13: 72 (2014).

Notes: van den Boom and Etayo (2014) established the new genus based on material from the Iberian Peninsula growing on Megaspora verrucosa. The new genus differs from Zwackhiomyces in having lageniform ascomata, with large and granulose papillae, larger asci and ascospores, and normally 3-septate smooth-walled ascospores (van den Boom and Etayo 2014). Later, Etayo and Berger (in Berger and Zimmerman 2016) combined Pyrenidium hyalosporum growing in Placopsis gelida into Zwackhiomacromyces based on ascoma and ascospore characters. This genus tentatively placed in the order Collemopsidiales and Xanthopyreniaceae until molecular data is available.


Economic and ecological significance

Members of the Xanthopyreniaceae are usually inconspicuous and have been often overlooked. Most of the lichenicolous species do not cause harm to the hosts, and many of the interactions may be considered commensalistic. Marine species of Collemopsidium with boring capacity may have relevant ecological significance since they actively produce bioweathering in littoral rocky substrates, boosting the deteriorative action of waves. Furthermore, those species are able to colonize and modified mollusk and barnacle shells, mediating mimicry in some species (Espoz et al. 1995).


Dyfrolomycetales K.L. Pang, K.D. Hyde and E.B.G. Jones.

Index Fungorum number: IF 805281; Facesoffungi number: FoF 07670.

Dyfrolomycetales with the type genus Dyfrolomyces was introduced to accommodate marine fungi and now includes terrestrial, wood-inhabiting taxa which are characterized by immersed, ostiolate, clypeate, papillate ascomata, bitunicate, cylindrical, short pedicellate asci, with a distinct ocular chamber along with a ring-like subapical ring, and overlapping uni-seriate, broadly fusiform, symmetrical, hyaline, multi-septate ascospores. Only the single family Pleurotremataceae is accepted in this order. The divergence time for Dyfrolomycetales is estimated as 171 MYA (stem age, Hongsanan et al. 2020) (Fig. 29).

Fig. 29
figure 29

Phylogram generated from maximum likelihood analysis (RAxML) of Dyfrolomycetales based on LSU and SSU sequence data. Maximum likelihood bootstrap values equal or above 70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/PP) are given at the nodes. An original isolate number is noted after the species name. The tree is rooted to Anisomeridium phaeospermum (MPN539) and A. ubianum (94). The ex-type and reference strains are indicated in bold. Hyphen (-) represents support values below 70% MLBS and 0.90 PP

Accepted families: Pleurotremataceae.


Pleurotremataceae Walt. Watson, New Phytol. 28: 113 (1929).

Index Fungorum: IF 81192; Facesoffungi number: FoF 01911, 47 species.

= Dyfrolomycetaceae K.D. Hyde, K.L. Pang, Alias, Suetrong & E.B.G. Jones, in Pang et al., Cryptog. Mycol. 34(3): 227 (2013).

Saprobic on wood in terrestrial and aquatic habitats. Sexual morph: Ascomata perithecial, gregarious or solitary, immersed or erumpent throughout the host, dark-brown to black, ovoid to subglobose, carbonaceous to membranaceous, clypeate, ostiolate, papillate. Peridium comprising of dark pigmented cells of textura angularis or epidermoidea. Hamathecium comprising numerous, filamentous, hyaline, septate, branched or unbranched, narrow cellular pseudoparaphyses. Asci 8-spored, bitunicate, clavate to cylindrical, short pedicellate, apical ring present, J-. Ascospores uni-seriate, ellipsoidal to cylindrical or fusiform, hyaline, septate, smooth-walled, mostly with guttules. Asexual morph: Undetermined.

Type: Pleurotrema Müll.

Notes: Pleurotremataceae is typified by Pleurotrema with Pleurotrema polysemum as the type species. Subsequently, the nomenclature and placement of this family has been the subject of discussion until Maharachchikumbura et al. (2015) placed it under Chaetosphaeriales, Sordariomycetes. Maharachchikumbura et al. (2016) examined the isotype of P. polysemum and found that Dyfrolomycetaceae (Dyfrolomycetales) is a synonym of Pleurotremataceae, thus removed Pleurotremataceae from Sordariomycetes to be a single family of Dyfrolomycetales. This family comprises three genera, and most members are characterized by perithecial, ostiolate, glabrous, ovoid to subglobose ascomata, clavate to cylindrical, unitunicate, short pedicellate asci with a J- apical ring, and hyaline, ellipsoidal to cylindrical, septate, guttulate ascospores that are uni-seriately arranged in the asci.


Pleurotrema Müll. Arg., Bot. Jb. 6: 388 (1885).

Index Fungorum number: IF 4251; Facesoffungi number: FoF 07671; 14 morphological species (Species Fungorum 2020), molecular data unavailable.

Type species: Pleurotrema polysemum (Nyl.) Müll. Arg.

≡ Parathelium polysemum Nyl., Bot. Ztg. 20: 279 (1872).

Notes: Pleurotrema was introduced as lichenized by Müller (1885). The placement of this genus has been the subject of controversy as it has been placed in different families viz., Pyrenulaceae, Pleurotremataceae and Ascomycota genera incertae sedis, based on its morphological characters (Barr 1994; Kirk et al. 2001; Lumbsch and Huhndorf 2010). Maharachchikumbura et al. (2016) re-examined the isotype of Pleurotrema polysemum, and synonymized Dyfrolomycetaceae under Pleurotremataceae.


Other genera included

Dyfrolomyces K.D. Hyde, K.L. Pang, Alias, Suetrong & E.B.G. Jones, in Pang et al., Cryptog. Mycol. 34(3): 227 (2013).

Index Fungorum number: IF 804660; Facesoffungi number: FoF 07672; – 9 morphological species (Species Fungorum 2020), 6 species with molecular data.

Type species: Dyfrolomyces tiomanensis K.L. Pang, Alias, K.D. Hyde, Suetrong & E.B.G. Jones, in Pang et al., Cryptog. Mycol. 34(3): 228 (2013).

Notes: Dyfrolomyces was established based on morphological and molecular analyses. Members of this genus are characterized by relatively large, immersed, globose or subglobose, clypeate, ostiolate, papillate ascomata, bitunicate, fissitunicate, cylindrical asci and broadly fusiform, symmetrical, hyaline, septate ascospores, with or without a mucilaginous sheath.


Dyfrolomyces distoseptatus M. Niranjan and V.V Sarma, sp. nov.

Index Fungorum number: IF 556726; Facesoffungi number: FoF 06625; Fig. 30

Fig. 30
figure 30

Dyfrolomyces distoseptatus (AMH 9984, holotype). a Host plant. b Papillate necks. c, d, i Vertical section of ascomata. e Peridium. f Pseudoparaphyses. g, h Asci j, k Cultures in Petri plates from single spore isolations. lo Ascospores. Scale bars: d = 100 µm, e–i = 50 µm, l–o = 10 µm

Etymology: In reference to the distoseptate ascospores.

Saprobic on undetermined decaying twig. Sexual morph: Ascomata 550–630 high × 450–600 μm wide, perithecial, immersed in periderm, erumpent neck with pseudoparaphyses, clypeate, ostiolate, papillate. Peridium 40 μm, with two strata, outer thick, carbonaceous and inner brown and hyaline cells of textura angularis to epidermoidea cells. Hamathecium comprising 1.8–2.1 μm wide, filamentous, septate, unbranched, dense, narrow cellular pseudoparaphyses, longer than asci. Asci (123.1)126.7–146.2 (148.6) × (4.1)4.7–6.3 (6.5) μm (\( \bar{x} \) = 136.8 × 5.6, n = 25), 8–spored, bitunicate, cylindrical, apical ends obtuse, smooth-walled, short pedicellate, persistent. Ascospores (19.4)19.7–24.9 × (4.1) 4.3–5 μm (\( \bar{x} \) = 21.9 × 4.7, n = 25), uni-seriate, fusoid, obtuse ends, hyaline, 3-distoseptate, apical ends slightly bent. Asexual morph: Undetermined.

Material examined: India, Andaman and Nicobar Islands, South Andaman, Manjery, (11˚52’25.7”N 92˚64’89.9”E), on unidentified twig, 10 December 2017, M. Niranjan (AMH 9984, holotype), extype living culture NFCCI: 4377.

GenBank numbers: ITS: MK024391, LSU: MH971236.

Notes: Dyfrolomyces distoseptatus clustered with D. thamplaensis with strong bootstrap support (100%) in ML analysis. It also forms a sister relationship clade with D. sinensis (Fig. 29). However, D. distoseptatus has several distinguishable characters. It is closely related to D. thamplaensis in having distoseptate ascospores with acute ends, but the ascospores in the latter have prominent guttules. In addition, the asci of D. distoseptatus are smaller than D. thamplaensis (123.1–148.6 × 4.1–6.3 μm vs. 114–160 × 6–8.5 μm) (Zhang et al. 2017a). Furthermore, D. distoseptatus produces ascospores that are 3-septate when compared to 6–7 septate ascospores in D. sinensis (Hyde et al. 2018). Hence, based on the above-mentioned morphological differences (Fig. 30) and molecular sequence analyses (Fig. 29), a new species, Dyfrolomyces distoseptatus, is introduced.


Melomastia Nitschke ex Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 90 (1875).

Index Fungorum number: IF 3118; Facesoffungi number: FoF 07673; – 24 morphological species (Species Fungorum 2020), 2 species with molecular data.

Type species: Melomastia mastoidea (Fr.) J. Schröt., in Chon, Krypt. -Fl. Schlesien (Breslau) 3.2(3): 320 (1894) [1908].

≡ Sphaeria mastoidea Fr., K. svenska Vetensk-Akad. Handl., ser. 3 38: 267 (1817).

Notes: Melomastia was introduced by Saccardo (1875) to accommodate the type species M. friesii. Schröter (1894) synonymized M. friesii under M. mastoidea. Species of Melomastia are characterized by immersed, black, ostiolate ascomata, cylindrical, thick-walled, pedicellate asci with a J- subapical ring, and ovoid, hyaline, 2-septate, guttulate ascospores (Norphanphoun et al. 2017).


Economic significance

Only three genera are accepted in Pleurotremaceae, and most of the taxa in this family are saprobes in aquatic, mangrove and terrestrial habitats (Pang et al. 2013; Norphanphoun et al. 2017; Zhang et al. 2017a).


Eremithallales Lücking & Lumbsch.

Index Fungorum number: IF 540500; Facesoffungi number: FoF 07674.

Eremithallales was introduced for a single species Eremithallus costaricensis (Lücking et al. 2008). Previously Eremithallales was reported close to Lichinomycetes. However, both phylogenetic and morpholocal analysis confirmed the placement of this order in Dothideomycetes with strong support and later Melaspileaceae was included (Ertz and Diederich 2015). In Hongsanan et al. (2020), Eremithallales froms a distinct clade within Dothideomycetes. The divergence time for Eremithallales is estimated as 238 MYA (stem age Hongsanan et al.