Redisposition of acremonium-like fungi in Hypocreales
Abstract
Acremonium is acknowledged as a highly ubiquitous genus including saprobic, parasitic, or endophytic fungi that inhabit a variety of environments. Species of this genus are extensively exploited in industrial, commercial, pharmaceutical, and biocontrol applications, and proved to be a rich source of novel and bioactive secondary metabolites. Acremonium has been recognised as a taxonomically difficult group of ascomycetes, due to the reduced and high plasticity of morphological characters, wide ecological distribution and substrate range. Recent advances in molecular phylogenies, revealed that Acremonium is highly polyphyletic and members of Acremonium s. lat. belong to at least three distinct orders of Sordariomycetes, of which numerous orders, families and genera with acremonium-like morphs remain undefined. To infer the phylogenetic relationships and establish a natural classification for acremonium-like taxa, systematic analyses were conducted based on a large number of cultures with a global distribution and varied substrates. A total of 633 cultures with acremonium-like morphology, including 261 ex-type cultures from 89 countries and a variety of substrates including soil, plants, fungi, humans, insects, air, and water were examined. An overview phylogenetic tree based on three loci (ITS, LSU, rpb2) was generated to delimit the orders and families. Separate trees based on a combined analysis of four loci (ITS, LSU, rpb2, tef-1α) were used to delimit species at generic and family levels. Combined with the morphological features, host associations and ecological analyses, acremonium-like species evaluated in the present study are currently assigned to 63 genera, and 14 families in Cephalothecales, Glomerellales and Hypocreales, mainly in the families Bionectriaceae, Plectosphaerellaceae and Sarocladiaceae and five new hypocrealean families, namely Chrysonectriaceae, Neoacremoniaceae, Nothoacremoniaceae, Pseudoniessliaceae and Valsonectriaceae. Among them, 17 new genera and 63 new combinations are proposed, with descriptions of 65 new species. Furthermore, one epitype and one neotype are designated to stabilise the taxonomy and use of older names. Results of this study demonstrated that most species of Acremonium s. lat. grouped in genera of Bionectriaceae, including the type A. alternatum. A phylogenetic backbone tree is provided for Bionectriaceae, in which 183 species are recognised and 39 well-supported genera are resolved, including 10 new genera. Additionally, rpb2 and tef-1α are proposed as potential DNA barcodes for the identification of taxa in Bionectriaceae.
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23 © 2023 Westerdijk Fungal Biodiversity Institute. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/bync-nd/4.0/). Studies in Mycology Redisposition of acremonium-like fungi in Hypocreales L.W. Hou1, A. Giraldo2, 3, J.Z. Groenewald2, T. Rämä4, R.C. Summerbell5,6, G.Z. Huang7, L. Cai1*, P.W. Crous2,8,9,10* 1State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing, 100101, China; 2Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, Utrecht, 3584 CT, The Netherlands; 3Netherlands Institute for Vectors, Invasive plants and Plant health (NIVIP), NVWA, Wageningen Netherlands; 4The Norwegian College of Fishery Science, Department at Faculty of Biosciences, Fisheries and Economics, UiT The Arctic University of Norway, Tromsø, Norway; 5Sporometrics, Toronto, ON, Canada; 6Dalla Lana School of Public Health, University of Toronto, Toronto, ON, Canada; 7State Key Laboratory of Integrated Management of Pest Insects and Rodents, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China; 8Microbiology, Department of Biology, Utrecht University, Padualaan 8, Utrecht, 3584 CH, The Netherlands; 9Department of Biochemistry, Genetics and Microbiology, Forestry and Agricultural Biotechnology Institute (FABI), Faculty of Natural and Agricultural Sciences, University of Pretoria, Private Bag X20, Hatfield, Pretoria, 0028, South Africa; 10Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, Wageningen, 6708 PB, The Netherlands *Corresponding authors: P.W. Crous, [email protected].nl; L. Cai, [email protected] Abstract: Acremonium is acknowledged as a highly ubiquitous genus including saprobic, parasitic, or endophytic fungi that inhabit a variety of environments. Species of this genus are extensively exploited in industrial, commercial, pharmaceutical, and biocontrol applications, and proved to be a rich source of novel and bioactive secondary metabolites. Acremonium has been recognised as a taxonomically difficult group of ascomycetes, due to the reduced and high plasticity of morphological characters, wide ecological distribution and substrate range. Recent advances in molecular phylogenies, revealed that Acremonium is highly polyphyletic and members of Acremonium s. lat. belong to at least three distinct orders of Sordariomycetes, of which numerous orders, families and genera with acremonium-like morphs remain undefined. To infer the phylogenetic relationships and establish a natural classification for acremonium-like taxa, systematic analyses were conducted based on a large number of cultures with a global distribution and varied substrates. A total of 633 cultures with acremonium-like morphology, including 261 ex-type cultures from 89 countries and a variety of substrates including soil, plants, fungi, humans, insects, air, and water were examined. An overview phylogenetic tree based on three loci (ITS, LSU, rpb2) was generated to delimit the orders and families. Separate trees based on a combined analysis of four loci (ITS, LSU, rpb2, tef-1α) were used to delimit species at generic and family levels. Combined with the morphological features, host associations and ecological analyses, acremonium-like species evaluated in the present study are currently assigned to 63 genera, and 14 families in Cephalothecales, Glomerellales and Hypocreales, mainly in the families Bionectriaceae, Plectosphaerellaceae and Sarocladiaceae and five new hypocrealean families, namely Chrysonectriaceae, Neoacremoniaceae, Nothoacremoniaceae, Pseudoniessliaceae and Valsonectriaceae. Among them, 17 new genera and 63 new combinations are proposed, with descriptions of 65 new species. Furthermore, one epitype and one neotype are designated to stabilise the taxonomy and use of older names. Results of this study demonstrated that most species of Acremonium s. lat. grouped in genera of Bionectriaceae, including the type A. alternatum. A phylogenetic backbone tree is provided for Bionectriaceae, in which 183 species are recognised and 39 well-supported genera are resolved, including 10 new genera. Additionally, rpb2 and tef-1α are proposed as potential DNA barcodes for the identification of taxa in Bionectriaceae. Key words: Bionectriaceae, multi-locus, new taxa, phylogeny, taxonomy, Sarocladiaceae, soilborne. Taxonomic novelties: New families: Chrysonectriaceae L.W. Hou, L. Cai & Crous, Neoacremoniaceae L.W. Hou, L. Cai & Crous, Nothoacremoniaceae L.W. Hou, L. Cai & Crous, Pseudoniessliaceae L.W. Hou, L. Cai & Crous, Valsonectriaceae L.W. Hou, L. Cai & Crous. New genera: Bionectriaceae: Alloacremonium L.W. Hou, L. Cai & Crous, Gossypinidium L.W. Hou, L. Cai & Crous, Monohydropisphaera L.W. Hou, L. Cai & Crous, Musananaesporium L.W. Hou, L. Cai & Crous, Paragliomastix L.W. Hou, L. Cai & Crous, Proliferophialis L.W. Hou, L. Cai & Crous, Proxiovicillium L.W. Hou, L. Cai & Crous, Ramosiphorum L.W. Hou, L. Cai & Crous, Verruciconidia L.W. Hou, L. Cai & Crous, Waltergamsia L.W. Hou, L. Cai & Crous; Clavicipitaceae: Subuliphorum L.W. Hou, L. Cai & Crous; Neoacremoniaceae: Neoacremonium L.W. Hou, L. Cai & Crous; Nothoacremoniaceae: Nothoacremonium L.W. Hou, L. Cai & Crous; Plectosphaerellaceae: Allomusicillium L.W. Hou, L. Cai & Crous, Parafuscohypha L.W. Hou, L. Cai & Crous; Pseudoniessliaceae: Pseudoniesslia L.W. Hou, L. Cai & Crous; Sarocladiaceae: Polyphialocladium L.W. Hou, L. Cai & Crous. New species: Bionectriaceae: Alloacremonium ferrugineum L.W. Hou, L. Cai & Crous, Al. humicola L.W. Hou, L. Cai & Crous, Acremonium aerium L.W. Hou, L. Cai & Crous, A. brunneisporum L.W. Hou, L. Cai & Crous, A. chlamydosporium L.W. Hou, L. Cai & Crous, A. ellipsoideum L.W. Hou, Rämä, L. Cai & Crous, A. gamsianum L.W. Hou, L. Cai & Crous, A. longiphialidicum L.W. Hou, L. Cai & Crous, A. multiramosum L.W. Hou, Rämä, L. Cai & Crous, A. mycoparasiticum L.W. Hou, L. Cai & Crous, A. stroudii K. Fletcher, F.C. Küpper & P. van West, A. subulatum L.W. Hou, L. Cai & Crous, A. synnematoferum L.W. Hou, Rämä, L. Cai & Crous, Bulbithecium ammophilae L.W. Hou, L. Cai & Crous, B. ellipsoideum L.W. Hou, L. Cai & Crous, B. truncatum L.W. Hou, L. Cai & Crous, Emericellopsis brunneiguttula L.W. Hou, L. Cai & Crous, Gliomastix musae L.W. Hou, L. Cai & Crous, Gossypinidium sporodochiale L.W. Hou, L. Cai & Crous, Hapsidospora stercoraria L.W. Hou, L. Cai & Crous, H. variabilis L.W. Hou, L. Cai & Crous, Mycocitrus odorus L.W. Hou, L. Cai & Crous, Nectriopsis ellipsoidea L.W. Hou, L. Cai & Crous, Paracylindrocarpon aurantiacum L.W. Hou, L. Cai & Crous, Pn. foliicola Lechat & J. Fourn., Paragliomastix rosea L.W. Hou, L. Cai & Crous, Proliferophialis apiculata L.W. Hou, L. Cai & Crous, Protocreopsis finnmarkica L.W. Hou, L. Cai, Rämä & Crous, Proxiovicillium lepidopterorum L.W. Hou, L. Cai & Crous, Ramosiphorum echinoporiae L.W. Hou, L. Cai & Crous, R. polyporicola L.W. Hou, L. Cai & Crous, R. thailandicum L.W. Hou, L. Cai & Crous, Verruciconidia erythroxyli L.W. Hou, L. Cai & Crous, Ve. infuscata L.W. Hou, L. Cai & Crous, Ve. quercina L.W. Hou, L. Cai & Crous, Ve. siccicapita L.W. Hou, L. Cai & Crous, Ve. unguis L.W. Hou, L. Cai & Crous, Waltergamsia alkalina L.W. Hou, L. Cai & Crous, W. catenata L.W. Hou, L. Cai & Crous, W. moroccensis L.W. Hou, L. Cai & Crous, W. obpyriformis L.W. Hou, L. Cai & Crous; Chrysonectriaceae: Chrysonectria crystallifera L.W. Hou, L. Cai & Crous; Nectriaceae: Xenoacremonium allantoideum L.W. Hou, L. Cai & Crous; Neoacremoniaceae: Neoacremonium distortum L.W. Hou, L. Cai & Crous, N. flavum L.W. Hou, L. Cai & Crous; Nothoacremoniaceae: Nothoacremonium subcylindricum L.W. Hou, L. Cai & Crous, No. vesiculophorum L.W. Hou, L. Cai & Crous; Myrotheciomycetaceae: Trichothecium hongkongense L.W. Hou, L. Cai & Crous; Plectosphaerellaceae: Brunneomyces polyphialidus L.W. Hou, L. Cai & Crous, Parafuscohypha proliferata L.W. Hou, L. Cai & Crous; Sarocladiaceae: Chlamydocillium acaciae L.W. Hou, L. Cai & Crous, C. antarcticum L.W. Hou, L. Cai & Crous, C. guttulatum L.W. Hou, L. Cai & Crous, C. lolii L.W. Hou, L. Cai & Crous, C. soli L.W. Hou, L. Cai & Crous, C. terrestre L.W. Hou, L. Cai & Crous, Parasarocladium chondroidum L.W. Hou, L. Cai & Crous, Polyphialocladium Available online at www.studiesinmycology.org StudieS in Mycology 105: 23–203 (2023).
24 Hou et al. INTRODUCTION The Hypocreales (Pezizomycotina, Ascomycota) is one of the largest orders of the class Sordariomycetes (Hyde et al. 2020b), currently comprising approximately 300 genera in 14 families, with some unresolved genera, such as Bulbithecium, Hapsidospora and Stanjemonium that are regarded as incertae sedis (Lumbsch & Huhndorf 2007, Hyde et al. 2020b). For many years the order Hypocreales was considered to consist of a single family, Hypocreaceae (Rogerson 1970). However, molecular phylogenetic studies suggested that the Clavicipitales should be included within the Hypocreales, and at least three major lineages were recognised, namely the Clavicipitaceae, Hypocreaceae, and Nectriaceae (O’Donnell 1993, Spatafora & Blackwell 1993, 1994, Glenn et al. 1996, Gams et al. 1998). Rossman et al. (1999) examined all available type specimens of the types in 199 genera that were considered to belong in Hypocreales, and two additional families were recognised, i.e. the Bionectriaceae that includes the Bionectria clade distinguished by Rehner & Samuels (1995), and the Niessliaceae for which no molecular data were available at the time. Afterwards, Calcarisporiaceae, Cocoonihabitaceae, Cordycipitaceae, Flammocladiellaceae, Myrotheciomycetaceae, Ophiocordycipitaceae, Stachybotryaceae, Sarocladiaceae and Tilachlidiaceae were added to the order (Sung et al. 2007a, Crous et al. 2014, 2015a, 2018a,b, Lombard et al. 2015, Sun et al. 2017, Zhuang & Zeng 2017). The Bionectriaceae was once ranked as the largest family in Hypocreales, including 26 perithecial and five cleistothecial genera (Rossman et al. 1999). This family is characterised by having white, pale tan orange or brown, uniloculate, perithecial, rarely cleistothecial ascomata, generally not changing colour in KOH (Rossman et al. 1999, 2001). The most common hyphomycetous asexual morphs of Bionectriaceae are acremonium-like or gliocladium-like, characterised by having phialidic conidiophores, and unicellular, ellipsoid, fusoid or subfusoid, hyaline to greenish conidia (Rossman et al. 1999). Many of the genera now recognised in the Bionectriaceae are based on species initially described in the broadly conceived genus Nectria (Rossman et al. 1999, 2001). With the addition of several recently introduced genera, namely Geonectria, Laniatria, Lasionectriella, Paracylindrocarpon, Periantria, Stromatonectria, Verrucostoma and Xanthonectria (Hirooka et al. 2010, Jaklitsch & Voglmayr 2011, Crous et al. 2016b, Lechat & Fournier 2016b, Lechat et al. 2016a, 2018b, Döbbeler & Davison 2017), and genera that were formerly regarded as incertae sedis (Crous et al. 2016a, Lin et al. 2016), the Bionectriaceae was consolidated in recent studies with 39 genera currently accepted (Wijayawardene et al. 2018). The genus Acremonium was first described by Link (1809) for a new species that was considered to produce single spores at the ends of its fertile cells. This name consists of “acro-” which means situated at the top, and “mono-” which means single (Link 1809, Summerbell & Scott 2015). Gams (1968) examined Link’s fungarium material and found that the type, A. alternatum, does not produce single conidia, but has conidia arranged in chains from thin, tapering phialides. The genus Acremonium has always been one of the most simply structured fungi of all filamentous asexual fungi. Characteristics of the genus include the production of septate hyphae giving rise to narrow, tapered, mostly lateral fusisporum L.W. Hou, L. Cai & Crous, Sarocladium agarici L.W. Hou, L. Cai & Crous, S. citri L.W. Hou, L. Cai & Crous, S. ferrugineum L.W. Hou, L. Cai & Crous, S. fuscum L.W. Hou, L. Cai & Crous, S. theobromae L.W. Hou, L. Cai & Crous; Valsonectriaceae: Valsonectria crystalligena L.W. Hou, L. Cai & Crous, V. hilaris L.W. Hou, L. Cai & Crous. New combinations: Bionectriaceae: Acremonium purpurascens (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous, Bulbithecium arxii (Malloch) L.W. Hou, L. Cai & Crous, Bu. borodinense (Tad. Ito et al.) L.W. Hou, L. Cai & Crous, Bu. pinkertoniae (W. Gams) L.W. Hou, L. Cai & Crous, Bu. spinosum (Negroni) L.W. Hou, L. Cai & Crous, Emericellopsis exuviara (Sigler et al.) L.W. Hou, L. Cai & Crous, E. fimetaria (Pers.) L.W. Hou, L. Cai & Crous, E. fuci (Summerb. et al.) L.W. Hou, L. Cai & Crous, E. moniliformis (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, E. salmonea (W. Gams & Lodha) L.W. Hou, L. Cai & Crous, E. tubakii (Gams) L.W. Hou, L. Cai & Crous, Fusariella arenula (Berk. & Broome) L.W. Hou, L. Cai & Crous, Hapsidospora chrysogena (Thirum. & Sukapure) L.W. Hou, L. Cai & Crous, H. flava (W. Gams) L.W. Hou, L. Cai & Crous, H. globosa (Malloch & Cain) L.W. Hou, L. Cai & Crous, H. inversa (Malloch & Cain) L.W. Hou, L. Cai & Crous, Hydropisphaera aurantiaca (C.A. Jørg.) L.W. Hou, L. Cai & Crous, Lasionectria atrorubra (Lechat & J. Fourn.) L.W. Hou, L. Cai & Crous, L. bisepta (W. Gams) L.W. Hou, L. Cai & Crous, L. castaneicola (Lechat & Gardiennet) L.W. Hou, L. Cai & Crous, L. cerealis (P. Karst.) L.W. Hou, L. Cai & Crous, L. olida (W. Gams) L.W. Hou, L. Cai & Crous, Lasionectriopsis dentifera (Samuels) L.W. Hou, L. Cai & Crous, Lasionectriella arenuloides (Samuels) L.W. Hou, L. Cai & Crous, La. marigotensis (Lechat & J. Fourn.) L.W. Hou, L. Cai & Crous, Monohydropisphaera fusigera (Berk. & Broome) L.W. Hou, L. Cai & Crous, Musananaesporium tectonae (R.F. Castañeda) L.W. Hou, L. Cai & Crous, Mycocitrus zonatus (Sawada) L.W. Hou, L. Cai & Crous, Nectriopsis microspora (Jaap) L.W. Hou, L. Cai & Crous, Ovicillium asperulatum (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, O. variecolor (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, Paracylindrocarpon multiloculatum (Samuels) L.W. Hou, L. Cai & Crous, Pn. multiseptatum (Samuels) L.W. Hou, L. Cai & Crous, Paragliomastix chiangraiensis (J.F. Li et al.) L.W. Hou, L. Cai & Crous, Px. luzulae (Fuckel) L.W. Hou, L. Cai & Crous, Px. znieffensis (Lechat & J. Fourn.) L.W. Hou, L. Cai & Crous, Protocreopsis rutila (W. Gams) L.W. Hou, L. Cai & Crous, Proxiovicillium blochii (Matr.) L.W. Hou, L. Cai & Crous, Stanjemonium dichromosporum (Gams & Sivasith.) L.W. Hou, L. Cai & Crous, Verruciconidia persicina (Nicot) L.W. Hou, L. Cai & Crous, Ve. verruculosa (W. Gams & Veenb.-Rijks) L.W. Hou, L. Cai & Crous, Waltergamsia citrina (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. dimorphospora (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. epimycota (Samuels) L.W. Hou, L. Cai & Crous, W. fusidioides (Nicot) L.W. Hou, L. Cai & Crous, W. hennebertii (W. Gams) L.W. Hou, L. Cai & Crous, W. parva (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. pilosa (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. zeylanica (Petch) L.W. Hou, L. Cai & Crous; Cephalothecaceae: Phialemonium thermophilum (W. Gams & J. Lacey) L.W. Hou, L. Cai & Crous; Clavicipitaceae: Subuliphorum camptosporum (W. Gams) L.W. Hou, L. Cai & Crous; Coniochaetaceae: Coniochaeta psammospora (W. Gams) L.W. Hou, L. Cai & Crous; Nothoacremoniaceae: Nothoacremonium exiguum (W. Gams) L.W. Hou, L. Cai & Crous; Neoacremoniaceae: Neoacremonium minutisporum (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous; Ne. taiwanense (K.L. Pang et al.) L.W. Hou, L. Cai & Crous; Ne. vitellinum (W. Gams) L.W. Hou, L. Cai & Crous; Plectosphaerellaceae: Allomusicillium domschii (W. Gams) L.W. Hou, L. Cai & Crous, Brunneomyces pseudozeylanicus (W. Gams) L.W. Hou, L. Cai & Crous; Pseudoniessliaceae: Pseudoniesslia minutispora (W. Gams et al.) L.W. Hou, L. Cai & Crous; Sarocladiaceae: Chlamydocillium curvulum (W. Gams) L.W. Hou, L. Cai & Crous, Parasarocladium funiculosum (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous; Valsonectriaceae: Valsonectria inflata (C.H. Dickinson) L.W. Hou, L. Cai & Crous, V. roseola (G. Sm.) L.W. Hou, L. Cai & Crous. Epitype (basionym): Sphaeria violacea J.C. Schmidt ex Fr. Neotype (basionym): Mastigocladium blochii Matr. Citation: Hou LW, Giraldo A, Groenewald JZ, Rämä T, Summerbell RC, Zang P, Cai L, Crous PW (2023). Redisposition of acremonium-like fungi in Hypocreales. Studies in Mycology 105: 23–203. doi: 10.3114/sim.2023.105.02 Received: 18 October 2022 ; Accepted: 16 May 2023; Effectively published online: 2 June 2023 Corresponding editor: Robert A. Samson
25www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales phialides with unicellular conidia arranged in mucoid heads or unconnected chains, and differentiated conidiophores with or without verticillate branches may be observed in some species (Gams 1971, 1975, Domsch et al. 2007, Perdomo et al. 2011, Summerbell et al. 2011). Species of Acremonium sensu lato (s. lat.) are cosmopolitan and distributed throughout a broad range of environments. They are mainly soilborne saprobes or weak to virulent, facultative or obligate pathogens of plants, other fungi, animals or humans (Gams 1971, 1975, Alfaro-García et al. 1996, Novicki et al. 2003, Lin et al. 2004, Zuccaro et al. 2004, Domsch et al. 2007, Perdomo et al. 2011, Mirtalebi et al. 2017, Summerbell et al. 2018, de Hoog et al. 2020, Pérez-Cantero & Guarro 2020, Kim et al. 2021). They are regarded of great importance in agro-forestry, food storage and preservation, industry, clinical mycology and pharmaceutical industries. Acremonium species have proven to be rich sources of novel and bioactive secondary metabolites. To date more than 350 metabolites have been isolated from Acremonium spp. (Tian et al. 2017). Although most of these metabolites are known from saprobic species, an increasing number of interesting metabolites have also been reported from endophytic or marine-derived fungi. For example, Sarocladium strictum (basionym A. strictum) derived from the ocean, is known to produce beta-lactam antibiotic cephalosporins that have been marketed (Burton & Abraham 1951, Gams 1971, Hamilton-Miller 2000). Some species of Acremonium are effective biological control agents for fungal plant pathogens (Auer & Ludwig-Müller 2014, Bobeck & Pearce 2017, Sutton & Mason 2017), or help to protect plants against environmental stress, hence promoting growth and productivity (Bobeck & Pearce 2017, Sutton & Mason 2017). Some Acremonium species are also commonly mentioned as being among the spoiling microorganisms on diverse foods, including fish meals, fruits, noodles, nuts, peas, apples and stored wheat (Gams 1971, Abdel-Hafez 1987, Fernández-Trujillo 1997, Fujikawa 1997, Summerbell & Scott 2015, Summerbell et al. 2018). In addition, species of Acremonium are highly relevant in the clinical field. Patients commonly develop onychomycoses, or following traumatic inoculation of the fungus infections, resulting in fungemia, ocular infections (keratitis), cutaneous and subcutaneous infections and mycetoma (Gupta et al. 2000, Perdomo 2011, Summerbell et al. 2018, Pérez-Cantero & Guarro 2020). Locally invasive infections such as arthritis, osteomyelitis, peritonitis, sinusitis, and less frequently central nervous system infections have also been frequently reported in recent years (Guarro et al. 1997, 2009, Gupta et al. 2000, Das et al. 2010, de Hoog et al. 2020, Pérez-Cantero & Guarro 2020). However, due to the reduced and little-differentiated morphology, the wide ecological distribution and substrate range, as well as the overlapping features among species, species identification in Acremonium is particularly difficult and the names are frequently misapplied. Although the morphology of the asexual morph is relatively plesiomorphic, Acremonium is perceived to be a heterogeneous taxon because large numbers of morphologically distinct sexual genera of ascomycetes have acremonium-like asexual morphs. Most of the sexuallytypified Acremonium members were identified as Nectria species (Gams 1971, Samuels 1973, 1976a, b, Lowen 1995), but the many genera of Hypocreales known from their sexual morphs, such as Emericellopsis, Epichloe, Hydropisphaera, Hypocrea, Hypomyces, Ijuhya, Lasionectria, Mycoarachis, Nectriopsis, Nigrosabulum, Ochronectria, and Pronectria (Malloch & Cain 1970, Gams 1971, Morgan-Jones & Gams 1982, Samuels 1988, Lowen 1995, Rossman et al. 1999), even Chaetomium and Thielavia of Sordariales, are also characterised by the production of acremonium-like asexual morphs (Morgan-Jones & Gams 1982). The main morphotaxonomic groundwork for Acremonium was conceived in the late 20th century by Gams (1971) in his monograph “Cephalosporium-artige Schimmelpilze (Hyphomycetes)”. A total of 82 Acremonium species were studied based on a meticulous morphological observation scheme. In this concept species of the genus Acremonium were classified into three sections based on their morphological characters: sect. Simplex (rarely branched conidiophores that mostly developed as orthophialides); sect. Gliomastix (darkly pigmented conidia or hyaline conidia and the appearance of chondroid hyphae, which make the cultures tough and difficult to cut through); sect. Nectrioidea (conidiophores that are basitonously branched several times, or unbranched but with multiple septa, mostly chromophilic and granular at the base, with small collarette at the tip) (Gams 1971). Later, two additional sections were proposed, sect. Albolanosa (grass endophytes), and sect. Chaetomioidea (short-aculeate to lageniform phialides with sexual morph in Chaetomiaceae) (Morgan-Jones & Gams 1982). Lowen (1995) added the section Lichenoidea, in which most of the lichenicolous species are included. However, due to the little-differentiated morphology and overlapping features among different sections, species classification is particularly difficult, and the classifications proposed by Gams (1971) seems to be artificial and to not reflect the natural evolutionary history of this group. The first major molecular phylogenetic study of Acremonium was carried out by Glenn et al. (1996), based on partial nuclear ribosomal small subunit sequences (SSU). Results of the preliminary phylogenetic study demonstrated that Acremonium was highly polyphyletic and recognised Acremonium members of the five sections in at least three distinct orders of Sordariomycetes, including Hypocreales, Microascales and Sordariales (Glenn et al. 1996). As a consequence, members of Acremonium were allocated to different families: type of the sections Acremonium, Nectrioidea, and Gliomastix resided in Hypocreaceae of Hypocreales; the clavicipitaceous grass endophytes of sect. Albolanosa were moved to the newly established genus Neotyphodium based on their unique morphology, ecology and obligate parasitism of the Clavicipitaceae; and the section Chaetomioidea which has Thielavia and Chaetomium sexual morphs grouped within Sordariales, and were thus excluded from Acremonium (Glenn et al. 1996). However, the taxonomic status of Acremonium s. lat. remained unresolved. With the introduction of more taxa to the dataset and additional genetic loci to phylogenetic studies, subsequent research further confirmed that Acremonium is highly polyphyletic. A molecular phylogenetic overview based on SSU and LSU sequences was generated, which grouped over 100 species of Acremonium and related taxa into five groups across different classes of Ascomycota (Coniochaetales, Hypocreales, Microascales, Sordariales and Cephalothecaceae; Summerbell et al. 2011). The bulk of Acremonium species falls into the Hypocreales. Epitypification of the type, A. alternatum, linked Acremonium sensu stricto to the Bionectriaceae (Hypocreales; Summerbell et al. 2011), which also accommodates several other, sexually typified genera with an acremonium-like asexual morph, such as Bulbithecium, Emericellopsis, Hapsidospora, Mycoarachis and Nigrosabulum (Gams 1971, Summerbell et al. 2011). However, the genus Acremonium still proved to be polyphyletic distributed in two major clades with at least 20 smaller clades across Hypocreales, including the Sarocladium, “curvulum-clade”, and “breve-clade”, that led to the proposal that the remaining acremonium-like species
26 Hou et al. should be allocated to other genera or families (Summerbell et al. 2011). Nine of the named Acremonium species in these analyses belong to the Plectosphaerellaceae (Microascales; Summerbell et al. 2011), which were recently revised by Giraldo & Crous (2019). The other species belonging to the Sordariales, Coniochaetales and the Cephalothecaceae had also been revised in subsequent studies (Perdomo et al. 2013). From these results, it became evident that the acremonium-like morphology had evolved multiple times. Therefore, the taxonomy and phylogeny of the acremoniumlike taxa urgently required revision. Multiple gene markers had been used for studies of acremonium-like species, including ITS, SSU, LSU. The rapidly evolving ribosomal internal transcribed spacer (ITS) is difficult to align in Acremonium as the genus is highly heterogeneous and distributed across the ascomycetes (de Hoog et al. 2000). However, ITS makers are alignable and frequently used in restricted Acremonium subgroups (Summerbell et al. 2011). The relatively slowly evolving genes, such as partial nuclear ribosomal small subunit RNA gene (SSU) and the partial nuclear ribosomal large subunit RNA gene (LSU), have the advantage of being alignable among a broad range of Acremonium groups, but always result in unresolved relationships (Summerbell et al. 2011). Combined ITSLSU or SSU-LSU have also been used in studies of acremoniumlike species and related sexual genera (Summerbell et al. 2011, Giraldo et al. 2012, Lechat et al. 2016), but still failed to provide sufficiently clear genera and species boundaries for acremoniumlike species. In recent years, protein-coding genes were frequently applied in the phylogenetic analysis for a few acremonium-like groups. The rpb2, tef-1α and tub2 genes were used for studies of Emericellopsis species (Giraldo et al. 2017, Gonçalves et al. 2020) Hagestad et al. 2021, ITS and act1 sequences were used in Parasarocladium and Sarocladium species (Giraldo et al. 2015, Gonçalves et al. 2020), and the combined ITS, LSU, rpb2 and tef1α were used for the analysis of Acremonium species belonging to the Plectosphaerellaceae (Giraldo & Crous 2019). These results provided a clearer picture of relationships among the Acremonium groups that were imperfectly resolved in LSU and SSU analyses. Given the importance of the genus Acremonium to agriculture, industry, and medicine, four objectives were defined in the present study: 1) to resolve the phylogenetic and taxonomic placement of species of Acremonium s. lat. across different genera, families and orders within Ascomycota; 2) to clarify the circumscription of Acremonium s. str. by updating and expanding currently available DNA sequence datasets; 3) to assign names to presently undescribed taxa based on morphological observations, and revisit the old species circumscriptions and reconsider their taxonomy based on a re-examination of original material and DNA sequence data; and 4) to construct a phylogenetic overview that delineates the phylogenetic lineages and generic boundaries of the Bionectriaceae via a polyphasic approach. MATERIALS AND METHODS Isolates All the cultures of acremonium-related fungi included in this study were obtained from the culture collection (CBS) of the Westerdijk Fungal Biodiversity Institute (WI) in Utrecht, The Netherlands, the working collection of Pedro W. Crous (CPC), housed at the Westerdijk Institute, the working collection of Teppo Rämä (TR) at UiT the Arctic University of Norway, the CABI Genetic Resource Collection in the UK (IMI), the Canadian Collection of Fungal Cultures (DAOMC) and the BIOTEC Culture Collection in Thailand (BCC). Most of these cultures were identified as species in the genus Acremonium or associated genera according to the morphological study of Gams (1971) and the study of Summerbell et al. (2011) (Supplementary Table S1). Representative cultures of the new species described in this study were deposited in the CBS culture collection. DNA extraction, PCR amplification and sequencing Total genomic DNA was extracted from fungal colonies growing on malt extract agar (MEA) or oatmeal agar (OA; Crous et al. 2019) using the Wizard® Genomic DNA purification kit (Promega, Madison, USA), according to the manufacturers’ protocols. Four loci were amplified, including the internal transcribed spacer regions 1 and 2 and 5.8S nuclear ribosomal RNA gene (ITS), partial large subunit nrRNA gene (28S nrDNA; LSU), the protein-coding genes translation elongation factor 1-alpha (tef-1α) and partial DNA-directed RNA polymerase II second largest subunit (rpb2) gene with the primer pairs ITS5/ITS4 (White et al. 1990, de Hoog & Gerrits van den Ende 1998), LR0R/LR5 (Vilgalys & Hester 1990, Vilgalys & Sun 1994), EF983F/EF-2218R (Rehner & Buckley 2005), and RPB2-5F2/RPB27cR (Liu et al. 1999, Sung et al. 2007b), respectively. The PCR amplifications were performed in a total volume of 25 μL containing 2.5 μL 10× EasyTaq Buffer (Bioline, Luckenwalde, Germany), 50 μM dNTPs, 0.1 μM of each primer, 0.75 U Taq DNA polymerase and 1–10 ng genomic DNA. The PCR amplifications of ITS, LSU and tef1α were set as follows: an initial denaturation at 95 °C for 5 min, followed by 35 cycles of denaturation, annealing and extension, and a final extension step at 72 °C for 10 min. For the LSU amplification, the 35 cycles consisted of 45 s at 95 °C, 45 s at 48 °C and 2 min at 72 °C; for the ITS 30 s at 95 °C, 30 s at 48 °C and 80 s at 72 °C; and for the tef-1α region 30 s at 95 °C, 30 s at 52 °C and 80 s at 72 °C. The procedures for amplifying and sequencing the rpb2 were performed as described in Hou et al. (2020). PCR products for four loci were purified and sequenced in both directions using an Applied Biosystems 3730xl DNA Analyzer (Thermo Fisher Scientific) as explained in Crous et al. (2013). The consensus sequences of each culture were assembled from forward and reverse sequences using Seqman Pro v. 10.0.1 (DNASTAR, Madison, USA). Novel sequences generated in this study were deposited in GenBank (http://www.ncbi. nlm.nih.gov, Supplementary Table S1). Phylogenetic analyses Sequence alignments of the four individual loci (LSU, ITS, rpb2, tef-1α) were generated with MAFFT v. 7 using the default settings (http://mafft.cbrc.jp/alignment/server/index.html) and were then manually edited in MEGA v. 7.0.21 (Kumar et al. 2015). Both Maximum Likelihood (ML) and Bayesian analysis (BA) were used for phylogenetic inferences of individual sequence alignments, followed by the concatenated alignments. The best substitution model of evolution for each of the four data partitions were estimated using jModeltest v. 2.1.4 (Darriba 2012) before the Bayesian analyses. Bayesian analyses were performed using MrBayes v. 3.2.6 (Ronquist et al. 2012) as described by Hou et al. (2020). Markov Chain Monte Carlo sampling (MCMC) analyses of four chains were started in parallel from a random tree topology. Four simultaneous Markov chains were run for 10 M generations with a sampling frequency set to the 1 000th generation (resulting in 10 000 total trees per parallel run) or until the run was stopped automatically when the average standard deviation of
27www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales split frequencies fell below 0.01. The burn-in fraction was set to 0.25, and the remaining trees were used to calculate posterior probabilities (Chen et al. 2015). Maximum Likelihood analyses include 1 000 bootstrap replicates and were conducted using RAxML v. 7.2.6 (Stamatakis 2014). A general time reversible (GTR) model was applied with a gamma-distributed rate variation. The resulting trees were viewed using FigTree v. 1.4.2 (http://tree.bio. ed.ac.uk/software/figtree). Alignments and the phylogenetic trees derived from this study were uploaded to figshare (doi: 10.6084/ m9.figshare.22258765). Morphology Cultures were cultivated on fresh oatmeal agar (OA), malt extract agar (MEA), potato dextrose agar (PDA) and synthetic nutrientpoor agar (SNA; Crous et al. 2019) and incubated at 25 °C in the dark for 4 wk. Colony diameter and characters were measured after 14 d of incubation. Colony colours (upper surface and reverse) were rated following the colour charts of Rayner (1970). Micromorphological observations of reproductive structures were carried out from cultures grown on OA after 14 d of incubation. Clear lactic acid was used as mounting medium for the observation of micromorphological structures of mature ascomata/conidiomata, ascospores/conidia and asci/conidiophores (Gams 1971). Observations were performed with a Nikon Eclipse 80i compound microscope with differential interference contrast (DIC) optics, and a Nikon AZ100 dissecting microscope. Photomicrographs and measurements were taken with a Nikon DS-Ri2 high-definition colour digital camera using the NIS-elements D software v. 4.50. At least 30 measurements were made for all morphologically informative features. Descriptions of novelties and taxonomic recombinations were deposited in MycoBank, and the fungarium specimens for the novel taxa represented by dried sporulating cultures were deposited in the CBS Fungarium, Utrecht, The Netherlands. RESULTS Phylogeny A total of 922 isolates resembling Acremonium spp. were included in this study. An overview phylogeny based on ITS, LSU and rpb2 sequences was conducted in order to position the isolates in the treated order, family and genera (Dataset 1). In order to portray more precise phylogenetic relationships of Acremonium and its related species, more inclusive analyses based on DNA sequence data from four loci (ITS, LSU, rpb2 and tef-1α) were carried out for three focused families from the overview tree separately (datasets 2–4), including all available cultures belonging to these families. Datasets 2–4 were analysed with the same phylogenetic methods applied to Dataset 1. Dataset 1 consisted of a concatenated alignment of three loci (ITS, LSU, and rpb2) that contained 392 taxa to represent several families belonging to the Hypocreales and its related orders, which was used to infer delimitation at the family and order levels. Saccharata proteae (CBS 115206; Saccharataceae, Botryosphaeriales, Dothideomycetes) was used as outgroup (Fig. 1). The final alignment of ITS, LSU and rpb2 contained a total length of 2 770 characters including alignment gaps (gene boundaries ITS: 1–1 071, 1 071 bp; LSU: 1 072–1 889, 818 bp; rpb2: 1 890– 2 770, 881 bp). Among those, 858 characters were conserved sites and 1 794 were variable sites, including 287 characters that were parsimony-uninformative and 1 507 characters that were parsimony informative. According to the result of jModeltest, a GTR+I+G model was proposed for Bayesian analysis of ITS, LSU and rpb2. The Bayesian analysis of the concatenated three-locus alignment lasted for 9 885 000 generations and a total of 14 852 trees were generated after the BI analysis reached the stop value of 0.01, with the first 25 % of trees discarded as the burn-in phase. The posterior probabilities (PP) were calculated from the remaining trees. The ML tree confirmed the same tree topology and the clades as those presented in the Bayesian phylogeny. Therefore, only the ML tree based on the combined dataset was presented here with the bootstrap support values of the ML analysis (MLBS) and relevant Bayesian posterior probabilities (BPP) shown at the nodes (Fig. 1; MLBS > 50 %, BPP > 0.90 shown). Five main supported clades were represented in the tree (Fig. 1), corresponding to four known orders: Cephalothecales, Coniochaetales, Glomerellales, Hypocreales and the outgroup clade. A total of 21 subclades were recognised in the strongly supported clade representing the Hypocreales (clades A–V), of which 16 represented existing families, and five are proposed here as new, Nothoacremoniaceae (clade I), Neoacremoniaceae (clade K), Chrysonectriaceae (clade L), Pseudoniessliaceae (clade M), and Valsonectriaceae (clade U). The Clade A (BPP < 0.9, MLBS = 90 %), representing those species classified in Clavicipitaceae, contains a new genus Subuliphorum proposed for a species with curved conidia, S. camptosporum (basionym: Acremonium camptosporum). The Clade G & Clade H comprised species of the genera Acremoniopsis, Collarina, Cylindromonium, Eucasphaeria, Myrtacremonium, Neoeucasphaeria, Niesslia, Phialoseptomonium, Rosasphaeria, Trichonectria, and Trichosphaerella representing the family Niessliaceae. Several strains previously recognised as Acremonium and Cephalosporium species also clustered in Niessliaceae, including ex-type strain of A. guillematii (CBS 766.69), A. cavaraeanum (CBS 101149), A. incrustatum (CBS 159.70), A. nigrosclerotium (CBS 154.72) and Cephalosporium ballagii (CBS 134.33). However, phylogeny of Niessliaceae was not well resolved and several genera remain polyphyletic, i.e. Niesslia, Cylindromonium, Trichonectria. Therefore, because of the unresolved phylogeny of Niessliaceae, the taxonomy of these Acremonium and Cephalosporium taxa has not been resolved. The new family Nothoacremoniaceae (Clade I; BPP = 1, MLBS = 100 %) grouped in one clade, which corresponds to the type genus Nothoacremonium. This clade encompassed the type species No. exiguum (basionym: A. exiguum) and two new species with acremonium-like asexual morphs, No. subcylindricum and No. vesiculophorum. The clade K (BPP = 1, MLBS = 95 %), which consists of several isolates previously recognised in Acremonium, is proposed as the new family Neoacremoniaceae. The type genus Neoacremonium is proposed for Ne. flavum, Ne. minutisporum (basionym: Cephalosporium minutisporum), Ne. taiwanense (basionym: Sedecimiella taiwanensis), Ne. vitellinum (basionym: A. vitellinum), and the type species Ne. distortum. The ex-type strain of Parapyrenis maritima (CBS 538.93) together with the strain CBS 795.69 that previously recognised as “A. minutisporum”, formed a subclade in Neoacremonium. The taxonomy and phylogeny of Parapyrenis maritima awaits further confirmation. The new family Chrysonectriaceae (clade L; BPP = 1, MLBS = 100 %) comprises the type species of Chrysonectria (C. finisterensis) and a novel species C. crystallifera.
28 Hou et al. Metarhizium huainamdangense BCC 44270 Trichoderma deliquescens CBS 121131 T Hypomyces lactifluorum TAAM 170476 Nigelia aurantiaca BCC 19475 Trichoderma sp. CBS 123792 Metacordyceps pseudoatrovirens TNS-F16380 Cocoonihabitus sinensis HMAS 254524 Claviceps paspali ATCC 13892 Subuliphorum camptosporum CBS 890.85 Subuliphorum camptosporum CBS 835.91 Metacordyceps atrovirens TNM-F10184 Ophiocordyceps coenomyia NBRC 106964 Trichoderma spinulosum CBS 121272 Trichoderma spinulosum CBS 121280 Ophiocordyceps acicularis OSC 110987 Cocoonihabitus sinensis HMAS 254523 Trichoderma aerugineum CBS 120541 T Tolypocladium paradoxum NBRC 106958 Metarhizium chaiyaphumense BCC 19021 Subuliphorum camptosporum CBS 756.69 T Metapochonia gonioides CBS 891.72 T Metapochonia rubescens CBS 464.88 T Claviceps purpurea AEG 97-2 Subuliphorum camptosporum CBS 757.69 Metarhizium huainamdangense BCC 32190 Metarhizium huainamdangense BCC 7672 Tolypocladium album CBS 393.89 Metarhizium takense BCC 30940 Pochonia chlamydosporia CBS 504.66 Trichoderma britannicum CBS 253.62 T Metarhizium guizhouense CBS 258.90 T Conoideocrella luteorostrata NHJ 11343 Ophiocordyceps heteropoda NBRC 100642 Hypomyces australasiaticus CBS 127152 T Metapochonia suchlasporia CBS 248.83 Orbiocrella petchii NHJ 6240 Metapochonia suchlasporia CPC 26469 Subuliphorum camptosporum CBS 677.74 Tolypocladium ophioglossoides NBRC 8992 Metarhizium pingshaense CBS 257.90 T Claviceps purpurea SA cp11 Trichoderma viride CBS 119325 T Claviceps fusiformis ATCC 26019 Ophiocordyceps acicularis OSC 110988 Conoideocrella tenuis NHJ 6293 Nigelia aurantiaca BCC 19950 Metapochonia bulbillosa CBS 145.70 Purpureocillium takamizusanense BCC 49261 Trichoderma spinulosum CBS 311.50 Metapochonia suchlasporia CBS 251.83 Pochonia chlamydosporia CBS 101244 1/61 1/100 1/100 0.95/50 0.75/83 1/100 1/90 1/100 1/88 1/100 1/97 1/93 1/100 1/100 0.96/99 0.85/93 1/100 1/98 1/85 -/59 1/88 -/90 0.92/74 1/100 -/83 1/99 -/73 1/100 1/96 1/100 1/98 1/100 1/- 1/100 1/97 1/99 A. Clavicipitaceae B. Hypocreaceae D. Ophiocordycipitaceae C. Cocoonihabitaceae Hypocreales 0.3 Fig. 1. Phylogenetic tree inferred from a Maximum Likelihood analysis based on a concatenated alignment of LSU, ITS and rpb2 sequences of 392 strains representing Hypocreales and related orders (Cephalothecales, Coniochaetales, Glomerellales). The RAxML bootstrap support values (MLBS) above 50 % and Bayesian posterior probabilities (BPP) above 0.90 are given at the nodes (BPP/MLBS). Some of the basal branches were shortened to facilitate layout. The scale bar represents the expected number of changes per site. Families are delimited in coloured boxes, with the family name indicated to the right. Strains with special status are indicated with a superscript letter after the accession number (T: ex-type). New species are printed in red font and new combinations in blue font. The tree is rooted to Saccharata proteae CBS 115206.
29www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Calcarisporium cordycipiticola CGMCC 3.17905T Niesslia exilis CBS 357.70 Myrtacremonium eucalypti CBS 142161T Beauveria bassiana ARSEF 2567T Akanthomyces sulphureus TBRC 7248T Niesslia xanthorrhoeae CBS 287.93T Simplicillium lanosoniveum CBS 531.72 Niesslia mucida CBS 404.66T Niesslia rhizomorpharum CBS 642.85T Calcarisporium arbusculum CBS 900.68T Neoeucasphaeria eucalypti CBS 145075T Niesslia grisescens CBS 599.88T Lecanicillium muscarium CBS 143.62T Lecanicillium lecanii CBS 101247 Cordyceps farinosa CBS 111113T Simplicillium lanosoniveum MFLUCC 18-1385 Eucasphaeria capensis CBS 120027T Niesslia tenuis CBS 113275 “Cephalosporium ballagii” CBS 134.33T Simplicillium lamellicola CBS 101912 Cordyceps fumosorosea CBS 107.10T “Acremonium nigrosclerotium” CBS 154.72T Niesslia ilicifolia CBS 460.74 Niesslia loricata CBS 778.69T Eucasphaeria rustici CBS 142085T Simplicillium lamellicola CBS 116.25T Niesslia artocarpi CBS 582.73T Niesslia exilis CBS 358.70 Niesslia exilis CBS 426.66 Simplicillium lanosoniveum CBS 123.42T Niesslia clarkii CBS 170.74T Beauveria bassiana CBS 534.81 Niesslia ligustica CBS 684.95T Niesslia tenuis CBS 203.70 “Acremonium potronii” CBS 443.77 Akanthomyces waltergamsii TBRC 7252T Rosasphaeria moravica CBS 124270T Calcarisporium arbuscula VB3 Niesslia cladoniicola CBS 960.73T Niesslia ilicifolia CBS 459.74 Simplicillium lamellicola CBS 101911 Lecanicillium lecanii CBS 102067T Eucasphaeria capensis CBS 120028 Calcarisporium cordycipiticola CGMCC 3.17904 Cordyceps javanica CBS 134.22T Niesslia aeruginosa CBS 264.89T Simplicillium lanosoniveum CBS 321.72 Simplicillium lanosoniveum CBS 322.72 Niesslia tenuis CBS 432.66T 1/96 0.99/- 1/100 1/85 1/97 1/100 1/58 1/100 1/100 1/98 1/100 1/100 1/100 0.97/67 1/100 0.99/- 0.99/- 1/100 1/100 1/100 1/100 -/56 0.99/71 1/100 1/100 0.99/74 1/- -/58 1/100 1/100 0.84/63 1/96 1/68 1/100 0.98/53 1/100 1/100 E. Cordycipitaceae F. Calcarisporiaceae G. Niessliaceae Hypocreales 0.3 Fig. 1. (Continued).
30 Hou et al. Fusarium subluratum CBS 189.34T “Acremonium persicinum” CBS 330.80 Phialoseptomonium eucalypti CBS 145542T Xenocylindrocladium subverticillatum CBS 113660T Neocosmospora vasinfecta CBS 517.71 Ilyonectria capensis CBS 132815T Cylindromonium rhabdosporum CBS 438.66 Thelonectria olida CBS 215.67T Trichonectria setadpressa AF29617 Mariannaea samuelsii CBS 125515T Cylindrocladiella australiensis CBS 129567T “Acremonium cavaraeanum” CBS 101149T Neocosmospora illudens CBS 126406 Nothoacremonium exiguum CBS 587.73T Penicillifer pulcher CBS 560.67T Thelonectria discophora CBS 125153 Nothoacremonium subcylindricum CBS 416.68T Ilyonectria destructans CBS 264.65 “Acremonium persicinum” CBS 110646 Cylindrocladiella elegans CBS 338.92 Penicillifer penicilliferi CBS 423.88T Trichosphaerella ceratophora CBS 130.82 Neonectria neomacrospora CBS 198.62 “Acremonium potronii” CBS 589.90 Fusarium venenatum CBS 458.93T Neocosmospora illudens CBS 119605 “Acremonium potronii” CBS 433.88 Acremoniopsis suttonii CBS 138708T Dactylonectria estremocensis CBS 129085T Nothoacremonium subcylindricum CBS 611.95 Cylindromonium lichenicola CBS 188.70 “Acremonium guillematii” CBS 766.69T Collarina aurantiaca CBS 138274T Neocosmospora vasinfecta CBS 562.70 Trichonectria setadpressa AF28886 Dactylonectria macrodidyma CBS 112615T Trichonectria rectipila CBS 132.87T Mariannaea punicea CBS 105.66 Monocillium griseo-ochraceum CBS 683.71T Nothoacremonium subcylindricum CBS 781.69 Nothoacremonium vesiculophorum CBS 397.70BT Nothoacremonium subcylindricum CBS 190.70 “Acremonium guillematii” CBS 110650 Neonectria tsugae CBS 788.69T “Acremonium cavaraeanum” CBS 111656 Mariannaea punicea CBS 239.56T Cylindrocladiella hawaiiensis CBS 129569T “Acremonium incrustatum” CBS 159.70T Xenocylindrocladium serpens CBS 128439T Neocosmospora vasinfecta CBS 325.54 Cylindromonium lichenicola CBS 303.70 Cylindromonium eugeniicola CBS 146075T 1/97 0.99/80 0.99/57 1/95 1/100 1/100 1/- 1/100 1/100 1/100 1/100 1/100 1/56 1/100 1/83 1/100 1/85 1/100 1/61 1/53 1/100 1/86 0.93/49 1/100 1/98 1/100 1/75 -/75 1/95 1/100 1/90 1/100 1/100 1/100 1/100 1/65 1/100 0.99/67 1/100 1/100 1/100 1/100 I. Nothoacremoniaceae H. Niessliaceae J. Nectriaceae Hypocreales 0.3 Fig. 1. (Continued).
31www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Cosmospora berkeleyana CBS 239.70 Paracremonium inflatum CBS 485.77 T Pseudonectria foliicola CBS 122566 Pseudocosmospora rogersonii CBS 133981 T Neoacremonium flavum CBS 452.70 T Paracremonium binnewijzendii CBS 143277 T Neoacremonium distortum CBS 291.70B Neoacremonium distortum CBS 710.73 Cosmospora khandalensis CBS 356.65 T Xenoacremonium brunneosporum CBS 362.76 Coccinonectria pachysandricola CBS 501.63 Cosmospora berkeleyana CBS 234.70 Xenoacremonium falcatum CBS 400.85 T Neoacremonium distortum CBS 706.73 Xenoacremonium allantoideum CBS 505.94 T Neoacremonium taiwanense CY5100 T Thyronectria pyrrhochlora CBS 125131 Coccinonectria rusci CBS 126108 T Nectria cinnabarina CBS 125165 T Volutella minima CBS 122767 “Parapyrenis maritima” CBS 795.69 Neoacremonium distortum CBS 315.72 Fusicolla violacea CBS 634.76 T Neoacremonium distortum CBS 314.72 T Pseudocosmospora eutypellae CBS 133966 T Thyronectria rhodochlora CBS 136006 T Nectria polythalama CBS 129240 T Neoacremonium vitellinum CBS 792.69 T Nectria polythalama CBS 128672 Nectria nigrescens CBS 125148 T Thyronectria rhodochlora NP4 Neoacremonium distortum CBS 146.72 Pseudonectria foliicola CBS 123190 T Sarcopodium circinatum CBS 587.92 Stylonectria wegeliniana CBS 125490 Neoacremonium distortum CBS 665.75 Paracremonium contagium CBS 110348 T “Parapyrenis maritima” CBS 538.93 T Neoacremonium minutisporum CBS 147.62 T Cosmospora butyri CBS 301.38 T “Tilachlidium brachiatum” CBS 397.66 Cosmospora berkeleyana CBS 235.70 Paracremonium variiforme LC5806 T Stylonectria applanata CBS 125489 Xenoacremonium recifei CBS 137.35 T Xenoacremonium brunneosporum MFLU 16-1204 T Paracremonium binnewijzendii CBS 698.86 Coccinonectria pachysandricola CBS 128674 Nectria polythalama CBS 117713 Neoacremonium flavum CBS 398.70 Neoacremonium distortum CBS 291.70A Sarcopodium flavolanatum CBS 128370 Volutella ciliata CBS 483.61 Neoacremonium vitellinum CBS 793.69 1/92 1/100 1/100 1/100 1/100 1/- 1/100 1/100 1/93 1/94 0.98/85 0.92/60 1/100 1/100 1/100 1/100 1/84 1/61 1/100 0.99/89 1/80 1/94 1/99 1/68 1/87 0.99/94 0.99/99 1/100 1/100 0.99/95 0.95/50 1/84 1/95 1/100 1/100 1/100 1/97 1/100 0.92/69 1/100 0.97/- 1/100 1/93 0.98/62 1/100 1/100 1/100 0.95/- 1/100 J. Nectriaceae K. Neoacremoniaceae Hypocreales 0.3 Fig. 1. (Continued).
38 Hou et al. Verruciconidia quercina CBS 355.77 Verruciconidia erythroxyli CBS 378.70D Verruciconidia persicina CBS 120889 Lasionectria cerealis TR055cI1.1 Lasionectria cerealis CBS 215.69 Lasionectria bisepta CBS 119908 Verruciconidia unguis CBS 378.70E Verruciconidia persicina CBS 439.66 Lasionectria bisepta CBS 753.69 Lasionectria bisepta CBS 750.69T Lasionectria antillana CBS 122797T Lasionectria cerealis TR055cII1.1 Lasionectria cerealis CBS 216.69 Verruciconidia verruculosa CBS 989.69T Verruciconidia persicina CBS 295.70A Lasionectria olida CBS 799.69T Verruciconidia quercina CBS 469.67T Lasionectria cerealis CBS 208.70 Verruciconidia persicina CBS 169.65 Verruciconidia verruculosa CBS 299.81B Verruciconidia unguis CBS 424.93T Lasionectria olida CBS 798.69 Verruciconidia persicina CBS 310.59T Verruciconidia siccicapita CBS 378.70AT Verruciconidia erythroxyli CBS 728.87T Lasionectria cerealis CBS 206.65 Lasionectria bisepta CBS 751.69 Verruciconidia persicina CBS 101712 Verruciconidia persicina CBS 116385 Lasionectria antillana CBS 114748 Lasionectria cerealis CBS 393.66 Verruciconidia persicina CBS 128826 Lasionectria castaneicola CBS 122792T Lasionectria cerealis CBS 461.88 Lasionectria cerealis CBS 227.66 Verruciconidia quercina CBS 183.78 Lasionectria cerealis CBS 207.70 Lasionectria cerealis CBS 207.65 Verruciconidia infuscata CBS 100888T Lasionectria mantuana A.R. 4029 Lasionectria mantuana CBS 114291 Lasionectria boothii CBS 129747 Lasionectria cerealis CBS 209.70 Lasionectria atrorubra CBS 123502T Lasionectria krabiense MFLUCC 15-0673T Lasionectria sylvana CBS 566.76 Lasionectria sp. CBS 102040 Verruciconidia persicina CBS 113716 Verruciconidia persicina CBS 218.96 Lasionectria bisepta CBS 752.69 Verruciconidia verruculosa CBS 990.69 Verruciconidia persicina CBS 127298 Verruciconidia persicina CBS 102349 Lasionectria cerealis CBS 144938 Verruciconidia persicina CBS 295.70B 1/91 1/100 1/100 1/100 1/75 1/100 1/98 1/100 1/96 1/93 1/97 1/100 1/100 1/100 1/87 1/100 1/100 1/100 1/94 1/88 1/99 1/79 1/83 1/99 1/96 1/90 1/100 -/54 1/92 1/79 1/100 1/75 1/100 1/100 1/100 1/99 1/100 1/100 O17. Lasionectria O18. Verruciconidia 0.1 Fig. 2. (Continued).
39www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Dataset 2 consists of 419 ingroup isolates that formed a wellsupported clade representing Bionectriaceae, with Flammocladiella aceris (CBS 138906), F. anomiae (CBS 142775), F. decora (CBS 142776), Tilachlidium brachiatum (CBS 363.97 and CBS 505.67) serving as outgroup (Fig. 2). The final alignment consists of 3 107 characters, including alignment gaps (gene boundaries ITS: 1–690, 690 bp; LSU: 691–1 478, 788 bp; rpb2: 1 479–2 280, 802 bp; tef-1α: 2 281–3 107, 827 bp). Among those, 1 494 characters were conserved sites and 1 555 were variable sites, including 196 characters that were parsimony-uninformative and 1 359 characters that were parsimony-informative. The jModeltest results recommended that the Bayesian analysis should use Dirichlet base frequencies for all data partitions. The GTR+I+G model was proposed for ITS, LSU, rpb2 and tef-1α. The sequence dataset Protocreopsis caricicola CBS 140572T Ramosiphorum thailandicum CBS 101914T Lasionectriella arenuloides CBS 576.76T Lasionectriopsis pteridii CBS 182.78 Ramosiphorum polyporicola CBS 100282 Ramosiphorum echinoporiae CBS 115288T Ramosiphorum polyporicola CBS 109.87 Lasionectriopsis pteridii CBS 579.90 Protocreopsis phormiicola CBS 567.76T Lasionectriopsis pteridii CBS 738.69 Ochronectria calami CBS 445.96 Protocreopsis rutila CBS 378.70C Protocreopsis rutila CBS 394.70 Protocreopsis finnmarkica CBS 147427 Ramosiphorum polyporicola CBS 123779T Protocreopsis finnmarkica TR 042cE1.2 Protocreopsis finnmarkica CBS 147428T Protocreopsis rutila CBS 263.89 Ochronectria thailandica MFLUCC 15-0140T Lasionectriopsis pteridii CBS 393.70 Protocreopsis rutila CBS 226.70 Lasionectriopsis pteridii CBS 102155 Lasionectriopsis pteridii CBS 784.69 Lasionectriopsis dentifera CBS 650.75 Lasionectriopsis pteridii CBS 437.66 Protocreopsis rutila CBS 396.66T Protocreopsis finnmarkica CBS 147429 Protocreopsis rutila CBS 118526 Protocreopsis pertusa CBS 568.76 Protocreopsis rutila CBS 395.66 Lasionectriopsis germanica CBS 143538T Lasionectriopsis dentifera CBS 574.76T Lasionectriopsis pteridii CBS 782.69T Protocreopsis rutila CBS 229.70 Lasionectriella marigotensis CBS 131606T Protocreopsis finnmarkica TR 041aN1.1 Ochronectria calami CBS 134535 Protocreopsis euphorbiae CPC 38896T Lasionectriopsis pteridii CBS 737.69 Ochronectria calami CBS 125.87 Protocreopsis freycinetiae CBS 573.76T Lasionectriopsis pteridii CBS 783.69 Lasionectriella herbicola CBS 140156T Lasionectriella rubioi CBS 140157T 1/95 1/100 1/99 1/96 1/100 1/97 1/55 1/80 1/100 1/100 1/100 0.97/100 1/95 1/100 0.90/60 1/96 1/100 1/93 1/100 1/90 1/100 1/100 1/100 1/100 1/100 1/100 1/66 1/100 1/100 1/100 1/100 0.94/97 1/100 4x 4x 2x 2x 2x 1/100 1/100 O19. Lasionectriopsis O20. Ochronectria O21. Lasionectriella O22. Ramosiphorum O23. Protocreopsis 0.98/- 0.1 Fig. 2. (Continued).
40 Hou et al. Nectriopsis microspora CBS 933.69 Clonostachys spinulosispora CBS 133762T Clonostachys phyllophila CBS 921.97T Emericellopsis fuci TR090bU1.1 Nectriopsis lindauiana CBS 897.70T Mycocitrus odorus CBS 100104T Emericellopsis fuci CBS 120529 Nectriopsis rexiana CBS 305.70C Emericellopsis fuci TR058aW2.1 Nectriopsis candicans CBS 424.64 Emericellopsis fuci TR080bE3.1 Bionectria grammicospora CBS 102566 Emericellopsis fuci CBS 485.92 Emericellopsis fuci TR090aU1.2 Stephanonectria keithii CBS 943.72 Nectriopsis violacea CBS 914.70T Nectriopsis candicans CBS 627.72 Nectriopsis candicans CBS 701.79T Stephanonectria keithii CBS 100007 Nectriopsis violacea CBS 849.70 Emericellopsis fuci CBS 112868T Emericellopsis fuci TR043cU1.1 Emericellopsis fuci TR070cS1.1 Nectriopsis candicans CBS 440.65 Bionectria ochroleuca CBS 114056 Mycocitrus phyllostachydis CBS 330.69 Mycocitrus zonatus CBS 400.70 Emericellopsis fuci TR061aN1.1 Emericellopsis fuci CBS 120530 Emericellopsis fuci CBS 484.92 Nectriopsis fuliginicola CBS 400.82T Emericellopsis fuci CBS 113888 Nectriopsis rexiana CBS 305.70A Emericellopsis fuci CBS 550.86 Nectriopsis rexiana CBS 542.92 Nectriopsis ellipsoidea CBS 358.78T Emericellopsis fuci CBS 113889 Emericellopsis fuci CBS 113887 Emericellopsis fuci TR058cD1.2 Nectriopsis microspora CBS 355.70 Emericellopsis fuci TR090cD1.2 Nectriopsis sporangiicola CBS 166.74T Emericellopsis fuci CBS 116467 Emericellopsis fuci TR061cN1.1 Clonostachys rogersoniana CBS 102564 Bionectria ochroleuca AFTOL-ID 187 Emericellopsis fuci TR061aS1.1 Mycocitrus odorus CBS 120610 Mycocitrus odorus CBS 232.75B Emericellopsis fuci CBS 120611 Emericellopsis fuci TR061aN1.2 Stephanonectria keithii CBS 434.70 Emericellopsis fuci TR085aW1.2 0.97/72 1/70 1/100 1/100 1/100 1/89 1/100 1/72 1/100 0.95/72 1/56 1/100 0.98/89 1/100 1/98 1/100 1/100 1/100 1/99 1/76 1/99 1/100 0.99/100 1/- 1/100 1/100 1/100 1/99 1/- 1/100 0.98/91 4x 2x 2x O24. Nectriopsis O25. Clonostachys O26. Stephanonectria O27. Mycocitrus O28. Emericellopsis 0.1 Fig. 2. (Continued).
41www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales did not show conflict in the tree topologies for the 70 % reciprocal bootstrap trees, which allowed us to combine the four genes for the multi-locus analysis. The Bayesian analysis lasted for 9 590 000 generations and a total of 14 388 trees were generated after the analysis reached the stop value of 0.01. The first of 25 % trees were discarded as the burn-in phase, and posterior probabilities (PP) were calculated from the remaining trees. The topology of the BI tree was confirmed by ML analysis and the posterior probabilities Emericellopsis fimetaria CBS 621.85 Stanjemonium fuscescens CBS 264.96T Emericellopsis glabra CBS 119.40T Emericellopsis moniliformis CBS 864.73 Emericellopsis salmonea CBS 721.71T Emericellopsis fimetaria CBS 382.62 Emericellopsis humicola CBS 180.56T Emericellopsis fimetaria CBS 628.85 Emericellopsis stolkiae CBS 159.71T Emericellopsis pallida CBS 624.73 Stanjemonium dichromosporum CBS 638.73T Emericellopsis brunneiguttula CBS 111360T Emericellopsis moniliformis CBS 139051T Proliferophialis apiculata CBS 303.64T Emericellopsis terricola CBS 120.40T Emericellopsis robusta CBS 105.70T Emericellopsis fimetaria CBS 115625 Stanjemonium ochroroseum CBS 656.79T Emericellopsis moniliformis CBS 463.92 Emericellopsis exuviara CBS 113360T Emericellopsis tubakii CBS 824.69 Proliferophialis apiculata CBS 397.78 Emericellopsis fimetaria CBS 511.67 Emericellopsis mirabilis CBS 176.53 Emericellopsis tubakii CBS 790.69T Emericellopsis microspora CBS 380.62T Emericellopsis maritima CBS 491.71T Emericellopsis fimetaria CBS 146.62 Emericellopsis stolkiae CBS 139531 Stanjemonium grisellum CBS 655.79T Emericellopsis alkalina CBS 127350T Emericellopsis salmosynnemata CBS 182.56T Emericellopsis pallida CBS 490.71T Emericellopsis minima CBS 190.55T Proliferophialis apiculata CBS 542.79 Emericellopsis tubakii CBS 822.70 Emericellopsis tubakii CBS 555.74 Emericellopsis fimetaria CBS 117.84 Emericellopsis pusilla CBS 226.62T Emericellopsis fimetaria CBS 176.60 Emericellopsis alkalina CBS 120049 Emericellopsis mirabilis CBS 177.53T Emericellopsis tubakii CBS 791.69 Emericellopsis fimetaria CBS 558.84 Proliferophialis apiculata CBS 365.64 Emericellopsis donezkii CBS 489.71T 1/100 0.99/70 1/99 1/100 1/100 0.91/47 1/85 1/93 1/99 1/100 1/99 1/100 -/51 1/99 1/100 0.91/96 1/100 1/100 1/100 1/100 0.91/80 1/59 1/100 1/98 1/86 1/100 -/63 1/92 -/74 1/92 1/95 1/80 1/100 1/84 0.97/77 1/78 0.94/- 1/100 4x 4x O29. Stanjemonium O30. Proliferophialis O28. Emericellopsis 1/- 1/- 0.1 Fig. 2. (Continued).
42 Hou et al. Acremonium sordidulum CBS 385.73T Acremonium cf. egyptiacum CBS 286.70B Acremonium multiramosum CBS 147436T Acremonium alternatum CBS 407.66T Acremonium egyptiacum CBS 124.42 Acremonium acutatum CBS 829.73 Acremonium synnematoferum CBS 147432 Acremonium charticola CBS 402.66 Acremonium charticola CBS 547.86 Acremonium synnematoferum CBS 147430 Acremonium subulatum CBS 588.73AT Acremonium purpurascens CBS 149.62T Acremonium ellipsoideum CBS 147433T Acremonium mycoparasiticum CBS 188.80T Acremonium brunneisporum CBS 142823 Acremonium chlamydosporium CBS 414.76T Acremonium acutatum CBS 140.62 Acremonium synnematoferum TR070aS1.2 Acremonium longiphialidicum CBS 451.70T Acremonium egyptiacum CBS 403.66 Acremonium synnematoferum CBS 147431T Acremonium egyptiacum CBS 391.89 Acremonium cf. egyptiacum CBS 109041 Acremonium ellipsoideum CBS 147435 Acremonium aerium CBS 379.70C Acremonium ellipsoideum CBS 147434 Acremonium egyptiacum CBS 740.69 Acremonium acutatum CBS 682.71T Acremonium brunneisporum CBS 413.76T Acremonium synnematoferum CPC 40326 Acremonium cf. egyptiacum CBS 159.61 Acremonium brachypenium CBS 866.73T Acremonium charticola CBS 117.25 Acremonium purpurascens CBS 780.69 Acremonium gamsianum CBS 145769 Acremonium psychrophilum CBS 139.93 Acremonium egyptiacum CBS 114785T Acremonium stroudii CBS 138820T Acremonium subulatum CBS 115996 Acremonium sordidulum CBS 102413 Acremonium egyptiacum CBS 114321 Acremonium egyptiacum CBS 121405 Acremonium mycoparasiticum CBS 684.71 Acremonium egyptiacum CBS 734.69 Acremonium aerium CBS 189.70T Acremonium egyptiacum CBS 500.82 Acremonium gamsianum CBS 881.73T Acremonium cf. egyptiacum CBS 270.86 Acremonium egyptiacum CBS 629.73 Acremonium cf. egyptiacum CBS 113719 Acremonium cf. egyptiacum CBS 112783 1/73 1/100 1/100 1/100 1/99 1/100 1/100 0.98/76 1/100 1/100 1/100 -/53 1/70 0.96/73 1/100 1/95 1/100 1/100 1/98 1/100 1/66 -/96 1/100 1/100 1/90 1/74 -/56 1/99 1/100 1/98 1/100 1/88 1/100 1/100 1/100 1/100 1/81 1/100 1/100 2x 2x 2x O31. Acremonium 0.1 Fig. 2. (Continued).
43www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales (BPP) from the BI inference were mapped with MLBS at the tree nodes on the ML tree (Fig. 2; MLBS > 50 %, BPP > 0.90 shown). The phylogenetic trees based on dataset 2 (Fig. 2), which were generated with Bayesian and Maximum Likelihood methods, distributed the phylogenetic species into 39 well-supported genera (Clades O1–O39) in Bionectriaceae. Clade O1 (BPP = 1, MLBS = 100 %) accommodated type species of the genus Gliomastix (G. murorum) and five other species with melanised conidia (G. masseei, G. musae, Bulbithecium arxii CBS 737.84T Waltergamsia pilosa CBS 511.82 Waltergamsia citrina CBS 758.69 Waltergamsia alkalina CBS 741.94T Bulbithecium pinkertoniae CBS 158.70 Ovicillium oosporum CBS 110151T Waltergamsia citrina CBS 384.96T Waltergamsia epimycota CBS 265.89 Geosmithia pallidum CBS 260.33T Waltergamsia zeylanica CBS 735.73A Waltergamsia obpyriformis CBS 115960 Ovicillium asperulatum CBS 426.95 Waltergamsia dimorphospora CBS 139050T Ovicillium asperulatum CBS 130362T Waltergamsia catenata CBS 102462T Waltergamsia pilosa CBS 124.70T Waltergamsia moroccensis CBS 512.82T Waltergamsia zeylanica CBS 746.73 Bulbithecium truncatum CBS 113718T Waltergamsia obpyriformis CBS 595.73T Bulbithecium borodinense CBS 101148T Waltergamsia parva CBS 381.70AT Geosmithia microcorthyli CCF 3861T Waltergamsia fusidioides CBS 705.86 Waltergamsia hennebertii CBS 768.69T Waltergamsia zeylanica CBS 736.73B Bulbithecium hyalosporum CBS 318.91T Bulbithecium ellipsoideum CBS 993.69T Ovicillium attenuatum CBS 399.86T Ovicillium subglobosum CBS 101963T Proxiovicillium blochii CBS 324.33 Bulbithecium spinosum CBS 391.66 Waltergamsia epimycota CBS 562.86 Bulbithecium pinkertoniae CBS 157.70T Waltergamsia fusidioides CBS 840.68T Waltergamsia zeylanica CBS 111659 Waltergamsia epimycota CBS 127459 Geosmithia lavendula CBS 344.48T Proxiovicillium blochii CBS 427.93T Waltergamsia zeylanica CBS 734.73 Bulbithecium spinosum CBS 136.33T Ovicillium variecolor CBS 130360T Proxiovicillium lepidopterorum CBS 101239T Waltergamsia parva CBS 831.97 Waltergamsia zeylanica CBS 735.73B Waltergamsia fusidioides CBS 113.69 Bulbithecium spinosum CBS 915.85 Bulbithecium ammophilae CBS 178.78T 1/100 1/78 1/100 1/100 1/100 1/100 1/100 1/75 1/100 1/100 1/83 1/100 1/100 1/96 1/100 1/93 1/100 1/100 1/93 1/99 1/100 1/57 1/100 -/55 1/100 1/100 1/68 1/100 1/100 1/100 1/100 1/100 1/100 1/100 1/80 -/61 1/100 -/56 2x 2x 2x 2x 2x 2x O32. Waltergamsia O33. Geosmithia O34. Bulbithecium O35. Ovicillium O36. Proxiovicillium 1/- 1/- 0.1 Fig. 2. (Continued).
44 Hou et al. G. polychroma, G. roseogrisea and G. tumulicola). Clade O2 (BPP = 1, MLBS = 100 %) comprised six Paracylindrocarpon species with cylindrocarpon-like conidia and ascospores with three or more septa, Paracylindrocarpon aloicola, Pn. multiloculatum (basionym: Nectria multiloculata), Pn. multiseptatum (basionym: Nectria multiseptata), Pn. nabanheensis, Pn. pandanicola, Pn. xishuangbannaensis and two novel species, Pn. aurantiacum and Pn. foliicola. Clade O3 (BPP = 1, MLBS = 100 %) included four species of the genus Fusariella, F. atrovirens, F. concinna, F. curvata, F. hughesii, and another species of Hydropisphaera which was recombined into this genus, F. arenula (syn.: H. arenula). Clades O4, O5 and O7 each comprised a single strain, representing Selinia, Roumegueriella, Synnemellisia, respectively. Clade O6 (BPP = 1, MLBS = 100 %) encompassed the ex-type strains of the two species of Verrucostoma, V. freycinetiae and V. martinicense. The proposed new monotypic genera Musananaesporium (Clade O8) and Gossypinidium (Clade O9) included one strain of the type, respectively, M. tectonae (basionym: Acremonium tectonae), and G. sporodochiale. The Monohydropisphaera clade (Clade O11) was recognised as a novel monophyletic genus, containing the type species of this genus, M. fusigera (basionym: Monotospora fusigera). Clade 12 (BPP = 1, MLBS = 93 %) comprised five species of Hydropisphaera, H. cyatheae, H. fungicola, H. peziza, H. suffulta, and another species of Heleococcum which was recombined into this genus, H. aurantiaca (basionym: Heleococcum aurantiacum; syn.: Heleococcum japonense). Clades O13, O15 and O16 each comprised a single strain, representing Geonectria (G. subalpina), Septofusidium (S. berolinense) and Pseudoacremonium (Pm. sacchari), respectively. The new genus Paragliomastix (clade O14; BPP = 1, MLBS = 99 %) included Px. chiangraiensis (basionym: Acremonium chiangraiense), Px. luzulae (basionym: Torula luzulae), Px. znieffensis, and a novel species Px. rosea. Clade O17 (BPP = 1, MLBS = 100 %) included five known species of the genus Lasionectria (L. antillana, L. boothii, L. krabiense, L. mantuana, and L. sylvana), and another five species were recombined into this genus, L. atrorubra (basionym: Nectriella atrorubra), L. bisepta (basionym: A. biseptum), L. castaneicola (basionym: Hydropisphaera castaneicola), L. cerealis (basionym: Coniosporium cerealis), and L. olida (basionym: A. olidum). Clade O18 (BPP = 1, MLBS = 100 %) comprised seven species accommodated in a novel genus Verruciconidia, i.e. Ve. persicina (syn.: A. persicinum), Ve. verruculosa (basionym: A. verruculosum) and five novel species with verrucose conidia, Ve. erythroxyli, Ve. infuscata, Ve. quercina, Ve. siccicapita and Ve. unguis. Clade O19 (BPP = 1, MLBS = 100 %) accommodated the genus Lasionectriopsis with its type species, L. germanica, and two other species, L. pteridii and L. dentifera (basionym: Nectria dentifera). Clade O20 (BPP = 1, 0.1 Flammocladiella anomiae CBS 142775 Hapsidospora chrysogena CBS 401.65 Stilbocrea macrostoma CBS 114375 Hapsidospora irregularis CBS 510.70T Hapsidospora flava CBS 597.70 Hapsidospora chrysogena CBS 899.85 Hapsidospora variabilis CBS 100549T Alloacremonium ferrugineum CBS 102877T Hapsidospora inversa CBS 517.70T Flammocladiella aceris CBS 138906T Hapsidospora globosa CBS 511.70 Hapsidospora stercoraria CBS 217.84 Hapsidospora flava CBS 142.71 Hapsidospora flava CBS 596.70T Tilachlidium brachiatum CBS 363.97 Alloacremonium humicola CBS 613.82T Hapsidospora globosa CBS 110041 Flammocladiella decora CBS 142776 Hapsidospora flava CBS 963.87 Hapsidospora chrysogena CBS 144.62T Hapsidospora flava CBS 316.72 Stilbocrea macrostoma CBS 141849 Tilachlidium brachiatum CBS 505.67 Stilbocrea walteri CBS 144627T Hapsidospora globosa CBS 513.70 Hapsidospora stercoraria CBS 516.70T Hapsidospora globosa CBS 514.70 Hapsidospora globosa CBS 416.73 Hapsidospora chrysogena CBS 779.69 Hapsidospora globosa CBS 512.70T 1/100 1/100 1/100 1/100 1/100 1/100 1/87 1/100 1/100 1/98 1/95 1/100 1/100 1/100 1/73 1/100 1/100 -/54 1/100 1/100 1/100 1/100 1/97 1/99 1/99 4x 2x 2x 1/100 O37. Hapsidospora O38. Alloacremonium O39. Stilbocrea 2x Fig. 2. (Continued).
45www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales V1. Sarocladium Sarocladium agarici CBS 113717T Sarocladium gamsii CBS 425.73 Sarocladium fuscum CBS 334.80T Sarocladium summerbellii CBS 430.70T Sarocladium ochraceum CBS 428.67T Sarocladium bactrocephalum CBS 749.69T Sarocladium junci CBS 148277T Sarocladium gamsii CBS 707.73T Sarocladium glaucum CBS 796.69T Sarocladium dejongiae CBS 694.93 Sarocladium strictum CBS 346.70T Sarocladium dejongiae CBS 144929T Sarocladium subulatum MUCL 9939T Sarocladium pseudostrictum CBS 137660T Sarocladium spinificis BCRC 34941T Sarocladium terricola CBS 243.59T Sarocladium agarici CBS 126941 Sarocladium dejongiae CBS 699.93 Sarocladium implicatum CBS 959.72T Sarocladium zeae CBS 800.69T Sarocladium kiliense CBS 289.70 Sarocladium kiliense CBS 122.29T Sarocladium strictum CBS 334.77 Sarocladium kiliense CBS 400.52 Sarocladium zeae CBS 801.69 Sarocladium strictum CBS 405.80 Sarocladium strictum CBS 152.72 Sarocladium theobromae CBS 113440T Sarocladium implicatum CBS 128323 Sarocladium attenuatum CBS 399.73T Sarocladium bacillisporum CBS 425.67T Sarocladium kiliense CBS 129746 Sarocladium oryzae CBS 180.74T Sarocladium strictum CBS 102039 Sarocladium hominis CBS 137659T Sarocladium bifurcatum CBS 137658T Sarocladium bacillisporum CBS 787.69 Sarocladium spinificis CBS 170.89 Sarocladium citri CBS 145044T Sarocladium ferrugineum CBS 102673T Sarocladium brachiariae CGMCC 2192T Sarocladium spinificis CBS 102676 1/100 0.97/71 1/100 1/100 1/53 1/90 1/100 1/100 1/100 1/99 1/100 1/100 1/100 1/88 1/100 1/96 1/96 -/81 0.98/98 -/59 1/57 1/92 1/100 1/100 1/100 1/100 1/100 1/97 0.92/- 1/100 1/100 1/72 Sarocladiaceae 0.07 Fig. 3. Phylogenetic tree inferred from a Maximum Likelihood analysis based on a concatenated alignment of LSU, ITS, rpb2 and tef-1α sequences of 77 strains representing Sarocladiaceae and outgroups. The RAxML bootstrap support values (MLBS) above 50 % and Bayesian posterior probabilities (BPP) above 0.90 are given at the nodes (BPP/MLBS). The scale bar represents the expected number of changes per site. Genera are delimited in coloured boxes, with the genus name indicated to the right. Strains with special status are indicated with a superscript letter after the accession number (T: ex-type). New species are printed in red font and new combinations in blue font. The tree is rooted to Acremonium egyptiacum CBS 124.42, A. alternatum CBS 407.66, and Paracremonium binnewijzendii CBS 698.
46 Hou et al. MLBS = 100 %) contained four isolates of Ochronectria, including the generic type, O. calami. Clade O21 (BPP = 1, MLBS = 100 %) accommodated the genus Lasionectriella, including the generic type, La. rubioi. Clade O22 (BPP = 1, MLBS = 100 %) contained three novel species that were accommodated in a new genus proposed below, Ramosiphorum, namely R. echinoporiae, R. polyporicola and R. thailandicum. Clade O23 accommodated the genus Protocreopsis, including five known species, one novel species Pt. finnmarkica, and an Acremonium species that was recombined to the genus as Pt. rutila (basionym: A. rutilum). The genus Nectriopsis (clade O24; BPP = 1, MLBS = 100 %) comprised eight species, including the generic type N. violacea. Clades O25 to O26 encompassed two genera: Bionectria/ Clonostachys (clade O25; BPP = 1, MLBS = 100 %) and Stephanonectria (clade O26; BPP = 1, MLBS = 100 %). Clade O27 (BPP = 1, MLBS = 100 %) comprised three species, Mycocitrus odorus, M. phyllostachydis, and M. zonatus (basionym: A. zonatum). Emericellopsis (Clade O28; BPP = 1, MLBS = 100 %) was represented by 15 previously described species (including the type species E. terricola), the new species E. brunneiguttula, and another six species, E. exuviara (basionym: A. exuviarum), E. fimetaria (syn. Stilbella fimetaria), E. fuci (basionym: A. fuci), E. moniliformis (basionym: A. moniliforme), E. salmonea (basionym: A. salmoneum), and E. tubakii (basionym: A. tubakii). Clade O29 (BPP = 1, MLBS = 100 %) comprised the type species, Stanjemonium grisellum, and three other species, S. dichromosporum (basionym: A. dichromosporum), S. fuscescens and S. ochroroseum. Clade O30 (BPP = 1, MLBS = 100 %) comprised four isolates previously received as Acremonium and characterised by producing abundant phialides with apical percurrent proliferation, representing one species, which belong to a newly introduced genus Proliferophialis, namely Pro. apiculata. Clade O31 (BPP = 1, MLBS = 100 %) accommodated the genus Acremonium s. str., which is represented by eight previously described species (including the generic type species A. alternatum), the 21 isolates that were previously identified as Acremonium spp., representing 10 novel species, namely A. aerium, A. brunneisporum, A. chlamydosporium, A. ellipsoideum, A. gamsianum, A. longiphialidicum, A. subulatum, A. V2. Parasarocladium V3. Chlamydocillium V4. Polyphialocladium 0.07 Chlamydocillium cyanophilum CBS 273.80 Polyphialocladium fusisporum CBS 406.66T Chlamydocillium lolii CBS 214.70T Parasarocladium breve CBS 150.62T Chlamydocillium cyanophilum CBS 102685 Chlamydocillium cyanophilum CBS 599.93 Chlamydocillium cyanophilum CBS 699.73 Chlamydocillium guttulatum CBS 104.78T Parasarocladium aestuarinum CMG 31 Parasarocladium aestuarinum CMG 30T Acremonium alternatum CBS 407.66T Parasarocladium breve CBS 102443 Parasarocladium gamsii CBS 726.71T Chlamydocillium acaciae CBS 523.72T Chlamydocillium cyanophilum CBS 246.74AT Parasarocladium alavariense CMG 32T Chlamydocillium terrestre CBS 110514T Chlamydocillium cyanophilum CBS 632.73 Paracremonium binnewijzendii CBS 698.86 Parasarocladium fusiforme CMG 36T Parasarocladium chondroidum CBS 652.93T Polyphialocladium fusisporum CBS 114602 Parasarocladium debruynii CBS 144942T Parasarocladium breve CBS 387.96 Parasarocladium breve CBS 303.96 Chlamydocillium antarcticum CBS 120502T Chlamydocillium curvulum CBS 229.75 Chlamydocillium curvulum CBS 430.66T Parasarocladium radiatum CBS 142.62T Parasarocladium breve CBS 427.96 Acremonium egyptiacum CBS 124.42T Parasarocladium funiculosum CBS 141.62T Chlamydocillium cyanophilum CBS 102681 Chlamydocillium soli CBS 347.76T Chlamydocillium guttulatum CBS 384.70C 1/100 0.99/93 1/100 0.99/97 1/96 1/100 1/100 0.99/74 0.99/90 0.98/91 0.90/64 1/82 1/82 1/99 1/100 1/100 1/89 0.98/88 1/100 1/100 -/56 1/100 1/98 1/100 1/100 0.98/62 5× 2× 2× 4× 2× 2× 4× 4× Sarocladiaceae Fig. 3. (Continued).
47www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales synnematoferum, A. multiramosum, and A. mycoparasiticum. One Cephalosporium species clustered within the Acremonium clade, phylogenetically close to A. aerium, and was thus recombined to this genus, as A. purpurascens (basionym: C. purpurascens). The majority of isolates that clustered in clade O32 (BPP = 1, MLBS = 100 %) were identified as “Acremonium” spp., and a new generic name Waltergamsia is introduced for this clade, which comprised 12 accepted species, four novel species, W. alkalina, W. catenata, W. moroccensis, W. obpyriformis, and eight species that were received as Acremonium, Cephalosporium or Nectriopsis species including W. citrina (basionym: A. citrinum), W. dimorphospora (basionym: A. dimorphosporum), W. epimycota (basionym: N. epimycota), W. fusidioides (syn. A. fusidioides), W. hennebertii (basionym: A. hennebertii), W. parva (basionym: A. parvum), W. pilosa (basionym: A. pilosum), and W. zeylanica (basionym: C. zeylanicum). Clade O33 (BPP = 1, MLBS = 100 %) encompassed the ex-type strains of three species of Geosmithia, G. lavendula, G. microcorthyli and G. pallidum. Bulbithecium formed a well-supported clade (clade O34; BPP = 1, MLBS = 100 %) and included the generic type species B. hyalosporum, four species that were received as Acremonium spp. or Leucosphaerina spp., namely B. arxii (basionym: L. arxii), B. borodinense (basionym: A. borodinense), B. pinkertoniae (basionym: A. pinkertoniae), B. spinosum (basionym: Cephalosporium spinosum) and three novel species B. ammophilae, B. ellipsoideum and B. truncatum. Clade O35 (BPP = 1, MLBS = 100 %) comprised six isolates of Ovicillium, representing five species, including its type species, O. attenuatum, and two species previously assigned to other genera, O. variecolor (basionym: A. variecolor) and O. asperulatum (basionym: A. asperulatum). The new genus Proxiovicillium (clade O36; BPP = 1, MLBS = 100 %) comprised Pr. blochii (basionym: Mastigocladium blochii, syn. Acremonium blochii), the type species, and the new species Pr. lepidopterorum. Clade O37 (BPP = 1, MLBS = 100 %) comprised seven species which belong to Hapsidospora. This clade contains two species initially classified as Acremonium and Cephalosporium species; and two species from the two genera from the same family with Hapsidospora, which were recombined as H. globosa (basionym: Nigrosabulum globosum) and H. inversa (basionym: Mycoarachis inversa). Clade O38 (BPP = 1, MLBS = 100 %) comprised two isolates representing two species, which belong to a new genus introduced here, Alloacremonium, namely Al. ferrugineum and Al. humicola. The clade corresponding to Stilbocrea (clade O39; BPP = 1, MLBS = 100 %), included two species, S. macrostoma and S. walteri. Dataset 3 consisted of 77 strains belonging to Sarocladiaceae, with Acremonium egyptiacum (CBS 124.42), A. alternatum (CBS 407.66) , and Paracremonium binnewijzendii CBS 698.86 as outgroup (Fig. 3). The final alignment consisted of 3 087 characters including gaps (gene boundaries ITS: 1–593, 593 bp; LSU: 594–1 382, 789 bp; rpb2: 1 383–2 156, 774 bp; tef-1α: 2 157–3 087, 931 bp). Among those, 1 694 characters were conserved sites and 1 235 were variable sites, including 190 characters that were parsimony-uninformative and 1 045 characters that were parsimony-informative. The jModeltest results recommended that the Bayesian analysis should use Dirichlet base frequencies for all data partitions. The GTR+I+G model was proposed for ITS, LSU, rpb2 and tef-1α. The sequence dataset did not show conflict in the tree topologies for the 70 % reciprocal bootstrap trees, which allowed us to combine the four genes for the multi-locus analysis. The BA lasted for 5 550 000 generations and a total of 8 287 trees were generated after the BI analysis reached the stop value of 0.01. The first 25 % trees were discarded as the burn-in phase, and posterior probabilities (PP) were calculated from the remaining 6 215 trees. The topology of the BI tree was confirmed by ML analysis and therefore, the posterior probabilities (PP) from the BI inference were mapped with MLBS at the tree nodes on the ML tree (Fig. 3; MLBS > 50 %, BPP > 0.90 shown). The families that belong to Sarocladiaceae, represented by the species grouping in clades V1–V4, clustered in a strongly supported clade (BPP = 1, MLBS = 100 %). Clade V1 was represented by 21 accepted species of Sarocladium (BPP = 1; MLBS = 100 %), and six strains that were labelled as Acremonium spp. were placed in five independent and well-supported clades/lineages, representing five novel species, S. agarici, S. citri, S. ferrugineum, S. fuscum, and S. theobromae. Clade V2 (BPP = 1, MLBS = 89 %) contained seven accepted species of Parasarocladium and two undescribed lineages, representing two species, Par. funiculosum (basionym: Cephalosporium acremonium var. funiculosum) and Par. chondroidum. Clade V3 (BPP = 1, MLBS = 100 %) encompassed one subclade representing Chlamydocillium cyanophilum, and seven subclades comprising seven cultures that were labelled as A. curvulum, including the ex-type culture of A. curvulum. Clade V1 (BPP = 1, MLBS = 100 %) was basal to Clades V1–V3, containing two “A. alternatum” cultures and forming an independent lineage, which represented the novel genus Polyphialocladium. Dataset 4 consisted of ITS, LSU, rpb2 and tef-1α sequences of 90 strains representing members of Plectosphaerellaceae, with Monilochaetes infuscans (CBS 869.66 and CBS 379.77) as outgroups (Fig. 4). The final alignment consisted of 3 036 characters including gaps (gene boundaries ITS: 1–614, 614 bp; LSU: 615–1448, 834 bp; rpb2: 1449–2268, 820 bp; tef-1α: 2 269–3 036, 768 bp). Among those, 1 855 characters were conserved sites and 1 134 were variable sites, including 1 009 characters that were parsimony-informative. The jModeltest results recommended that the Bayesian analysis should use Dirichlet base frequencies for all data partitions. The GTR+I+G model was proposed for ITS, LSU, rpb2 and tef-1α. The sequence dataset did not show conflict in the tree topologies for the 70 % reciprocal bootstrap trees, which allowed us to combine the four genes for the multi-locus analysis. The BA lasted for 295 000 generations and a total of 439 trees were generated after the BI analysis reached the stop value of 0.01, with the first 25 % trees discarded as the burn-in phase. The posterior probabilities (PP) were calculated from the remaining 330 trees. The topology of the BI tree was confirmed by ML analysis and therefore, the posterior probabilities (PP) from the BI inference were mapped with MLBS at the tree nodes on the ML tree (Fig. 4; MLBS > 50 %, BPP > 0.90 shown). A total of 23 well-supported clades represent 23 known genera in the phylogenetic tree based on dataset 4: Acremoniisimulans (BPP < 0.90; MLBS = 97 %), Acrostalagmus (BPP = 1; MLBS = 100 %), Brunneochlamydosporium (BPP = 1; MLBS = 100 %), Brunneomyces (BPP = 1; MLBS = 100 %), Chlamydosporiella (BPP = 1; MLBS = 100 %), Chordomyces (BPP = 1; MLBS = 100 %), Furcasterigmium (BPP = 1; MLBS = 100 %), Fuscohypha (BPP = 1; MLBS = 100 %), Gibellulopsis (BPP = 1; MLBS = 100 %), Lectera (BPP = 1; MLBS = 100 %), Musicillium (BPP = 1; MLBS = 100 %), Musidium (BPP = 1; MLBS = 100 %), Nigrocephalum (BPP = 1; MLBS = 100 %), Paramusicillium (BPP = 1; MLBS = 100 %), Paragibellulopsis (BPP = 1; MLBS = 100 %), Phialoparvum, Plectosphaerella (BPP = 1; MLBS = 100 %), Sayamraella (BPP = 1; MLBS = 74 %), Sodiomyces (BPP = 1; MLBS = 100 %), Stachylidium (BPP = 1; MLBS = 97 %), Summerbellia (BPP = 1; MLBS = 100 %), Theobromium (BPP = 1; MLBS = 93 %), and Verticillium (BPP = 1; MLBS = 99 %). Two clades with species identified as Acremonium s. lat. were reassigned to new genera: Allomusicillium (BPP = 1; MLBS = 96 %), and Parafuscohypha (BPP = 1; MLBS = 78 %), respectively.
54 Hou et al. Additional materials examined: Germany, Kiel, Hortus Botanicus, on decaying wood in heated greenhouse, 1965, W. Gams No. 656, CBS H-8330, culture CBS 190.70; unknown substrate, collection date and collector, isol. H.I. Nirenberg, No. 1.9/1V2(1), culture CBS 611.95. USA, Florida, from deformed toenail, unknown collection date and collector, isol. 1968 by N. Zaias, No. 1485, CBS H-8328, culture CBS 781.69. Notes: Cultures representing Nothoacremonium subcylindricum were initially identified as Acremonium potronii. There are no ex-type cultures of A. potronii that have survived. Although type materials of A. potronii were not examined, observation of the conidiogenous structures in culture revealed No. subcylindricum to have conidiophores with basitonous side branches, longer phialides and conidia without a tapered base, which differ from the description of A. potronii (Gams 1971). This species clade is fully supported by the phylogenetic analysis (Fig. 1; BPP/MLBS = 1/100 %). Nothoacremonium vesiculophorum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845806. Fig. 8. Etymology: Referring to the production of phialides with a vesicular structure at the apex. Fig. 7. Nothoacremonium subcylindricum (ex-type culture CBS 416.68). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, G–J. Branched or unbranched conidiophores. E. Conidiophores radiating out from coils formed by the mycelium. F. Conidial heads. K. Conidia. Scale bars = 10 μm.
55www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1–1.5 μm wide, mycelial coils formed. Conidiophores solitary, erect, straight or curved, arising directly from submerged or superficial hyphae, or radiating out from mycelial coils, unbranched or poorly branched, proliferating sympodially, showing conidiogenous cells as short lateral, cylindrical, asymmetrical projections, or bearing 1–2 phialides per node, up to 67 μm long, 1-septa at base, hyaline, smooth-walled, cell walls usually thicker than those of vegetative hyphae. Phialides lateral, acicular, hyaline, thick-, smooth-walled, (14.3–)20.7–57 μm long, 1–2.3 μm wide at base, occasionally extending into a 1.3–1.7 μm wide vesicular expansion at apex, with inconspicuous periclinal thickening and collarette at conidiogenous loci, commonly with a percurrent proliferation; polyphialides with two conidiogenous loci occasionally present. Conidia aseptate, fusoid, straight, with thickened, truncate base (hilum) at both ends, hyaline, thick-, smooth-walled, 3.4–5.2 × 1.3–1.8 μm, eguttulate, arranged in long chains, soon collapsing into conidial heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 15–16 mm diam, flat, sparse aerial mycelium, dusty, white Fig. 8. Nothoacremonium vesiculophorum (ex-type culture CBS 397.70B). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D–H, K. Conidiophores. I, J. Phialides extending into vesicular expansion at apex (arrows). L. Conidia. Scale bars = 10 μm.
56 Hou et al. to pale salmon, margin entire, reverse rosy buff; On MEA reaching 16–18 mm diam, flat, with moderate aerial mycelium, moist, hairy, white, margin crenate, reverse saffron, with buff radial lines; On PDA reaching 17–18 mm diam, flat, sparse aerial mycelium, dusty, dirty white or pale salmon, margin entire, reverse pale saffron; On SNA reaching 15–18 mm diam, flat, membranous without aerial mycelium, white, margin entire, reverse white. Lacking odour on all media. Typus: Netherlands, North Holland Province, De Vuntus near Loosdrecht, on dying leaf of Cladium mariscus (Cyperaceae), unknown collection date and collector, isol. Jun. 1968 by W. Gams, No. 1406 (holotype CBS H-24637, ex-type culture CBS 397.70B). Notes: The culture CBS 397.70B was labelled as “Acremonium implicatum (currently Sarocladium implicatum)”. However, initial molecular work based on LSU/SSU sequences indicated that it is not a Sarocladium but a member of the “A. exiguum” clade (Summerbell et al. 2011). In our present study, the culture clusters in Nothoacremonium according to the phylogenetic analysis based on the combined four genes and represents a monocillium-like species producing vesicular as well as acicular phialides. However, it is not congeneric with Monocillium or Niesslia (Fig. 1). This is the only monocillium-like morphology observed in Nothoacremonium. Morphologically, it differs from the other two species in the genus Nothoacremonium in the production of vesicular structure at the apex on phialides and conidia arranged in long chains, while No. exiguum and No. subcylindricum have cylindrical or subulate phialides and conidia arranged in slimy heads. Based on a blastn search of NCBIs GenBank nucleotide database, the closest hit using the LSU sequence of No. vesiculophorum is a sequence from the ex-isotype culture of No. subcylindricum CBS 416.68 [Genbank HQ232097.1; Identity = 769/778 (99 %), no gaps]. The closest hit using the ITS sequence is CBS 587.73 [GenBank NR_159619.1; Identity = 444/493 (90 %), 11 gaps (2 %)], a sequence from the ex-type culture of A. exiguum (currently No. exiguum). Clade J Nectriaceae Tul. & C. Tul. [as ‘Nectriei’], Select. Fung. Carpol. (Paris) 3: 3. 1865. Classification: Hypocreales, Sordariomycetes. Type genus: Nectria (Fr.) Fr. Xenoacremonium L. Lombard & Crous, Stud. Mycol. 80: 234. 2015. Stromata erumpent, orange to brown, pseudoparenchymatous, cells forming textura angularis, intergrading with ascomatal wall. Ascomata superficial, aggregated in groups of 2–3, brown, subglobose to globose. Peridium 50–90 μm thick, comprises two regions, outer region intergrading with stroma, cells forming textura angularis, inner region elongate, thin-walled, hyaline cells, forming textura prismatica. Paraphyses 2–3 μm wide, persistent, aseptate, not branched, tapering towards the apex. Asci 8-spored, unitunicate, mainly uniseriate to partly biseriate, with inconspicuous ring at apex, clavate. Ascospores ellipsoidal to fusoid with rounded ends, straight or slightly curved, aseptate, hyaline, smoothwalled. Mycelium consisting of hyaline, septate, branched hyphae. Conidiophores either as lateral phialidic pegs or arising laterally from somatic hyphae, erect, cylindrical to subcylindrical, unbranched, branched, or repeatedly verticillate towards the apex, aseptate or septate, smooth, hyaline to pale brown. Conidiogenous cells lateral or terminal, monophialidic, hyaline, smooth, elongateampulliform or subcylindrical, tapering towards the apex, with periclinal thickening and inconspicuous collarette, with short sterile outgrowths in some species. Conidia aseptate, allantoid, fusoid to ellipsoidal to cylindrical, slightly or strongly curved, forming slimy heads on the conidiophore (emended from Lombard et al. 2015). Type: Xenoacremonium recifei (Leão & Lôbo) L. Lombard & Crous Other accepted species with available sequences: Xenoacremonium allantoideum L.W. Hou, L. Cai & Crous, X. brunneosporum Dayar., E.B.G. Jones & K.D. Hyde, X. falcatum L. Lombard & Crous Xenoacremonium allantoideum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845808. Fig. 9. Etymology: Referring to the allantoid shape of its conidia. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, 1.5–3 μm wide, mycelial ropes formed. Sporulation abundant, phalacrogenous, nematogenous, plectonematogenous. Conidiophores mostly aggregated, erect, straight, arising directly from aerial or substratal mycelium, or from ropes and coils formed by mycelium, unbranched, basitonously branched, or repeatedly verticillate towards the apex, bearing 1–4 levels with 2–3 phialides per node, occasionally chondroid at base, 30.5–98.5 μm long, with 1–2 septa, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides solitary and in divergent whorls of 2–3, lateral, rarely terminal, subcylindrical, hyaline, thick-, smooth-walled, 17.5–67.5 μm long, 1.4–2.5 μm wide at base, commonly with conspicuous periclinal thickening and cylindrical collarette at conidiogenous loci, with short sterile outgrowths; polyphialides not observed. Conidia aseptate, allantoid, with rounded ends, curved, hyaline, thin-, smooth-walled, eguttulate, 3.6–6 × 1.3–2 μm, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 50–53 mm diam, flat, membranous at centre, thinly felty at periphery, dirty white, margin entire, pale salmon in old cultures, reverse buff; On MEA reaching 48 mm diam, raised, with abundant aerial mycelium, felty, or short hairy, white, with ochreous edge, margin entire, reverse apricot; On PDA reaching 48–49 mm diam, flat, with moderate aerial mycelium, floccose at centre, thinly felty at periphery, white with dirty white and entire margin , reverse dirty white; On SNA reaching 53–55 mm diam, flat, with sparse aerial mycelium, dusty, white, margin entire, reverse concolourous. Lacking odour on all media. Typus: Unknown country, substrate, collection date and collector, dep. G. Seehann (holotype CBS H-24635, ex-type culture CBS 505.94). Notes: The ex-type culture of Xenoacremonium allantoideum was previously labelled as Acremonium cf. curvulum, which is probably based on its curved conidia. Previous phylogenetic analysis demonstrated that CBS 505.94 clustered with the ex-type of Acremonium recifei (currently Xenoacremonium recifei), distant from ex-type of A. curvulum (Summerbell et al. 2011). In this study, the phylogenetic results agreed with the previous results of Lombard et al. (2015), which placed culture CBS 505.94 in a separate lineage related to X. recifei (Fig. 1). Morphologically, culture CBS 505.94
57www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales differs from X. recifei in producing longer phialides (17.5–67.5 μm long vs 15–55 μm long) and allantoid conidia, while X. recifei has strongly curved club-shaped conidia. In addition, CBS 505.94 differs from X. recifei by the lack of chlamydospores (Lombard et al. 2015) and can be distinguished from X. falcatum in its repeatedly basitonously or verticillately branched conidiophores, and the phialides with conspicuous periclinal thickening and cylindrical flared collarette at the conidiogenous loci (Lombard et al. 2015). Clade K Neoacremoniaceae L.W. Hou, L. Cai & Crous, fam. nov. MycoBank MB 845809. Classification: Hypocreales, Sordariomycetes. Mycelium consisting of branched, septate, smooth-, thin-walled hyphae. Conidiophores solitary or aggregated, erect, arising directly from submerged or superficial hyphae, or ropes formed by mycelium, occasionally basitonously branched or unbranched, proliferating sympodially, hyaline, septate, smooth-walled, cell walls usually thicker than those of vegetative hyphae. Conidiogenous cells enteroblastic, monoor polyphialidic, lageniform, subcylindrical, subulate, hyaline, thinor thick-, smooth-walled, with conspicuous or inconspicuous periclinal thickening and cylindrical collarette at conidiogenous loci; polyphialides with 2–3 conidiogenous loci occasionally present. Conidia aseptate, cylindrical, ellipsoid, ovoid, wide fusoid, spindleshaped, straight, hyaline, thin-, smooth-walled, eguttulate, arranged in chains or slimy heads. Chlamydospores laterally on short stalks, single, globose to sub-globose, hyaline, smooth-, thick-walled. Sexual morph not observed. Fig. 9. Xenoacremonium allantoideum (ex-type culture CBS 505.94). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores with conidial heads. E. Mycelial ropes with conidiophores. F–I. Conidiophores. J. Conidia. Scale bars = 10 μm.
58 Hou et al. Type: Neoacremonium L.W. Hou, L. Cai & Crous Notes: The family Neoacremoniaceae presently only includes Neoacremonium, based on Ne. distortum. Five acremonium-like species together with Parapyrenis maritima formed a fully supported clade basal to the Nectriaceae, separated from all known families in the Hypocreales (Fig. 1). Neoacremonium L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845810. Etymology: Referring to its morphological similarity to the genus Acremonium. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae. Conidiophores solitary or aggregated, erect, arising directly from submerged or superficial hyphae, or ropes formed by mycelium, with 1–2 irregularly basitonous side branches, or unbranched, proliferating sympodially, showing conidiogenous cells as short lateral, cylindrical asymmetrical projections, hyaline, smooth-walled, cell walls usually thicker than those of vegetative hyphae. Conidiogenous cells enteroblastic, monoor polyphialidic, lateral or terminal, lageniform, subcylindrical, subulate, hyaline, thinor thick-, smooth-walled, with conspicuous or inconspicuous periclinal thickening and cylindrical collarette at conidiogenous loci; polyphialides with 2–3 conidiogenous loci occasionally present; adelophialides present in some species. Conidia aseptate, often variable in shape and size, cylindrical, ellipsoid, ovoid, wide fusoid, spindle-shaped, straight, hyaline, thin-, smooth-walled, eguttulate, arranged in chains or slimy heads. Chlamydospores laterally on short stalks, single, globose to sub-globose, hyaline, smooth-, thick-walled. Sexual morph not observed. Type species: Neoacremonium distortum L.W. Hou, L. Cai & Crous Other accepted species with available sequences: Neoacremonium flavum L.W. Hou, L. Cai & Crous, Ne. minutisporum (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous, Ne. taiwanense (K.L. Pang, Alias & E.B.G. Jones) L.W. Hou, L. Cai & Crous, Ne. vitellinum (W. Gams) L.W. Hou, L. Cai & Crous Notes: Neoacremonium currently accommodates five species of Acremonium s. lat. However, they are not congeneric with the type of Acremonium s. str. based on the phylogenetic analysis, which shows that Neoacremonium forms an independent lineage with high support values in Hypocreales (Fig. 1). Although species in this genus are morphologically variable and share few characters, we prefer not to introduce more genera but retain them in one genus, until more materials are available to clarify and stabilise the taxonomy of these taxa. Neoacremonium distortum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845811. Fig. 10. Etymology: Named after the distorted phialides produced by this fungus. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, becoming roughand thick-walled with age, 1.6– 3.2 μm wide. Conidiophores solitary or aggregated, erect, arising directly from submerged or superficial hyphae, or from ropes formed by mycelium, mostly with 1–2 irregularly basitonous side branches, bearing 1–3 levels with 1–2 phialides per node, often proliferating sympodially, showing conidiogenous cells as short lateral and cylindrical, asymmetrical projections, or unbranched, 16–57 μm long, 1–2-septate at base and upper part, hyaline, smooth-walled, cell walls usually thicker than those of vegetative hyphae. Phialides lateral or terminal, subulate, distorted, hyaline, thick-, smooth-walled, 10.5–35.5 μm long, 1.5–2.5 μm wide at base, with conspicuous periclinal thickening and cylindrical collarette at the conidiogenous loci, commonly with a percurrent subterminal proliferation; polyphialides with two conidiogenous loci occasionally present. Conidia aseptate, ellipsoid, broad fusoid, apiculate at both ends, hyaline, thin-, smooth-walled, eguttulate, 2.8–4.7 × 1.5–2.6 μm, arranged in chains, collapse into heads with age. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 35–38 mm diam, dusty, white, with buff pigment, margin entire, reverse olivaceous buff; On MEA reaching 45–47 mm diam, flat, dusty, slightly hairy at centre, mycelium arranged in radially lines, white at centre, colourless at periphery, margin entire, reverse luteous; On PDA reaching 65 mm diam, flat, dusty, white, with creamy white and entire margin, reverse dirty white, with buff radial lines. Typus: Netherlands, Flevoland Province, Zuidelijk Flevoland, from soil under Phragmites australis (Poaceae), unknown collection date and collector, isol. J.W. Veenbaas-Rijks, No. 711123/4 (holotype CBS H-6647, ex-type culture CBS 314.72 = IPO 1110). Additional materials examined: Canada, Manitoba, Winnipeg, unknown substrate, collection date and collector, isol. J. Reid, No. UM 185, culture CBS 706.73. Netherlands, Baarn, Cantonspark, from leaf sheath of Musa sp., 11 Oct. 1967, W. Gams, CBS H-8468, culture CBS 291.70A; Flevoland Province, Zuidelijk Flevoland, from soil, unknown collection date and collector, isol. J.W. Veenbaas-Rijks, No. 711013/933, culture CBS 146.72; Flevoland Province, Zuidelijk Flevoland, from soil, unknown collection date and collector, isol. J.W. Veenbaas-Rijks, No. 720111/361, culture CBS 315.72 = IPO 1115; Gelderland Province, Wageningen, from air, unknown collection date and collector, isol. J. van der Spek, culture CBS 291.70B; Utrecht Province, Baarn, aerial contaminant, unknown collection date and collector, isol. 1975 by W. Gams, culture CBS 665.75; Utrecht Province, Maartensdijk, greenhouse, from dead leaf sheath of Musa sapientum (Musaceae), unknown collection date, isol. Jul. 1973, coll. and isol. by W. Gams, culture CBS 710.73. Notes: Eight cultures labelled Acremonium distortum and A. vitellinum were examined, including the culture CBS 314.72, which was recorded as ex-type culture of A. distortum in the database of the CBS culture collection. However, the name A. distortum had never been published. According to the phylogenetic inference in the present study, all cultures form a fully supported and independent clade that is distant from the ex-type culture of A. vitellinum (CBS 792.69) and other species in Neoacremonium (Fig. 1). Thus, a new species is described here as Ne. distortum. Morphologically, Ne. distortum differs from Ne. flavum in its longer conidiophores with asymmetrical projections (16–57 μm vs up to 36.3 μm), and longer conidia (2.8–4.7 μm vs 2.2–3.3 μm). Neoacremonium flavum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845947. Fig. 11. Etymology: From Latin flavus, yellow, due to the yellow colony of this fungus. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.5–2.3 μm wide. Sporulation abundant, phalacrogenous, nematogenous. Conidiophores aggregated or
59www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales solitary, (sub-)erect, mostly curved, irregular bend at base, arising directly from submerged and superficial hyphae, densely arranged, forming sporodochia-like structure in old cultures, mostly with 1–2 irregularly basitonous side branches, or unbranched, up to 36.3 μm long, 1–2(–4)-septate at base and middle, hyaline, smooth-walled, cell walls usually thicker than those of vegetative hyphae. Phialides lateral, subulate or lageniform, hyaline, thick-, smooth-walled, 5.7–24.6 μm long, 1.1–2.5 μm wide at base, with conspicuous periclinal thickening and minute collarette at conidiogenous loci; polyphialides with two conidiogenous loci occasionally present. Conidia aseptate, ellipsoid or ovoid, hyaline, thin-, smoothwalled, eguttulate, 2.2–3.3 × 1.9–2.5 μm, arranged in long chains. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA flat, spreading, felty and sulphur yellow at centre, membranous without aerial mycelium and greenish yellow at periphery, margin entire, abundant yellow pigment, reverse sulphur yellow; On MEA reaching 25–29 mm diam, flat, felty, granulose, slightly hairy at centre, white, with dirty white and entire margin, reverse umber; On PDA reaching 38–39 mm diam, flat, felty, slightly short hairy at centre, dirty white, margin entire, reverse pale olivaceous buff at centre, buff at periphery; On SNA reaching 32 mm diam, flat, sparse aerial mycelium, dusty, white, margin entire, reverse white. With strong geosmin odour on all media. Typus: Netherlands, Utrecht Province, Baarn, Cantonspark, from dead petiole of Chamaerops humilis (Arecaceae), unknown collection date and collector, isol. W. Gams, No. 1269 (holotype CBS H-8470, ex-type culture CBS 398.70). Additional material examined: Netherlands, Utrecht Province, Baarn, Cantonspark, from stem of Musa sp. (Musaceae), unknown collection date and collector, isol. Oct. 1967 by W. Gams, No. 1257, CBS H-8469, culture CBS 452.70. Fig. 10. Neoacremonium distortum (ex-type culture CBS 314.72). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Unbranched conidiophores. F–I. Branched conidiophores with monoor polyphialides. J. Conidia. Scale bars = 10 μm.
60 Hou et al. Notes: The two cultures representing Neoacremonium flavum were formerly identified as Acremonium vitellinum (currently Ne. vitellinum). However, they fall in a separate branch in Neoacremonium, phylogenetically distant from the branch bearing the ex-type culture of Ne. vitellinum, which also shows low sequence similarity (97 % on ITS, 98 % on LSU, 85 % on rpb2) (Fig. 1). Neoacremonium flavum can be easily distinguished from Ne. vitellinum based on its shorter phialides (5.7–24.6 μm) and conidia arranged in long chains, while Ne. vitellinum has longer phialides (32–80 μm) and conidia arranged in slimy heads (this study). Besides, Ne. flavum has greenish yellow colonies on OA, while those of Ne. vitellinum are dirty white to salmon. Neoacremonium minutisporum (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845948. Fig. 12. Basionym: Cephalosporium minutisporum Sukapure & Thirum., Mycologia 55: 566. 1963. Synonym: Acremonium minutisporum (Sukapure & Thirum.) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 51. 1971. Mycelium consisting of branched, septate, smooth, hyaline, thinwalled hyphae, up to 2.7 μm wide. Conidiophores solitary, erect, straight, arising from submerged or superficial hyphae, unbranched or branched at lower part, up to ca. 38.6 μm long, 1.2–3 μm wide at base, normally with 1–2-septa at base, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, subcylindrical or lageniform, hyaline, thin-, smooth-walled, often borne on short cylindrical subtending cells, 9.3–35.5 μm long, 1–3 μm wide at base, with inconspicuous minute collarette and periclinal thickening at conidiogenous loci; adelophialides present, 10–17 × 1.5–2.5 μm; polyphialides with up to three conidiogenous loci occasionally present. Conidia aseptate, short cylindrical, ovoid, both ends rounded, hyaline, thin-, Fig. 11. Neoacremonium flavum (ex-type culture CBS 398.70). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores with conidial chains. F–L. Conidiophores. M. Conidia. Scale bars = 10 μm.
61www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales smooth-walled, 2.5–4.3 × 1.2–1.7 μm, arranged in slimy heads. Chlamydospores laterally on short stalks, single, globose to subglobose, hyaline, smooth-, thick-walled, 2.5–3.7 × 2.5–3.4. Sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 65–70 mm diam, flat, membranous with sparse aerial mycelium, dirty white, margin entire, reverse concolourous; On MEA reaching 75–80 mm diam, flat, radially folded, hairy, dirty white, margin fimbriate, reverse ochreous, with radial lines; On PDA reaching 73– 80 mm diam, flat, membranous with sparse aerial mycelium, slightly hairy at centre, dirty white, margin fimbriate, reverse concolourous, with buff radial lines; On SNA reaching 75–80 mm diam, flat, membranous without aerial mycelium, colourless, margin fimbriate, reverse colourless. With geosmin odour on all media. Typus: India, Bombay, Sewri, from salt-marsh soil, 11 Jul. 1958, M.J. Thirumalachar, CBS H-6794 (holotype of Cephalosporium minutisporum HACC 108, ex-type cultures CBS 147.62 = ATCC 14612 = IMI 091576). Notes: Cephalosporium minutisporum was isolated from a soil sample collected in a salt marsh at the sea coast, an area which turns to clay in the summer, with few fungi surviving in this habitat (Sukapure & Thirumalachar 1963). This species was later transferred to the genus Acremonium as one of the species in Acremonium section Simplex (Gams 1971). According to the phylogenetic analysis, the ex-type of C. minutisporum (CBS 147.62) clusters with the new genus Neoacremonium (Fig. 1), and it is therefore recombined as Ne. minutisporum. This species is morphologically characterised by the production of single chlamydospores laterally borne on short stalks, a feature that distinguishes it from other species in this genus. Morphologically, characters of the ex-type Fig. 12. Neoacremonium minutisporum (ex-type culture CBS 147.62). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores with conidial heads. E–G. Monophialidic conidiophores. H, I. Adelophialide. J. Polyphialides. K, L. Chlamydospores. M. Conidia. Scale bars = 10 μm.
62 Hou et al. culture are similar to the description available in the literature (Gams 1971). Based on a blastn search of NCBIs GenBank nucleotide database, the closest hit using the LSU sequence of N. minutisporum is a sequence from the ex-isotype culture of Sedecimiella taiwanensis [currently Ne. taiwanense, CY5100; GenBank HM451496.1; Identity = 775/777 (99.74 %), no gaps]. However, only LSU sequences of the culture CY5100 were available for comparison, and there is only 2 bp difference between the LSU sequences of the ex-type cultures of Ne. minutisporum and Ne. taiwanense. Morphological comparison is difficult as Ne. taiwanense was only described based on a sexual morph (Pang et al. 2010), while Ne. minutisporum was only observed as an asexual morph in culture. Therefore, whether they are conspecific still awaits to be confirmed with more molecular data and morphological comparisons. Furthermore, if the two species proved to be conspecific, the Ne. minutisporum should be adopted as this species was published earlier than S. taiwanensis. Neoacremonium taiwanense (K.L. Pang et al.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 848117. Basionym: Sedecimiella taiwanensis K.L. Pang et al., Bot. Mar. 53: 495. 2010. Description and illustration: Pang et al. (2010). Typus: China, Taiwan, Chunan, on a twig of Kandelia obovata (Rhizophoraceae), 12 Aug. 2008, K.L. Pang (holotype BBH26390, BIOTEC Bangkok Herbarium, dried wood). Additional material examined: China, Shenzhen, Futian Nature Reserve, on a twig of unidentified mangrove wood, 14 Mar. 2006, K.L. Pang, exisotype culture CY5100-CY5101. Notes: This species was described based on a sexual morph that is characterised by producing 16 globose ascospores in asci; its asexual morph is unknown (Pang et al. 2010). Only LSU (HM451496) and SSU (HM451495) sequences are available, and the phylogenetic tree based on limited taxa in the original article shows this species is close to Niesslia exilis (CBS 357.70), on which basis it was placed in Niessliaceae (Pang et al. 2010). However, its LSU sequence is 99 % similar to CBS 147.62 (ex-type culture of Cephalosporium minutisporum, currently Neoacremonium minutisporum) based on the blastn search. Phylogenetically, Ne. taiwanense has a close phylogenetic affinity to Ne. minutisporum and distant from Niesslia exilis (Fig. 1). The habitat of the two species shares some similarities: Ne. taiwanense was described from a twig of a mangrove plant (Kandelia obovata) that grows in coastal areas, while Ne. minutisporum was from a soil sample collected in salt marsh areas on the sea coast (Sukapure & Thirumalachar 1963). Therefore, Ne. taiwanense is possibly the first species of Neoacremoniaceae with a known sexual morph. However, considering the close phylogenetic affinity, whether Ne. minutisporum and Ne. taiwanense are conspecific awaits to be clarified with more sequences or cultures. Neoacremonium vitellinum (W. Gams) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845949. Fig. 13. Basionym: Acremonium vitellinum W. Gams, Cephalosporiumartige Schimmelpilze: (Stuttgart): 65. 1971. Mycelium consisting of branched, septate, smooth, hyaline, thin-walled hyphae, up to 2 μm wide. Sporulation abundant, phalacrogenous, nematogenous, plectonematogenous. Conidiophores solitary or aggregated, erect, arising from submerged or superficial hyphae, straight or slightly curved, unbranched or branched, often with short sterile basal outgrowths, up to 82 μm long, 1–3 μm wide at base, normally with 1–2 septa at base, sometimes up to 4 septa, inflated at base, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal, lateral, cylindrical, hyaline, thick-, smooth-walled, often borne on short cylindrical subtending cells, 32–80 μm long, 1–2.2 μm wide at base, with cylindrical collarette and inconspicuous periclinal thickening at conidiogenous loci. Conidia aseptate, broad ellipsoid, ovoid, spindleshaped, often with an apiculate basal end and obtuse apices,, hyaline, thin-, smooth-walled, 2.8–4.9 × 1.6–2.5 μm, arranged in slimy heads. Crystals are usually abundantly formed. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 18 mm diam, flat, dusty, salmon at centre, white at periphery, margin entire, reverse pale saffron at centre, dirty white at periphery; On MEA reaching 22 mm diam, flat, abundant aerial mycelium, hairy, dirty white, buff at centre, margin entire, reverse orange at centre, saffron at periphery; On PDA reaching 22 mm diam, flat, abundant aerial mycelium, hairy, white, margin fimbriate, reverse pale orange at centre, dirty white at periphery; On SNA reaching 16 mm diam, flat, membranous without aerial mycelium, white, margin fimbriate, reverse white. Lacking odour on all media. Typus: Netherlands, Utrecht Province, Baarn, Cantonspark, greenhouse, from dead leaf of Phoenix dactylifera (Arecaceae), unknown collection date and collector, isol. Oct. 1967 by W. Gams, No. 1273, CBS H-6673 (holotype CBS 792.69 preserved as metabolically inactive culture, ex-type culture CBS 792.69). Additional material examined: Netherlands, Utrecht Province, Baarn, Cantonspark, greenhouse, from old petiole of Phoenix dactylifera (Arecaceae), unknown collection date and collector, isol. Oct. 1967 by W. Gams, No. 1274, CBS H-8472, culture CBS 793.69. Notes: Gams (1971) reported conidia of Acremonium vitellinum to be arranged in chains, and tends to form sporodochium based on culture CBS 452.70, which proved to be a different species (Fig. 1). However, in the present study, the ex-type culture of A. vitellinum (CBS 792.69) was observed to produce longer phialides [32–80 μm vs 15–25(–30) μm] and conidia arranged in slimy heads. In addition, adelophialides were not observed as described in Gams (1971). Parapyrenis maritima Aptroot, Nova Hedwigia 60: 354. 1995. Description and illustration: Aptroot (1995). Typus: Papua New Guinea, Madang Province, Laing Island in Hansa Bay near Bogia, alt. 1 m, from wood in coastal forest on coral island, 20 Jul. 1992, A. Aptroot (holotype CBS H-5923, ex-holotype culture CBS 538.93 = Aptroot No. 30166). Additional material examined: Japan, Wakayama, from seashore sand, unknown collection date, K. Tubaki (No. V-I), CBS H-8246, culture CBS 795.69. Notes: Parapyrenis maritima is a member of Parapyrenis originally placed in Requienellaceae, Xylariales. It is characterised by the production of globose ascomata that are immersed in a stroma, ellipsoid, medium brown ascospores, 1-septate, without spinulose ornamentation, and with a thick endosporium leaving a cordate to sexangular lumen. The asexual morph of this species is unknown (Aptroot 1995). The ex-type culture CBS 538.93, examined in
63www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales the present study, clustered within the Neoacremonium clade, with an acremonium-like species, Ne. vitellinum. Morphologically, culture CBS 538.93 was only observed as an acremoniumlike asexual morph, and therefore cannot be compared with the protologue. However, it is unknown whether Pa. maritima truly had the acremonium-like asexual morph or the culture was swapped or contaminated by an acremonium-like species at some point before or after it was deposited. DNA sequences derived from the holotype material of Pa. maritima (CBS H-5923; if possible) would help resolve this problem. Clade L Chrysonectriaceae L.W. Hou, L. Cai & Crous, fam. nov. MycoBank MB 845950. Classification: Hypocreales, Sordariomycetes. Sexual morph: ascomata superficial, non-stromatic, sub-globose, not collapsing upon drying, pale orange, changing colour in 3 % KOH and lactic acid, overlain by golden yellow hyphal elements; asci 8-spored, cylindrical to fusoid, with a refractive apical apparatus; ascospores subfusoid to narrowly clavate, two-celled, smooth. Asexual morph: mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae. Conidiophores solitary or aggregated, (sub-)erect, mostly curved, irregularly wavy, arising directly from submerged or superficial hyphae, verticillately branched, bearing 1–3 whorls of 1–3 phialides per node, rarely unbranched and reduced to single phialides, septate at base and middle, hyaline, thickand smooth-walled, cell walls usually thicker than those of vegetative hyphae. Phialides lateral or terminal, cylindrical or subulate, straight or curved at base, hyaline, Fig. 13. Neoacremonium vitellinum (ex-type culture CBS 792.69). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores with conidial heads. F. Unbranched conidiophore. G, H. Conidiophores with sterile basal outgrowths. I–K. Branched conidiophores. L. Conidia. Scale bars = 10 μm.
70 Hou et al. clade and is phylogenetically closely related to Pn. foliicola (Fig. 2). Morphologically, Pn. aurantiacum differs from H. erubescens in producing pale yellow to orange and non-ostiolate ascomata, and differs from Pn. foliicola in its larger asci [(53–)59–81(–83) × 8–11.5 μm vs 50–60 × 8–10 μm], and larger ascospores with 3 septa (19– 26.5 × 3.5–5 μm, 3-septate vs (14–)15–17 (–18) × 4–4.7(–5) µm 1–3-septate) (Rossman et al. 1999, Lechat & Fournier 2017). Paracylindrocarpon foliicola Lechat & J. Fourn., sp. nov. MycoBank MB 845815. Basionym: Hydropisphaera foliicola Lechat & J. Fourn., Ascomycete.org 9: 6. 2017. (nom. inval., Art. F.5.1 (Shenzhen); the identifier cited in the protologue, MB 815589, was not issued for that name). Etymology: “foliicola” referring to its habitat on a dead leaf. Description and illustration: Lechat & Fournier (2017). Typus: France, Martinique, Fort-de-France, forêt de Colson, PlateauPerdrix, on dead leaf of Pouteria pallida (Sapotaceae), 20 Aug. 2015, C. Lechat (holotype CLLM15128 in LIP, ex-type culture CBS 140758). Notes: Hydropisphaera foliicola was described based on the culture from Pouteria pallida (Sapotaceae) of France (Lechat & Fournier 2017). However, when this species was published, the identifier cited in the protologue was not issued for that name. Thus, the epithet is invalid based on the International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code). Here, this species is validated by describing it as a new species of Paracylindrocarpon. Phylogenetically, the ex-type strain CBS 140758 clustered within Paracylindrocarpon and is closely related to Pn. aurantiacum (Fig. 2). Despite the similarity with Hydropisphaera species in the production of perithecia that are collapsing and cupulate when Fig. 17. Paracylindrocarpon aurantiacum (ex-type culture CBS 135909). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Ascomata on pine needle. E. Ascoma. F–H. Asci. I. Ascospores. Scale bars: E = 100 μm; F–I = 10 μm.
71www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales dry, its 1–3-septate ascospores match better with the generic characters of Paracylindrocarpon, while species in Hydropisphaera s. str. produce 1-septate ascospores. Paracylindrocarpon multiloculatum (Samuels) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845816. Basionym: Nectria multiloculata Samuels, New Zealand J. Bot. 16: 78. 1978. Synonym: Hydropisphaera multiloculata (Samuels) Rossman & Samuels, Stud. Mycol. 42: 31. 1999. Description and illustration: Samuels (1978). Typus: New Zealand, Auckland, Waitemata County, Waitakere Ranges, Lucy Cranwell Track, vic. Kitakiki Stream, on dead leaf of Astelia sp. (Asteliaceae), 30 May 1973, J.M. Dingley, G.J. Samuels & S. Haydon, G.J.S. 73-98 (holotype PDD 31786, ex-type culture CBS 339.77). Additional materials examined: New Zealand, Little Barrier Island, track to summit of Mt. Hauturu, from dead leaf of Astelia sp. (Asteliaceae), 22 Feb. 1976, G.J. Samuels, G.J.S. 76-8, PDD 34940, culture CBS 340.77. Notes: Paracylindrocarpon multiloculatum was originally described as Nectria multiloculata by Samuels (1978) from a dead leaf of Astelia sp. in New Zealand. It was subsequently transferred to Hydropisphaera (Rossman 1999). The original materials examined by Samuels (1978), i.e., CBS 339.77 and CBS 340.77 are included in the present study. According to our phylogenetic inference, the ex-type of N. multiloculata CBS 339.77 falls in a fully supported clade representing the genus Paracylindrocarpon. Paracylindrocarpon multiseptatum (Samuels) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845817. Basionym: Nectria multiseptata Samuels, New Zealand J. Bot. 16: 77. 1978. Synonym: Hydropisphaera multiseptata (Samuels) Rossman & Samuels, Stud. Mycol. 42: 31. 1999. Description and illustration: Samuels (1978). Typus: New Zealand, Auckland, Waitakere Range, vic. Piha, Marowhara Loop Track, from leaf of Phormium tenax (Asphodelaceae), 17 Dec. 1974, J.M. Dingley, G.J. Samuels & S. Francis, G.J.S. 74-136, PDD 34937, culture CBS 337.77. Additional materials examined: New Zealand, Auckland, Waitakere Range, vic. Piha, Marowhara Loop Track, from leaf of Phormium tenax, 19 Jul. 1973, G.J. Samuels, G.J.S. 73-141, PDD 32423, culture CBS 336.77; Wenderholm Scenic Reserve, from midrib of Rhopalostylis sapida (Arecaceae), 26 Sep.1973, J.M. Dingley et al., G.J.S. 73-205, PDD 32658, culture CBS 333.77. Clade O3 Fusariella Sacc., Atti Ist. Veneto Sci. Lett. Arti., Sér. 6 2: 463. 1884. Colonies compact or effuse, greyish green, blackish green or black. Mycelium superficial and immersed. Stroma none. Setae and hyphopodia absent. Conidiophores semi-macronematous, mononematous, branched irregularly or sometimes dichotomously or trichotomousIy, flexuous, colourless or pale brown, smooth or verruculose. Conidiogenous cells monophialidic, integrated and terminal, or discrete, determinate, often curved, cylindrical, subulate or lageniform, with collarettes. Conidia catenate, acrogenous, semiendogenous, developing in basipetal succession and frequently hanging together in slipped chains, the tip of each conidium except the apical one being deflected laterally, simple, straight, bent or flexuous, often fusiform pointed at the apex blunt at the base but sometimes cylindrical, dumb-bell-shaped, clavate or obclavate, pale to mid olive brown, olive green or greyish green, pale greyish green, blackish green or black in mass, usually smooth, 1–3-septate (Ellis 1971). Type: Fusariella atrovirens (Berk.) Sacc. Other accepted species with available sequences: Fusariella arenula (Berk. & Broome) L.W. Hou, L. Cai & Crous, F. concinna (Syd.) S. Hughes, F. curvata C.G. Lin, Yong Wang bis & K.D. Hyde, F. hughesii Chab.-Frydm. Notes: The genus Fusariella is characterised by semito macronematous, mononematous conidiophores, with cylindrical, subulate or lageniform phialidic conidiogenous cells, which produce catenate, septate, curved to straight, subhyaline to brown conidia (Hughes 1949, Chabelska-Frydman 1964, Roy & Rai 1968, Ellis 1971, 1976, Seifert et al. 2011, Lin et al. 2016). Fusariella has been known for more than 130 years since it was established by Saccardo (1884), and its classification remained uncertain because of the lack of molecular data and comprehensive taxonomic treatment (Lin et al. 2016). Recently, a phylogenetic analysis based on SSU, LSU, tef1-α and rpb2 sequence data indicated that the genus Fusariella belongs to the family Bionectriaceae (Lin et al. 2016). Our study agrees with this conclusion that Fusariella clusters in Bionectriaceae, closely related to Paracylindrocarpon (Fig. 2). In addition, two Hydropisphaera species, H. cirsii and H. arenula, clustered within the Fusariella clade (Fig. 2, Supplementary Fig. S1). Both species have fusoid or subfusoid ascospores and are finely spinulose (Samuels 1978, Lechat & Fournier 2020), while most species of Hydropisphaera s. str. have ellipsoid, smoothwalled ascospores (Rossman et al. 1999). However, H. cirsii only had ITS and LSU sequences data available (Supplementary Fig. S2), and therefore the phylogenetic position of this species could not be confirmed. Fusariella arenula (Berk. & Broome) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845818. Basionym: Sphaeria arenula Berk. & Broome, Ann. Mag. Nat. Hist., Ser. 2, 9: 320. 1852. Synonyms: Nectria arenula (Berk. & Broome) Berk., Outl. Brit. Fung. (London): 394. 1860. Dialonectria arenula (Berk. & Broome) Cooke, Grevillea 12(no. 64): 110. 1884. Cucurbitaria arenula (Berk. & Broome) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 460. 1898. Hydropisphaera arenula (Berk. & Broome) Rossman & Samuels, Stud. Mycol. 42: 30. 1999. Description and illustration: Samuels (1978). Materials examined: New Zealand, Auckland, Waitemata County, Riverhead State Forest, 2 m NE of Riverhead, from leaf of Phormium tenax (Asphodelaceae), 23 Jun. 1973, G.J. Samuels, C.S. Samuels & D.R.W. Watson, G.J.S. 73-125, PDD 31886, culture CBS 329.77; idem. G.J.S. 73131, PDD 31887, culture CBS 330.77. Notes: This species was originally introduced as Sphaeria arenula (Berkeley & Broome 1852). The taxonomic history of this species is complex and has been addressed by multiple authors (Berkeley 1860, Cooke 1884, Booth 1959, Samuels 1978). Booth (1959)
72 Hou et al. examined the co-type specimen and described the ascospores as fusoid to ellipsoid, occasionally slightly curved, and hyaline with faint longitudinal striations when mature, measuring 15–18 × 3–5 μm. Samuels (1978) recombined Sphaeria arenula as Nectria arenula, but did not examine the holotype specimen. Later it was transferred to Hydropisphaera based on morphology (Rossman et al. 1999), and the recent phylogenetic analysis based on LSU sequences supported the result of Rossman et al. (1999), showing that this species clustered in Hydropisphaera (Lechat & Fournier 2020). Although most Hydropisphaera species were included in the study of Lechat & Fournier (2020), a limited number of strains/species of its related genera, especially Acremonium, Fusariella and Paracylindrocarpon, were included in the phylogenetic analysis (Lechat & Fournier 2020). In the present study, this species appears phylogenetically distant from Hydropisphaera s. str. as circumscribed in this study (Fig. 2), clustering within the Fusariella clade. Morphologically, its curved and multi-septate conidia are in agreement with the generic characters of Fusariella, and it differs from other species in its conidia that are not constricted at their septa, while conidia of most Fusariella species are commonly constricted at their conidial septa. Fusariella atrovirens (Berk.) Sacc., Atti Reale Ist. Veneto Sci. Lett. Arti ser. 6, 2: 463 (1884) Basionym: Fusisporium atrovirens Berk., Engl. Fl., Fungi 5(2): 351. 1836. Synonym: Fusarium atrovirens (Berk.) Mussat, Syll. Fung. (Abellini) 15: 144. 1901. Material examined: Algeria, Western Sahara, near Béni-Abbès (Saoura), from desert soil, Oct. 1972, unknown collector, isol. J. Nicot & J. Mouchacca, culture CBS 311.73 = IMI 171130 = LCP 2177. Fusariella hughesii Chab.-Frydm., Canad. J. Bot. 42: 1485. 1964. Description and illustration: Chabelska-Frydman (1964). Material examined: Netherlands, Flevoland Province, Oostelijk Flevoland, from agricultural soil, under permanent potato cultivation, unknown collection date and collector, isol. Oct. 1969 by J.W. Veenbaas-Rijks, culture CBS 435.70. Fusariella sp. Material examined: China, Hubei, Shennongjia, alt. 1 800 m, rotten twig, 17 Sep. 2006, W.Y. Zhuang & N. Ye, 5805, culture CBS 128364. Notes: The culture was isolated from the rotten twig collected from Shennongjia in China and was originally labelled as Hydropisphaera erubescens (Nong & Zhuang 2005). However, the phylogeny based on sequence obtained from the ITS, LSU, rpb2 and tef-1α place this strain in a sister branch to Fusariella hughesii (Fig. 2). Unfortunately, the culture was sterile and morphological comparison with other Fusariella species was impossible. This is probably a novel species and awaits further study. Clade O8 Musananaesporium L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845821. Etymology: Name derived from the banana-shaped (Musa nana) conidia. Mycelium consisting of branched, septate, (sub-)hyaline, thin-, smooth-walled hyphae, brown and thick-walled in old cultures. Conidiophores arising from submerged mycelium or ropes formed by the mycelium, erect, with multiple septa, unbranched or branched, commonly repeatedly proliferating sympodially. Conidiogenous cells enteroblastic, monoto polyphialidic, solitary, lateral, cylindrical or subulate, hyaline to (sub-)hyaline, thick-, smooth-walled, with inconspicuous collarette and periclinal wall thickening at conidiogenous loci , with percurrent or subterminal proliferations. Polyphialides with up to four conidiogenous loci occasionally present. Conidia aseptate, becoming 1–4-septate with age, cylindrical, obclavate, fusoid, with a protuberant and curved, basal abscission scar, straight to curved, thick-, smoothwalled, hyaline, produced in slimy heads. Typical chlamydospores absent, but hyphae disintegrate into arthrospore-like fragments in old cultures. Sexual morph not observed. Type: Musananaesporium tectonae (R.F. Castañeda) L.W. Hou, L. Cai & Crous Notes: This monotypic genus, Musananaesporium, is established to accommodate Acremonium tectonae, since it is not congeneric with Acremonium s. str. based on A. alternatum. Musananaesporium tectonae is represented by a single lineage in the phylogenetic analysis (Fig. 2). The morphology of this species is unique in producing branched, repeatedly sympodially proliferating conidiophores and phialides, and cylindrical, obclavate, fusoid conidia, with a protuberant, curved basal abscission scar, rendering it different from other known genera in Bionectriaceae (Fig. 2). Musananaesporium tectonae (R.F. Castañeda) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845822. Fig. 18. Basionym: Acremonium tectonae R.F. Castañeda [as ‘Acromoniun’], Fungi Cubenses II (La Habana): 2. 1987. Mycelium consisting of branched, septate, (sub)hyaline, thin-, smooth-walled hyphae, brown and thick-walled in old cultures, 3–4 μm wide. Sporulation phalacrogenous , plectonematogenous, and nematogenous. Conidiophores (suberect, straight, arising from submerged hyphae or ropes of hyphae, unbranched or basitonously branched, bearing 1–2 phialides per node, often repeatedly proliferating sympodially, up to 268 μm long, 2.7–6.2 μm wide at base, with 1–7 septa at basal, middle and apical part, with cell walls usually thicker than those of vegetative hyphae. Phialides solitary, lateral, cylindrical or subulate, hyaline to (sub-)hyaline, thick-, smooth-walled, (15–) 25–76(–120) long, 1.7–6.2 μm wide at base, with inconspicuous collarette and periclinal wall thickening at conidiogenous loci; polyphialides terminally and subterminally proliferating, with up to four conidiogenous loci occasionally present. Conidia aseptate, becoming 1–4-septate with age, ellipsoidal, cylindrical, obclavate and fusoid, straight, curved at base, with a protuberant and curved basal abscission scar, thick-, smooth-walled, hyaline, (9–)11.5–25.5(–28.5) × 2.5–4.5 μm, produced in slimy heads. Typical chlamydospores absent, but catenate hyphae disintegrate into arthroconidium-like fragments in old cultures, hyaline, thick-, smooth-walled, 4.2–9.5 long, 3–4 μm diam. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 20 mm diam, flat, with sparse aerial mycelium, dusty, becoming felty at some zones with age, pale saffron, margin entire, with brown pigment produced on old culture, reverse concolourous; On MEA reaching 11–14 mm diam, raised, with sparse aerial mycelium, cerebriform, peach, margin dendritic, reverse apricot; On PDA reaching 10–13 mm diam, raised, cerebriform, with sparse aerial mycelium, dusty, pale saffron, margin crenate, reverse ochreous,
73www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales with slightly pale brown pigment; On SNA reaching 15–17 mm diam, flat, membranous without aerial mycelium, colourless, margin entire, reverse colourless. Typus: Cuba, Matanzas Province, San Miguel de Los Baños, from living leaf of Tectona grandis (Lamiaceae), isol. 23 Jan. 1987, R.F. Castañeda & G. Arnold, CBS H-24609 (holotype INIFAT C87/32, ex-type culture CBS 725.87). Notes: Musananaesporium tectonae was originally described as A. tectonae by Castañeda (1987) from a living leaf of Tectona grandis in Cuba. The ex-type culture was examined in this study, and morphologically matched the original description except for a few characters. In its original description, the conidiophores were described as macronematous and mononematous, phialides were monophialidic, terminal or lateral, conidia were unicellular, and no mention was made of the production of moniliform hyphae and the septate conidia. In the present study, the septate and thick-walled conidia are commonly produced in old cultures, becoming up to 4-septate. In addition, arthroconidium-like hyphae are present, which are much thicker than normal hyphae. This species differs from the other acremonium-like taxa by producing longer, septate conidia and abundant macronematous conidiophores. According to the phylogenetic inference, the ex-type Fig. 18. Musananaesporium tectonae (culture CBS 725.87) A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores with conidia arranged in slimy heads. F. Unbranched conidiophore. G, H. Percurrently proliferated conidiophores. I. Polyphialide. J. Branched conidiophore. K. Septate mycelium. L. Conidia. Scale bars = 10 μm.
74 Hou et al. culture CBS 725.87 falls in a separate lineage, representative of a novel genus in Bionectriaceae (Fig. 2). Clade O9 Gossypinidium L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845819. Etymology: From Latin, gossypina (= cottony), referring to the cottony sporodochia produced by the type of this genus. Mycelium consisting of branched, septate, hyaline at first, pale brown in old cultures, slightly warted and thick-walled hyphae. Crystals absent. Sporulation abundant from sporodochia, rarely from conidiophores formed directly on substratal or aerial mycelium. Conidiophores in aerial mycelium erect, straight, slightly curved, repeatedly verticillate towards the apex, bearing 1–3(–4) whorls of 1–3 phialides, rarely unbranched, septate, hyaline, smooth-walled. Conidiogenous cell from solitary conidiophores enteroblastic, monophialidic, mostly lateral, sub-cylindrical to subulate, hyaline, thick-, smooth-walled, with inconspicuous minute collarette and periclinal thickening at conidiogenous loci. Conidiophores in pale pink or orange cottony sporodochia mostly branched, hyaline, smooth-walled. Sporodochial phialides enteroblastic, monophialidic, mostly lateral, aculeate or subulate, tapering at top, hyaline, thick-, smooth-walled, with minute collarette and inconspicuous conspicuous periclinal thickening at conidiogenous loci. Conidia from solitary conidiophores and sporodochial conidiophores not significantly different, aseptate, short ellipsoid, hyaline, thin-, smooth-walled, arranged in chains. Chlamydospores and sexual morph not observed. Type: Gossypinidium sporodochiale L.W. Hou, L. Cai & Crous Notes: Gossypinidium presents a distinct lineage on the 4-locus (ITS-LSU-rpb2-tef-1α) phylogenetic tree (Fig. 2). It produces abundant cottony sporodochia with abundant conidia arranged in chains, a character that clearly distinguishes it from other genera of Bionectriaceae. Gossypinidium sporodochiale L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845820. Fig. 19. Etymology: Named after the abundant sporodochia produced in culture by this species. Mycelium consisting of branched, septate, hyaline at first, pale brown in old cultures, slightly warted, thick-walled hyphae, up to 4 μm wide. Crystals absent. Sporulation abundant from sporodochia, rarely from conidiophores formed directly on the aerial or substratal mycelium. Conidiophores from aerial mycelium erect, straight or slightly curved, repeatedly verticillate towards the apex, bearing up to 4 whorls of 1–3 phialides, or unbranched, up to 60 μm long, 1–2.4 μm wide at base, with 1–3 septa, hyaline, smooth-walled. Phialides from solitary conidiophores mostly lateral, sub-cylindrical to subulate, hyaline, thick-, smooth-walled, 10–25 μm long, 1.3– 2.3 μm wide at base, with minute collarette and inconspicuous periclinal thickening at conidiogenous loci. Conidiophores in pale pink or orange cottony sporodochia mostly branched, bearing multiple levels with 1–3 phialides per node, hyaline, smoothwalled. Sporodochial phialides mostly lateral, aculeate or subulate, tapering at top, hyaline, thick-, smooth-walled, 10–15 μm long, 1.5–2 μm wide at base, with minute collarette and inconspicuous conspicuous periclinal thickening at conidiogenous loci. Conidia from solitary conidiophores and sporodochial conidiophores not significantly different, aseptate, short ellipsoid, hyaline, thin-, smooth-walled, 3–4.6 × 2–2.8 μm, eguttulate, arranged in chains. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 46–47 mm diam, flat, floccose, granulose at centre, salmon at centre, white at periphery, margin crenate, reverse buff at centre, dirty white at periphery; On MEA reaching 30–40 mm diam, raised, felty or woolly, rosy buff at centre, buff at periphery, margin lobulated, reverse umber, pale umber at periphery; On PDA reaching 75 mm diam, flat, felty, white, buff or creamy white at periphery, margin fimbriate, reverse pale brown at centre, buff at periphery. Typus: Puerto Rico, Luquillo National Forest, El Yunque Trail, from dead rachis of Praestoea montana (Arecaceae), Jun. 1998, W. Gams (holotype CBS H-24608, ex-type culture CBS 101694). Notes: The strain CBS 101694 formed a distinct branch and was originally identified as A. persicinum based on the LSU and SSU phylogenetic analysis for the majority of Acremonium (Summerbell et al. 2011), although phylogenetically different from other A. persicinum strains, including the ex-type CBS 310.59 (Summerbell et al. 2011). Phylogenetic analysis based on more loci (ITS-LSU-rpb2-tef-1α) shows that CBS 101694 forms a distinct branch in Bionectriaceae (Fig. 2). It differs from closely related genera by producing repeatedly verticillate conidiophores and cottony sporodochia with abundant ellipsoid conidia arranged in long chains. Based on a blastn search of NCBIs GenBank nucleotide database, the closest hits using the ITS sequence are Fusariella curvata from a leaf of decaying Quercus sp. in Thailand [culture MFLUCC 15-0844; GenBank KX025152; Identity = 473/522 (91 %), 11 gaps (2 %)] and F. atrovirens [culture CBS 311.73; GenBank MH860688.1; Identity = 472/522 (90 %), 11 gaps (2 %); Lin et al. 2016]; the closest hit using the LSU sequence is Acremonium persicinum [culture CBS 469.67; GenBank MH870741.1; Identity = 760/778 (98 %), 3 gaps (0 %)]; the closest hit using the rpb2 sequence is Hydropisphaera peziza [culture CBS 102038; GenBank DQ522444.1; Identity = 664/756 (98 %), no gaps]; the closest hit using the tef-1α sequence is Fusariella curvata [culture MFLUCC 15-0844; GenBank KX025155.1; Identity = 734/771 (95 %), 2 gaps (0 %)]. Clade O11 Monohydropisphaera L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845823. Etymology: Referring to its similarity with Hydropisphaera. Mycelium with hyphae branching, septate, hyaline to pale brown, smooth. Perithecia solitary or crowded in groups of 2–10, superficial, subglobose, reddish brown, collapsing cupulate when dry, not changing colour in 3 % KOH or lactic acid. Perithecial apex with short, acute papilla, margin with fasciculate, thick-walled hairs, arising from cells of ascomatal wall. Hairs brownish orange, cylindrical, slightly flexuous, thick-walled, rounded at tips, septate. Perithecial wall composed of two regions: outer region of globose to ellipsoid, thick-walled cells, inner region of elongate, flattened, thin-walled cells. Asci unitunicate, clavate, apices rounded, without ring, with eight biseriate ascospores. Ascospores aseptate, fusoid, hyaline, striate with striations finely verrucose. Asexual morph: conidiophores borne on aerial hyphae, macronematous, mononematous, unbranched, elongate, erect, straight to flexuous,
75www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales hyaline to light brown, surface smooth to very faintly roughened. Conidiogenous cells integrated, monophialidic, terminal, subulate towards apex, at times with a minutely flared collarette. Conidia solitary or catenulate in short chains, obpyriform to fusiform with apices rounded to somewhat acute and bases having a prominent, almost apiculate hilum, aseptate, walls smooth to verrucose, hyaline, becoming dark brown, paler and dark brown to black in mass (emended from Lechat et al. 2010). Type: Monohydropisphaera fusigera (Berk. & Broome) L.W. Hou, L. Cai & Crous Notes: Monohydropisphaera is proposed to accommodate the single species M. fusigera, which was originally received as Hydropisphaera fusigera (Lechat et al. 2010), but distinct from Hydropisphaera s. str. in the multi-locus phylogenetic tree in this study with more strains and loci included for phylogenetic inferences (Fig. 2). Morphologically, the ascospores of Monohydropisphaera are aseptate and coarsely striate with striae somewhat wavy, which are distinct characters that differentiate it from other genera in Bionectriaceae. Monohydropisphaera fusigera (Berk. & Broome) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845824. Basionym: Monotospora fusigera Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74): 99. 1873 (1875). Synonyms: Gliomastix fusigera (Berk. & Broome) C.H. Dickinson, Mycol. Pap. 115: 7. 1968. Acremonium fusigerum (Berk. & Broome) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 94. 1971. Hydropisphaera bambusicola Lechat, Mycotaxon 111: 96. 2010. Hydropisphaera fusigera (Berk. & Broome) Rossman et al., Stud. Mycol. 80: 242 (2015). Fig. 19. Gossypinidium sporodochiale (ex-type culture CBS 101694). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Sporodochia on SNA. E. Sporodochia. F. Unbranched conidiophore. G–I. Branched conidiophores. J. Conidia. Scale bars = 10 μm.
76 Hou et al. Description and illustration: Lechat et al. (2010). Material examined: French West Indies, Martinique, Prêcheur, Anse Couleuvre, sentier de la cascade Couleuvre, in dead tops of Bambusa vulgaris (Poaceae), 25 Aug. 2008, C. Lechat (holotype of Hydropisphaera bambusicola CLL8323 in LIP, ex-type culture CBS 124147). Notes: This species was originally linked to the sexual morph H. bambusicola that characterised by the production of ascomata that often collapse upon drying to form cupulate perithecia (Lechat et al. 2010), and was transferred to Hydropisphaera, as H. fusigera (Lombard et al. 2015). However, species in Hydropisphaera s. lat. had proved to be highly polyphyletic (Rossman et al. 2001). In the present study, H. fusigera forms a separate lineage that is distinct from the Hydropisphaera s. str. clade (Fig. 2), and also remote from any known genera in Bionectriaceae. Morphologically, H. fusigera produces aseptate ascospores that are coarsely striate with somewhat wavy striae, which is rare in Hydropisphaera (Lechat et al. 2010). And although H. fusigera resembles some species of Protocreopsis, it differs from the latter by lacking white to tan hyphae that envelop the ascomatal wall (Doi 1977). The asexual morph of H. fusigera was reported to belong to the genus Gliomastix as G. fusigera (Lechat et al. 2010), which is characterised by producing dark brown conidia in chains occurring on members of the Arecaceae and Poaceae throughout the tropics (Lechat et al. 2010). However, it is distinct from other Gliomastix species in having much larger conidia that are longer than 12 µm which is rare in Gliomastix, and therefore further supports it as a different genus. Clade O12 Hydropisphaera Dumort., Comment. Bot. (Tournay): 89. 1822. Synonyms: Nectria subgen. Hyphonectria Sacc., Syll. Fung. (Abellini) 2: 501. 1883. Neohenningsia Koord., Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., sect. 2, 13: 164. 1907. Perrotiella Naumov, Zap. Ural’sk. Obshch. Lyubit. Estestv. 35(11– 12, Champ. Oural.): 25. 1916. Heleococcum C.A. Jørg., Bot. Tidsskr. 37: 417. 1922. Neuronectria Munk, Dansk bot. Ark. 17(1): 56 (1957). Hyphonectria (Sacc.) Petch, J. Bot. 75: 220 (1937). Ascomata superficial, perithecial, rarely cleistothecial, nonstromatic, pale yellow, orange or umber, KOH-, globose to subglobose or doliiform, usually collapsed and deeply cupulate, smooth or with fasciculate hairs. Ascomatal wall generally over 25 μm thick, of two regions; outer region of thin-walled, globose cells. Asci clavate. Ascospores ellipsoid, 1-septate, rarely multiseptate, hyaline, generally finely to coarsely striate, rarely smooth or spinulose (emended from Rossman et al. 1999). Type: Hydropisphaera peziza (Tode) Dumort. Other accepted species with available sequences: H. aurantiaca (C.A. Jørg.) L.W. Hou, L. Cai & Crous, H. cyatheae (Dingley) Rossman & Samuels, H. fungicola Rossman et al., H. suffulta (Berk. & M.A. Curtis) Rossman & Samuels Notes: The genus Hydropisphaera was established by Dumortier based on H. peziza (Dumortier 1822). This genus comprised a number of species with nectria-like sexual morphs and had previously been placed in the Nectria peziza group (Booth 1959, Samuels 1976b, Rossman 1983). Hydropisphaera was long considered a synonym of the genus Nectria until Rossman et al. (1999) resurrected it as a distinct genus in Bionectriaceae (Rossman et al. 1999, Lechat et al. 2010). Hydropisphaera s. lat. is characterised by producing ascomata with walls that are generally over 25 μm thick, deeply collapsed, cupulate perithecia upon drying, and oneto multi-septate ascospores that are often finely to coarsely striate, spinulose or smooth (Rossman et al. 1999). The asexual morph of Hydropisphaera has been recognised and placed in diverse genera including acremonium-like, cylindrocarpon-like, Cephalosporium or Gliomastix (Samuels 1978, Rossman et al. 2008, Lechat et al. 2010, Lechat & Fournier 2016a, 2017). Although more than 30 species have been described in Hydropisphaera, this genus has been revealed to be highly polyphyletic (Rossman et al. 2001). In our present phylogenetic analysis based on combined ITS, LSU, rpb2 and tef-1α genes, Hydropisphaera species were distributed in at least eight clades that could be differentiated from each other based on their phylogenetic and morphological characters (Fig. 2). Hydropisphaera is here restricted to those taxa related to the type H. peziza; they produce only an acremonium-like asexual morph, and have thick-walled ascomata with 1-septate, finely to coarsely striate ascospores. Furthermore, the cleistothecial genus Heleococcum is synonymised under Hydropisphaera based on the phylogenetic placement of the type, He. aurantiacum. Hydropisphaera aurantiaca (C.A. Jørg.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845825. Basionym: Heleococcum aurantiacum C.A. Jørg., Bot. Tidsskr. 37: 417. 1922. Synonym: Heleococcum japonense Tubaki, Trans. Mycol. Soc. Japan 8: 5. 1967. Description and illustration: Jørgensen (1922). Typus: Denmark, Botanical Garden of the University of Copenhagen, in the door, on the moist soil, 1921, L.K. Rosenvinge (C; NY, slides of holotype). Unknown, unknown substrate, unknown collection date and collector, BPI 691939 (recorded in BPI as type). Additional materials examined: Japan, Hokkaido, from wood panel of Abies firma (Pinaceae) in seawater, Oct. 1966, K. Tubaki (holotype of Heleococcum japonense CBS H-13245, ex-type culture CBS 397.67 = ATCC 18157 = IFO 8643). Unknown, from mushroom compost, unknown collection date and collector, dep. F.C. Wood, culture CBS 201.35. Notes: Although Heleococcum has cleistothecial ascomata and lacks a nectrioid centrum, its fleshy, bright-coloured ascomata, 1-septate, hyaline ascospores and acremonium-like asexual morph suggest that Heleococcum fits well in Hypocreales and it was therefore placed in Bionectriaceae (Rossman et al. 1999). In the present study, the reference culture of He. aurantiacum (CBS 201.35) and the ex-type culture He. japonense (CBS 397.67) had identical sequences on four genes and grouped in a fully supported clade in the Hydropisphaera clade, closely related to Hy. peziza (Fig. 2), based on which we concluded that they are conspecific, with He. aurantiacum having priority. Although the morphological features of the two species are not always constant, the discrepancies could be influenced by external factors. Hydropisphaera cirsii Lechat & J. Fourn., Ascomycete.org 12: 39. 2020. Description and illustration: Lechat & Fournier (2020). Typus: Germany, North Rhine-Westphalia, MTB 4506/441 Duisburg, cityforest, Uhlenhorstweg, S of cultural monument ‘Steinbruch’, 51°41’27” N, 6°80’41” E, on Cirsium arvense (Asteraceae), 17 Feb. 2013, leg. K.
77www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Müller, comm. K. Siepe (holotype CLL 13012 in LIP, ex-type culture CBS 135615). Notes: When this species was initially described, the only available LSU sequences placed it within the genus Hydropisphaera, which formed a long branch with low support values (Lechat & Fournier 2020). Hydropisphaera cirsii was observed to be different from known species of Hydropisphaera in having erect, glassy hairs scattered on its lateral ascomatal wall, finely spinulose ascospores and its occurrence on Cirsium arvense (Lechat & Fournier 2020). Based on the phylogenetic analysis performed on the four-gene alignment and more available cultures, this species forms a single lineage in Fusariella (Supplementary Fig. S2). The phylogenetic position of this species remains unresolved pending more cultures and DNA sequence data. Hydropisphaera cyatheae (Dingley) Rossman & Samuels, Stud. Mycol. 42: 30. 1999. Basionym: Nectria cyatheae Dingley [as ‘cyathea’], Trans. Roy. Soc. New Zealand. 83: 652. 1956. Illustration: Samuels (1976b). Typus: New Zealand, Auckland, on stipes of Cyathea medullaris (Cyatheaceae), Apr. 1984, J.M. Dingley (holotype PDD 6201). Material examined: New Zealand, Auckland, Waitakere Ranges, Rangemore Track, on rachis of Cyathea medullaris (Cyatheaceae), 10 Jul. 1974, G.J. Samuels, G.J.S. 74-103, PDD 32563, culture CBS 575.76. Hydropisphaera erubescens (Roberge ex Desm.) Rossman & Samuels, Stud. Mycol. 42: 30. 1999. Basionym: Sphaeria erubescens Roberge ex Desm., Ann. Sci. Nat., Bot., sér. 3, 6: 72. 1846. Synonyms: Calonectria erubescens (Roberge ex Desm.) Sacc., Michelia 1 (no. 3): 309. 1878. Nectria erubescens (Roberge ex Desm.) W. Phillips & Plowr., Grevillea 10 (no. 54): 70. 1881. Dialonectria erubescens (Roberge ex Desm.) Cooke, Grevillea 12: 111. 1884. Calonectria umbelliferarum Seaver, Mem. New York Bot. Gard. 6: 507. 1916. Amphinectria erubescens (Roberge ex Desm.) Sacc. ex Speg., Bol. Acad. Nac. Ci. Córdoba 26(2–4): 347. 1921. Calonectria venezuelensis Syd., Ann. Mycol. 33: 88. 1935. Calonectria crescentiae Seaver & Waterston, Mycologia 32: 404. 1940. Dimerosporiella guarapiensis (Speg.) Rossman & Samuels, Stud. Mycol. 42: 23. 1999. Descriptions: Rossman (1983), Samuels (1978). Typus: France, on old leaves of Ilex aquifolium (Aquifoliaceae), 1846, J.B.H.J. Desmazières, lectotype of Sphaeria erubescens, NY No.1766. Notes: This species was originally described by Desmazières (1846) as Sphaeria erubescens from old leaves of Ilex aquifolium in France based on the sexual morph. Our present study includes three cultures (CBS 333.77 = PDD 32658, CBS 334.77 = PDD 32478, CBS 335.77 = PDD 34939) that were examined and demonstrated to have identical morphological characters to type specimens of Sphaeria erubescens (currently H. erubescens; Samuels 1978). Based on the phylogenetic analysis, the three strains are genetically heterogeneous. Sequences obtained from CBS 334.77 are identical to the ex-type strain of Acremonium strictum (currently Sarocladium strictum, CBS 346.70; Fig. 3), suggesting that CBS 334.77 was swapped or contaminated by A. strictum at some point before or after it was deposited, since the morphology does not match what was described and illustrated as protologue of S. erubescens (Samuels 1978). The sequences obtained from the culture CBS 333.77 were identical to H. multiseptata (currently Paracylindrocarpon multiseptatum, CBS 337.77; Fig. 2), possibly due to the inaccurate identification. The third culture CBS 335.77 has identical sequences with the ex-type culture of Pn. aloicola (CBS 141300; Fig. 2). Unfortunately, culture CBS 335.77 proved to be sterile, and we were unable to compare its morphology with that of the type description. Hydropisphaera fungicola Rossman et al., Fungal Planet 24: 2. 2008. Description and illustration: Rossman et al. (2008). Typus: USA, Idaho, Lapwai Canyon, in the riparian community of Lapwai Creek, on Ulocladium atrum (Pleosporaceae) associated with Melampsora rust on decaying leaves of Populus trichocarpa (Salicaceae), unknown collection date, G. Newcombe (holotype BPI 878275, ex-type culture AR 4170 = CBS 122304). Hydropisphaera peziza (Tode) Dumort., Comment. Bot. (Tournay): 90. 1822. Basionym: Sphaeria peziza Tode, Fung. Mecklenb. Sel. (Lüneburg) 2: 46. 1791. Synonyms: Nectria peziza (Tode) Fr., Summa veg. Scand., Sectio Post. (Stockholm) 2: 388. 1849. Dialonectria peziza (Tode) Cooke, Grevillea 12: 110. 1884. Cucurbitaria peziza (Tode) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 461. 1898. Neuronectria peziza (Tode) Munk, Dansk Bot. Ark. 17: 58. 1957 [nom. inval., Art. 41.5 (Melbourne)]. Illustrations: Booth (1959, fig. 32, as Nectria peziza); Rossman et al. (1999). Typus: Sweden, on rotten wood, Sclerom. Suec. 24 no. 235, 1882 [(BPI, in Sbarbaro collections in bound volumes 1–3), lectotype designated by Rossman et al. (1999)]. Materials examined: Austria, from Juglans sp. (Juglandaceae), unknown collection date, W. Jaklitsch, culture CLL 13025 = CBS 135908. France, Ile de Ré, Saint-Martin, from dead wood, 11 Oct. 2014, isol. 14 Oct. 2014, coll. and isol. C. Lechat, culture CLL 14063 = CBS 139487. UK, Yorkshire, Mulgrave Woods, from Polyporus squamosus (Polyporaceae), unknown collection date and collector, isol. 1953 by C. Booth, CBS H-15060, culture CBS 296.65 = IMI 053559. Hydropisphaera suffulta (Berk. & M.A. Curtis) Rossman & Samuels, Stud. Mycol. 42: 32. 1999. Basionym: Nectria suffulta Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10(no. 46): 378. 1868 (1869). Synonyms: Lasionectria suffulta (Berk. & M.A. Curtis) Cooke, Grevillea 12(64): 112. 1884. Cucurbitaria suffulta (Berk. & M.A. Curtis) Kuntze, Revis. Gen. Pl. (Leipzig) 3(3): 461. 1898. Neohenningsia suffulta (Berk. & M.A. Curtis) Höhn. ex Petch, Trans. Brit. Mycol. Soc. 21: 268. 1938. Description and illustration: Samuels et al. (1976a).
78 Hou et al. Typus: Cuba, on palm leaves, unknown date, C. Wright (ex herb. M.J. Berkeley) (holotype of Nectria suffulta Berk. & M.A. Curtis 1868, K(M) 36225). Material examined: Indonesia, Sulavesi, Eastern Dumoga-Bone Nat. Park, Komangaan, Limestone Caves, alt. 400 m, on inflorescence of Cocos nucifera (Arecaceae), 24 Oct. 1985, G.J. Samuels, G.J.S. 85-194, representative culture CBS 122.87 = G.J.S. 2363. Clade O14 Paragliomastix L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845826. Etymology: Morphologically resembling the genus Gliomastix, but phylogenetically distinct. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae. Conidiophores solitary or aggregated, erect, arising directly from vegetative hyphae or ropes formed by the mycelium, usually reduced to single phialides, unbranched or poorly branched, hyaline or pale brown, smooth-, or roughenand finely spinulosewalled, septate at base, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral, terminal, subulate, hyaline at first, becoming pale brown with age, thick-, rough-walled, with conspicuous or inconspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides with up to two conidiogenous loci occasionally present. Conidia aseptate, ovoid, fusiform, with symmetrically apiculate base, having prominent, darker, truncate hilum at one end or both ends, hyaline at beginning, becoming olivaceous green, dark brown with age, or with darker equatorial belt, thick-walled, smooth, spinulose or wrinkled, arranged in dry, long chains. Chlamydospores and sexual morph not observed. Type: Paragliomastix luzulae (Fuckel) L.W. Hou, L. Cai & Crous Other accepted species with available sequences: Paragliomastix chiangraiensis (J.F. Li et al.) L.W. Hou, L. Cai & Crous, Px. rosea L.W. Hou, L. Cai & Crous, Px. znieffensis (Lechat & J. Fourn.) L.W. Hou, L. Cai & Crous Notes: Paragliomastix has a close morphological similarity with Gliomastix by having darkly pigmented ameroconidia. However, Paragliomastix produces spinulose, rough-walled phialides and conidia arranged in long, dry chains, while most Gliomastix species produce smooth-walled phialides and conidia arranged in slimy heads or chains. The phylogenetic analysis of the combined ITS, LSU, rpb2 and tef-1α dataset revealed that species of Paragliomastix clustered distant from Gliomastix in Bionectriaceae (Fig. 2). Paragliomastix chiangraiensis (J.F. Li et al.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845827. Basionym: Acremonium chiangraiense J.F. Li et al., Fungal Diversity 100: 197. 2020. Description and illustration: Hyde et al. (2020a). Typus: Thailand, Chiang Rai Province, Khun Korn Waterfall, on dead moistened leaf of palm, 9 Jan. 2014, J.F. Li, H-10b (holotype MFLU 140202, isotype KUN-HKAS, ex-type culture MFLUCC 14-0397). Material examined: Colombia, Cundinamarca, near Fómeque, from dead leaf of Zea mays (Poaceae), 5 Dec. 1979, W. Gams, Col 161c, culture CBS 277.80B. Notes: Acremonium chiangraiense was described by Hyde et al. (2020a), which was collected from Khun Korn Waterfall in Thailand and was isolated from a dead, moist palm leaf. This fungus is characterised by producing ampulliform conidiophores with phialidic conidiogenous cells and bearing hyaline to greenyellow conidia in chains (Hyde et al. 2020a), which morphologically correlated with the generic characteristics of Paragliomastix. Phylogenetically, it falls in a well-supported lineage closed to the type of Paragliomastix, Px. luzulae (Fig. 2). Paragliomastix luzulae (Fuckel) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845828. Fig. 20. Basionym: Torula luzulae Fuckel, Fungi Rhenani Exsiccati, Supplementi Fasc. 2: no. 1624. 1866. Synonyms: Gliomastix luzulae (Fuckel) E.W. Mason, Nat. Hist. Scarborough Distr. 1: 154. 1953. [nom. inval., Art. 41.5 (Melbourne)]. Gliomastix luzulae (Fuckel) E.W. Mason ex S. Hughes, Canad. J. Bot. 36: 769. 1958. Acremonium luzulae (Fuckel) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 92. 1971. Sagrahamala luzulae (Fuckel) Subram., Curr. Sci. 41: 48. 1972. Fusidium viride Grove, J. Bot., Lond. 23: 164. 1885. Description based on culture CBS 494.67: Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.2–2.9 μm wide. Conidiophores solitary or aggregated, erect, straight, arising directly from vegetative hyphae or from a granule formed by mycelium, usually reduced to single phialides, unbranched or poorly branched, up to 44 μm long, 1.8–3.5 μm wide at base, hyaline at beginning, turning pale brown with age, roughened, septate at base, with warts or spinulate at the upper part, rarely pigmented at the tip, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral, terminal, subulate, hyaline at first, becoming pale brown with age, thick-, roughwalled, 22–35 μm long, 2–3.5 μm wide at base, with inconspicuous collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, ovoid or fusoid, with symmetrically truncate base at both ends, having prominent, darker, apiculate hilum at one end or both ends, hyaline at beginning, becoming olivaceous green with age, mostly with a distinct darker equatorial belt, sometimes with distinct darker zones at one side of conidia, and darker zones enlarged, irregularly encrusted with pigment, thick-, rough-walled, 4.9–7.5 × 1.9–2.8 μm, arranged in chains. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 12–14 mm diam, raised, felty, granular at centre, olivaceous black at centre, white at periphery, margin crenate, brown pigment, reverse pale olivaceous; On MEA reaching 16 mm diam, raised, felty, granular at centre, olivaceous with abundant liquid exudate at centre, olivaceous black at periphery, margin crenate, reverse olivaceous, with buff edge; On PDA reaching 21–22 mm diam, flat, felty, moist with abundant liquid exudate, olivaceous black with buff edge, with fawn pigment at agar, margin lobate, reverse concolourous with fawn pigment; On SNA reaching 16–17 mm diam, flat, dusty, greenish olivaceous, white at periphery, margin lobate, reverse concolourous. Lacking odour on OA, MEA, SNA media; with strong geosmin odour on PDA. Materials examined: Germany, Kr. Rendsburg, Gut Schierensee, from decaying wood of Picea sp. (Pinaceae), together with many other fungi, unknown collection date, W. Gams, CBS H-8241 & CBS H-24595, culture CBS 494.67 = IAM 14654; Spessart, Lochmühle near Bieber, from decaying wood of Fagus sylvatica (Fagaceae), Sep. 1969, W. Gams, No. 1614, CBS
79www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales H-8243, culture CBS 935.69. Russia, Novgorod, Saint Sophia Cathedral, work of art, unknown collection date and collector, isol. D. Kudritsina, dep. L.A. Belyakova, CBS H-8030, culture CBS 495.67 = ATCC 18665 = IMI 133983 = VKM F-1168. Notes: This species was originally described as Torula luzulae from leaves of Luzula maxima (Fuckel 1866), and later transferred to Gliomastix (Hughes 1958). However, Gams (1971) demonstrated that it belonged to Acremonium in the Luzulae-series because of its pigmented conidia. According to our multi-locus phylogenetic analyses, this species is not congeneric with Gliomastix s. str. nor Acremonium s. str., but clustered with a separate clade representing the novel genus Paragliomastix in Bionectriaceae (Fig. 2). Morphologically, Px. luzulae is characterised by its conidial shape, having truncate ends, and with a dark belt in the middle of conidia, based on which it is differentiated from other species in Paragliomastix. Paragliomastix rosea L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845829. Fig. 21. Etymology: Epithet derived from the colony colour on PDA. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.5–2.8 μm wide. Conidiophores solitary or aggregated, erect, straight, arising directly from vegetative hyphae or ropes formed by the mycelium, usually reduced to single phialides, unbranched or poorly branched, up to 34.5 μm long, 2–3.4 μm wide at base, with 1–2 septa at base and middle part, initially hyaline, becoming pale brown with age, roughened, warty or spinulate at the upper part, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral, terminal, subulate, hyaline at first, becoming pale brown with age, thick-, rough-walled, 19–31.5 μm long, 2–3.5 μm wide at base, with cylindrical collarette and inconspicuous periclinal thickening at conidiogenous loci; Fig. 20. Paragliomastix luzulae (culture CBS 494.67). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial chains on pine needle. E–H. Conidiophores. I, J. Conidia. Scale bars = 10 μm.
86 Hou et al. 602, CBS H-8115, culture CBS 208.70; Schleswig-Holstein, Kiel-Kitzeberg, from wheat field soil, unknown collection date and collector, isol. 1964 by W. Gams, No. C 985, culture CBS 393.66 = MUCL 9408. Norway, Troms County, Nordkapp municipality, Skarsvåg, Mefjorden, from deciduous driftwood in splash zone, 17 Aug. 2010, T. Rämä, culture TR 055cII1.1, ibid. culture TR 055cI1.1. Netherlands, from agricultural soil, unknown collection date and collector, isol. 10 Jan. 1969 by J.H. van Emden, No. 369, culture CBS 216.69; Gelderland Province, Eibergen, from soil, Mar. 2017, T. Hilhorst (holotype of Lasionectria hilhorstii CBS H-23747, extype culture CBS 144938 = JW85024). UK, England, Cheshire, Delamere Forest, from leaf litter of Pinus sylvestris (Pinaceae), unknown collection date and collector, isol. Dec. 1956 by W.B. Kendrick, CBS H-8114, culture CBS 206.65; Cheshire, Delamere Forest, from needle of Pinus sylvestris (Pinaceae), unknown collection date and collector, isol. Dec. 1956 by W.B. Kendrick, culture CBS 227.66 = MUCL 9407; Northumberland, Bamburgh, from sand dune soil, unknown collection date and collector, isol. Mar. 1955 by J.C. Frankland, No. D 1889 (ex-type culture of Gliomastix guttuliformis CBS 207.65 = IAM 14644 = IMI 061279 = MUCL 9443); Ireland, Co. Westmeath, Tyrellspass, from esker soil, unknown collection date and collector, isol. Feb. 1966 by E.M. Rix, No. G 23, culture CBS 215.69 = IMI 137295. Notes: Lasionectria cerealis was originally described as the basionym Coniosporium cerealis, collected from leaves of Secalis cerealis in Finland (Karsten 1887). Later this species was transferred to the genus Gliomastix as G. cerealis (Dickinson 1968). This species is morphologically characterised by producing drop-shaped conidia, a green-black colony on malt agar and phialides occasionally with flared collarettes (Dickinson 1968). Although not all cultures used in this study match the original host and location of L. cerealis, their morphological characters agree well with the description of L. cerealis from literature (Gams 1971). Phylogenetically, all cultures formed a fully supported clade nestled in the same clade with other known Lasionectria species (Fig. 2), and the strain CBS 208.70 is considered a good representative of the species based on its morphological characteristics. Lasionectria hilhorstii was described from soil in the Netherlands, morphologically resembling Lasionectria cerealis (Gams 1971), but differed in the production of the phialides that lack flared collarettes and basal swelling (Crous et al. 2018a). In the present study, the ex-type strain of L. hilhorstii (CBS 144938) together with CBS 393.66 clustered slightly apart from other L. cerealis cultures based on 4 bp changes over the available sequences of three genes. Therefore, we suggest that the lack of flared collarette in L. hilhorstii should be considered strain variation among cultures of L. cerealis. The presence of a collarette may not be informative at the species levels. This idea also corresponded well with the comments of Dickinson et al. (1968), stating that “The collarette may be seen only with difficulty in some collections”. Lasionectria mantuana (Sacc.) Cooke, Grevillea 12 (no. 64): 112. 1884. Basionym: Nectria mantuana Sacc., Michelia 1 (no. 1): 52. 1877. Descriptions & Illustrations: Weese (1916), Rossman et al. (1999). Typus: Italy, Mantova, Migliaretto, on decorticated poplar wood, Feb. 1873, A. Magnaguti-Rondinini (holotype PAD S00014). Materials examined: Finland, Lempäälä, Innilä, from decorticated wood, possibly Populus sp. (Salicaceae), 27 Sep. 2003, U. Söderholm, BPI 843540, culture CBS 114291 = AR4029. Note: This species is the type of Lasionectria. Lasionectria olida (W. Gams) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845836. Fig. 25. Basionym: Acremonium olidum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 108. 1971. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, up to 2.6 μm wide. Conidiophores solitary, (sub-) erect, arising from submerged or superficial hyphae, unbranched or basitonously branched, bearing 1–2 levels with 1–2 phialides per node, occasionally with short sterile outgrowths, up to ca. 92 μm long, 2–4.5 μm wide at base, with 1–3 septa at base or upper part, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, cylindrical or subulate, hyaline, thick-, smooth-walled, 17.5–50(–63) μm long, 1.5–3 μm wide at base, with inconspicuous collarette and periclinal thickening at conidiogenous loci. Conidia aseptate, ellipsoidal with rounded ends, or with a weak apiculate base, hyaline, thin-, or thick-, smooth-walled, 3.5–5.7 × 2–2.75 μm, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 39 mm diam, flat, membranous with sparse aerial mycelium, dirty white, margin entire, reverse concolourous; On MEA reaching 27–30 mm diam, flat, spirally rugose, membranous, short hairy, orange at centre, buff to dirty white at periphery, margin entire, reverse orange; On PDA reaching 30 mm diam, flat, floccose, with greyish rose, salmon, rosy buff concentric rings, margin entire, with brown pigment, reverse with rust, apricot and saffron rings. Strong, pleasantly aromatic smell. Typus: Poland, Augustow, from Phellinus sp. (Hymenochaetaceae), together with Ceratocystis sp. (Ceratocystidaceae) and Hypocrea pulvinata (Hypocreaceae), unknown collection date and collector, isol. Sep. 1966 by W. Gams, No. 633, CBS H-6660 (holotype CBS 799.69 preserved as metabolically inactive culture, ex-type culture CBS 799.69). Additional material examined: Austria, Tirol, Kranebitter Klamm, from Fomitopsis pinicola (Fomitopsidaceae) on Picea abies (Pinaceae), unknown collection date and collector, isol. Dec. 1965 by W. Gams, No. 590, CBS H-8268, culture CBS 798.69. Notes: Based on the description provided by Gams (1971), the fungus formerly known as Acremonium olidum clustered in the genus Lasionectria (BPP/MLBS = 1/100 %), supported by phylogenetic inference in this study (Fig. 2). Therefore, a new combination is provided in the genus Lasionectria. Lasionectria sylvana (Mouton) Rossman & Samuels, Stud. Mycol. 42: 37. 1999. Fig. 26. Basionym: Nectria sylvana Mouton, Bull. Soc. Roy. Bot. Belgique. 39: 49. 1900. Synonym: Calonectria fimbriata Seaver & Waterston, Mycologia 32: 404. 1940. Illustrations: Samuels (1976a, fig. 14; 1976b, fig. 18, both as N. sylvana); Seaver & Waterston (1940, fig. 3 lower as C. fimbriata); Rossman et al. (1999). Description based on culture CBS 566.76: Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, up to 2.5 μm wide. Conidiophores solitary, (sub-)erect, arising from submerged or superficial hyphae, unbranched, rarely branched, occasionally with a basally sterile short outgrowth, up to ca. 55 μm long, 2.5–2.5 μm wide at base, with 1–2 septa
87www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales at base, hyaline, smooth-walled, rough-walled at basal part, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, cylindrical or aculeate, hyaline, thick-, smooth-walled, 16.5–40.7 μm long, 1.8–3.3 μm wide at base, with inconspicuous collarette and periclinal thickening at conidiogenous loci. Conidia aseptate, cylindrical, without visible basal abscission scar, hyaline, thin-, smooth-walled, (2.5–)4– 8.5 × 1.7–2.6 μm, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 36–37 mm diam, flat, membranous with sparse aerial mycelium, dusty at periphery, dirty white at centre, white at periphery, margin entire, reverse concolourous; On MEA reaching 30–35 mm diam, flat, radially folded, felty, saffron at centre, buff to creamy white at periphery, margin entire, reverse pale orange at centre, pale luteous at periphery, with buff radial lines; On PDA reaching 22–24 mm diam, flat, membranous with sparse aerial mycelium, buff, margin entire, reverse buff, with creamy white radial lines. Typus: Belgium, near Liege, on stems of Angelica sylvestris (Apiaceae), unknown collection date and collector (holotype of N. sylvana NY01013209). Material examined: New Zealand, Westland Province, Westland County, vic. Lake Ianthe, Lake Ianthe State Forest, SF 42, from rachis of Cyathea smithii (Cyatheaceae), unknown collection date, G.J. Samuels, G.J.S. 7475, CBS H-15071, culture CBS 566.76. Clade O18 Verruciconidia L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845837. Fig. 25. Lasionectria olida (ex-type culture CBS 799.69). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Slimy conidial heads. E–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
88 Hou et al. Etymology: Referring to the verrucose conidia produced by species in this genus. Mycelium consisting of branched, septate, hyaline or pale brown, rough-, thin-walled hyphae, forming bundles or coils. Conidiophores (sub-)erect, straight or curved, arising from vegetative hyphae or from ropes formed by the mycelium, unbranched, poorly branched, septate at base or middle part, hyaline, smoothor rough-walled, with cell walls usually thicker than those of vegetative hyphae. Conidiogenous cell monophialidic, terminal or lateral, cylindrical or subulate, hyaline, thick-, smoothwalled, with conspicuous or inconspicuous collarette and periclinal thickening at conidiogenous loci; adelophialides occasionally present in some species, differentiating intercalary as short lateral protrusions usually from submerged hyphae. Conidia aseptate, ovoid, (broad) ellipsoid, subglobose, rounded at both ends, or with apiculate end(s), straight, hyaline, thickand rough-walled, verrucose, or thinand smoothwalled, eguttulate or guttulate, arranged in conidial dry heads or long chains. Chlamydospores and sexual morph not observed. Type: Verruciconidia verruculosa (Nicot) L.W. Hou, L. Cai & Crous Other accepted species with available sequences: Verruciconidia erythroxyli L.W. Hou, L. Cai & Crous, Ve. infuscata L.W. Hou, L. Cai & Crous, Ve. persicina (Nicot) L.W. Hou, L. Cai & Crous, Ve. quercina L.W. Hou, L. Cai & Crous, Ve. siccicapita L.W. Hou, L. Cai & Crous, Ve. unguis L.W. Hou, L. Cai & Crous Notes: The genus Verruciconidia is proposed here for a group of cultures clustering in a fully supported clade in Bionectriaceae. Most of these cultures were previously identified as A. persicinum and A. verruculosum based on morphological characters. However, they are phylogenetically heterogeneous and distributed in different clades within the large family clade (Fig. 2). Thirteen “A. persicinum” strains, including the ex-type CBS 310.59 form Fig. 26. Lasionectria sylvana (culture CBS 566.76). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, F–J. Conidiophores. E. Conidial heads. K. Conidia. Scale bars =10 μm.
89www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales a well-supported lineage, representing A. persicinum s. str. The other strains are located in other clades representing five different species (Fig. 2). Species in Verruciconidia are different from other genera in Bionectriaceae by producing warty conidia, except for Ve. erythroxyli, Ve. persicina and Ve. unguis, which produce smoothwalled conidia. Verruciconidia erythroxyli L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845838. Fig. 27. Etymology: Referring to the host, Erythroxylum areolatum, from which the holotype was collected. Mycelium consisting of branched, septate, hyaline, rough-, thinwalled hyphae, up to 3 μm wide, forming bundles. Conidiophores solitary or aggregate, (sub-)erect, arising from submerged and superficial hyphae, or from ropes formed by mycelium, unbranched, poorly branched, bearing 1–3 phialides per node, up to ca. 45 μm long, 1.8–3.3 μm wide at base, with 1–3 septa, hyaline, smoothwalled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, cylindrical or subulate, hyaline, thick-, smooth-walled, (15–)19.5–31.5(–37) μm long, 2–3 μm wide at base, with inconspicuous collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, broad ellipsoid, occasionally with a slightly apiculate bases and rounded apices, hyaline, thin-, smooth-walled, 3.5–4.5 × 2–3 μm, with minute guttules, arranged in dry chains, soon collapsing, forming conidial heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 36–39 mm diam, flat, felty or dusty, white, margin Fig. 27. Verruciconidia erythroxyli (ex-type culture CBS 728.87). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial chains. E–H. Unbranched conidiophores. I, J. Branched conidiophores. K. Conidia. Scale bars = 10 μm.
90 Hou et al. fimbriate, reverse concolourous; On MEA reaching 28–30 mm diam, flat, radially folded, shortly hairy, white, margin crenate, reverse pale orange, with buff radial lines; On PDA reaching 35–38 mm diam, flat, felty, hairy at centre, white, margin filiform, reverse buff, brown at centre. Typus: Cuba, Matanzas Province, San Miguel de los Baños, from leaf of Erythroxylum areolatum (Erythroxylaceae), unknown collection date and collector, isol. 22 Jan. 1987 by R.F. Castañeda, C87/10 (holotype CBS H-24612, ex-type culture CBS 728.87). Additional material examined: Zaire, Bas-Zaire, Nsangi, from leaf of Urena lobata (Malvaceae), isol. 1964, isol. G.L. Hennebert, CBS H-24689, culture CBS 378.70D = MUCL 6269. Notes: Verruciconidia erythroxyli is represented by two strains from different host plants, Erythroxylum areolatum (Erythroxylaceae) and Urena lobata (Malvaceae). This species is phylogenetically close to another new species, Ve. unguis described in this study (Fig. 2), and their morphological differences are discussed under the latter species. Although both species differ from other Verruciconidia species in their smooth-walled conidia, they are phylogenetically congeneric within Verruciconidia. Verruciconidia infuscata L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845839. Fig. 28. Etymology: Derived from the brown hyphae produced by this species. Mycelium consisting of branched, septate, hyaline, rough-, thinwalled hyphae, brown and thick-walled in older culture, up to 2.5 μm wide, forming bundles and coils. Conidiophores solitary or aggregate, (sub-)erect, arising from submerged and superficial Fig. 28. Verruciconidia infuscata (ex-type culture CBS 100888). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores with conidia heads arising from mycelial ropes. E–H. Conidiophores. I. Brown hyphae. J, K. Conidia. Scale bars = 10 μm.
91www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales hyphae, or from ropes formed by the mycelium, unbranched or rarely branched, up to ca. 64.5 μm long, 1.6–2.8 μm wide at base, with 1–2 septa in basal part, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides monophialidic, mostly lateral, rarely terminal, subcylindrical or subulate, hyaline, thick-, smooth-walled, 38–50.5 μm long, 1.8–2.5 μm wide at base, with conspicuous cylindrical collarette and periclinal thickening at conidiogenous loci. Conidia aseptate, ellipsoid, subglobose, rounded at both ends, or with an inconspicuous truncate bases, hyaline, thick-, rough-walled, warty, 3–5 × 2–2.8 μm, with 1–2 large guttules, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 47–48 mm diam, flat, sparse aerial mycelium, membranous, pale olivaceous buff at centre, with dirty white margin, margin entire, pale brown pigment present in old cultures, reverse olivaceous buff, buff at centre; On MEA reaching 43 mm diam, flat, radially folded, with moderate aerial mycelium, felty, creamy white, margin lobulated, pale brown pigment present in old cultures, reverse pale orange, with buff radial lines; On PDA reaching 57–58 mm diam, flat, sparsely floccose, dirty white, few shallow striations over the colony surface, margin filiform, pale brown pigment present in old cultures, reverse creamy white; On SNA reaching 50 mm diam, flat, membranous without aerial mycelium, colourless, margin entire, reverse colourless. Typus: Japan, Kanagawa Pref., from air, unknown collection date and collector, isol. M. Sakamoto, No. 94 (holotype CBS H-24613, ex-type culture CBS 100888). Notes: Verruciconidia infuscata is placed on a single branch, nestling in the same clade with Ve. erythroxyli, Ve. siccicapita and Ve. unguis (Fig. 2). The species can be morphologically distinguished from Ve. erythroxyli and Ve. unguis by producing warty conidia arranged in moist slimy heads, while the last two species produce smooth-walled conidia that are arranged in chains but later collapse into conidial heads; Ve. infuscata differs from Ve. siccicapita by producing longer conidiophores (up to 64.5 μm vs up to 41 μm) and phialides (38–50.5 μm vs 25–36 μm). In addition, conidia of Ve. infuscata are arranged in moist slimy heads while Ve. siccicapita produces conidia arranged in dry conidial heads. Verruciconidia persicina (Nicot) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845840. Fig. 29. Basionym: Paecilomyces persicinus Nicot, Bull. Soc. Mycol. France 74: 222. 1958. Synonym: Acremonium persicinum (Nicot) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 75. 1971. Mycelium consisting of branched, septate, hyaline, rough-, thinwalled hyphae, up to 3.5 μm wide, forming bundles. Sporulation abundant, phalacrogenous, nematogenous, plectonemtogenous. Conidiophores solitary or aggregate, (sub-)erect, straight to flexuous, irregularly bent at lower part, arising from submerged and superficial hyphae, or ropes formed by mycelium, unbranched or repeatedly branched, bearing 1–3 phialides per node, commonly with short sterile outgrowths, up to ca. 61(–118) μm long, 1.8–3 μm wide at base, with 1–3 septa in basal, middle and apical part, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, cylindrical or subulate, irregularly bent, hyaline, thick-, smooth-walled, 12–36 μm long, 1.8–2.7 μm wide at base, with minute collarette and inconspicuous periclinal thickening at conidiogenous loci, polyphialides not observed. Conidia aseptate, ellipsoid, ovoid, occasionally with slightly apiculate basal end, hyaline, thin-, smooth-walled, 4.2–6.2 × 2.4–3 μm, eguttulate or with one large guttule, arranged in dry chains, soon collapsing. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 40–42 cm diam, flat, with sparse aerial mycelium, dusty, white, pale brown with age, margin entire, reverse concolourous; On MEA reaching 33–35 diam, flat, radially fold, with moderate aerial mycelium, floccose or dusty, salmon at centre, rosy buff at periphery, margin crenate, reverse saffron, with buff radial lines; On PDA reaching 40–42 cm diam, flat, with moderate aerial mycelium, felty, granular or dusty, dirty white or rosy buff, margin entire, reverse creamy white; On SNA reaching 42–45 cm diam, flat, with sparse aerial mycelium, dusty, colourless, margin entire, reverse colourless. Without odour on all media. Description and illustration: Gams (1971). Typus: France, Arcachon, from coastal sand under Ammophila arenaria (Poaceae), unknown collection date, J. Nicot, CBS H-6661 (ex-isotype culture of Paecilomyces persicinus CBS 310.59 = ATCC 18551 = IAM 14656 = IMI 069374 = LCP 56.1537). Additional materials examined: Iran, Golestan Province, Gorgan, endophyte in Festuca ovina (Poaceae), unknown collection date and collector, isol G. Dehghampour, culture CBS 116385; Āz arbāyjān-e Sharqī Province, Tabriz, from Pleurotus ostreatus (Pleurotaceae), 2002, M.R. Asef, culture CBS 113716. Italy, from leaf of Vitis vinifera (Vitaceae), inoculated with zoospores of Plasmopara viticola (Peronosporaceae), unknown collection date, S. Burruano, Inst. Patol. Veget., Palermo, culture CBS 101712; from soil, unknown collection date and collector, isol. M.A. Pisano, No. 10C, culture CBS 169.65. Papua New Guinea, Bougainville Island, from tent canvas, unknown collection date and collector, isol. 1944, CBS H-8280, culture CBS 295.70A = QM 1b; Madang, Jais Aben, from soil along coral reef coast, Nov. 1995, A. Aptroot, isol. Nov. 1995 by A. van Iperen, No. A 152, culture CBS 218.96. USA, Hawaii, (Milepost 43) Hgw. 200 Pu’u La’au, subalpine dry forest, from a white mycelium growth of a wood decay fungus on the undersurface of a dead hardwood branch, 4 Nov. 2002, D.T. Wicklow, isol. 30 May 2002, MYC 1794, culture CBS 120889; Iowa, Steele Prairie, Cherokee County, T93N R40W S16, from soil, 1982, unknown collector, culture CBS 128826 = RMF 7540; Kansas, Konza Prairie Research Natural Area, Long Term Ecological Research site (LTER), near Manhattan, from soil, 1986, unknown collector, culture CBS 127298 = RMF 8134; Minnesota, Steele, from cyst of Heterodera glycines (nematode, Heteroderidae), unknown date, Fajun Chen, S2-81, culture CBS 102349; unknown location, from canvas legging, unknown collection date and collector, isol. 1959, CBS H-8281, culture CBS 295.70B = QM 89c. USSR, from pasture soil, unknown collection date and collector, isol. Institute of Microbiology, Moskva, CBS H-8275 & CBS H-8276, culture CBS 439.66 = VKM F-888. Notes: Based on the multi-locus phylogenetic analyses, 13 cultures labelled as Acremonium persicinum, including its ex-isotype strain (CBS 310.59), form a fully supported clade within the clade representing the new genus Verruciconidia (Fig. 2). Acremonium persicinum is therefore transferred to Verruciconidia, and a new combination is proposed. Our morphological description of the extype culture slightly differs from the one in Gams (1971) by having repeatedly branched conidiophores and lacking elongated crystals. Some cultures are described as having ochreous brown to reddish brown mycelium in Gams (1971), while we only observed a white or rosy buff colour on OA and PDA media and salmon colour on MEA plates. These differences may be due to intraspecies variation or are related to culture conditions.
92 Hou et al. Verruciconidia quercina L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845841. Fig. 30. Etymology: Referring to Quercus, the host genus from which the ex-type culture of this fungus was isolated. Mycelium consisting of branched, septate, hyaline, rough-, thinwalled hyphae, up to 2.7 μm wide, forming bundles. Conidiophores solitary or aggregate, (sub-)erect, straight, arising from submerged and superficial hyphae, or from ropes formed by the mycelium, unbranched or poorly branched, up to ca. 49 μm long, 1.9–3.7 μm wide at base, with 1–3-septa in basal part, hyaline, smoothwalled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, cylindrical or subulate, hyaline, thick-, smooth-walled, 12.8–43.5 μm long, 1.3–2.7 μm wide at base, with inconspicuous collarette and periclinal thickening at conidiogenous loci; polyphialides not observed; adelophialides present, differentiating intercalary as short lateral protrusions usually from submerged hyphae, more or less cylindrical, up to 16 μm diam. Conidia aseptate, ellipsoid, without obviously apiculate base, hyaline, thick-, rough-walled, verrucose, 2.8–5.2 × 2.5–3.5 μm, eguttulate, arranged in dry heads, conidial heads becoming pale brown in older culture. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 40–42 mm diam, flat, dusty, dirty white at centre, creamy white at periphery, margin entire, reverse saffron at centre, buff at periphery; On MEA reaching 29–30 mm diam, flat, radially folded, short hairy, white, margin filiform, reverse saffron with buff radial lines; On PDA reaching 43–45 mm diam, flat, thinly felty, dirty white, with pale greenish zones, margin entire, reverse buff. Fig. 29. Verruciconidia persicina (ex-type culture CBS 310.59). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial chains. E–I. Conidiophores. J. Conidia. Scale bars = 10 μm.
93www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Typus: Unknown, from leaf litter of Quercus sp. (Fagaceae), unknown collection date and collector, isol. 1965 by T. Hering, No. 642 (holotype CBS H-8278, culture CBS 469.67). Additional materials examined: Netherlands, North Brabant Province, Haren, from agricultural soil, unknown collection date and collector, isol. G. Jager, No. M41, culture CBS 355.77; South Holland Province, Katwijk, Lysimeter 1, from sand dune soil, 95 cm depth in lysimeter at upper limit of art, Jan. 1978, W. Gams, No. 1/95 19, CBS H-1038, culture CBS 183.78. Notes: The three cultures of Verruciconidia quercina form a distinct basal clade of Verruciconidia on the multi-locus tree (Fig. 2). Morphologically, it differs from all other species in Verruciconidia in producing short lateral adelophialides. This is the first report of acremonium-like species with verrucose conidia from Quercus spp . Verruciconidia siccicapita L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845842. Fig. 31. Etymology: Latin, siccus = dry, caput = head, referring to the dry conidial heads produced by this fungus. Mycelium consisting of branched, septate, hyaline, rough-, thin-walled hyphae, up to 1.7 μm wide, forming thick bundles. Sporulation abundant, phalacrogenous, nematogenous, plectonematogenous. Conidiophores solitary or aggregate, (sub-) erect, straight to flexuous, curved at base part, arising from submerged and superficial hyphae, or from ropes formed by the mycelium, unbranched, up to 41 μm long, 1.3–3.2 μm wide at base, with single septum at base, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides most lateral, rarely terminal, cylindrical or subulate, hyaline at Fig. 30. Verruciconidia quercina (ex-type culture CBS 469.67). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores with conidial heads. F–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
94 Hou et al. first, becoming pale brown with age, thick-, smooth-walled, 25– 36 μm long, 1.2–2.6 μm wide at base, with flared collarette and inconspicuous periclinal thickening at conidiogenous loci. Conidia aseptate, ellipsoid or subglobose, rounded at both ends, hyaline, thickand finely verruculose, 3.5–5 × 2.3–3 μm, guttulate, arranged in dry conidial heads, hyaline, becoming pale brown with age. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 45 cm diam, flat, with sparse aerial mycelium, dusty or membranous, dirty white, with some hairy mycelium, become fawn with age, margin entire, reverse pale fawn, later become fawn, without any odour; On MEA reaching 43 diam, flat, with moderate aerial medium, hairy, rosy buff, later become dusty or hairy and brick, margin entire, reverse pale apricot, with buff radial lines, with strong pungent odour; On PDA reaching 60 mm diam, flat, with sparse aerial mycelium, dusty, dirty white, few shallow striations over the colony surface, buff at periphery, margin fimbriate, reverse pale olivaceous buff, strong “rotten wood” odour; On SNA reaching 40–42 cm diam, flat, with sparse aerial mycelium, dusty, colourless, margin entire, reverse colourless, without any odour. Typus: Thailand, from soil, unknown collection date and collector, isol. J. Nicot, No. BSA3 (holotype CBS H-8298, ex-type culture CBS 378.70A = LCP 2149). Notes: Verruciconidia siccicapita is represented by a single culture isolated from soil in Thailand and differs from all other species according to the multi-locus tree (Fig. 2). It is phylogenetically different from the closely related species Ve. infuscata (98.2 % sequence similarity on ITS, 99.7 % on LSU, 91.8 % on rpb2, 98.2 % on tef-1α). Their morphological differences are discussed under Ve. infuscata. Fig. 31. Verruciconidia siccicapita (ex-type culture CBS 378.70A). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial heads. E. Conidial heads. F. Conidiophores arising from mycelial ropes. G, H. Conidiophores. I. Conidia. Scale bars = 10 μm.
95www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Verruciconidia unguis L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845843. Fig. 32. Etymology: Referring to the substrate from which the holotype culture was collected. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, up to 3.2 μm wide, forming bundles. Conidiophores (sub-)erect, arising from submerged and superficial hyphae, or from ropes of hyphae, unbranched, up to ca. 40.5 μm long, 1.5–3.5 μm wide at base, with 1–2 septa in basal part, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal or lateral, cylindrical or subulate, occasionally acuminate at top, hyaline, thick-, smooth-walled, (20.5–)22–34(– 39) μm long, 2–3 μm wide at base, with inconspicuous collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, obovoid, subglobose, ellipsoid, with an inconspicuous apiculate bases, and a rounded apices, hyaline, thin-, smooth-walled, with 1–2 large guttules, 3.2–4 × 2.3–2.9 μm, arranged in dry chains and soon collapse as conidial heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 43–45 mm diam, flat, dusty, granular, dirty white, margin fimbriate, reverse creamy white; On MEA reaching 43–45 mm diam, raised, radially folded, finely floccose, dirty white, margin crenate, reverse pale orange, with buff radial lines; On PDA reaching 45 mm diam, flat, felty or dusty, dirty white, pale fawn at centre, margin fimbriate, reverse orange at centre, buff at periphery; On SNA reaching 48 mm diam, flat, dusty, with concentric rings, dirty white, margin entire, reverse concolourous. Fig. 32. Verruciconidia unguis (ex-type culture CBS 424.93). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores and conidia chains. F–I. Conidiophores J. Conidia. Scale bars = 10 μm.
102 Hou et al. by lacking of crystal, while R. thailandicum produces abundant, globose or irregular shaped, brown crystal. Ramosiphorum thailandicum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845851. Fig. 37. Etymology: Named after the location where the fungus was collected, Thailand. Mycelium consisting of branched, septate, hyaline, rough-, thin-walled hyphae, up to 3 μm wide. Sporulation abundant, phalacrogenous, nematogenous, rarely plectonematogenous. Conidiophores (sub-) erect, straight or irregularly curved and flexuous, arising from submerged and superficial hyphae, mostly repeatedly verticillately, basitonously branched, bearing up to 5 whorls of 1–2(–3) phialides, aggregated as sporodochia-like structures, rarely unbranched, up to ca. 114 μm long, 1.5–3.1 μm wide at base, with 1–3 septa in basal, middle or upper part, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides monophialidic, lateral or terminal, subulate, hyaline, thick-, smooth-walled, 9–35 μm long, 1–2.3 μm wide at base, with conspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, subglobose to ellipsoid, hyaline, thick-, rough-walled, 2.5–4 × 1.8–2.3 μm, arranged in moist slimy heads, becoming salmon with age. Crystals abundant, globose or irregular shaped, brown. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: colonies on OA reaching 35–36 mm diam, flat, with moderate aerial mycelium, floccose or felty, with mycelial ropes and granule, white, margin crenate, reverse pale olivaceous buff, with strong geosmin odour; On MEA reaching 28–29 Fig. 36. Ramosiphorum polyporicola (ex-type culture CBS 123779). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial heads. E–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
103www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales mm diam, flat, radially folded, with moderate aerial mycelium, felty, white, margin filiform, reverse pale orange, with buff radial lines; On PDA reaching 39–40 mm diam, flat, with moderate aerial mycelium, felty or floccose, with mycelial ropes and granule, with concentric rings, dirty white, creamy white and pale fawn at centre, margin filiform, reverse olivaceous buff; On SNA reaching 25–26 mm diam, flat, with sparse aerial mycelium, dusty, with mycelial ropes and granule, white, margin entire, reverse concolourous. Without odour on all media. Typus: Thailand, from bark of dead tree, unknown collection date, G.J. Samuels, G.J.S. 97-71 (holotype CBS H-24598, isotype BPI 745614, extype culture CBS 101914). Note: See notes under Ramosiphorum echinoporiae and R. polyporicola. Clade O23 Protocreopsis Yoshim. Doi, Kew Bull. 31: 551. 1977. Ascomata superficial on substrata, densely gregarious, less commonly solitary, surrounded by white to tan hyphae arising from ascomatal wall with few to many free ends visible, thus appearing hypocrea-like, often extensive; ascomata hyaline to orange, KOH-; cells at ascomatal surface completely obscured by investing hyphae; ascomatal wall more than 20 μm thick, comprising a single region of small, brick-like cells. Asci clavate to fusoid, apex simple or with an obscure ring; ascospores bito pluriseriate. Ascospores ellipsoid to fusoid, 1-septate, hyaline, typically striate, also smooth, punctatestriate, or tuberculate. Asexual morph where known, acremoniumFig. 37. Ramosiphorum thailandicum (ex-type culture CBS 101914). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidial heads. E. Conidiophores and conidial heads. F–H. Branched conidiophores. I, J. Unbranched conidiophores. K. Conidia. Scale bars = 10 μm.
104 Hou et al. like. On dead monocotyledonous substrata (Doi 1977, Rossman et al. 1999, Lechat et al. 2016b). Type: Protocreopsis fusigera (Berk. & Broome) Yoshim. Doi Other accepted species with available sequences: Pt. caricicola Lechat & J. Fourn., Pt. euphorbiae Crous, Pt. finnmarkica L.W. Hou, L. Cai, Rämä & Crous, Pt. freycinetiae (Samuels) Samuels & Rossman, Pt. pertusa (Pat.) Samuels & Rossman, Pt. phormiicola (Samuels) Samuels & Rossman, Pt. rutila (W. Gams) L.W. Hou, L. Cai & Crous Notes: The genus Protocreopsis was introduced by Doi (1977) for a group of nectria-like species that have ascomata completely enclosed in long, white to tan or green, flexuous hyphal stroma, ascomatal walls that are more than 20 μm thick, typically striate ascospores and acremonium-like asexual morphs (Rossman et al. 1999, Lechat et al. 2016b). Although molecular sequences of the type Pt. fusigera are not available in GenBank, phylogenetical analysis of four Protocreopsis species including Pt. caricicola, Pt. freycinetiae, Pt. pertusa and Pt. phormiicola demonstrate that this genus belongs to Bionectriaceae. Protocreopsis caricicola Lechat & J. Fourn., Ascomycete.org 8: 30. 2016. Description and illustration: Lechat et al. (2016b). Typus: Germany, Westmecklenburg, 5 km NE of Rhena, meadow pond 1 km E-SE of Strohkirchen, from last year’s leaf residues of Carex acutiformis (Cyperaceae), 25 May 2015, T. Richter, isol. 26 May 2016 by C. Lechat (holotype CLL 15081 in LIP, ex-type culture CBS 140572). Protocreopsis finnmarkica L.W. Hou, L. Cai, Rämä & Crous, sp. nov. MycoBank MB 845852. Etymology: Name reflects the most north-eastern county of Norway, Finnmark, where the species was collected. Culture sterile. Protocreopsis finnmarkica differs from its closest phylogenetic neighbour Pt. euphorbiae (culture: CPC 38896) by unique fixed alleles in three loci based on alignments of the concatenated three loci deposited in Figshare (doi: 10.6084/ m9.figshare.22258765): ITS position shares similarity: 97(C), 152(C), 180(C), 183(G), 443(T), 444(G), 513(T), 528(T), 529(C), 535(gap), 540(C), 560(C), 618(A), 621(T); LSU position: 1 459 (C); tef-1α position: 2 376(G), 2 403(C), 2 415(A), 2 439(T), 2 504(C), 2 508 (C), 2 546 (G), 2 575 (C), 2 576 (T), 2 578 (T), 2 590 (C), 2 656 (T), 2 665 (C), 2 674 (T), 2 681 (C), 2 684 (T). Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 35 mm diam, flat, hairy at centre, felty at periphery, white, margin entire, reverse creamy white. On MEA reaching 23 mm diam, raised, felty, white, margin entire, reverse orange. On PDA reaching 27 mm diam., raised, hairy and buff at centre, flat, felty and white at periphery, margin entire, reverse pale luteous at centre, buff at periphery. On SNA reaching 28 mm diam, flat, aerial mycelium sparse, white, margin fimbriate, reverse concolourous. Typus: Norway, Troms og Finnmark county, Kvænangen municipality, Alteidet, from old dock made of Pinus sylvestris (Pinaceae) in the sea, 9 Aug. 2010, T. Rämä 041cS1.2 (holotype CBS H-24724, ex-type culture CBS 147428 = CPC 40316 = TR041cS1.2). Additional materials examined: Norway, Troms County, Kvænangen municipality, Alteidet, from old dock of P. sylvestris in the sea, 9 Aug. 2010, T. Rämä, culture CBS 147427 = CPC 40315 = TR041bE1.1; ibid., culture CBS 147429 = CPC 40318 = TR042bE1.2; ibid., culture TR042cE1.2, ibid., culture TR041aN1.1. Notes: This fungus was isolated from wood as explained in Rämä et al. (2014). The strains from an old dock of Pinus sylvestris in the Norwegian Sea were examined in this study. They cluster on a separate branch close to Pt. euphorbiae, from which they are phylogenetically distinct, which supports this species as unique (Fig. 2). However, all strains remained sterile in all culture media tested in this study. Considering that it is phylogenetically distinct, a new species, Pt. finnmarkica, is introduced here and the molecular differences based on the sequence data are provided. Protocreopsis freycinetiae (Samuels) Samuels & Rossman, Stud. Mycol. 42: 65. 1999. Basionym: Nectria freycinetiae Samuels, New Zealand J. Bot. 14: 243. 1976. Description and illustration: Samuels (1976b). Typus: New Zealand, Auckland, Thames County, Coromandel Forest Park, Kauaeranga Valley, vic. Thames, from leaf of Freycinetia banksia (Pandanaceae), 17 Aug. 1974, J.M. Dingley et al., isol. G.J. Samuels, CBS H-718 (holotype of Nectria freycinetiae PDD 32577, ex-isotype culture CBS 573.76 = ATCC 34044 = IMI 208153). Note: Protocreopsis freycinetiae clustered within the Protocreopsis clade but with low support values. Protocreopsis pertusa (Pat.) Samuels & Rossman, Stud. Mycol. 42: 66. 1999. Basionym: Nectria pertusa Pat., Bull. Soc. Mycol. France. 11: 227. 1895. Synonym: Cucurbitaria pertusa (Pat.) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 461. 1898. Description and illustration: Rossman et al. (1999). Material examined: New Zealand, Auckland, Waitakere Ranges, West Coast Road, Marguerite Track, from rachis of Cyathea medullaris (Cyatheaceae), 21 Aug. 1974, J.M. Dingley & Haydon, isol. G.J. Samuels, PDD 32584, culture CBS 568.76. Protocreopsis phormiicola (Samuels) Samuels & Rossman, Stud. Mycol. 42: 67. 1999. Basionym: Nectria phormiicola Samuels, New Zealand J. Bot. 14: 244. 1976. Description and illustration: Rossman et al. (1999). Typus: New Zealand, Auckland, Waitemata County, Waitakere Ranges, vic. Piha, Marowhara Loop Track, from dead leaf of Phormium tenax (Asphodelaceae), 17 Dec. 1974, J.M. Dingley et al., G.J.S. 74-133 (holotype of Nectria phormiicola PDD 32684, ex-type culture CBS 567.76 = ATCC 34049 = IFO 30281 = IMI 208158). Protocreopsis rutila (W. Gams) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845853. Fig. 38. Basionym: Acremonium rutilum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 105. 1971. Description based on culture CBS 394.70: Mycelium consisting of branched, septate, hyaline, pale brown in old cultures, smooth-, thin-walled hyphae, up to 2 μm wide, often formed mycelial coils. Sporulation abundant, phalacrogenous, nematogenous,
105www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales rarely plectonematogenous. Conidiophores (sub-)erect, straight to flexuous, arising from submerged and superficial hyphae, unbranched or poorly branched, up to ca. 68 μm long, 1.3–3.8 μm wide at base, with short sterile outgrowths, with 1–2(–3)-septa at basal or upper part, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae; degenerated to very short conidiophores with age. Phialides monophialidic, lateral, subulate, always narrowed at base, hyaline, thin-or thick-, smooth-walled, 10.5–60.5 μm long, 1.5–3 μm wide at base, with inconspicuous collarette and conspicuous periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, long cylindrical, with an inconspicuous truncate hilum at basal end, hyaline, thin-, smooth-walled, with two large inconspicuous guttules, 3.5–8.7 × 2.2–3 μm, arranged in moist slimy heads, mostly confluent, salmon. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: colonies on OA reaching 30–32 mm diam, flat, thinly felty, white at centre and periphery, pale salmon in middle, margin entire, reverse buff; On MEA reaching 25–27 mm diam, flat, radially folded, felty, dirty white, undulant margin, reverse ochreous, with radially buff lines; On PDA reaching 27–30 mm diam, flat, thinly felty, sienna at centre, peach at middle, salmon at periphery, margin entire, with yellowish pigment, reverse orange, with buff edge; On SNA reaching 32–33 mm diam, flat, dusty, white, margin entire, reverse concolourous. Description and illustration: Gams (1971). Typus: Germany, Kiel-Kitzeberg, from moist wall in greenhouse, unknown collection date and collector, isol. 1966 by W. Gams, No. 599, CBS H-24701 (holotype CBS 396.66 preserved as metabolically inactive culture, ex-type culture CBS 396.66 = IAM 14661). Fig. 38. Protocreopsis rutila (strain CBS 394.70). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidial heads on OA media. E. Conidiophores and conidial heads. F–H. Monophialides. I, J. Degraded conidiophores. K. Conidia. Scale bars = 10 μm.
106 Hou et al. Additional materials examined: Belgium, Haasrode, Meerdael Bos, from soil, unknown collection date and collector, isol. Nov. 1963 by G.L. Hennebert, CBS H-8300, culture CBS 378.70C = MUCL 6094. Denmark, Zealand, Graese, from agricultural soil, coll. 17 Nov. 2004, isol. 22 Dec. 2004, coll. and isol. S. Ronhede, Gr 161, culture CBS 118526. Germany, Kiel-Kitzeberg, from moist wall in greenhouse, unknown collection date and collector, isol. W. Gams, No. 602, culture CBS 229.70; Kiel-Kitzeberg, from scab of Malus domestica (Rosaceae), unknown collection date and collector, isol. W. Gams, No. 564, culture CBS 226.70; near Braunschweig, from agricultural löss soil, unknown collection date and collector, isol. H. Nirenberg, No. 208, culture CBS 263.89; from wheat field soil, unknown date and collector, culture CBS 395.66. Netherlands, Utrecht Province, Baarn, from humus rich soil, unknown collection date and collector, isol. Apr. 1964 by G.L. Hennebert, CBS H-8368, culture CBS 394.70 = MUCL 6903. Notes: According to the phylogenetic analysis (Fig. 2), the extype strain of Acremonium rutilum (CBS 396.66) is placed in Protocreopsis, and it is therefore recombined as Pt. rutila. Eight cultures representing Pt. rutila form two subclades: subclade I contains two strains from the moist wall in the greenhouse in Germany, including the ex-type CBS 396.66; subclade II comprises six strains mainly from soil of various European countries. Morphologically, the size of the structures observed in CBS 394.70 (subclade II) differs slightly from those described in the protologue of Pt. rutila (Gams 1971): phialides and conidia are larger than those of Pt. rutila [phialides 10.5–60.5 μm × 1.5–3 μm vs (15–)20– 40(–55) × 2.0–2.5(–3.0); conidia: 3.5–8.7 × 2.2–3 μm vs 3.0–6.0 × 2.0–3.3 μm]. However, we consider that these morphological differences may reflect intraspecific variability and are insufficient to allow confident splitting of this group into more than one species. Clade O24 Nectriopsis Maire, Ann. Mycol. 9: 323. 1911. (nom. cons.). Synonym: Dasyphthora Clem., Gen. Fung. (Minneapolis): 45. 1909. Perithecia superficial or immersed in substratum, generally not conspicuously stromatic, generally less than 200 μm diam., nearly white to pale yellow or orange, rarely violet or purple, KOH-. Ascomatal wall less than 20 μm thick, usually comprising of a single region of small thin-walled, non-descript cells; wall cells at the surface forming textura epidermoidea. Asexual morph where known, acremonium-like, gliocladium-like, or verticillium-like. On free-living fungi, lichens, and myxomycetes, less frequently on herbaceous substrata (Samuels 1988, Rossman et al. 1999). Type: Nectriopsis violacea (J.C. Schmidt ex Fr.) Maire Other accepted species with available sequences: Nectriopsis candicans (Plowr.) Maire, N. ellipsoidea L.W. Hou, L. Cai & Crous, N. rexiana (Sacc.) Rossman, L. Lombard & Crous, N. lindauiana (Bubák) Zare & W. Gams, N. fuliginicola Zare & W. Gams, N. microspora (Jaap) L.W. Hou, L. Cai & Crous, N. sporangiicola (Samuels) Samuels Notes: Nectriopsis was originally established for four hypocrealean fungi with ascomata in a byssoid stroma, which were considered intermediate between Nectria and Hypomyces (Maire 1911; Rossman et al. 1999). Samuels (1988) described and illustrated 43 species of this genus that were recognised three intergrading subgroups. This genus can be easily distinguished from Nectria s. str. in its thin perithecial wall with textura epidermoidea at the surface, combined with yellow pigmentation, small perithecial and a fungicolous habit (Samuels 1988). The monotypic genus Peloronectriella was introduced for a fungus (Pe. sasae) on bamboo having elongate, tuberculate stroma with nectria-like ascomata, and 1-septate ascospores (Doi 1968). Rossman et al. (1999) regarded it as synonym of Nectriopsis based on morphological characters. However, the ex-type strain of the type Pe. sasae (CBS 333.69) clustered with Cosmospora in Nectriaceae, which are distant from Nectriopsis s. str. (Supplementary Fig. S1). Therefore, it would seem that it is not a synonym of Nectriopsis. Nectriopsis candicans (Plowr.) Maire, Ann. Mycol. 9: 324. 1911. Fig. 39. Basionym: Hypomyces candicans Plowr., Grevillea 11: 50. 1882. Synonyms: Nectria candicans (Plowr.) Samuels, Mycologia 65: 412. 1973. Hypolyssus candicans (Plowr.) Kuntze, Revis. Gen. Pl. 3: 488. 1898. Description based on ex-epitype culture CBS 701.79: Perithecia superficial, solitary, gregarious or scattered, globose, sub-globose or broadly pyriform, pale yellow, becoming orange, dull, smooth, uniloculate, covered with abundant white, cylindrical, septate, unbranched hyphae, woolly except the ostiolum, 121–197 × 104– 165 μm. Ostiole single, central, papillate, formed of thin-walled, septate hyphae; hyphae arising from upper half of perithecial wall, branching, forming a network around papillae. Perithecial wall 8.5– 16 μm wide, composed of several layers of thick-walled, hyaline to pale brown, cylindrical cells, forming textura angularis. Paraphyses not observed. Asci 8-spored, unitunicate, cylindrical, apex simple, 37–55 × 3–4.5 μm. Ascospores uniseriate, with overlapping ends, oblong, cylindrical, rounded at both ends, 1-septate, not constricted at septa, hyaline, smooth-walled, eguttulate, without mucilaginous sheath, 4.5–6.8 × 2–2.7 μm. Asexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 42–47 mm diam, flat, felty, granular, dirty white, margin dendritic, reverse pale olivaceous buff; On MEA reaching 38–39 mm diam, flat, radially fold at centre, felty, dirty white, pale olivaceous buff at centre, with concentric rings, margin dendritic, reverse pale ochreous, with dirty white radial lines; On PDA reaching 49 mm diam, flat, thinly felty, granulose, dirty white, creamy white at periphery, margin dendritic, reverse pale olivaceous buff; On SNA reaching 18 mm diam, flat, sparse aerial mycelium, felty, white, margin dendritic, reverse concolourous. Other illustrations: Maire (1911), Samuels et al. (1973). Typus: UK, Great Britain, England, Bathford Down, on Mycetozoa, Aug. 1880, Leziate (K, lectotype in 1973). Additional materials examined: Germany, former West-Germany, from Fuligo septica (Protozoa), unknown collection date and collector, isol. W. Gams, culture CBS 440.65; Holzdorf, from Fuligo septica (Protozoa), unknown collection date and collector, isol. 15 Sep. 1961 by G. Arnold (MW m 43), culture CBS 424.64. Netherlands, Gelderland Province, Doorwerth, from Leocarpus fragilis (Protozoa) on Prunus laurocerasus (Rosaceae), 7 Aug. 1972, N.E. Nannenga-Bremekamp, culture CBS 627.72. ; Flevoland Province, Noordoostpolder, Kuinderbos, L perceel KB 72, from Fuligo sp. (Protozoa), Sep. 1979, W. Gams, CBS H-15093, culture CBS 701.79. Notes: According to the original literature, Hypomyces candicans was initially isolated from a kind of Mycetozoa in the UK described as having globose, gregarious perithecia, 200 × 150 μm, cylindrical asci 50–60 × 3–5 μm, ovoid to oblong, uniseptate ascospores measuring 8 × 3 μm, blunt at both ends, and surrounded by white,
107www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales floccose mycelium (Plowright 1882). Later, it was transferred to Nectriopsis as N. candicans (Maire 1911). No holotype specimen was cited in the original description. Samuels et al. (1973) designated the collection from Bathford Downs in England as lectotype, because it agreed in all respects with the original description of H. candicans. In the present study, four strains received as N. candicans and isolated from Mycetozoa Fuligo spp. and Leocarpus fragilis in Europe were examined. One of the strains, CBS 701.79, morphologically matches the original description by producing comparable sized and shaped ascomata, asci and ascospores. Therefore, CBS 701.79 is considered a representative culture of H. candicans, and typification of this species awaits further collections. Nectriopsis ellipsoidea L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845854. Fig. 40. Etymology: Referring to the ellipsoid conidia produced by this fungus. Mycelium consisting of branched, septate, hyaline, rough-, thin-walled hyphae, up to 2.3 μm wide. Sporulation abundant, phalacrogenous or nematogenous. Conidiophores (sub-)erect, arising from submerged and superficial hyphae, orthogonally, unbranched or basitonously branched, bearing 1–3 levels with 1–3 phialides per node, commonly proliferating sympodially, up to ca. 254 μm long, 1.6–3.4 μm wide at base, 1–2-septate, hyaline, smooth-walled, coarse in lower part. Phialides monoor polyphialidic, lateral or terminal, orthotropic, cylindrical or subulate, hyaline, thick-, smooth-walled, (9.0–)15.2–51.8 μm long, 1–2.1 μm wide at base, with conspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides with up to three conidiogenous loci commonly present; with percurrent Fig. 39. Nectriopsis candicans (ex-epitype culture CBS 701.79). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Ascomata on OA. E, F. Ascomata. G, H. Asci. I. Ascospores. Scale bars: E, F = 20 μm; G–I = 10 μm.
108 Hou et al. or subterminal proliferations. Conidia aseptate, ellipsoid, with obviously apiculate bases and rounded apices, hyaline, thin-, smoothor rough-walled, 4.3–7 × 2.5–3.5 μm, guttulate, arranged in moist slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 85 mm diam, flat, felty, slightly floccose in some sections, dirty white, margin filiform, reverse colourless; On MEA reaching 85 mm diam, flat, woolly at centre, felty at periphery, mycelial ropes abundant, creamy white, aerial mycelium arranged radially, margin filiform, reverse luteous; On PDA reaching 85 mm diam, flat, aerial mycelium arranged radially, floccose, mycelial ropes abundant, dirty white, margin filiform, reverse buff; On SNA reaching 52 mm diam, flat, membranous without aerial mycelium, white, margin fimbriate, reverse concolourous. With strong geosmin odour on PDA and MEA media, without odour at OA and SNA media. Typus: USA, Louisiana, along Chappepeela Creek, Tangipahoa Parish, from Polyporus pargamenus (Polyporaceae) on log, unknown collection date, isol. 5 Jun. 1976, coll. and isol. C.T. Rogerson (holotype CBS H-24628, ex-type culture CBS 358.78 = C.T.R. 76-39). Notes: Nectriopsis ellipsoidea was associated with Polyporus pargamenus on a log of an unknown tree. It is phylogenetically different from the closely related species N. rexiana. Morphologically, it differs from N. rexiana in producing longer and ellipsoid conidia (4.3–7 × 2.5–3.5 μm), while conidia of the latter are shorter and ovoid (5 × 3 μm) (Saccardo 1882). Nectriopsis fuliginicola Zare & W. Gams, Mycol. Progr. 15: 1027. 2016. Description and illustration: Zare & Gams (2016). Fig. 40. Nectriopsis ellipsoidea (ex-type culture CBS 358.78). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores and conidial heads. F. Percurrently proliferated conidiophores. G. Unbranched conidiophore. H–K. Branched conidiophores. L. Conidia. Scale bars = 10 μm.
109www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Typus: Russia, St. Petersburg region, Puschkin, from Fuligo septica (Protozoa), unknown collection date and collector, isol. 11 Oct. 1969 by G. Arnold, No. 11 (holotype CBS H-22407, ex-type culture CBS 400.82). Nectriopsis lindauiana (Bubák) Zare & W. Gams, Mycol. Progr. 15: 1023. 2016. Basionym: Verticillium lindauianum Bubák, Ann. Mycol. 12: 210. 1914. Description and illustration: Zare & Gams (2016). Typus: Germany, Kr. Plön, Schüttbrehm, from Fuligo septica (Protozoa), 30 Aug. 1965, W. Gams (neotype CBS H-22408, ex-neotype culture MUCL 7926 = CBS 897.70). Nectriopsis microspora (Jaap) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845855. Basionym: Verticillium microsporum Jaap, Verh. Bot. Ver. Prov. Brandenb. 58: 38. 1916. Synonyms: Sesquicillium microsporum (Jaap) Veenb.-Rijks & W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 226. 1971. Tolypocladium microsporum (Jaap) Bissett, Canad. J. Bot. 61: 1318. 1983. Gliocladium microsporum (Jaap) Arx, Mycotaxon 25: 157. 1986. (nom. illegit.) Sesquicillium parvulum Veenb.-Rijks, Acta Bot. Neerl. 19(3): 323 (1970) Description and illustration: Gams (1971). Material examined: Canada, Ontario, Petawawa, from forest soil under Populus tremuloides (Salicaceae), Oct. 1968, G.C. Bhatt, No. PET 160, CBS H-18229, culture CBS 355.70 = DAOM 175216. Netherlands, Flevoland Province, Oostelijk Flevoland, from wheat field soil, 12 Mar. 1969, J.W. Veenbaas-Rijks, No. 223 (holotype of Sesquicillium parvulum CBS H-7751, isotypes CBS H-1022, CBS H-7753, CBS H-7754 & CBS H-7755, ex-type culture CBS 933.69 = ATCC 18932 = DAOM 175219 = IMI 152204 = IPO 738). Notes: According to Gams (1971), the type material of Verticillium capitatum Ehrenb. in B no longer shows the original fungus. The substrate was indicated as “small dead insects and larvae, and then transferred to rotten wood”, which is why identity with the present fungus is unlikely. In the present study, the representative strain and type strain of its synonym (Sesquicillium microsporum) clustered within the genus Nectriopsis. Nectriopsis rexiana (Sacc.) Rossman et al., Stud. Mycol. 80: 243 (2015) Basionym: Verticillium nanum subsp. rexianum Sacc., Michelia 2: 577. 1882. Synonyms: Verticillium rexianum (Sacc.) Sacc., Syll. Fung. 4: 153. 1886. Hypomyces exiguus Pat., Bull. Soc. Mycol. France. 18: 180. 1902. Nectriopsis exigua (Pat.) W. Gams, Netherlands J. Pl. Pathol. 88: 73. 1982. Verticillium niveostratosum Lindau, Verh. Bot. Vereins Prov. Brandenburg. 45: 158. 1904 (1905). Nectria myxomyceticola Samuels, Mycologia 65: 409. 1973. Description: Saccardo (1882). Materials examined: Germany, Kr. Rendsburg, Enkendorfer Gehölz, from Arcyria sp. (Protozoa), unknown collection date, W. Gams, No. 1090, culture CBS 305.70C. UK, England, Derbyshire, Haddon Hall, from Physarum nutans (Protozoa), unknown collection date, W. Gams, No. 1075, culture CBS 305.70A. Nectriopsis sporangiicola (Samuels) Samuels, Mem. New York Bot. Gard. 48: 49. 1988. Basionym: Nectria sporangiicola Samuels, Mycologia 65: 416. 1973. Descriptions: Samuels (1973, 1988). Typus: USA, New Jersey, Cape May County, 5 km SE of Woodbine, from Physarum polycephalum (Protozoa), 14 Sep. 1967, C.T. Rogerson (isotypes CBS H-7415, CBS H-7416 & CBS H-7417, ex-isotype culture CBS 166.74 = ATCC 26542 = C.T.R. 67-136). Nectriopsis violacea (J.C. Schmidt ex Fr.) Maire, Ann. Mycol. 9: 323. 1911. Fig. 41. Basionym: Sphaeria violacea J.C. Schmidt ex Fr., Syst. Mycol. (Lundae) 2: 441. 1823. Synonyms: Nectria violacea (J.C. Schmidt ex Fr.) Fr., Summa Veg. Scand., Sectio Post. (Stockholm) 1: 388. 1849. Hypomyces violaceus (J.C. Schmidt ex Fr.) Tul. & C. Tul, Ann. Sci. Nat., Bot., sér. 4 13: 14. 1860. Peckiella violacea (J.C. Schmidt ex Fr.) Sacc., Syll. Fung. (Abellini) 9: 945. 1891. Hypolyssus violaceus (J.C. Schmidt ex Fr.) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 488. 1898. Byssonectria violacea (J.C. Schmidt ex Fr.) Seaver, Mycologia 2: 65. 1910. Hyphonectria violacea (J.C. Schmidt ex Fr.) Petch, J. Bot. 74: 220. 1937 (1936). Description based on ex-epitype culture CBS 914.70: Sexual morph: perithecia immersed in mycelium, becoming collabent when dry, solitary, gregarious or scattered, globose, sub-globose or broadly pyriform, violet, vinaceous, uniloculate, covered with abundant cylindrical, septate, hyaline hyphae, 148.5–304 × 126.5–238.5 μm. Ostiole single, central, papillate, formed of thickwalled, septate, unbranched hyphae; hyphae extending outwardly as hairs, 25–55 μm long, 3–4 μm diam at rounded tips, forming a fringe around papillae. Perithecial wall 14–19 μm wide, composed of several layers of thick-walled, cylindrical, hyaline to pale brown cells forming textura angularis. Paraphyses not observed. Asci unitunicate, 8-spored, cylindrical, apex simple, 46–70.5 × 3–4.5 μm. Ascospores obliquely uniseriate with overlapping ends, oblong, cylindrical, rounded at both ends, 1-septate, not constricted, hyaline, spinulose, without mucilaginous sheath, 4.7–8 × 2–2.7 μm. Asexual morph: sporulation abundant, phalacrogenous or nematogenous. Conidiophores (sub-)erect, arising from submerged and superficial hyphae, unbranched or branched, bearing 1–3 levels with 1–3 phialides per node, up to ca. 355 μm long, with 1–2 septa in basal and upper part, hyaline, smooth-walled. Phialides monophialidic, lateral or terminal, cylindrical or subulate, hyaline, thin-, smoothwalled, 18–55 μm long, 1.3–2.3 μm wide at base, with conspicuous cylindrical collarette and periclinal thickening at conidiogenous loci. Conidia variable in size, aseptate, ellipsoid, cylindrical or oblong, rounded at both ends, or with a truncate basal end, hyaline, thin-, smooth-walled, 2.8–9.3 × 1.9–2.8 μm, guttulate, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 70–72 mm diam, flat, felty, with some floccose zones, dirty white, margin entire, reverse buff; On MEA reaching 60–62 mm diam, flat, radially fold at centre, floccose and white at centre, felty and pale luteous at periphery, margin entire, reverse orange at centre, pale
110 Hou et al. Fig. 41. Nectriopsis violacea (ex-epitype culture CBS 914.70). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Ascomata on OA. E. Ascoma. F. Ostiole formed of thick-walled hyphae. G–I. Asci. J–L. Conidiophores. M. Conidia. Scale bars: E = 50 μm; F–G = 20 μm; H–M = 10 μm.
111www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales luteous at periphery, with white radial lines; On PDA reaching 65 mm diam, flat, felty, slightly floccose at centre, white, dirty white and filiform margin, reverse buff; On SNA reaching 33 mm diam, flat, sparse aerial mycelium, membranous, colourless, margin entire, reverse colourless. Typus: Germany, Sachsen, Bernstadt, Fuligo septica (Protozoa), 1817, G. Kunze (holotype UPS: BOT: F-004721 as Sphaeria violacea); Eifel, Gerolstein, Gerolsteiner Wald, from Fuligo septica (Protozoa), 15 Sep. 1970, W. Gams (epitype designated here, CBS H-15101, MBT10009440, ex-epitype culture CBS 914.70). Additional material examined: Germany, Eifel, Pelmer Wald bei Gerolstein, from Fuligo septica (Protozoa), 18 Sep. 1970, W. Gams, CBS H-15102, CBS H-15104 & CBS H-15105, culture CBS 849.70. Notes: Several nectria-like species had been reported to develop on sporangia, commonly of Fuligo and Stemonitis, including N. candicans, N. hirsuta, N. myxomyceticola, N. sporangiicola and N. violacea (Samuels et al. 1973). Nectria violacea, N. candicans and N. myxomyceticola are closely related, having cylindrical or oblong ascospores which are less than 10 μm long, cylindrical asci averaging less than 60 μm long, and thin perithecial walls with modified hyphae or hairs. Nectriopsis violacea (basionym: Sphaeria violacea) was originally reported on Fuligo septica in Germany (Fries 1823), and is characterised by having violet to purple perithecia. In the present study, two cultures isolated from Fuligo septica in Germany were examined. Culture CBS 914.70, which is from the same host and country and agreed well in morphology with the protologue (Fig. 41), is herewith designated as ex-epitype culture, and the specimen CBS H-15101 as epitype of Sphaeria violacea. This species formed a fully supported clade in Bionectriaceae (BPP/MLBS = 1/100 %; Fig. 2). Clade O27 Mycocitrus Möller, Bot. Mitt. Trop. 9: 297. 1901. Stroma well-developed, buff to rufous, clasping and surrounding the substratum. Ascomata immersed, with apices barely visible, densely gregarious, forming a single layer. Asci cylindrical, ascal apex simple. Ascospores ellipsoid, 1-septate, hyaline, spinulose. Asexual morph acremonium-like. On living stems of bamboo (Rossman et al. 1999). Type: Mycocitrus aurantium Möller Other accepted species with available sequences: Mycocitrus odorus L.W. Hou, L. Cai & Crous, M. phyllostachydis (Syd. & P. Syd.) Yoshim, M. zonatus (Sawada) L.W. Hou, L. Cai & Crous Notes: Mycocitrus was originally described based on the type M. aurantium from culms of living bamboo (Guadua) and on Microstachys sp. (Euphorbiaceae) in the city of Blumenau, Santa Catarina, in southern Brazil (Möller 1901). This genus is characterised by its large fleshy orange stromata that clasp and surround bamboo stems, and perithecial ascomata partially to fully immersed in the upper region of the stromata. Mycocitrus was originally placed in “Hypocreaceen, Didymosporae” (Möller 1901), or in Hypocreaceae (Doi 1967). Rossman et al. (1999) included Mycocitrus in the newly introduced family Bionectriaceae based on morphology. Recently, two bamboo-inhabiting Mycocitrus aurantium strains (BAFC 3843 and BAFC 51693) found in South America were discovered to form an independent lineage within Hypocreales based on ITS phylogenetic tree and limited taxa. This lineage was distinct from Bionectriaceae, Nectriaceae, Cordycipitaceae, Clavicipitaceae and Hypocreaceae, and suggesting Mycocitrus represents an additional lineage within Hypocreales (Leite et al. 2018). However, only ITS sequences were available for these strains, and may affect the support values of clades and result in unreliable phylogenies, we therefore excluded these strains from the dataset for the phylogenetic analysis of Bionectriaceae (Fig. 2). A dataset that included more strains was used to construct a more complete phylogeny, showing that the two strains of M. aurantium clustered in a well-supported lineage together with strain CBS 330.69 that representing Mycocitrus phyllostachydis. This cluster lies within the Bionectriaceae (Supplementary Fig. S2), a placement which conforms to the proposition of Rossman et al. (1999). Mycocitrus odorus L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845856. Fig. 42. Etymology: Referring to the prominent odour this fungus produces in culture. Mycelium consisting of branched, septate, hyaline, rough-, thin-walled hyphae, up to 2.5 μm wide. Sporulation abundant, phalacrogenous or nematogenous. Conidiophores (sub-)erect, straight or curved, arising from submerged and superficial hyphae, or radiating out from the coils formed by mycelium, unbranched or basitonously, verticillately branched, bearing 1–4 levels with 2–3 phialides per node, up to ca. 133 μm long, 2–3.8 μm wide at base, with 1–2(–3) septa at basal, middle or upper part, coarse at lower part, hyaline, smooth-walled. Phialides monophialidic, lateral, cylindrical or subulate, hyaline, thick-, smooth-walled, 14.5–60 μm long, 1.5–2.8 μm wide at base, with conspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, oblong, symmetrically rounded, hyaline, thin-, smooth-walled, 3.5–7.5 × 2–4.5 μm, with 2–4 minute guttules, arranged in slimy heads, mostly confluent. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 78–80 mm diam, flat, moderate aerial mycelium, thinly felty, abundant mycelial granula, dirty white, margin entire, reverse dirty white, strong fresh-leaf odour (aromatic hydrocarbon); On MEA reaching 78–80 mm diam, raised, abundant aerial mycelium, floccose, granular, abundant mycelial ropes, white at centre and periphery, with a buff ring, margin entire, reverse luteous; strong fresh-leaved odour (aromatic hydrocarbon); On PDA reaching 80 mm diam, flat, moderate aerial mycelium, floccose at centre, felty at periphery, dirty white, margin filiform, reverse concolourous, strong geosmin odour; On SNA reaching 80 mm diam, flat, with sparse aerial mycelium, membranous, white, margin filiform, reverse white (no obvious odour). Typus: Netherlands, North Holland Province, Amsterdam, Slotervaart Hospital, on onychomycosis (human), unknown collection date, W.C. van Dijk & W. Pauw (holotype CBS H-24690, ex-type culture CBS 100104). Additional material examined: Netherlands, human skin, unknown collection date and collector, culture CBS 120610. Sweden, Stockholm, from human skin, unknown collection date and collector, No. BF 220/74c, culture CBS 232.75B. Notes: Three cultures from human skin or nails formed a fully supported clade sister to Mycocitrus zonatus, representing
118 Hou et al. Sagadahoc County, Bailey Island, from wrack of eelgrass (probably Zostera marina, Zosteraceae, seagrasses), 10 Aug. 2005, unknown collector, culture CBS 120611; Maine, Kittery Point, York County, from Ascophyllum nodosum (Fucaceae, brown algae), 10 Aug. 2005, D. Malloch, ACR 8-1, culture CBS 120530; Massachusetts, Paine’s Creek Beach, from wrack of Fucus vesiculosus, 9 Aug. 2005, D. Malloch, ACR 3-7, culture CBS 120529. Notes: Emericellopsis fuci is characterised by its relatively large, ovate conidia with a truncate basal hilum and a slightly flattened apex (Zuccaro et al. 2004). Zuccaro et al. (2004) stated that the basionym, A. fuci, was the second Acremonium species with distinctively large conidia to have been described from the marine environment. Phylogenetically, the algal-associated A. fuci clustered within Emericellopsis, representing a different ecological niche for this genus (Zuccaro et al. 2004). In this study, 11 strains from diverse algae (including the type of A. fuci) and 16 strains from the coast of Norway formed a fully supported clade in the “marine clade” of Emericellopsis. Therefore, a new combination is introduced, E. fuci (Fig. 2). Emericellopsis glabra (J.F.H. Beyma) Backus & Orpurt, Mycologia 53: 75. 1962 (1961). Basionym: Emericellopsis terricola var. glabra J.F.H. Beyma, Antonie van Leeuwenhoek 6: 266. 1940 (1939–1940). Description and illustration: van Beyma (1940). Typus: Netherlands, Utrecht Province, Baarn, from soil, 1937, F.H. van Beyma, isol. 1937 by N. Schierbeek, CBS H-12388 (holotype culture CBS 119.40 preserved as metabolically inactive culture, ex-type culture CBS 119.40 = IAM 14674 = IAM 14675 = JCM 10470 = MUCL 11486). Emericellopsis microspora Backus & Orpurt, Mycologia 53: 67. 1962 (1961). Description and illustration: Backus & Orpurt (1961). Typus: USA, Wisconsin, from wet prairie soil, summer 1953, unknown collector, isol. P.A. Orpurt, CBS H-12400 (holotype WSF 47 in WIS, exisotype culture CBS 380.62 = ATCC 14645 = IFO 9241 = IMI 092625 = MUCL 11494 = VKM F-1301 = WSF 47). Emericellopsis minima Stolk, Trans. Brit. Mycol. Soc. 38: 419. 1955. Description and illustration: Stolk (1955). Typus: Mozambique, Inhaca, from mangrove soil, unknown collection date, isol. Jul. 1953, coll. and isol. H.J. Swart, CBS H-7056 (holotype K(M) 36100, ex-type culture CBS 190.55 = JCM 10472 = VKM F-1484). Emericellopsis mirabilis (Malan) Stolk, Trans. Brit. Mycol. Soc. 38: 421. 1955. Basionym: Peyronellula mirabilis Malan, Mycopathol. Mycol. Appl. 6: 173. 1952. Typus: Italy, from damp soil, dead protonema of moss, isol. 1952 by C.E. Malan, No. 1 (ex-syntype of Peyronellula mirabilis CBS 177.53 = JCM 10471 = MUCL 11482); unknown substrate and collection date, isol. C.E. Malan (ex-syntype culture of Peyronellula mirabilis CBS 176.53). Emericellopsis moniliformis (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 846143. Basionym: Acremonium moniliforme A. Giraldo et al., Mycol. Progr. 16: 357. 2017. Description and illustration: Giraldo et al. (2017). Typus: Spain, Aragón, Huesca Province, Ordesa y Monte Perdido National Park, from forest soil, 15 Mar. 2011, A. Giraldo, M. Hernández, & J. Capilla, isol. A. Giraldo (holotype CBS H-22022, dried culture on OA, ex-type culture CBS 139051 = FMR 11785). Additional materials examined: Chile, from volcanic ash soil, unknown collection date and collector, isol. J. Grinbergs, No. 520/73, culture CBS 864.73. Netherlands, Overijssel Province, Kloosterhaar, from human nail, unknown collection date and collector, isol. Streeklab. voor de Volksgezondheid, Groningen, No 1984, culture CBS 463.92. Emericellopsis pallida Beliakova, Mikol. Fitopatol. 8: 386. 1974. Description and illustration: Beljakova (1974). Typus: Ukraine, Crimea, Black Sea, from water, unknown collection date and collector, isol. L.A. Belyakova (holotype CBS 490.71 preserved as metabolically inactive culture, ex-type culture CBS 490.71 = VKM F-925). Additional material examined: Canada, Manitoba, Delta, University Field Station, from organic soil under Phragmites communis (Poaceae), unknown collection date, J. Reid, CBS H-12403, culture CBS 624.73 = UM 194. Emericellopsis salmonea (W. Gams & Lodha) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845862. Basionym: Acremonium salmoneum W. Gams & Lodha, Trans. Brit. Mycol. Soc. 64: 399. 1975. Description and illustration: Gams (1975). Typus: India, Rajasthan, Jaipur, from dung of deer, unknown collection date, B.C. Lodha, CBS H-6667 (holotype CBS 721.71 preserved as metabolically inactive culture, isotype IMI 185379 = ATCC 32185, ex-type culture CBS 721.71). Emericellopsis salmosynnemata Grosklags & Swift, Mycologia 49: 305. 1957. Description and illustration: Grosklags & Swift (1957). Typus: USA, Michigan, laboratory contaminant, unknown collection date and collector, isol. 1953 by J.M. Roberts, (holotype MDH 3590A, ex-type culture VKM F-1260 = CBS 182.56 = ATCC 11661 = DAOM 64321 = IFO 9239 = IMI 058330 = NRRL 2271). Emericellopsis stolkiae D.E. Davidson & M. Chr., Trans. Brit. Mycol. Soc. 57: 385. 1971. Description and illustration: Davidson & Christensen (1971). Typus: USA, South East Wyoming, from mud in saline lake, unknown collection date, M. Christensen, CBS H-7059 (holotype IMI 155476, exholotype culture CBS 159.71 = ATCC 22761 = IFO 9604 = RMF 3002). Additional material examined: USA, from soil in shore of permanent alkali lake, unknown collection date and collector, culture CBS 139531. Emericellopsis terricola J.F.H. Beyma, Antonie van Leeuwenhoek 6: 265. 1940 (1939). Description and illustration: Beyma (1940). Typus: Netherlands, Utrecht Province, Baarn, from soil, unknown collection date, J.F.H. Beyma isol. 1937 by H. Schierbeek, CBS H-12407
119www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales (holotype CBS 120.40 preserved as metabolically inactive culture, ex-type culture CBS 120.40 = JCM 10474). Emericellopsis tubakii (Gams) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845863. Basionym: Acremonium tubakii W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 55. 1971. Synonyms: Cephalosporium polyaleurum Tubaki, Mycologia 65: 939. 1973. Acremonium polyaleurum (Tubaki) E.B.G. Jones et al., Fungal Diversity 35: 147. 2010. Description and illustration: Gams (1971). Typus: Japan, coast near Wakayama City, from marine sediment, collection date unknown, isol. 1967, coll. and isol. K. Tubaki, No. VII-4, CBS H-8458 (holotype of Acremonium tubakii CBS 790.69 preserved as metabolically inactive culture, ex-type culture CBS 790.69 = IAM 14664 = IFO 874). Additional materials examined: India, Andhra Pradesh, Hyderabad, from Oryza sativa (Poaceae), rhizosphere, unknown collection date, B.K. Vaidehi, CBS H-8462, culture CBS 791.69 = IMI 131943. Japan, coast of Wakayama City, from seashore, unknown collection date and collector, isol. 1967 by K. Tubaki, No. IV-8, CBS H-8464 & CBS H-8465, culture CBS 824.69 = IFO 788; Wakayama Bay, from coastal mud, unknown collection date, K. Tubaki, MM-199 (holotype of Cephalosporium polyaleurum IFO-H11637, ex-type culture CBS 555.74 = IFO 9394). Netherlands, Flevoland Province, Zuidelijk Flevoland, from soil, unknown collection date and collector, isol. Sep. 1970 by G.M. Tichelaar, No. 295, culture CBS 822.70. Notes: Emericellopsis tubakii was originally reported as Acremonium tubakii from marine sediment in Japan, near Wakayama City, which is characterised by its cylindrical conidia (3.5–5.7 × 1.5–2.3 μm), arranged in heads, and sub-globose to ovoid chlamydospores covered with chromophilic warts and arranged in chains (Gams 1971). Two years later Cephalosporium polyaleurum was described from the same kind of substrates and country as E. tubakii during a study of Emericellopsis species from the marine environment (Tubaki 1973). Morphologically, C. polyaleurum is similar with E. tubakii in its shape and size of conidia (3–6 × 1.5–2 μm) and chlamydospores (Tubaki 1973). In our molecular analyses, the extype of C. polyaleurum (CBS 555.74) and A. tubakii (CBS 790.69) are phylogenetically identical, grouping in a fully supported lineage in Emericellopsis (Fig. 2). Therefore, they are conspecific with E. tubakii having priority, and a new combination was introduced. Clade O29 Stanjemonium W. Gams et al., Canad. J. Bot. 76: 1579. 1999 (1998). Colonies slow growing, densely woolly to felty. Sporulation abundant. Conidiophores on single or slightly fasciculate aerial hyphae. Conidiogenous cells flask-shaped or cylindrical, scattered along subtending cells, either singly or in pseudowhorls (i.e., whorls not concentrated near a septum), each producing a single conidium, or producing multiple conidia arranged in slimy heads; cells collapsing soon after maturation of conidia, with short low projections on subtending hyphae. Conidia with rotational symmetry, ellipsoid to cylindrical, with apiculate base, hyaline to distinctly pigmented. Chlamydospores and sexual morph absent. (emended from Gams 1998). Type: Stanjemonium grisellum W. Gams et al. Other accepted species with available sequences: Stanjemonium dichromosporum (Gams & Sivasith.) L.W. Hou, L. Cai & Crous, S. fuscescens W. Gams, Schroers & Abdullah, S. ochroroseum W. Gams et al. Notes: Stanjemonium is an asexual morph genus, and not known to be associated with any sexual morph. This genus is quite different from Acremonium and is known to have a close relationship with Emericellopsis (Fig. 2), as shown by Gonçalves et al. (2020). It was characterised by producing slow-growing colonies and pigmented conidia from flask-shaped conidiogenous cells which collapse soon after conidial maturation, leaving low projections on the subtending hyphae (Gams et al. 1998). In addition, each conidiogenous cell produces a single conidium (Gams et al. 1998). Based on the published molecular data and phylogenetic analysis, Gams et al. (1998) demonstrated the affinity between Stanjemonium and Emericellopsis. Our phylogenetic analysis results agree well with the previous study showing that Stanjemonium formed a fully supported clade in Bionectriaceae, showing a close relationship with Emericellopsis (Fig. 2). Stanjemonium dichromosporum (Gams & Sivasith.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845864. Basionym: Acremonium dichromosporum W. Gams & Sivasith., Trans. Brit. Mycol. Soc. 64: 397. 1975. Synonym: Gliomastix dichromospora (W. Gams & Sivasith.) Subram., Kavaka 5: 98. 1978 (1977). Description and illustration: Gams (1975). Typus: Australia, Western Australia, Nedlands, from rhizosphere of Triticum aestivum (Poaceae), unknown collection date and collector, dep. K. Sivasithamparam, CBS H-6604 (holotype of Acremonium dichromosporum CBS 638.73 preserved as metabolically inactive culture, isotype IMI 185377, ex-type culture CBS 638.73 = ATCC 32181 = IMI 185377). Notes: This species was originally reported as Acremonium dichromosporum in section Gliomastix (Gams 1975). The multilocus phylogenetic analysis placed the ex-type strain (CBS 638.73) in the genus Stanjemonium (Fig. 2). Morphologically, S. dichromosporum differs from other known Stanjemonium species in its cylindrical conidiogenous cells and abundant conidia arranged in orange, slimy heads (Gams 1975), while the others have flask-shaped conidiogenous cells giving rise to only a single conidium (Gams et al. 1998). The uniconidial blastic cells or aphanophialides proved to coexist with typical phialides even within individual species (Zare & Gams 2001), and thus appear to be less important at the generic level. The described morphology of A. dichromosporum is consistent with the generic description of Stanjemonium in its unbranched, densely arranged phialides, and pigmented conidia (Gams 1975). Stanjemonium fuscescens W. Gams, Schroers & Abdullah, Canad. J. Bot. 76: 1580. 1999 (1998). Description and illustration: Gams et al. (1998). Typus: Iraq, from desert soil, unknown collection date and collector, isol. S. Abdullah on actidione medium, dep. J. Guarro (holotype of Stanjemonium fuscescens CBS H-6076, isotypes CBS H-5711 & CBS H-8778, ex-type culture CBS 264.96 = NRRL 26546).
120 Hou et al. Stanjemonium grisellum W. Gams et al., Canad. J. Bot. 76: 1580. 1999 (1998). Description and illustration: Gams et al. (1998). Typus: USA, Wyoming, ca. 11 km west of Rock Springs, Sweetwater County, alt. 1 921 m, soil from native Artemisia tridentata (Asteraceae) grassland, Sep. 1978, M. Christensen, RMF 05 (holotype CBS H-6077, isotype CBS H-8777, ex-type culture CBS 655.79 = NRRL 26538). Stanjemonium ochroroseum W. Gams et al., Canad. J. Bot. 76: 1580. 1999 (1998). Description and illustration: Gams et al. (1998). Typus: USA, Wyoming, approximately 11 km west of Rock Springs, Sweetwater County, alt. 1 921 m, soil from native Artemisia tridentata (Asteraceae) grassland, Sep. 1978, M. Christensen, RMF G27 (holotype CBS 656.79 preserved as metabolically inactive culture, ex-type culture CBS 656.79 = NRRL 26539). Clade O30 Proliferophialis L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845865. Etymology: Referring to the proliferation of conidiophores and phialides of the type. Mycelium consisting of branched, septate, hyaline, smoothor rough-, thin-walled hyphae. Conidiophores solitary or aggregated, erect, usually reduced to single phialides, arising from aerial, substratal mycelium, or from mycelial ropes, unbranched, basitonously branched, repeatedly proliferating sympodially or percurrently, straight or curved, hyaline, smoothor rough-walled, with cell walls usually thicker than those of vegetative hyphae. Conidiogenous cells enteroblastic, monoand polyphialidic, terminal or lateral, subulate, hyaline, thick-, smooth-walled, commonly with apical percurrent proliferation, and with inconspicuous periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, ovoid, obpyriform, or fusoid, with hilar spot at both apices and bases, hyaline, thick-, smooth-walled, arranged in long chains, often irregularly collapsing into conidial heads. Chlamydospores and sexual morph not observed. Type: Proliferophialis apiculata L.W. Hou, L. Cai & Crous Notes: The monotypic genus Proliferophialis is proposed here to accommodate four cultures clustering in a fully supported lineage in Bionectriaceae (Fig. 2). Three cultures were formerly identified as Acremonium egyptiacum and one was received as A. potronii. However, they are phylogenetically distant from the ex-type of A. egyptiacum and A. potronii, and form an independent clade that differs from all other known genera in Bionectriaceae (Fig. 2). Morphologically, Proliferophialis is characterised by producing abundant phialides with repeatedly apical percurrent proliferation. Proliferophialis apiculata L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845866. Fig. 45. Etymology: Referring to its terminal conidia, which are apiculate at both ends. Mycelium consisting of branched, septate, hyaline, smoothor rough-, thin-walled hyphae, 1–2.5 μm wide. Conidiophores solitary or aggregated, erect, straight or irregularly curved, arising directly from aerial or substratal mycelium, or from ropes formed by mycelium, usually reduced to single phialides, unbranched, or basitonously branched, repeatedly proliferating sympodially or percurrently, up to 162 μm long, 1.8–3.2 μm wide at base, hyaline, smoothor roughwalled 1–2-septate, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral or terminal, subulate, hyaline, thick-, smooth-walled, 17–94 μm long, 1–2 μm wide at base, commonly with apical percurrent proliferation, and with minute collarette and inconspicuous periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, ovoid, obpyriform or fusoid, with apiculate thicken hilar spot at both ends, hyaline, thick-, smooth-walled, 3.2–6.6(–7.3) ×1.9–2.7 μm, arranged in long chains, often irregularly collapsing in conidial heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 26–27 mm diam, flat, moderate aerial mycelium, thinly felty, dusty, with mycelium ropes at centre, white, margin entire, reverse creamy white; On MEA reaching 25–26 mm diam, raised, radially folded, moderate aerial mycelium, felty, white, margin entire, reverse ochreous, with buff radial lines; On PDA reaching 30 mm diam, flat, felty, white, margin entire, reverse olivaceous at centre, rosy buff at periphery; On SNA reaching 22–23 mm diam, flat, membranous without aerial mycelium, white, margin entire, reverse concolourous. Typus: France, Rennes, from Triticum aestivum (Poaceae), stored grain, unknown collection date and collector, isol. Sep. 1962 by J. Pelhâte, No. X3 (holotype CBS H-8155, ex-holotype culture CBS 303.64). Additional materials examined: France, near Rennes, from Triticum aestivum (Poaceae), stored grain, unknown collection date and collector, isol. Sep. 1962 by J. Pelhâte, No. X4, CBS H-8157, culture CBS 365.64. USA, Wisconsin, from Acer saccharum (Sapindaceae), leaf litter, unknown collection date and collector, isol. G.A. Kuter, No. 1175, culture CBS 397.78 = IAM 14646; Wyoming, appr. 11 km west of Rock Springs, soil from native Artemisia tridentata (Asteraceae) grassland, unknown collection date and collector, isol. Jun. 1978 by M. Christensen, No. S 104, culture CBS 542.79. Notes: The four cultures of Proliferophialis apiculata form a distinct clade basal to Emericellopsis and Stanjemonium (Fig. 2). Morphologically, Pro. apiculata is characterised by the production of abundant phialides with repeatedly apical percurrent proliferation. Based on a blastn search of NCBIs GenBank nucleotide database, the closest hits using the ITS sequence of Pro. apiculata are sequences of Stanjemonium ochroroseum [culture CBS 125923; GenBank MH864099.1, identity = 469/516 (91 %), no gaps] (except culture CBS 127849, which was probably misidentified as “A. egyptiacum”, as the ITS sequences only shares 88.5 % similarity to the ex-type strain of A. egyptiacum CBS 114785). The closest hits using the LSU sequence is a culture of Acremonium brachypenium [culture CBS 866.73; GenBank MH872547.1, identities = 759/780 (97 %), 5 gaps (0 %)]. The closest hits using the rpb2 sequence is sequence of Fusarium solani [culture CBS 102429; GenBank KM232376.1, identity = 305/403 (75.68 %), 20 gaps (4 %)]. The closest hits using the tef sequence is sequence of Amphichorda guana [culture LC5815; GenBank KX855211.1, identities = 745/808 (92 %), no gaps]. Clade O31 Acremonium Link, Mag. Gesell. Naturf. Freunde, Berlin 3(1–2): 15. 1809.
121www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Synonyms: Monoconidia Roze, Bull. Soc. Mycol. France 13: 83. 1897. Mastigocladium Matr., C. R. Hebd. Séanc. Acad. Sci., Paris 152: 325. 1911. Pseudofusidium Deighton, Mycol. Pap. 118: 26. 1969. Mycelium consisting of branched, septate, hyaline, smooth-, thinor thick-walled hyphae, occasionally chondroid. Sporulation abundant, phalacrogenous, nematogenous, synnematogenous, plectonematogenous. Conidiophores solitary, erect, straight or irregularly curved, lateral or terminal, arising directly from aerial and substratal mycelium, or from ropes formed by mycelium, or radiating out from coils formed by mycelium, unbranched or repeatedly basitonously, verticillately or asymmetrically branched, bearing 1–10 levels with 1–4 phialides per node, proliferating sympodially percurrently in some species, 1–3(–10)-septate at apex, middle and base, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Conidiogenous cells monoor polyphialidic, lateral or terminal, awl-shaped, subulate, (sub-) cylindrical or acicular, hyaline, thinor thick-, smooth-walled, with distinct or inconspicuous collarette and periclinal thickening at conidiogenous loci, with percurrent or subterminal proliferations in some species; chromophilic on the basal septum in some species; polyphialides with two conidiogenous loci occasionally present in Fig. 45. Proliferophialis apiculata (ex-type culture CBS 303.64). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial chains. E–I. Branched or unbranched conidiophores and phialides with repeatedly apical percurrent proliferation (arrows). J. Conidia. Scale bars = 10 μm.
122 Hou et al. some species. Conidia aseptate, fusiform, short fusoid, cylindrical, bacilliform, ovoid, obovoid, tear-shaped, pyriform, ellipsoidal, (sub-) globose, occasionally shape and size vary between those borne internally and peripherally of conidial heads in some species, with or without hilum, truncate, apiculate or rounded at apices or bases, or both ends, thinor thick-, smooth-walled, hyaline or slightly pigmented, eguttulate or guttulate, arranged in slimy heads or in long dry chains, or arranged in long chains, soon collapsing into slimy heads. Crystals present in some species, elongated. Chlamydospores present in some species, intercalary or terminal, mostly in single chains, subglobose, hyaline, becoming pale brown with age, smooth-, thick-walled. Sexual morph not observed. Type: Acremonium alternatum Link Other accepted species with available sequences: Acremonium aerium L.W. Hou, L. Cai & Crous, A. acutatum W. Gams, A. brachypenium W. Gams, A. brunneisporum L.W. Hou, L. Cai & Crous, A. charticola (Lindau) W. Gams, A. chlamydosporium L.W. Hou, L. Cai & Crous, A. egyptiacum (J.F.H. Beyma) W. Gams, A. ellipsoideum L.W. Hou, Rämä, L. Cai & Crous, A. longiphialidicum L.W. Hou, L. Cai & Crous, A. gamsianum L.W. Hou, L. Cai & Crous, A. multiramosum L.W. Hou, Rämä, L. Cai & Crous, A. mycoparasiticum L.W. Hou, L. Cai & Crous, A. psychrophilum C. Möller & W. Gams, A. purpurascens (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous, A. sordidulum W. Gams & D. Hawksw., A. subulatum L.W. Hou, L. Cai & Crous, A. stroudii K. Fletcher, F.C. Küpper & P. van West, A. synnematoferum L.W. Hou, Rämä, L. Cai & Crous Acremonium acutatum W. Gams, Trans. Brit. Mycol. Soc. 64: 394. 1975. Synonym: Cephalosporium acremonium var. cereum Sukapure & Thirum., Sydowia 19: 173. 1966 (1965). Description and illustration: Gams (1975). Typus: India, Varanasi, mycoparasite on Cercospora atromarginalis (Mycosphaerellaceae) on Solanum nigrum, Jun. 1971, M.S. Pavgi, strain 3, dep. W. Gams, CBS H-6634 (holotype of Acremonium acutatum CBS 682.71 preserved as metabolically inactive culture, isotype IMI 185374 = ATCC 32209, ex-type culture CBS 682.71). Additional materials examined: India, Koyna valley, from soil, 21 May 1958, unknown collector, isol. 22 Jan. 1960 by R.S. Sukapure & M.J. Thirumalachar, dep. M.J. Thirumalachar, CBS H-24580 (holotype of Cephalosporium acremonium var. cereum HACC 117, ex-type culture CBS 140.62 = ATCC 14610 = IMI 091574 = MUCL 9730). Netherlands, South Holland Province, Rotterdam, from skin scraping of a patient with clinical eczema, unknown collection date, A. Notowicz, CBS H-8093, culture CBS 829.73. Notes: Cephalosporium acremonium var. cereum differs from C. acremonium in topography, texture of the colonies, and measurement of conidia and conidiophores (Sukapure & Thirumalachar 1965). Sequences of the ex-type of C. acremonium var. cereum (CBS 140.62) are identical to the ex-type of Acremonium acutatum (CBS 682.71) and form a fully supported lineage within the Acremonium s. str. clade (Fig. 2). However, their morphology differs in the shape and size of conidia: Acremonium acutatum produces fusiform conidia with symmetrically elongated and truncated ends, 4–6(–8) × 1.5–2 (–2.5) μm; C. acremonium var. cereum has oblong conidia, 2.7–3.7 × 1.2–1.5 μm (Sukapure & Thirumalachar 1965, Gams 1975). In Fig. 46. Acremonium aerium (ex-type culture CBS 189.70). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D–F. Conidiophores. G. Conidiophores radiating out from coils formed by the mycelia. H. Conidia. Scale bars = 10 μm.
123www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales addition, hyaline chlamydospores were observed in A. acutatum, while absent in C. acremonium var. cereum. These might result from the different media used for morphological observation. Acremonium aerium L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845867. Fig. 46. Etymology: Referring to the air sample, from which the type strain was collected. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, 1.1–2.2 μm wide. Conidiophores solitary, erect, straight, arising directly from aerial and substratal mycelium, rarely radiating out from coils formed by mycelium, usually reduced to single phialides, unbranched, straight, up to 38.5 μm long, 1.3–2 μm wide at base, hyaline, smooth-walled, often with single septum, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral or terminal, subulate, short, hyaline, thick-, smooth-walled, (7.3–)14.7–35.5 μm long, 1.3–1.8 μm wide at base, with inconspicuous periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, thin-, smooth-walled, hyaline, shape and size vary between those borne internally and peripherally of conidial heads: internal conidia (sub-)globose or ellipsoid, 3.5–5.6 × 2–3 μm; peripheral conidia oblong or cylindrical, 3–4.5 × 1.5–2.3 μm. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 21 mm diam, flat, membranous with sparse aerial mycelium, white, undulate margin, reverse buff; On MEA reaching 18–19 mm diam, flat, radially folded, rugose, felty, rosy buff, margin crenate, reverse saffron, with creamy white radial lines; On PDA reaching 30 mm diam, flat, radially folded, membranous, dirty white, margin entire, reverse creamy white, with buff radial lines. Typus: Morocco, Rabat, from air, unknown collection date and collector, isol. M. Chabert, No. 43-G, dep. J. Nicot (holotype CBS H-8331, ex-type culture CBS 189.70 = LCP 2130). Additional material examined: Netherlands, North Holland Province, Hilversum, from human toenail, unknown collection date and collector, isol. 1969 by J. Hoogendoorn, CBS H-8333, culture CBS 379.70C. Notes: Acremonium aerium is fully supported in the phylogenetic analysis (BPP/MLBS 1/100 %) and closely related with A. purpurascens (Fig. 2, BPP/MLBS 1/100 %). It differs from A. purpurascens in producing straight conidiophores, while the later occasionally producing curved conidiophores. Acremonium aerium and A. purpurascens are similar by producing variably shaped and sized conidia that differ within the same conidial head, however, the internal conidia of the conidial heads of A. aerium are longer than that of A. purpurascens (3.5–5.6 μm vs 2.7–3.6 μm), while conidia at the periphery of the heads of A. aerium are smaller than that of A. purpurascens [3–4.5 μm vs (3.2–)3.8–8.3 μm]. Acremonium alternatum Link, Mag. Gesell. Naturf. Freunde, Berlin 3(1–2): 15. 1809. Fig. 47. Descriptions: Link (1809), Gams (1971). Typus: Austria, Innsbruck, Stangensteig, from Hypoxylon deustum (Hypoxylaceae), Dec. 1965, isol. 22 Dec. 1965, coll. and isol. W. Gams, No. C 584 (epitype CBS H-20525, ex-epitype culture CBS 407.66; Summerbell et al. (2011)). Notes: Gams (1968) designated Acremonium alternatum as the lectotype of Acremonium and chose four strains as representative cultures of this species. Summerbell et al. (2011) designated CBS 407.66 with a dried culture as the ex-epitype for A. alternatum. Our present study confirms the results of Summerbell et al. (2011) that the culture CBS 407.66 groups with A. charticola, A. sclerotigenum and A. sordidulum, based on which the genus Acremonium was restricted in Bionectriaceae (Fig. 2). Acremonium brunneisporum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845868. Fig. 48. Etymology: Referring to the brown conidia produced by this fungus. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, 1–2 μm wide. Conidiophores solitary, erect, straight or curved, arising directly from aerial and substratal mycelium, unbranched, poorly branched at lower part, with up two septa at base, some with apical percurrent proliferation, slightly hackly, 27.5–61.5 μm long, 1.1–2.5 μm wide at base, hyaline, smooth-walled. Phialides mostly lateral, subcylindrical or acicular, hyaline, thick-, smoothwalled, 23–40 μm long, 1.5–2 μm wide at base, with inconspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, fusoid, with slightly thicken and truncate hilar spot at both apices and bases, thick-, smooth-walled, hyaline at first, becoming pale brown with age, 4–5.5 × 1.3–2 μm, eguttulate, arranged in long chains, soon collapsing into slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 25–26 mm diam, flat, felty, greenish olivaceous at centre, white at periphery, margin entire, reverse creamy white; On MEA reaching 24–25 mm diam, flat, radially folded, felty or hairy, with ropes of mycelial strands, dirty white, margin entire, reverse pale ochreous, with buff radial lines; On PDA reaching 33–35 mm diam, flat, felty, dusty at periphery, dark brick at centre, rosy buff at periphery, margin entire, reverse dark olivaceous at centre, pale umber to buff at periphery; On SNA reaching 27–28 mm diam, flat, felty, white, margin entire, reverse concolourous. Typus: India, Varanasi, on Colletotrichum dematium (Glomerellaceae) on pod of Albizzia lebbek (Leguminosae), unknown collector and collection date, dep. U.P. Singh (holotype CBS H-24591, ex-type culture CBS 413.76). Additional material examined: Russia, Moscow, from plaster, 13 Nov. 2013, V. Ponizovskaya, N. Rebrikova, CBS 142823. Notes: In our phylogenetical analysis, Acremonium brunneisporum grouped with A. gamsianum and A. stroudi. Morphologically, A. brunneisporum differs by producing fusoid, pale brown conidia arranged in long dry chains, while A. gamsianum and A. stroudi produce globose to oblong or cylindrical, hyaline conidia arranged in slimy heads (Fletcher et al. 2017). In addition, A. brunneisporum can be distinguished from the last two species by its conidia having distinct hilar spots at both ends. Acremonium charticola (Lindau) W. Gams, Cephalosporiumartige Schimmelpilze (Stuttgart): 46. 1971. Fig. 49. Basionym: Cephalosporium charticola Lindau, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.8: 107. 1904 (1907). Synonym: Cephalosporium malorum Kidd & Beaumont, Trans. Brit. Mycol. Soc. 10: 110. 1924.
124 Hou et al. Description: Kidd & Beaumont (1924) (based on CBS 117.25); Gams (1971). Materials examined: Germany, Kiel-Kitzeberg, from cellar wall, unknown collection date and collector, isol. Feb. 1966 by W. Gams, No. 610, culture CBS 402.66. UK, England, from rotten apple (Malus sp.), unknown collection date and collector, isol. M.N. Kidd, CBS H-24584 (ex-type culture of Cephalosporium malorum CBS 117.25 = MUCL 9710). Unknown, unknown host, collection date and collector, isol. J. Lacey, (171)1403, CBS H-24585, culture CBS 547.86. Notes: Acremonium charticola was originally described as Cephalosporium charticola from damp wallpaper by Lindau (1904). Gams (1971) examined the type specimen from the S herbarium, as well as several cultures from diverse substrates, and concluded that this species is generally distributed on mouldy wallpaper and damp cellar walls. Acremonium charticola is characterised by the multiply branched conidiophores, relatively short conidia and crystal formation (Gams 1971), which could be differentiated from its closely related species A. sordidulum and A. subulatum on the phylogenetic tree (Fig. 2). Acremonium sordidulum and A. subulatum produces unbranched or poorly branched conidiophores and lack of crystals (Gams 1975). Cephalosporium malorum together with C. ballagii were synonymised under A. charticola based on morphological characters (Gams 1971). Our multi-locus phylogenetic analysis agrees with the results of Gams (1971), showing that the ex-type culture of C. malorum is genetically identical to cultures of A. charticola (Fig. 2). However, phylogenetic analysis locates the ex-type culture of C. ballagii in Niessliaceae (Fig. 1), distant from Acremonium s. str., as demonstrated previously by Summerbell et al. (2011). Therefore, C. ballagii is resurrected as a separate species treated under Niessliaceae in future studies. Fig. 47. Acremonium alternatum (ex-type culture CBS 407.66). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
125www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Acremonium chlamydosporium L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845869. Fig. 50. Etymology: Referring to the chlamydospores produced by this species. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, occasionally chondroid, 1.8–2.5 μm wide. Conidiophores solitary, erect, straight, lateral, arising directly from aerial and substratal mycelium, unbranched or basitonously branched, bearing 1–3 levels with 1–2(–3) phialides per node, 10–107.5 μm long, 1.4–2.2 μm wide at base, 1–4-septate in lower and upper part, hyaline, smooth-walled. Phialides lateral or terminal, aciculate, cylindrical, hyaline, thick-, smooth-walled, 10.8–69(–84.5) μm long, 1.2–2.2 μm wide at base, with inconspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, cylindrical with inconspicuous truncate bases and obtuse apices, thin-, smooth-walled, hyaline, 3.3–5.8 × 1.5–2.4 μm, eguttulate, arranged in slimy heads. Chlamydospores intercalary or terminal, mostly in single chains, subglobose, hyaline, becoming pale brown with age, smooth-, thick-walled, 2.9–6 × 3–4.7 μm. Sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 20–21 mm diam, flat, dusty, white, margin entire, reverse dirty white; On MEA reaching 20–22 mm diam, radially folded, flat, membranous, rosy buff, margin entire, reverse saffron, with buff radial lines; On PDA reaching 26–27 mm diam, flat, radially folded at centre, thinly felty, creamy white, margin fimbriate, reverse creamy white, with buff radial lines; On SNA reaching 18–20 mm diam, flat, membranous without aerial mycelium, white, margin entire, reverse colourless. Fig. 48. Acremonium brunneisporum (ex-type culture CBS 413.76). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidiophores with conidial heads and chains. F, G. Unbranched conidiophores. H. Conidiophore with apical percurrent proliferation. I. Conidia. Scale bars = 10 μm.
126 Hou et al. Typus: India, Uttar Pradesh, Varanasi, on Colletotrichum dematium (Glomerellaceae) on pods of Albizia lebbek (Fabaceae), unknown collection date and collector, dep. U.P. Singh (holotype CBS H-24592, ex-type culture CBS 414.76). Notes: Both Acremonium chlamydosporium and its closely related species A. acutatum are facultative or obligate mycoparasitic fungi. Acremonium chlamydosporium is morphologically similar to A. acutatum in the shape of conidiophores and the size of conidia, but it could be distinguished by its multi-septate conidiophores (up to 4-septate vs 1-septate), longer phialides [10.8–69(–84.5) μm vs 20–40 μm], different shape of conidia (cylindrical with truncate bases and obtuse apices vs fusiform with more or less symmetrically elongated and minutely truncated ends) (Gams 1975). Acremonium egyptiacum (J.F.H. Beyma) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 64. 1971. Fig. 51. Basionym: Oospora egyptiaca J.F.H. Beyma, Centbl. Bakt. ParasitKde, Abt. II 89: 243. 1933. Synonyms: Cephalosporium sclerotigenum Moreau & R. Moreau ex Valenta, Acta Acad. Sci. Nat. Morav.-Siles. 20: 4. 1948. Acremonium sclerotigenum (Moreau & R. Moreau ex Valenta) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 45. 1971. Description and illustration: Gams (1971). Typus: Egypt, Cairo, from soil, unknown collection date and collector, isol. Sabet, G5, CBS H-24583, ex-type culture of Oospora egyptiaca CBS 114785 = IFO 4607 = NBRC 4607. Materials examined: Canada, Ontario, Toronto, from soft contact lens from inflamed human eye, unknown collection date and collector, isol. 1989 by Fig. 49. Acremonium charticola (culture CBS 117.25). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidial heads. E–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
127www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales R.C. Summerbell, culture CBS 109041. France, Dunkerque, fishmeal, unknown collection date and collector, isol. Aug. 1964 by G.L. Hennebert, culture CBS 740.69 = MUCL 6966; Nancy, from human toenail, unknown collection date and collector, isol. E. Kiffer, culture CBS 270.86; Pointe de Chemoulin, from dune sand, under Ammophila and Convolvulus sp. (Convolvulaceae), unknown collection date and collector, dep. F. Moreau, CBS H-24762 (ex-type culture of Cephalosporium sclerotigenum CBS 124.42 = ATCC 22616 = IAM 14662); Torbali, Izmir, from root of Lycopersicon esculentum (Solanaceae), unknown collection date and collector, isol. I. Karaca, No. 106B, CBS H-8158, culture CBS 734.69. Germany, former DDR, unknown substrate, collection date and collector, isol. G. Sörgel, No. 4081, CBS H-8377, culture CBS 286.70B; former West-Germany, Hamburg, from human toenail, unknown collection date and collector, isol. H. Listemann, culture CBS 391.89; Kiel-Kitzeberg, from mouldy cellar wall, unknown collection date and collector, isol. 4 Nov. 1965 by W. Gams, No. 568, CBS H-24581, culture CBS 403.66. Greece, Thessaloniki, from human peritoneal fluid, Dec. 2002, E. Bibashi, culture CBS 112783. Iran, Sarab, from leaf of Hordeum vulgare (Poaceae), unknown collection date, B. Askari, culture CBS 114321; unknown substrate, collection date and collector, dep. R. Zare, Department of Botany, Plant Pests & Diseases Research Institute (Tehran, Iran), culture CBS 113719. Netherlands, North Holland Province, Amsterdam, from bark of apple and pear trees (Rosaceae), unknown collection date and collector, isol. J. Scheepens, culture CBS 500.82. Syria, from grape vines, unknown collection date and collector, dep. K.A. Halim, No. Syrie 1, culture CBS 121405. USA, unknown substrate, collection date and collector, isol. Aug. 1960 by M.A. Pisano, No. A-29c, CBS H-24582, culture CBS 159.61. UK, Bristol, from human toenail, isol. 9 Oct. 1972 by M.P. English, CBS H-8389, culture CBS 629.73 = IMI 171213. Notes: Acremonium sclerotigenum and A. egyptiacum were frequently reported to be associated with food contamination and human disease. Acremonium sclerotigenum was synonymised to A. egyptiacum by Summerbell et al. (2018) based on the examination of 72 isolates on morphological characters and Fig. 50. Acremonium chlamydosporium (ex-type culture CBS 414.76). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores with conidial heads. E. Conidial heads. F–I. Conidiophores. J, K. Chlamydospores. L. Conidia. Scale bars = 10 μm.
134 Hou et al. Notes: Based on the phylogenetic analyses, Acremonium mycoparasiticum is represented by two strains from fungus and mites, respectively, forming a distinct clade related to A. alternatum, A. ellipsoideum and A. psychrophilum (Fig. 2). Morphologically, A. mycoparasiticum can be differentiated from all these species in its tear-shaped or obovoid conidia with one end slightly elongated and apiculate (2.7–5.3 × 1.6–2.3 μm) , arranged in dry conidial heads. The conidia of A. alternatum are elongated obovate, rounded at the apex, clearly apiculate at the base (4.3–5 × 1.9–2.1 μm) and arranged in chains (Gams 1971), those of A. ellipsoideum are subglobose to broadly ellipsoid or cylindrical (2.5–4.1 × 1.8–2.4 μm), and the conidia of A. psychrophilum cylindrical to ellipsoid (5–13.5 × 2–2.5 μm; Möller & Gams 1993). Acremonium psychrophilum C. Möller & W. Gams, Mycotaxon 48: 445. 1993. Description and illustration: Möller & Gams (1993). Material examined: Antarctica, King George Island, slope near Argentinian station Jubany (current name Carlini Base), from Turgidosculum complicatum (lichen, Verrucariaceae), Dec. 1991, C. Möller, No. 6/22, CBS H-5321, culture CBS 139.93. Notes: Acremonium psychrophilum isolated from the lichen Turgidosculum complicatum in Antarctica differs from other known lichen-inhabiting Acremonium species (including A. antarcticum, A. lichenicola, A. rhabdosporum, A. spegazzinii) based on morphological characteristics (Möller & Gams 1993). Acremonium psychrophilum was originally classified in Acremonium section Gliomastix because of the production of chondroid hyphae and thick-walled phialides (Möller & Gams 1993). Although the ex-type culture, CBS 732.92, is unavailable to us, the strain CBS 139.93 was examined, which had the same host, location and was identified by the original authors as Fig. 56. Acremonium mycoparasiticum (ex-type culture CBS 188.80). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Mycelial ropes with conidiophores and conidial heads. E–H. Conidiophores. I. Conidia. Scale bars = 10 μm.
135www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales this species (Möller & Gams 1993). It was recognised as a possible ex-type culture of A. psychrophilum based on the information recorded in the CBS culture collection. Phylogenetically, the strain CBS 139.93 clusters within the genus Acremonium, and is closely related to the type, A. alternatum (Fig. 2). Acremonium purpurascens (Sukapure & Thirum.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845878. Fig. 57. Basionym: Cephalosporium purpurascens Sukapure & Thirum., Bull. Torrey Bot. Club. 93: 310. 1966. Illustration: Sukapure & Thirumalachar (1966). Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, 1.2–1.6(–2.5) μm wide. Conidiophores solitary or aggregate, erect, straight or waved, arising directly from aerial and substratal mycelium, usually reduced to single phialides, unbranched, poorly branched, up to 40 μm long, 1.3–2.1 μm wide at base, hyaline, smooth-walled , 1-septate at base, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral or terminal, subulate, hyaline, thin-, smooth-walled, 12–31 μm long, 1.2–2 μm wide at base, with inconspicuous periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, thin-, smooth-walled, eguttulate, hyaline, arranged in slimy heads, shape and size vary between those borne internally and peripherally of conidial heads: internal conidia (sub-)globose or ellipsoid, 2.7–3.6 × 2.5–3.2 μm; peripheral conidia are cylindrical, (3.2–)3.8–8.3 × 1.7–2.4 μm; conidial heads confluent in old cultures. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 22 mm diam, flat, dusty or membranous with sparse aerial Fig. 57. Acremonium purpurascens (ex-type culture CBS 149.62). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D–H. Conidiophores. I. Conidia. Scale bars = 10 μm.
136 Hou et al. mycelium, white, pale salmon at centre, margin entire, reverse concolourous; On MEA reaching 27–29 mm diam, flat, radially and deeply folded, hairy and buff at centre, felty and rosy buff at periphery, margin slightly crenate, reverse pale ochreous, with buff radial lines at periphery; On PDA reaching 35–36 mm diam, flat, radially folded at centre, dusty, rosy buff at centre, dirty white towards periphery, margin fimbriate, reverse creamy white. Typus: India, Koyna Valley, Chiplun, from soil, unknown collection date and collector, isol. 1 Aug. 1957 by R.S. Sukapure & M.J. Thirumalachar, dep. W. Gams, CBS H-8273 (holotype HACC 104, ex-type culture CBS 149.62 = ATCC 14611 = IMI 091575 = MUCL 9709 = HACC 104). Additional material examined: Unknown, from human skin, unknown collection date and collector, isol. 1966 by Vink, No. 354, CBS H-8327, culture CBS 780.69. Notes: Sukapure & Thirumalachar (1966) described Cephalosporium purpurascens from soil in India. This species was once synonymised with Acremonium persicinum (Gams 1971). We re-examined the ex-type culture of C. purpurascens (CBS 149.62) and concluded that the size of the structures observed in this study differs slightly from those described in the protologue (Sukapure & Thirumalachar 1966). The conidiophores are slightly larger than those recorded in the protologue (up to 40 μm vs 15–25 μm). In addition, the shape and size of conidia differ among conidia held internally in the conidial heads and those held on the periphery. The conidia found internally are (sub-)globose, consistent with the original description; the external conidia are cylindrical, a finding not described in the protologue. According to the multi-locus phylogenetic analysis, the ex-type culture of C. purpurascens clustered within Acremonium s. str. (Fig. 2), distant from A. persicinum (currently Verruciconidia persicina). Thus, C. purpurascens is transferred to Acremonium as A. purpurascens. It differs from the closest species A. brachypenius in its conidia with variable shape and size internal and peripheral of the conidial heads. Morphologically differences of A. purpurascens and A. aerium were discussed under A. aerium. Acremonium sordidulum W. Gams & D. Hawksw., Trans. Brit. Mycol. Soc. 64: 392. 1975. Description and illustration: Gams (1975). Typus: India, Tamil Nadu, from old stem of Euphorbia tirukalli, Jan. 1973, W. Gams (holotype CBS H-6669, ex-type living culture CBS 385.73 = ATCC 32186 = IMI 185373). Additional material examined: Brazil, Fortaleza, from Ravenelia sp. (Raveneliaceae) on Stryphnodendron coriaceum (Fabaceae), 1999, F. Freire, culture CBS 102413. Note: Acremonium sordidulum has a strong resemblance to A. egyptiacum, but differs in the production of shorter phialides and grey pigmentation (Gams 1975). Acremonium stroudii K. Fletcher, F.C. Küpper & P. van West, sp. nov. MycoBank MB 848120. Etymology: Named after the collector (Stedson Stroud) of the initial field material. Description and illustration: Fletcher et al. (2017). Typus: UK, Ascension Island, Whale Point, from environmental sample taken from a seawater blow hole, 31 Aug. 2012, S. Stroud & J. Sim (holotype CBS 138820 preserved as metabolically inactive culture, extype culture CBS 138820). Notes: Acremonium stroudii was invalidly described (Fletcher et al. 2017), because the citation of the identifier issued for the name was not included, and the type of the name was not indicated in the protologue (International Code of Nomenclature for algae, fungi, and plants ShenZhen, Art. F.5.1. and Art. 40.1). The taxon is thus validated here. Acremonium subulatum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845879. Fig. 58. Etymology: Referring to the subulate shape of its phialides. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1–1.5 μm wide, mycelial ropes present. Sporulation abundant, phalacrogenous, nematogenous, plectonematogenous. Conidiophores solitary, erect, straight or curved, arising directly from aerial and substratal mycelium, or from ropes formed by mycelium, usually reduced to single phialides, unbranched or poorly basitonously branched, up to 37.5 μm long, 1.2–1.8 μm wide at base, hyaline, smooth-walled, 1-septate at base, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral or terminal, subulate, hyaline, thick-, smooth-walled, 13.7–35.5 μm long, 1.3–1.9 μm wide at base, with inconspicuous periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, thin-, smoothor rough-walled, short fusoid, with thick and truncate hila at apices and bases, or with rounded apices and apiculate bases, hyaline, 2.8–5.3 × 1.5–2.3 μm, arranged in long chains, irregularly collapsing into heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 23–25 mm diam, flat, dusty, salmon at centre, white at periphery, margin entire, reverse buff; On MEA reaching 15–18 mm diam, flat, radially folded, rugose, felty, salmon, creamy white at periphery, margin crenate, reverse pale ochreous, with buff radial lines; On PDA reaching 23–24 mm diam, flat, radially folded and hairy at centre, membranous at periphery, white, margin entire, reverse buff; On SNA reaching 20–22 mm, flat, dusty, white, margin entire, reverse white. Typus: Sri Lanka, Anuradhapura, from fruit of Annona reticulata (Annonaceae), unknown collection date, isol. Jan. 1973, coll. and isol. W. Gams (holotype CBS H-24587, ex-type culture CBS 588.73A). Additional material examined: South Africa, Western Cape Province, Paarl, Zandrift, from Vitis vinifera (Vitaceae), unknown collection date, isol. L. Mostert, LM 89, CBS H-24586, culture CBS 115996 = CPC 5434. Notes: Strain CBS 588.73A was initially received as ‘A. sordidulum’ based on morphological characters (Gams 1975). However, it formed an independent branch together with strain CBS 115996. This branch was found to be distinct from that bearing the ex-type culture of A. sordidulum (Fig. 2). Morphologically, A. subulatum differs from the closely related A. sordidulum by its colony that is salmon on OA and MEA, and whitish at PDA after two weeks. The colony of A. sordidulum is white to pale greenish grey. Although culture CBS 115996 has 33 bp (ITS 4 bp, LSU 3 bp, rpb2 10 bp, tef-1α 16 bp) differences with CBS 588.73A, the morphological characters of the cultures show no significant differences and the two are thus treated as conspecific.
137www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales Acremonium synnematoferum L.W. Hou, Rämä, L. Cai & Crous, sp. nov. MycoBank MB 845880. Fig. 59. Etymology: Referring to the conidiophores arranged in synnemata. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.1–2 μm wide, abundant mycelial ropes formed. Sporulation abundant, phalacrogenous, nematogenous, plectonematogenous, synnematogenous. Conidiophores solitary or aggregate, (sub-)erect, straight or slightly bended, lateral or terminal, arising directly from aerial and substratal mycelium, or from ropes formed by the mycelium, unbranched, poorly branched, bearing up to two phialides per node, forming synnemata-like ropes, 27–68.6(–99) μm long, 1.1–2.5 μm wide at base, 1–4-septate at base, hyaline, smooth-walled. Phialides mostly lateral, acicular, hyaline, thick-, smooth-walled, 20–51 μm long, 1.1–2 μm wide at base, with conspicuous cylindrical collarette and periclinal thickening at conidiogenous loci; polyphialides not observed. Conidia aseptate, oblong, apices rounded, with a slightly truncate hilum at bases, thin-, smooth-walled, hyaline, 2.8–5.8 × 1.6–2.3 μm, eguttulate, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 15 mm diam, flat, with moderate aerial mycelium, felty, short hairy, white, creamy white or salmon at centre, margin entire, reverse creamy white; On MEA reaching 14–16 mm diam, raised, radially folded, with moderate aerial mycelium, felty, rosy buff, margin crenate, reverse pale apricot at centre, saffron at periphery, with buff radial lines; On PDA reaching 17–19 mm diam, flat, with moderate aerial mycelium, felty, buff, margin entire, reverse buff; On SNA reaching 15–17 mm diam, flat, membranous without aerial mycelium, white, margin entire, reverse concolourous. Without odour on all media. Fig. 58. Acremonium subulatum (ex-type culture CBS 588.73A). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores with conidial heads and conidial chains. E–H. Conidiophores. I. Conidia. Scale bars = 10 μm.
138 Hou et al. Typus: Norway, Troms County, Vadsø municipality, Varangerfjorden, Ekkerøy, from construction wood in tidal zone, 10 Sep. 2010, T. Rämä (holotype CBS H-24727, CBS 147431 = CPC 40324 = TR090cU1.1). Additional materials examined: Norway, Troms County, Berlevåg municipality, Tanafjorden, Store Molvik, from driftwood (Pinus sp., Pinaceae) in tidal zone, 6 Sep. 2010, T. Rämä, culture TR070aS1.2; Troms County, Berlevåg municipality, Sandfjorden, from driftwood (Larix sp., Pinaceae) in tidal zone, 7 Sep. 2010, T. Rämä, culture CBS 147430 = CPC 40330 = TR076aD1.1; Troms County, Båtsfjord municipality, Ytre Syltevika, from driftwood (Pices sp.) in splash zone, 8 Sep. 2010, T. Rämä, culture CPC 40326 = TR079cD1.1; Troms County, Berlevåg municipality, Sandfjorden, from driftwood (Larix sp., Pinaceae) in tidal zone, 7 Sep. 2010, T. Rämä, culture CBS 147432 = CPC 40325 = TR076cU1.1. Notes: Acremonium synnematoferum is represented by five cultures from intertidal wood in Norway that were isolated as explained in Rämä et al. (2014). According to the phylogenetic inference, they fall into a separate clade and differ from other known species of Acremonium and from the other two novel species described from this location and habitat. Morphologically, sporulation of A. synnematoferum is abundant and synnematogenous, forming synnemata-like ropes, which is rare in Acremonium. Clade O32 Waltergamsia L.W. Hou, L. Cai & Crous, gen. nov. MycoBank MB 845881. Etymology: Named after Dr Walter Gams, in acknowledgement for the tremendous contribution that he made to the taxonomy of acremonium-like fungi. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, chondroid hyphae present. Conidiophores solitary Fig. 59. Acremonium synnematoferum (ex-type culture CBS 147431). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Mycelial ropes and conidiophores. F–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
139www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales or aggregated, erect or flexuous, arising directly from submerged or superficial hyphae, sometimes radiating out from sterile coils or ropes formed by mycelium, unbranched or poorly branched, septate, hyaline, smooth-walled. Phialides lateral or terminal, subulate, acicular, ampulliform, or sub-cylindrical, occasionally widened and flat at lower part, hyaline, thinor thick-, smoothwalled, with inconspicuous or conspicuous periclinal thickening at conidiogenous loci, percurrently proliferating phialides can be present; polyphialides present in some species. Conidia aseptate, pyriform, obpyriform, clavate, lanceolate, (narrowly) ovoid, obovoid, fusoid, ellipsoid, cylindrical, or sub-globose, hyaline or subhyaline, weakly pigmented, thinor thick-, smooth-walled, with apiculate or obtuse end(s), guttulate or eguttulate, arranged in slimy heads or dry chains, or arranged in chains at first, later collapsing soon as conidial heads. Chlamydospores and sexual morph unknown. Type: Waltergamsia fusidioides (Nicot) L.W. Hou, L. Cai & Crous Other accepted species with available sequences: Waltergamsia alkalina L.W. Hou, L. Cai & Crous, W. catenata L.W. Hou, L. Cai & Crous, W. citrina (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. dimorphospora (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. epimycota (Samuels) L.W. Hou, L. Cai & Crous, W. hennebertii (W. Gams) L.W. Hou, L. Cai & Crous, W. moroccensis L.W. Hou, L. Cai & Crous, W. obpyriformis L.W. Hou, L. Cai & Crous, W. parva (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. pilosa (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, W. zeylanica (Petch) L.W. Hou, L. Cai & Crous Notes: Waltergamsia is proposed to accommodate eight Acremonium s. lat. species that form a fully supported clade. Morphologically, most species in this genus produce conidia arranged in slimy heads or in chains at first but soon collapsing Fig. 60. Waltergamsia alkalina (ex-type culture CBS 741.94). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores and conidial heads. E, F. Conidiophores radiating from coils formed by mycelium. G–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
140 Hou et al. to form conidial heads. However, W. pilosa and W. zeylanica were only recorded to have conidia arranged in chains. Waltergamsia alkalina L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845882. Fig. 60. Etymology: Referring to the substrate from which the holotype culture was collected. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.7–2.4 μm wide. Conidiophores solitary or aggregated, (sub-)erect, straight or flexuous, arising directly from submerged or superficial hyphae, sometimes radiating from sterile coils formed by mycelium, usually reduced to single phialides, unbranched, poorly branched, bearing up to two phialides per node, 21.5–39.5 long, 1.5–2.5 μm wide at base, 1–2-septate, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal, lateral, subulate, hyaline, thin-, smooth-walled, 13–32.8 μm long, 1.3–2.5 μm wide at base, with inconspicuous periclinal thickening and cylindrical collarette at conidiogenous loci, occasionally with a percurrent proliferation; polyphialides not observed. Conidia aseptate, cylindrical, symmetrically rounded, hyaline, thin-, smooth-walled, 3–5.4 × 1.2–1.8 μm, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at ca. 25 °C: Colonies on OA reaching 30 mm diam, flat, felty to dusty, white, margin entire, reverse creamy white; On MEA reaching 28 mm diam, flat, felty, white, margin entire, reverse umber at centre, pale apricot at periphery; On PDA reaching 25 mm diam, flat, dusty, white at centre, hazel at periphery, margin entire, reverse greenish glaucous at centre, olivaceous buff at periphery. Typus: Indonesia, from alkaline soil, unknown collection date and collector, isol. K. Nagai, Drug Serendipity Research laboratories, Yamanouche Pharmaceutical Co., Tokyo, Japan, No. 577 (holotype CBS H-24603, extype culture CBS 741.94). Notes: Waltergamsia alkalina is represented by an isolate from alkaline soil collected in Indonesia. According to phylogenetic inference from the ITS, LSU, rpb2, and tef-1α loci (Fig. 2), W. alkaline occupies a single long branch that is clearly different from other species in Waltergamsia. Morphologically, W. alkalina differs from its closely related species, W. citrina, by producing cylindrical conidia with both ends rounded, arranged in slimy heads, while conidia of W. citrina are obovoid or ellipsoid, arranged in long chains (Giraldo et al. 2014). Waltergamsia catenata L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 848119. Fig. 61. Etymology: Referring to the conidial chains produced by this species. Mycelium consisting of branched, septate, hyaline, smooth-, thinwalled hyphae, 1.2–1.9 μm wide. Conidiophores solitary, (sub-)erect, straight, arising directly from submerged or superficial hyphae, usually reduced to single phialides, unbranched, poorly branched, bearing up to two phialides per node, up to 51 μm long, 1.6–2.8 μm wide at base, hyaline, smooth-walled, 1–2-septate at base, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal, lateral, subulate or cylindrical, hyaline, thin-, smooth-walled, 21–39 μm long, Fig. 61. Waltergamsia catenata (ex-type culture CBS 102462). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Monophialidic conidiophores. F. Branched conidiophore. G. Proliferating conidiophore. H. Conidia. Scale bars = 10 μm.
141www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales 1.5–2.5 μm wide at base, with inconspicuous periclinal thickening and conspicuous cylindrical collarette at conidiogenous loci, occasionally with a percurrent proliferation; polyphialides not observed. Conidia aseptate, ovoid or fusoid, with truncate base at both ends, hyaline, thin-, smooth-walled, 3.5–6.5 × 1.7–2.8 μm, eguttulate, arranged long conidial chains. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 15–17 mm diam, flat, dusty, white, margin entire, reverse white; On MEA reaching 16–17 mm diam, flat, felty, rosy buff at centre, dirty white at periphery, margin filiform, reverse pale orange; On PDA reaching 16–18 mm diam, flat, radially rugose, felty, rosy buff, margin dendritic, reverse concolourous; On SNA reaching 6 mm diam, flat, felty, white, margin dendritic, reverse concolourous. Typus: Netherlands, Horst, Forestry Dept., Frimochalk, unknown date, F.P. Geels (holotype CBS H-24605, ex-type culture CBS 102462). Notes: Waltergamsia catenata formed a distinct branch, being distant from other species in Waltergamsia. Waltergamsia catenata is morphologically different from W. moroccensis and W. zeylanica in its ovoid or fusoid conidia arranged in long chains, while that of W. moroccensis are ellipsoid arranged in slimy heads (Fig. 63), and W. zeylanica produces lanceolate or narrow-ovoid conidia arranged in chains (Fig. 65). Waltergamsia citrina (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845883. Basionym: Acremonium citrinum A. Giraldo et al., Mycologia 106: 334. 2014. Description & illustration: Giraldo et al. (2014). Typus: Papua New Guinea, Madang, Jais Aben, from decaying fruit along the coast, Nov. 1995, A. Aptroot, isol. Nov. 1995 by A. van Iperen, No. A 145 (holotype CBS H-21330, ex-type cultures CBS 384.96 = FMR 11427). Additional material examined: Netherlands, from human sputum, unknown collection date and collector, dep. A. Kikstra, Academic Hospital Groningen, culture CBS 758.69. Notes: This fungus was introduced based on the collection of decaying fruit in Madang, Papua New Guinea (Giraldo et al. 2014). It is morphologically characterised by the production of diffusible light yellow pigment and restricted growth on PDA media in which a diameter of 4–5 mm is attained after 14 d (Giraldo et al. 2014). Based on our phylogeny, the ex-type culture of A. citrinum nestled with full support in the Waltergamsia clade, and therefore the new combination W. citrina is introduced here. Waltergamsia dimorphospora (A. Giraldo et al.) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845884. Basionym: Acremonium dimorphosporum A. Giraldo et al., Mycol. Progr. 16: 356. 2017. Description & illustration: Giraldo et al. (2017). Typus: USA, Texas, from human bronchoalveolar lavage fluid, 15 Oct. 2008, D.A. Sutton (holotype CBS H-22021, ex-type culture CBS 139050 = UTHSC 08-3639 = FMR 10548). Notes: Waltergamsia dimorphospora is characterised by producing dimorphic conidia arranged in slimy heads and is morphologically similar to Bulbithecium borodinense (previously Acremonium borodinense; Ito et al. 2000, Giraldo et al. 2017). However, the extype culture of A. dimorphospora phylogenetically clustered within the genus Waltergamsia, which is distant from Bu. borodinense (Fig. 2). Waltergamsia epimycota (Samuels) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845885. Basionym: Nectriopsis epimycota Samuels, Mem. New York Bot. Gard. 48: 64. 1988. Description & illustration: Samuels (1988). Typus: French Guiana, Vic. Saül, alt. 200 m, on Kretzschmaria sp. (Xylariales), Feb. 1986, G.J. Samuels (holotype Samuels 3463 in NY). Additional materials examined: Germany, former West-Germany, from root of Triticum aestivum (Poaceae), unknown collection date and collector, isol. A. Walz, No. W2111, culture CBS 562.86; near Braunschweig, from agricultural löss soil, unknown collection date and collector, isol. H. Nirenberg, BBA, Inst.f. Mikrobiol., Berlin, No. 308, CBS H-24600, culture CBS 265.89. Puerto Rico, from Pyrenomycete in forest, unknown collection date, G.J. Samuels, G.J.S. 95-94, BPI 737695, culture CBS 127459. Waltergamsia fusidioides (Nicot) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845886. Basionym: Paecilomyces fusidioides Nicot, Cah. Maboké 6: 18. 1968. Synonym: Acremonium fusidioides (Nicot) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 70. 1971. Description & illustration: Gams (1971). Typus: Central African Republic, from dung of antelope, unknown collection date and collector, isol. R. Cailleux, CBS H-24601 (holotype of Paecilomyces fusidioides Culture no. 1964 in Mycothèque L.C. Paris, ex-type culture CBS 840.68 = IAM 14648 = LCP 66.1964). Additional materials examined: France, Toulouse, unknown substrate, unknown collection date, W. Gams, “Bouteille”, No. 50, CBS H-8187, culture CBS 705.86. Italy, Padova, Botanical Garden, from decaying leaf of Canna indica (Cannaceae), Dec. 1968, L. de Zoller, isol. Dec. 1968 by W. Gams, CBS H-8239, culture CBS 113.69. Notes: According to Gams (1971), all cultures of this species were observed to produce dimorphic conidia: i.e., globose and spindleshaped. According to our phylogenetic inference, the ex-type culture of Paecilomyces fusidioides (CBS 840.68) falls in a fully supported clade (BPP/MLBS = 1/100 %) in Waltergamsia, therefore the new combination W. fusidioides is proposed here. Waltergamsia hennebertii (W. Gams) L.W. Hou, L. Cai & Crous, comb. nov. MycoBank MB 845887. Fig. 62. Basionym: Acremonium hennebertii W. Gams, Cephalosporiumartige Schimmelpilze (Stuttgart): 79. 1971. Description & illustration: Gams (1971). Typus: Zaire, Kimuenza, Lovanium University Campus, from leaf rachis of Elaeis guineensis (Arecaceae), 20 Mar. 1968, G.L. Hennebert (holotype CBS H-6614, isotypes CBS H-6615 & CBS H-6616, ex-type culture CBS 768.69 = MUCL 11580 = MUCL 28812). Notes: Waltergamsia hennebertii was originally described as Acremonium hennebertii from a leaf rachis of Elaeis guineensis, which was treated as one of the species of Acremonium section Acremonium (Gams 1971). Based on the multi-locus phylogenetic analysis, the ex-type culture of A. hennebertii (CBS 768.69) clustered in the clade representing the genus Waltergamsia (Fig.
142 Hou et al. 2). Morphologically, W. hennebertii differs from W. alkalina in the production of inverted ovoid conidia, while that of W. alkaline are cylindrical with rounded base at both ends; W. hennebertii differs from W. obpyriformis by the absent of polyphialides (Gams 1971; Fig. 2). Acremonium hennebertii is therefore transferred to Waltergamsia, and a new combination is proposed. Waltergamsia moroccensis L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845888. Fig. 63. Etymology: Referring to Morocco, the country where this fungus was collected. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.3–2.5 μm wide. Conidiophores solitary or aggregated, erect, straight or curved, arising directly from aerial or substratal mycelium, usually reduced to single phialides, unbranched or basitonously branched, bearing 1–2 levels with 2–4 divergent phialides per node, up to 23.5 μm long, 1.3–1.9 μm wide at base, hyaline, smooth-walled, with a single basal septum, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral, subulate or ampulliform, slightly swollen and flat at lower part, narrowed at base, hyaline, thin-, smooth-walled, (3.5–)10.5– 21.5 μm long, 1–2.5(–3.2) μm wide at base, with inconspicuous periclinal thickening at conidiogenous loci; polyphialides with up to four conidiogenous loci occasionally present. Conidia aseptate, ovoid, ellipsoid, both ends rounded, hyaline, thin-, smoothwalled, 2.8–4 × 1.7–2.4 μm, eguttulate, arranged in slimy heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 20–24 mm diam, flat, thinly felty, slightly floccose, rosy buff at centre, white at periphery, margin entire, reverse creamy white; On MEA reaching 18–20 mm diam, raised, radially folded, felty, buff, margin crenate, reverse straw, pale ochreous at centre, with buff radial lines; On PDA reaching 19–20 mm diam, flat, short hairy and dirty white at centre, thinly felty and white at periphery, margin entire, reverse buff. Typus: Morocco, from fragmenting human nail, unknown collection date and collector (holotype CBS H-24602, ex-type culture CBS 512.82). Notes: Based on the multi-gene phylogenetic analysis, Waltergamsia moroccensis forms an independent branch, clearly separated from other species in Waltergamsia (Fig. 2). Morphologically, W. moroccensis differs from its most closely related species, W. obpyriformis, by producing shorter phialides [10.5–21.5 μm vs 16.8–36(–49.5) μm] and conidia with rounded ends, arranged in slimy heads, while W. obpyriformis has conidia with elongated hilum at basal ends, arranged in chains. Waltergamsia obpyriformis L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845889. Fig. 64. Etymology: Referring to the shape of conidia produced by this species. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.3–1.8 μm wide, abundant mycelial ropes and coils present. Sporulation abundant, phalacrogenous, Fig. 62. Waltergamsia hennebertii (ex-type culture CBS 768.69). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D–G. Conidiophores. H. Conidia. Scale bars = 10 μm.
143www.studiesinmycology.org Redisposition of acRemonium-like fungi in Hypocreales plectonematogenous, rarely nematogenous. Conidiophores solitary or aggregated, (sub-)erect, straight or slightly curved, arising directly from submerged or superficial hyphae, sometimes radiating out from coils or ropes formed by the mycelium, usually reduced to single phialides, unbranched or branched, bearing 1–2 levels with 1–2 phialides per node, 17.5–57(–64.5) μm long, 1.7– 2.7 μm wide at base, with 1–4 inconspicuous or conspicuous septa, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides terminal, lateral, subulate, hyaline, thin-, smooth-walled, 16.8–36(–49.5) μm long, 1–2.2 μm wide at base, with inconspicuous periclinal thickening and conspicuous cylindrical collarette at conidiogenous loci, occasionally with a percurrent or subterminal proliferation; polyphialides with two conidiogenous loci occasionally present. Conidia aseptate, obpyriform or obovoid with elongated apiculate bases and obtuse apices, hyaline, thin-, smooth-walled, 3–5 × 1.7–2.4 μm, arranged dry chains, often collapsing soon in dry heads. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 25–28 mm diam, flat, dusty at centre, hairy at periphery, white, margin entire, reverse buff; On MEA reaching 27 mm diam, flat, radially folded, felty, dirty white, margin entire, reverse saffron, with radial buff lines; On PDA reaching 35–38 mm diam, flat, felty, dirty white at centre, creamy white at periphery, margin entire, reverse pale saffron at centre, pale rosy buff at periphery; On SNA reaching 23 mm diam, flat, dusty, white, margin entire, reverse concolourous. Typus: India, Bangalore, Forestry Dept., from decaying pod of Delonix regia (Fabaceae), Jan. 1973, W. Gams (holotype CBS H-24604, ex-type culture CBS 595.73). Fig. 63. Waltergamsia moroccensis (ex-type culture CBS 512.82). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D, E. Conidial heads. F–J. Conidiophores. K. Conidia. Scale bars = 10 μm.
150 Hou et al. sequences were uploaded to GenBank (Supplementary Table S1). Bulbithecium truncatum L.W. Hou, L. Cai & Crous, sp. nov. MycoBank MB 845899. Fig. 68. Etymology: Referring to the broadly truncate conidia produced by the ex-type strain. Mycelium consisting of branched, septate, hyaline, smooth-, thin-walled hyphae, 1.3–2.4 μm wide, mycelial coils and ropes present. Sporulation sparse, phalacrogenous. Conidiophores solitary, (sub-)erect, straight or irregularly curved, flexuous, arising directly from submerged or superficial hyphae, or from coil formed by mycelium, mostly unbranched, reduced to single phialides, occasionally swollen at base, (25–)32–63.5(–74.5) μm long, 1.6–2.7 μm wide at base, 1–2(–5)-septate at base and middle, hyaline, smooth-walled, with cell walls usually thicker than those of vegetative hyphae. Phialides lateral, rarely terminal, cylindrical or subulate, irregularly curved, hyaline, thick-, smoothwalled, (12.8–)20–42(–54) μm long, 1.6–2.2 μm wide at base, with conspicuous periclinal thickening and minute collarette at conidiogenous loci; polyphialides not observed. Conidia aseptate, sub-globose or broadly ovoid, straight, with broadly truncate and elongated bases and obtuse apices, hyaline, thin-, smoothwalled, 2.6–4.1 × 1.8–2.8 μm, eguttulate or with one mediumsized guttule, arranged in chains and slimy heads at same colony. Chlamydospores and sexual morph not observed. Culture characteristics after 14 d at 25 °C: Colonies on OA reaching 23 cm diam, flat, membranous without aerial mycelium, white, margin entire, reverse concolourous; On MEA reaching 20–21 mm diam, flat, radially fold, moderate aerial mycelium, felty, white at centre, Fig. 68. Bulbithecium truncatum (ex-type culture CBS 113718). A–C. Colonies on OA, MEA and PDA, respectively, after 14 d at 25 °C. D. Conidiophores with conidial heads and conidial chains. E–H. Conidiophores. I. Conidiophores radiating out from coils formed by the mycelium. J. Conidia. Scale bars = 10 μm.
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