available online at www.studiesinmycology.org doi:10.3114/sim.2011.68.06 StudieS in Mycology 68: 139–162. 2011. Acremonium phylogenetic overview and revision of Gliomastix, Sarocladium, and Trichothecium R.C. Summerbell1, 2*, C. Gueidan3, 4, H-J. Schroers3, 5, G.S. de Hoog3, M. Starink3, Y. Arocha Rosete1, J. Guarro6 and J.A. Scott1, 2 1 Sporometrics, Inc. 219 Dufferin Street, Suite 20C, Toronto, Ont., Canada M6K 1Y9; 2Dalla Lana School of Public Health, University of Toronto, 223 College St., Toronto ON Canada M5T 1R4; 3CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; 4Department of Botany, The Natural History Museum, Cromwell Road, SW7 5BD London, United Kingdom; 5Agricultural Institute of Slovenia, Hacquetova 17, 1000 Ljubljana, Slovenia, Mycology Unit, Medical School; 6IISPV, Universitat Rovira i Virgili, Reus, Spain *Correspondence: Richard Summerbell, rsummerbell@sporometrics.com Abstract: Over 200 new sequences are generated for members of the genus Acremonium and related taxa including ribosomal small subunit sequences (SSU) for phylogenetic analysis and large subunit (LSU) sequences for phylogeny and DNA-based identiication. Phylogenetic analysis reveals that within the Hypocreales, there are two major clusters containing multiple Acremonium species. One clade contains Acremonium sclerotigenum, the genus Emericellopsis, and the genus Geosmithia as prominent elements. The second clade contains the genera Gliomastix sensu stricto and Bionectria. In addition, there are numerous smaller clades plus two multi-species clades, one containing Acremonium strictum and the type species of the genus Sarocladium, and, as seen in the combined SSU/LSU analysis, one associated subclade containing Acremonium breve and related species plus Acremonium curvulum and related species. This sequence information allows the revision of three genera. Gliomastix is revived for ive species, G. murorum, G. polychroma, G. tumulicola, G. roseogrisea, and G. masseei. Sarocladium is extended to include all members of the phylogenetically distinct A. strictum clade including the medically important A. kiliense and the protective maize endophyte A. zeae. Also included in Sarocladium are members of the phylogenetically delimited Acremonium bacillisporum clade, closely linked to the A. strictum clade. The genus Trichothecium is revised following the principles of unitary nomenclature based on the oldest valid anamorph or teleomorph name, and new combinations are made in Trichothecium for the tightly interrelated Acremonium crotocinigenum, Spicellum roseum, and teleomorph Leucosphaerina indica. Outside the Hypocreales, numerous Acremonium-like species fall into the Plectosphaerellaceae, and A. atrogriseum falls into the Cephalothecaceae. Key words: Acremonium, Cephalothecaceae, Gliomastix, holomorph concept, Leucosphaerina, nomenclature, Sarcopodium, Sarocladium, Trichothecium. Taxonomic novelties: Trichothecium sympodiale Summerbell, Seifert, & Schroers, nom. nov.; Gliomastix roseogrisea (S.B. Saksena) Summerbell, comb. nov., Gliomastix tumulicola (Kiyuna, An, Kigawa & Sugiy.) Summerbell, comb. nov., Sarocladium bacillisporum (Onions & Barron) Summerbell, comb. nov., Sarocladium bactrocephalum (W. Gams) Summerbell, comb. nov., Sarocladium glaucum (W. Gams) Summerbell, comb. nov., Sarocladium kiliense (Grütz) Summerbell, comb. nov., Sarocladium ochraceum (Onions & Barron) Summerbell, comb. nov., Sarocladium strictum (W. Gams) Summerbell, comb. nov., Sarocladium zeae (W. Gams & D.R. Sumner) Summerbell, comb. nov., Trichothecium crotocinigenum (Schol-Schwarz) Summerbell, Seifert, & Schroers, comb. nov., Trichothecium indicum (Arx, Mukerji & N. Singh) Summerbell, Seifert, & Schroers, comb. nov., Trichothecium ovalisporum (Seifert & Rehner) Seifert & Rehner, comb. nov. INTRODUCTION The genus Acremonium includes some of the most simply structured of all ilamentous anamorphic fungi. The characteristic morphology consists of septate hyphae giving rise to thin, tapered, mostly lateral phialides produced singly or in small groups. Conidia tend to be unicellular, produced in mucoid heads or unconnected chains. They can be hyaline or melanised, but the hyphae are usually hyaline. A preliminary study of the phylogenetic diversity of Acremonium by Glenn et al. (1996), based on partial nuclear ribosomal small subunit (SSU) sequences, showed that recognised members belonged to at least three groups in distinct orders of fungi. Most species including the type, A. alternatum, belong to the order Hypocreales. A smaller group of species, Acremonium section Chaetomioidea, belongs to the Sordariales. This section, typiied by the Acremonium alabamense anamorph of Thielavia terrestris, was conceived as including the Acremonium-like anamorphs of Chaetomium and Thielavia species (Morgan-Jones & Gams 1982). A recent study has placed several of these heretofore unnamed Acremonium-like anamorphs into the new genus Taifanglania (Liang et al. 2009), based on the type, T. hechuanensis. Another Acremonium species included by Glenn et al. (1996), A. furcatum, belongs to an order of uncertain identity. Subsequent publications have shown that A. furcatum is related to the well-known phytopathogen Verticillium dahliae and belongs to the recently established family Plectosphaerellaceae (Zare et al. 2007, Schoch et al. 2009), which groups together with the Glomerellaceae in a clade that forms a poorly deined, unnamed, ordinal-level sistertaxon to the Microascales. Several other Acremonium species such as the phytopathogen A. cucurbitacearum also have been shown to belong to the Plectosphaerellaceae (Zare et al. 2007). The simple structure of Acremonium has either convergently evolved in diverse fungal orders within the class Sordariomycetes or is symplesiomorphic at a very deep level. The diversity of fungi throughout the Ascomycota that produce Acremonium-like anamorphs is high, including genera such as Gabarnaudia (Microascales), Lecythophora (Coniochaetales), and Pseudogliomastix (Sordariales incertae sedis). The present study does not review the vast range of fungi producing simple phialidic conidiophores, but instead, focuses speciically on: 1) anamorphs that have been formally placed into the genus Acremonium, and 2) species and genera phylogenetically related to the named Acremonium species. The number of previously phylogenetically unstudied fungi is large. Currently, there are approximately 95 named Acremonium species with Copyright 2011 CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner speciied by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 139 SuMMerbell et al. traceable material (cultures or specimens in good condition), excluding endophyte species that were transferred to Neotyphodium by Glenn et al. (1996). In addition, there are an undetermined number of nectriaceous teleomorphs with unnamed Acremonium-like anamorphs plus about 15 named and unnamed Emericellopsis species with Acremonium anamorphs (Zuccaro et al. 2004). The preliminary phylogeny done by Glenn et al. (1996) includes only seven species that would currently be considered Acremonium, inclusive of the Acremonium berkeleyanum, anamorph of Cosmospora berkeleyana, formerly considered the anamorph of Cosmospora vilior, plus two Emericellopsis species. Clearly, further work is needed on the phylogeny of Acremonium. Because of the high biodiversity within Acremonium, relatively evolutionarily labile, rapidly evolving genes like the ribosomal internal transcribed spacer (ITS) are not alignable across the genus (de Hoog et al. 2000) or even within some of the individual orders that the genus spans. Because many Acremonium species are derived from relatively closely related families in the Sordariomycetes, relatively slowly evolving genes that are alignable such as the ribosomal large subunit (LSU) may yield considerable ambiguity about relationships. To address this problem, we performed an analysis of LSU and whole SSU sequences for a larger number of Acremonium isolates than has been examined previously. Based on these results, we chose six of the most phylogenetically distinctive species and included them in the Ascomycetous Tree of Life project (Schoch et al. 2009). The elegant phylogenetic analysis in that project was based on two nuclear ribosomal genes, one mitochondrial ribosomal gene, and portions of three protein-coding genes. These results permitted us to gain a clearer picture of relationships among the Acremonium groups that were imperfectly resolved in LSU and SSU analysis. In the present study we present the results of the LSU and small subunit (SSU) phylogenetic analyses for the majority of Acremonium species available in pure culture including described and undescribed species. This gives not only a phylogenetic overview of the genus, but also provides identiication-enabling sequences for Acremonium species that have not been sequenced previously. Taken with the Tree of Life studies, these results shed new light on the biodiversity of these morphologically simple fungi that have long been profoundly problematical in terms of accurate classiication and reliable species identiication. Medium (Raper & Raper 1972). The LSU region of ribosomal DNA (rDNA) was ampliied with primers V9G (de Hoog & Gerrits van den Ende 1998) and LR5 (Vilgalys & Hester 1990). The SSU region was ampliied with primers NS1 and NS24 and sequenced using primers NS1–NS4, NS6, NS24 (White et al. 1990; Gargas et al. 1992). The components for the PCR were used as described by Schroers (2000). The PCR program was 60 s at 94 °C (initial denaturation); 35 cycles of 35 s at 94 °C (denaturation), 50 s at 55 °C (annealing), and 120 s at 72 °C (elongation); and 6 min at 72 °C (inal elongation) followed by chilling to 4 °C. The PCR products were puriied with a GFX puriication kit (Amersham Pharmacia Biotech Inc., Roosendaal, The Netherlands) and visualised on an electrophoresis gel after ethidium bromide staining. The rDNA was sequenced with a BigDye terminator cycle sequencing kit (Applied Biosystems, Foster City, Calif.) and analysed on an ABI Prism 3700 instrument (Applied Biosystems) by using the standard conditions recommended by the vendor. The primers used in the sequence reaction were NL1 and NL4 (O’Donnell 1993), and LR5. Alignments and phylogenetic analyses Sequences were assembled and edited using SeqMan II software (DNAStar, Inc., Madison, Wis.). Manual alignments were performed using MacClade v. 4.08 (Maddison & Maddison 2003). Ambiguous regions (sensu Lutzoni et al. 2000) and introns were delimited manually and excluded from the alignments. Congruence was tested using a 70 % reciprocal bootstrap criterion (Mason-Gamer & Kellogg 1996, Reeb et al. 2004). Final phylogenetic analyses of the two-gene and one-gene datasets were performed using Stamatakis’s “randomised axelerated (sic) maximum likelihood for high performance computing” (RAxML VI-HPC, Stamatakis et al. 2005, 2008) on the Cipres Web Portal (http://www.phylo.org/ sub_sections/portal/). For the two-gene analysis, the maximum likelihood search followed a "GTRMIX" model of molecular evolution applied to two partitions, nucLSU and nucSSU. The same model was applied to the one-gene analysis without partition. Support values were obtained in RAxML with bootstrap analyses of 500 pseudoreplicates. The trees are labeled with the updated scientiic names. MATeRIAls AND MeThODs Two separate sets of data matrices were assembled (Table 1). The irst is a two-gene analysis that aims at investigating the phylogenetic position of Acremonium within the Sordariomycetes. The second is a one-gene analysis focusing on the Acremonium strains belonging to the order Hypocreales. The irst set includes the large and small subunits of the nuclear ribosomal RNA gene (nucLSU and nucSSU, respectively) and 166 taxa, including 56 strains of Acremonium, 105 reference taxa of Sordariomycetes, and ive species of Leotiomycetes as an outgroup (Botryotinia fuckeliana, Chalara aurea, Leotia lubrica, Microglossum rufum, and Pseudeurotium zonatum). The second set includes only the nucLSU for 331 taxa including 170 strains of Acremonium, 158 sequences of Hypocreales, and three outgroup species (Ceratocystis imbriata, Glomerella cingulata, and Ophiostoma piluliferum). DNA isolation and sequencing DNA was extracted with a FastDNA kit (Qbiogene, Heidelberg, Germany) from mycelium grown for 5–14 d in liquid Complete 140 ResUlTs DNA sequence alignments A total of 228 new sequences were generated for Acremonium, 192 nucLSU and 36 nucSSU (Table 1). For the two-gene dataset, one nucLSU and 41 nucSSU sequences were missing. After exclusion of ambiguous regions and introns, the two-gene dataset included 2 955 characters (1 250 nucLSU and 1 705 nucSSU). Among these, 1 739 were constant while 900 were parsimony-informative. After exclusion of ambiguous regions and introns, the one-gene dataset included 848 characters. Among these, 481 were constant while 260 were parsimony-informative. Phylogenetic inference As shown in Fig. 1, the species of Acremonium mostly fall into three groups, namely the Hypocreales, the Plectosphaerellaceae, and the Sordariales. The bulk of species fall into the Hypocreales. Gliomastix/Bionectria clade 99 “Acremonium hyalinulum” CBS 271.36 Roumegueriella rufula 75 Hydropisphaera erubescens 1 Acremonium spinosum CBS 136.33 T 99 85 Acremonium rutilum CBS 396.66 T Bionectria ochroleuca 2 99 “Acremonium blochii” CBS 427.93 Acremonium pinkertoniae CBS 157.70 T 97 Acremonium chrysogenum CBS 144.62 T chrysogenum 74 -clade Acremonium flavum CBS 596.70 T Acremonium sclerotigenum CBS 124.42 T Geosmithia lavendula 99 Geosmithia putterillii “Acremonium egyptiacum” CBS 303.64 sclerotigenum/Geosmithia clade Sarocladium bactrocephalum CBS 749.69 T Sarocladium strictum CBS 346.70 T strictum Sarocladium kiliense CBS 146.62 95 -clade Sarocladium kiliense CBS 122.29 T 99 Sarocladium zeae CBS 801.69 T Acremonium breve CBS 150.62 T Acremonium radiatum CBS 142.62 T breve/curvulum -clade Acremonium gamsii CBS 726.71 T 96 Acremonium curvulum CBS 430.66 T “Acremonium blochii” CBS 424.93 persicinum 98 Acremonium persicinum CBS 310.59 T -clade 96 99 Glomerellales Hypocreales Sarocladium clade Acremoniella lutzi T Petriella setifera Doratomyces stemonitis Microascales (1) Microascus longirostris 93 Microascus trigonosporus Aniptodera chesapeakensis 99 Halosphaeria appendiculata Lignincola laevis 99 Nimbospora effusa 96 Halosphaeriales Nohea umiumi 80 Corollospora maritima 95 Varicosporina ramulosa Ceriosporopsis halima Graphium penicillioides Microascales (2) Ambrosiella xylebori Microascales (3) Ceratocystis fimbriata Acremonium alcalophilum CBS 114.92 T Acremonium restrictum CBS 178.40 T Acremonium antarcticum CBS 987.87 92 89 Acremonium stromaticum CBS 863.73 T Acremonium furcatum CBS 122.42 T 75 Verticillium alboatrum (Acremonium apii T) CBS 130.51 Verticillium dahliae Acremonium nepalense CBS 971.72 T Plectosphaerella cucumerina 70 Acremonium cucurbitacearum CBS 683.88 96 Acremonium brunnescens CBS 559.73 T Glomerella cingulata “Acremonium alternatum” CBS 406.66 92 Sarocladium bacillisporum CBS 425.67 T bacillisporum -clade “Acremonium implicatum” CBS 243.59 Plectosphaerellaceae acremonium phylogeny A Fig. 1.A–C. The phylogenetic position of Acremonium and related fungi within the Sordariomycetes, as seen in combined analysis of the large and small subunits of the nuclear ribosomal RNA gene (LSU + SSU) analysed by maximum likelihood via RAxML VI-HPC following a GTRMIX model applied to two partitions. 100 % bootstrap values are indicated by a black dot on the relevant internode. www.studiesinmycology.org 141 71 96 Simplicillium lanosoniveum CBS 321.72 Cordycipitaceae Cordyceps cardinalis Acremonium camptosporum CBS 756.69 T Paecilomyces lilacinus 2 Elaphocordyceps capitata Ophiocordycipitaceae Elaphocordyceps ophioglossoides Balansia henningsiana Epichloe typhina Clavicipitaceae Claviceps purpurea Acremonium guillematii CBS 766.69 T Acremonium minutisporum CBS 147.62 T Acremonium vitellinum CBS 792.69 T Acremonium exiguum CBS 587.73 T “Acremonium potronii” CBS 416.68 Acremonium psammosporum CBS 590.63 T Acremonium recifei CBS 137.35 T Nectria cinnabarina Acremonium roseolum CBS 289.62 T Stachybotrys chartarum Stachybotrys/ Stachybotrys subsimplex Peethambara -clade Myrothecium roridum 1 Peethambara spirostriata Pseudonectria rousseliana Acremonium nigrosclerotium CBS 154.72 T 96 Hypocrea americana Hypocrea lutea Hypocreaceae Sphaerostilbella berkeleyana Niesslia exilis Nectria haematococca Chaetosphaerella phaeostroma 96 Melanospora tiffanii Melanospora zamiae Lindra thalassiae Lulworthiales Lulworthia grandispora “Acremonium alabamense” CBS 456.75 Cercophora terricola Apiosordaria verruculosa 97 Cercophora striata Podospora decipiens Bombardia bombarda Lasiosphaeria ovina 71 Sordariales Cercophora newfieldiana Gelasinospora tetrasperma 99 Neurospora crassa Immersiella immersa Lasiosphaeria hispida Podospora fibrinocaudata Strattonia carbonaria Camarops amorpha 84 Camarops microspora 98 Boliniales Camarops tubulina 94 Camarops ustulinoides 97 Chaetosphaeria ovoidea 89 Chaetosphaeriales Menispora tortuosa Melanochaeta hemipsila 74 Lasiosphaeriella nitida Linocarpon appendiculatum Fig. 1. (Continued). 142 Hypocreales SuMMerbell et al. B acremonium phylogeny 99 Cephalothecaceae “Acremonium cf. alternatum” CBS 109043 Acremonium atrogriseum CBS 544.79 75 Acremonium atrogriseum CBS 981.70 80 Acremonium atrogriseum CBS 774.97 Acremonium atrogriseum CBS 507.82 Acremonium atrogriseum CBS 733.70 86 Acremonium atrogriseum CBS 252.68 Acremonium atrogriseum CBS 604.67 T 92 Acremonium atrogriseum CBS 306.85 Cephalotheca sulfurea Acremonium thermophilum CBS 734.71 T 88 Albertiniella polyporicola Coniochaeta ostrea 99 Coniochaetales 98 Coniochaeta savoryi 70 Coniochaeta discoidea Annulatascus triseptatus 89 Fragosphaeria purpurea 91 Ophiostoma piliferum 1 Ophiostomatales 97 Ophiostoma stenoceras Papulosa amerospora Apiognomonia errabunda 92 Plagiostoma euphorbiae Cryptodiaporthe aesculi 99 Gnomonia gnomon Cryptosporella hypodermia 95 Melanconis alni Melanconis stilbostoma 89 99 81 Melanconis marginalis Chromendothia citrina Diaporthales 97 Endothia gyrosa 89 Chrysoporthe cubensis Diaporthe eres Diaporthe phaseolorum 84 Mazzantia napelli 88 Leucostoma niveum Valsella salicis Valsa ambiens Cryphonectria parasitica Anthostomella torosa Apiospora montagnei Diatrype disciformis 82 Eutypa lata Xylariales Graphostroma platystoma 77 Xylaria acuta Xylaria hypoxylon 90 Seynesia erumpens Leotia lubrica Microglossum rufum Pseudeurotium zonatum Chalara aurea Leotiomycetes (outgroup) Acremonium butyri (?) CBS 301.38 T Acremonium lichenicola CBS 425.66 T Botryotinia fuckeliana 0.01 substitutions/site C Fig. 1. (Continued). www.studiesinmycology.org 143 SuMMerbell et al. Table 1. List of Acremonium species included in this study as well as other novel or independently redone sequences of related fungi used for comparison. Sequences from GenBank of other comparison taxa are listed in Supplemental Table 1a - see online Supplementary Information. Collection and GenBank numbers are indicated and type strains (T) are mentioned. Sequences generated in this study are shown in bold. Dashes indicate missing data in the two-gene analysis. Isolates that cannot be assigned a phylogenetically conirmed name are listed under the name under which they are currently held in the CBS collection. Currently assigned species name Collection numbers nuclsU Acremoniella lutzi T Acremonium acutatum T Acremonium alabamense Acremonium alcalophilum T Acremonium alternatum T “Acremonium alternatum” “Acremonium cf. alternatum” Acremonium antarcticum Acremonium atrogriseum T Acremonium biseptum T “Acremonium blochii” Acremonium borodinense T Acremonium brachypeniumT Acremonium breve T Acremonium brunnescens T Acremonium butyri T Acremonium camptosporum T Acremonium cavaraeanum Acremonium cerealis Acremonium chrysogenum T Acremonium cucurbitacearum Acremonium curvulum T “Acremonium aff. curvulum” Acremonium egyptiacum Acremonium exiguum T Acremonium exuviarum T Acremonium lavum T Acremonium fuci Acremonium furcatum T 144 CBS 103.48 CBS 682.71 CBS 456.75 CBS 114.92 CBS 407.66 CBS 381.70A CBS 406.66 CBS 831.97 CBS 114602 CBS 109043 CBS 987.87 CBS 604.67 CBS 252.68 CBS 306.85 CBS 507.82 CBS 544.79 CBS 733.70 CBS 774.97 CBS 981.70 CBS 750.69 CBS 324.33 CBS 424.93 CBS 427.93 CBS 993.69 CBS 101148 CBS 866.73 CBS 150.62 CBS 559.73 CBS 301.38 CBS 756.69 CBS 677.74 CBS 757.69 CBS 835.91 CBS 890.85 CBS 758.69 CBS 207.65 CBS 215.69 CBS 144.62 CBS 683.88 CBS 430.66 CBS 104.78 CBS 214.70 CBS 229.75 CBS 333.92 CBS 384.70A CBS 384.70C CBS 523.72 CBS 761.69 CBS 898.85 CBS 110514 CBS 100551 CBS 113275 CBS 303.64 CBS 587.73 UAMH 9995 CBS 596.70 UAMH 6508 CBS 122.42 hQ231971 hQ231965 hQ231972 hQ231973 hQ231988 hQ231986 hQ231987 hQ231989 hQ231990 hQ231974 hQ231975 hQ231981 hQ231977 hQ231978 hQ231979 hQ231980 hQ231982 hQ231983 hQ231984 hQ231998 hQ231999 hQ232000 hQ232001 hQ232002 hQ232003 hQ232004 hQ232005 hQ231966 hQ231967 hQ232008 hQ232007 hQ232009 hQ232010 hQ232011 hQ232012 hQ232013 hQ232014 hQ232017 hQ231968 hQ232026 hQ232019 hQ232020 hQ232021 hQ232022 hQ232023 hQ232024 hQ232028 hQ232029 hQ232030 hQ232032 hQ232031 hQ232033 hQ232034 hQ232035 hQ232036 hQ232037 hQ232038 AY378154 nucssU Notes – Ex-type of Acremoniella lutzi – – Ex-type of Acremonium alcalophilum hQ232178 – – – – – – – – – – Ex-type of Phaeoscopulariopsis atrogrisea Ex-type of Acremonium biseptum hQ232181 hQ232182 Ex-type of Acremonium borodinense hQ232183 hQ232184 – hQ232186 Ex-type of Cephalosporium roseum var. breve Ex-type of Acremonium brunnescens Ex-type of Tilachlidium butyri; synonym of Cadophora malorum Ex-type of Acremonium camptosporum Ex-type of Gliomastix guttuliformis hQ232187 – hQ232188 Ex-type of Cephalosporium chrysogenum Previously identiied as Acremonium strictum Ex-type of Acremonium curvulum hQ232189 hQ232190 Ex-type of Acremonium exiguum hQ232191 Ex-type of Acremonium lavum hQ232192 Ex-type of Acremonium furcatum acremonium phylogeny Table 1. (Continued). Currently assigned species name Collection numbers nuclsU Acremonium fusidioides T Acremonium gamsii T Acremonium guillematii T Acremonium hansfordii Acremonium hennebertii T Acremonium hyalinulum “Acremonium implicatum” Acremonium incrustatumT Acremonium inlatum T Acremonium lichenicola T Acremonium longisporum “Acremonium luzulae” Acremonium minutisporum T Acremonium nepalense T Acremonium nigrosclerotium T Acremonium persicinum T “Acremonium persicinum” “Acremonium aff. persicinum” “Acremonium cf. persicinum” Acremonium pinkertoniae T “Acremonium potronii” Acremonium psammosporum T Acremonium pseudozeylanicum T Acremonium pteridii T Acremonium radiatum T Acremonium recifei T “Acremonium recifei” www.studiesinmycology.org CBS 840.68 CBS 726.71 CBS 766.69 CBS 390.73 CBS 768.69 CBS 271.36 CBS 243.59 CBS 397.70B CBS 159.70 CBS 212.69 CBS 439.70 CBS 403.70 CBS 425.66 CBS 113.69 CBS 495.67 CBS 579.73 CBS 147.62 det 267B CBS 971.72 CBS 154.72 CBS 310.59 CBS 169.65 CBS 295.70A CBS 295.70M CBS 330.80 CBS 378.70A CBS 378.70D CBS 378.70 E CBS 439.66 CBS 469.67 CBS 101694 CBS 102349 CBS 378.70C CBS 110646 CBS 203.73 CBS 263.89 CBS 102877 CBS 157.70 CBS 189.70 CBS 379.70F CBS 416.68 CBS 433.88 CBS 781.69 CBS 590.63 CBS 560.73 CBS 782.69 CBS 784.69 CBS 142.62 CBS 137.35 CBS 135.71 CBS 220.84 CBS 362.76 CBS 402.89 CBS 411.91 CBS 442.66 CBS 541.89 CBS 555.73 CBS 596.74 CBS 976.70 CBS 400.85 CBS 505.94 CBS 485.77 CBS 482.78 hQ232039 hQ232040 hQ232042 hQ232043 hQ232044 hQ232045 hQ232046 hQ232047 hQ232049 hQ232050 hQ232051 hQ231991 hQ231969 hQ232057 hQ232058 hQ232059 hQ232061 hQ232062 hQ231970 hQ232069 hQ232077 hQ232072 hQ232075 hQ232076 hQ232078 hQ232079 hQ232081 hQ232082 hQ232083 hQ232084 hQ232085 hQ232086 hQ232080 hQ232088 hQ232073 hQ232074 hQ232087 hQ232089 hQ232094 hQ232096 hQ232097 hQ232098 hQ232099 hQ232100 hQ232101 hQ232102 hQ232103 hQ232104 hQ232106 hQ232105 hQ232107 hQ232108 hQ232109 hQ232110 hQ232111 hQ232114 hQ232115 hQ232116 hQ232117 hQ232025 hQ232027 hQ232113 hQ232112 nucssU hQ232193 hQ232194 hQ232195 hQ232196 Notes Ex-type of Paecilomyces fusidioides Ex-type of Acremonium gamsii Ex-type of Acremonium guillematii Ex-type of Acremonium hennebertii Authentic strain of Fusarium terricola Ex-type of Gliomastix inlata – In CBS as Acremonium atrogriseum Ex-type of Acremonium lichenicola hQ232199 Ex-type of Cephalosporium minutisporum – hQ232200 hQ232201 Ex-type of Acremonium nepalense Ex-type of Acremonium nigrosclerotium Ex-type of Paecilomyces persicinus hQ232202 Ex-type of Acremonium pinkertoniae hQ232203 hQ232204 Ex-type of Acremonium psammosporum Ex-type of Acremonium pseudozeylanicum Ex-type of Acremonium pteridii hQ232205 hQ232206 Ex-type of Cephalosporium acremonium var. radiatum Ex-type of Cephalosporium recifei In CBS as Acremonium curvulum In CBS as Acremonium cf. curvulum 145 SuMMerbell et al. Table 1. (Continued). Currently assigned species name Collection numbers nuclsU CBS 110348 Acremonium restrictum T CBS 178.40 Acremonium rhabdosporum T CBS 438.66 Acremonium roseolum T CBS 289.62 Acremonium rutilum T CBS 396.66 Acremonium salmoneum T CBS 721.71 Acremonium sclerotigenum T CBS 124.42 CBS 270.86 CBS 281.80 CBS 384.65 CBS 786.69 CBS 100816 OMH F1648.97 OMH F2365.97 OMH F2969.97 OMH F3691.97 CBS 287.70O CBS 379.70D CBS 223.70 Acremonium sordidulum T CBS 385.73 Acremonium sp. CBS 314.72 Acremonium spinosum T CBS 136.33 “Acremonium strictum” CBS 106.23 CBS 147.49 Acremonium stromaticum T CBS 863.73 Acremonium tectonae T CBS 725.87 Acremonium thermophilum T CBS 734.71 Acremonium tsugae T CBS 788.69 Acremonium tubakii T CBS 790.69 “Acremonium tubakii” CBS 824.69 Acremonium verruculosum T CBS 989.69 Acremonium vitellinum T CBS 792.69 Acremonium zeylanicum CBS 746.73 Acremonium zonatum CBS 565.67 “Cephalosporium acremonium var. cereum” T CBS 140.62 “Cephalosporium acremonium funiculosum” T “Cephalosporium ballagii” T hQ232118 hQ232119 hQ232120 hQ232123 hQ232124 hQ232125 hQ232126 hQ232127 hQ232128 hQ232129 hQ232130 hQ232131 hQ232132 hQ232133 hQ232134 hQ232135 hQ232140 hQ232095 hQ231985 hQ232136 hQ232156 hQ232137 hQ232138 hQ232139 hQ232143 hQ232144 hQ232145 hQ232146 hQ232148 hQ232149 hQ232150 hQ232151 hQ232154 hQ232155 hQ232147 var. CBS 141.62 hQ232053 CBS 134.33 hQ232016 “Cephalosporium malorum” T CBS 117.25 hQ232015 “Cephalosporium purpurascens” T CBS 149.62 hQ232071 Cosmospora khandalensis T CBS 356.65 hQ231996 Cosmospora lavitskiaeT CBS 530.68 hQ231997 Gliomastix masseei Gliomastix murorum CBS 794.69 CBS 154.25 hQ232060 hQ232063 CBS 195.70 CBS 119.67 CBS 157.72 CBS 378.36 hQ232064 hQ232066 hQ232067 hQ232068 CBS 181.27 hQ232091 CBS 151.26 hQ232090 Gliomastix roseogrisea T CBS 617.94 CBS 134.56 hQ232093 hQ232121 Lanatonectria lavolanata CBS 279.79 CBS 213.69 CCFC 226570 CBS 211.69 CBS 230.31 hQ232122 hQ232092 AY283559 hQ232065 hQ232157 Gliomastix polychroma T 146 nucssU Notes – Ex-type of Verticillium dahliae f. restrictum Ex-type of Acremonium rhabdosporum Ex-type of Paecilomyces roseolus Ex-type of Acremonium rutilum Ex-type of Acremonium salmoneum Ex-type of Cephalosporium sclerotigenum hQ232207 hQ232208 hQ232209 In CBS as Acremonium strictum In CBS as Acremonium potronii In CBS as Acremonium alternatum Ex-type of Acremonium sordidulum hQ232210 Ex-type of Cephalosporium spinosum – Ex-type of Acremonium stromaticum Ex-type of Acremonium tectonae Ex-type of Acremonium thermophilum Ex-type of Acremonium tsugae Ex-type of Acremonium tubakii – hQ232212 Ex-type of Acremonium verruculosum Ex-type of Acremonium vitellinum Ex-type of Cephalosporium acremonium var. cereum. In CBS as Acremonium tubakii Ex-type of Cephalosporium acremonium var. funiculosum. In CBS as Acremonium kiliense Ex-type of Cephalosporium ballagii. In CBS as Acremonium charticola Ex-type of Cephalosporium malorum. In CBS as Acremonium charticola Ex-type of Cephalosporium purpurascens. In CBS as Acremonium persicinum Ex-type of Cephalosporium khandalense. In CBS as Acremonium berkeleyanum Ex-type of Gliomastix lavitskiae. In CBS as Acremonium berkeleyanum In CBS as Acremonium masseei Ex-type of Graphium malorum. In CBS as Acremonium murorum var. felina In CBS as Acremonium murorum var. felina In CBS as Acremonium murorum var. murorum In CBS as Acremonium murorum var. murorum Ex-type of Torula cephalosporioides. In CBS as Acremonium murorum var. murorum Ex-type of Oospora polychroma. In CBS as Acremonium polychromum Ex-type of Periconia tenuissima var. nigra. In CBS as Acremonium polychromum In CBS as Acremonium polychromum Ex-type of Cephalosporium roseogriseum. In CBS as Acremonium roseogriseum In CBS as Acremonium roseogriseum In CBS as Acremonium polychromum Identiied as Acremonium murorum var. felina In CBS as Acremonium murorum var. felina acremonium phylogeny Table 1. (Continued). Currently assigned species name Collection numbers nuclsU Lanatonectria locculenta Leucosphaerina arxii T Nalanthamala diospyri T Nectria rishbethii T Neocosmospora endophytica Paecilomyces lilacinus Pochonia bulbillosa Sarcopodium circinatum Sarcopodium circinosetiferum Sarcopodium vanillae Sarocladium attenuatum T Sarocladium bacillisporum T Sarocladium bactrocephalum T Sarocladium glaucum T Sarocladium kiliense T CBS 113461 CBS 737.84 CBS 560.89 CBS 496.67 AR 2674 CBS 101068 CBS 102853 CBS 376.81 CBS 587.92 CBS 114068 CBS 100251 CBS 100252 CBS 100998 CBS 101116 CBS 100582 CBS 399.73 CBS 425.67 CBS 749.69 NRRL 20583 CBS 796.69 CBS 122.29 CBS 146.62 hQ232158 hQ232159 hQ232160 hQ232162 U17411 hQ232163 hQ232164 hQ232167 hQ232168 hQ232169 hQ232170 hQ232171 hQ232172 hQ232173 hQ232174 hQ232165 hQ231992 hQ231994 hQ231995 hQ232041 hQ232052 hQ232048 CBS 155.61 hQ232054 CBS 156.61 hQ232055 CBS 157.61 Sarocladium ochraceum T Sarocladium oryzae Sarocladium strictum T “Sarocladium cf. strictum” Sarocladium zeae T CBS 428.67 CBS 180.74 CBS 346.70 JY03-006 CBS 801.69 KAS 965 Simplicillium lanosoniveum CBS 321.72 Simplicillium obclavatum T CBS 311.74 Trichothecium crotocinigenum T CBS 129.64 “Trichothecium indicum”/ Leucosphaerina CBS 123.78 indicaT Trichothecium roseum DAOM 208997 Trichothecium sympodiale ATCC 36477 Verticillium alboatrum CBS 130.51 Verticillium insectorum CBS 101239 Verticillium leptobactrum CBS 109351 hQ232214 hQ232179 hQ232180 hQ232198 hQ232197 hQ232056 hQ232070 hQ232166 hQ232141 hQ232142 hQ232152 hQ232153 hQ232006 hQ232175 hQ232018 AF096194 U69891 U69889 hQ231976 hQ248107 hQ231993 Nine of the named Acremonium species in this analysis belong to the Plectosphaerellaceae. The Sordariales are represented in Fig. 1 only by Acremonium alabamense, the only named Acremonium species in Acremonium section Chaetomioidea. Outside these groups Acremonium atrogriseum, represented by numerous conspeciic isolates, belongs to the family Cephalothecaceae (Fig. 1C), along with Albertiniella polyporicola and Cephalotheca sulfurea; this family is sister to the Coniochaetales. Another Acremonium species, A. thermophilum, falls into the Cephalothecaceae clade grouping with Albertiniella polyporicola. An isolate provisionally identiied as Acremonium alternatum, CBS 109043 is a member of the Cephalothecaceae. The complex status of A. alternatum is discussed below. The Acremonium species in the Hypocreales form an array of poorly to well distinguished clades, most of which do not correspond to previously recognised genera or suprageneric taxa. Included within the Hypocreales in the Sarocladium clade labeled the "strictum-clade" www.studiesinmycology.org nucssU Notes Ex-type of Leucosphaerina arxii Ex-type of Cephalosporium diospyri = Acremonium diospyri Ex-type of Nectria rishbethii Anamorph is Acremonium fungicola Atypical monophialidic isolate, Acremonium+E402-like Atypical isolate Ex-type of Sarocladium attenuatum Ex-type of Acremonium bacillisporum Ex-type of Acremonium bactrocephalum Ex-type of Acremonium glaucum Ex-type of Acremonium kiliense Ex-type of Cephalosporium incoloratum. In CBS as Acremonium incoloratum Ex-type of Cephalosporium incarnatum. In CBS as Acremonium kiliense Ex-type of Cephalosporium incarnatum var. macrospora. In CBS as Acremonium kiliense Ex-type of Cephalosporium infestans.In CBS as Acremonium kiliense Ex-type of Paecilomyces ochraceus. In CBS as Acremonium ochraceum hQ232211 Ex-type of Acremonium strictum hQ232213 Ex-type of Acremonium zeae hQ232185 Ex-type of Acremonium byssoides Ex-type of Acremonium obclavatum Ex-type of Acremonium crotocinigenum Ex-type of ‘Leucosphaera’ indica – In CBS as Spicellum roseum Ex-type of Cephalosporium apii, in CBS as Acremonium apii In CBS as Acremonium cf. bacillisporum is the well known soil fungus long known as Acremonium strictum (Fig. 1A). The soil fungus and human opportunistic pathogen traditionally called A. kiliense is also included as is the maize corn endophyte known as A. zeae. The corresponding clade in Fig. 2C based on LSU reveals that this group of fungi includes the rice pathogen Sarocladium oryzae as a saltatory morphological apomorph. No teleomorphs are known to be associated with this group. This clade consists of fungi forming conidia in mucoid heads; it is closely related to a clade of species forming catenulate conidia, namely the Acremonium bacillisporum clade including A. bacillisporum, A. glaucum, A. implicatum pro parte, and, in a separate subclade, A. ochraceum (Figs 1A, 2C). The "bacillisporum-clade" and "strictumclade" grouped together in an overall Sarocladium clade (Figs 1, 2). Two catenulate-conidial isolates labeled A. alternatum are also loosely associated with the A. bacillisporum clade in Fig. 2C. In Fig. 1A, one isolate CBS 406.66 is connected to the Sarocladium and A. breve/curvulum clades with a 96 % bootstrap value. 147 SuMMerbell et al. 85 94 T “Cephalosporium acremonium var. cereum” CBS 140.62 T Acremonium sclerotigenum CBS 223.70 Acremonium sclerotigenum CBS 379.70D Acremonium sclerotigenum OMH F2365.97 Acremonium sclerotigenum OMH F2969.97 Acremonium sclerotigenum OMH F1648.97 Acremonium sclerotigenum OMH F3691.97 Mycopepon smithii Pseudonectria sp. Acremonium sclerotigenum CBS 100816 sclerotigenum Acremonium sclerotigenum CBS 287.70O -clade Acremonium sclerotigenum CBS 384.65 Acremonium sclerotigenum CBS 281.80 Acremonium sclerotigenum CBS 270.86 89 Acremonium sclerotigenum CBS 124.42 T Acremonium sclerotigenum CBS 786.69 Acremonium alternatum CBS 407.66 T “Cephalosporium malorum” CBS 117.25 T Acremonium sordidulum CBS 385.73 T Acremonium brachypenium CBS 866.73 T brachypenium 83 “Cephalosporium purpurascens” CBS 149.62 T -clade “Acremonium potronii” CBS 189.70 Acremonium exuviarum UAMH 9995 T “Acremonium tubakii” CBS 824.69 Emericellopsis terricola 79 Acremonium fuci UAMH 6508 Emericellopsis “Acremonium potronii” CBS 379.70F Acremonium salmoneum CBS 721.71 T -clade 99 93 Stilbella fimetaria Acremonium tubakii CBS 790.69 T Stanjemonium grisellum Acremonium chrysogenum CBS 144.62 T 90 Acremonium flavum CBS 596.70 T chrysogenum 71 Hapsidospora irregularis -clade Nigrosabulum globosum 74 Mycoarachis inversa Geosmithia lavendula Geosmithia putterillii “Acremonium blochii” CBS 993.69 99 Acremonium borodinense CBS 101148 T pinkertoniae 96 Acremonium pinkertoniae CBS 157.70 T -clade Bulbithecium hyalosporum Leucosphaerina arxii T “Acremonium cf. persicinum” CBS 102877 “Acremonium alternatum” CBS 381.70A 85 “Acremonium alternatum” CBS 831.97 “Acremonium cavaraeanum” CBS 758.69 fusidioides Acremonium hennebertii CBS 768.69 T -clade Acremonium fusidioides CBS 840.68 T “Acremonium hansfordii” CBS 390.73 “Acremonium egyptiacum” CBS 303.64 99 “Acremonium blochii” CBS 427.93 96 “Acremonium blochii” CBS 324.33 Verticillium insectorum “Acremonium zeylanicum” CBS 746.73 Stilbocrea macrostoma sclerotigenum/Geosmithia-clade 75 Acremonium acutatum CBS 682.71 A Fig. 2.A–E. The phylogenetic position of Acremonium and related fungi within the Hypocreales, as seen in nucLSU analysed by maximum likelihood via RAxML VI-HPC following a GTRMIX model applied to a single partition. 100 % bootstrap values are indicated by a black dot on the relevant internode. 148 acremonium phylogeny Acremonium biseptum CBS 750.69 T cerealis Acremonium cerealis CBS 207.65 -clade Acremonium cerealis CBS 215.69 93 “Acremonium blochii” CBS 424.93 Acremonium persicinum CBS 378.70E Acremonium persicinum CBS 378.70D Acremonium persicinum CBS 378.70A Acremonium persicinum CBS 102349 Acremonium persicinum CBS 169.65 persicinum Acremonium persicinum CBS 295.70M Acremonium persicinum CBS 295.70A -clade Acremonium persicinum CBS 439.66 Acremonium persicinum CBS 101694 Acremonium persicinum CBS 469.67 91 Acremonium persicinum CBS 330.80 Acremonium verruculosum CBS 989.69 T Acremonium persicinum CBS 310.59 T Hydropisphaera erubescens 2 Hydropisphaera erubescens 1 Gliomastix masseei CBS 794.69 T Gliomastix murorum CBS 157.72 Gliomastix murorum CBS 195.70 Gliomastix murorum CBS 119.67 80 Gliomastix murorum CBS 378.36 92 Gliomastix murorum CBS 154.25 98 Gliomastix polychroma CBS 151.26 Gliomastix polychroma CBS 181.27 T Gliomastix polychroma CBS 617.94 Gliomastix roseogrisea CBS 211.69 Gliomastix roseogrisea CBS 213.69 Gliomastix roseogrisea CBS 134.56 T 99 Gliomastix roseogrisea CBS 279.79 Gliomastix roseogrisea CCFC 226570 Heleococcum japonicum 99 Hydropisphaera peziza 1 89 Hydropisphaera peziza 2 Roumegueriella rufula 72 Selinia pulchra “Acremonium hyalinulum” CBS 271.36 Nalanthamala squamicola “Acremonium aff. persicinum” CBS 203.73 “Acremonium luzulae” CBS 495.67 Nectria zonata Acremonium tectonae CBS 725.87 T “Acremonium longisporum” CBS 113.69 “Acremonium persicinum” CBS 378.70C 100 “Acremonium aff. persicinum” CBS 263.89 Acremonium rutilum CBS 396.66 T Acremonium pteridii CBS 784.69 pteridii 100 Acremonium spinosum CBS 136.33 T -clade Acremonium pteridii CBS 782.69 T Ochronectria calami 93 Bionectria grammicospora Bionectria ochroleuca 2 72 Nectria sesquicillii Bionectria pityrodes Stephanonectria keithii Nectriopsis sporangiicola 99 93 Nectriopsis violacea Sesquicillium microsporum Kallichroma glabrum Tilachlidium brachiatum 86 78 Fig. 2. (Continued). www.studiesinmycology.org Gliomastix/Bionectria-clade Gliomastix clade 85 B 149 SuMMerbell et al. Acremonium incrustatum CBS 159.70 T “Acremonium potronii” CBS 433.88 93 Linkosia fusiformis Acremonium guillematii CBS 766.69 T “Acremonium persicinum” CBS 110646 Acremonium sp. CBS 314.72 minutisporum Acremonium vitellinum CBS 792.69 T T Acremonium minutisporum CBS 147.62 100 -clade Acremonium minutisporum det 267B 100 “Acremonium alternatum” CBS 406.66 “Acremonium alternatum” CBS 114602 99 Sarocladium bacillisporum CBS 425.67 T bacillisporum 77 “Acremonium implicatum” CBS 243.59 -clade Sarocladium glaucum CBS 796.69 T 94 Sarocladium bactrocephalum CBS 749.69 T Sarocladium bactrocephalum NRRL 20583 84 Sarocladium cf. strictum JY03-006 83 Sarocladium strictum CBS 346.70 T “Acremonium zonatum” CBS 565.67 84 Sarocladium 100 Sarocladium attenuatum T strictum Sarocladium oryzae -clade 94 Sarocladium kiliense CBS 146.62 -clade Sarocladium kiliense CBS 122.29 T Sarocladium kiliense CBS 155.61 Sarocladium kiliense CBS 157.61 Sarocladium kiliense CBS 156.61 Sarocladium zeae CBS 801.69 T 98 Sarocladium zeae KAS 965 Sarocladium ochraceum CBS 428.67 T Acremonium breve CBS 150.62 T 76 Acremonium radiatum CBS 142.62 T breve 91 “Acremonium strictum” CBS 147.49 100 “Cephalosporium acremonium var. funiculosum” CBS 141.62 T -clade Acremonium gamsii CBS 726.71 T 83 Trichothecium crotocinigenum CBS 129.64 T “Trichothecium indicum”/Leucosphaerina indica T Trichothecium 100 Trichothecium sympodiale -clade 73 Trichothecium roseum 74 Acremonium curvulum CBS 430.66 T 75 Acremonium curvulum CBS 229.75 Acremonium curvulum CBS 384.70A Acremonium curvulum CBS 898.85 100 Acremonium curvulum CBS 761.69 curvulum Acremonium curvulum CBS 333.92 -clade Acremonium curvulum CBS 214.70 Acremonium curvulum CBS 110514 95 Acremonium curvulum CBS 384.70C 100 Acremonium curvulum CBS 104.78 Acremonium curvulum CBS 523.72 Valetoniellopsis laxa Acremonium rhabdosporum CBS 438.66 T Lanatonectria flavolanata 1 98 Lanatonectria flavolanata 2 Sarcopodium vanillae 94 Sarcopodium circinatum 2 Sarcopodium circinosetiferum 3 Lanatonectria Sarcopodium circinosetiferum 4 -clade 82 Lanatonectria flocculenta Sarcopodium circinosetiferum 2 84 Sarcopodium circinosetiferum 1 Pseudonectria rousseliana Fig. 2. (Continued). 150 C acremonium phylogeny Acremonium inflatum CBS 403.70 99 Acremonium inflatum CBS 439.70 76 Acremonium inflatum CBS 212.69 T 100 inflatum Nectriaceae/recifei-clade “Acremonium luzulae” CBS 579.73 -clade 100 Scopinella solani Acremonium roseolum CBS 289.62 T Peethambara sundara Albosynnema elegans 88 Didymostilbe echinofibrosa 98 Peethambara spirostriata Sarcopodium circinatum 3 Sarcopodium circinatum 1 Myrothecium cinctum Stachybotrys/Peethambara Myrothecium inundatum Myrothecium roridum 1 -clade Myrothecium roridum 2 Myrothecium leucotrichum 86 Myrothecium verrucaria Parasarcopodium ceratocaryi Stachybotrys chartarum 78 Chaetosphaeria aterrima 96 Didymostilbe matsushimae Melanopsamma pomiformis 76 Cosmospora lavitskiae CBS 530.68 T Cosmospora episphaeria 80 “Acremonium aff. curvulum” CBS 100551 Cosmospora 86 Cosmospora vilior Cosmospora khandalensis CBS 356.65 T Acremonium recifei CBS 400.85 Acremonium recifei CBS 976.70 Acremonium recifei CBS 411.91 Acremonium recifei CBS 596.74 Acremonium recifei CBS 505.94 Acremonium recifei CBS 220.84 Acremonium recifei CBS 555.73 Acremonium recifei CBS 362.76 88 Acremonium recifei CBS 137.35 T Acremonium recifei CBS 135.71 Acremonium recifei CBS 541.89 Acremonium recifei CBS 402.89 Acremonium recifei CBS 442.66 “Acremonium strictum” CBS 106.23 Fusarium dimerum 89 Fusarium domesticum 100 Fusarium oxysporum 1 91 Fusarium oxysporum 2 99 Fusarium Fusarium verticillioides Persiciospora africana -clade 81 Nectria rigidiuscula 87 Fusarium falciforme 70 Fusarium solani Fusarium lichenicola Neocosmospora endophytica Acremonium tsugae CBS 788.69 T Cylindrocarpon cylindroides 89 Heliscus lugdunensis Calonectria morganii Nectria radicicola Leuconectria clusiae Dematiocladium celtidis Nectria cinnabarina 90 “Acremonium recifei” CBS 485.77 “Acremonium recifei” CBS 482.78 “Acremonium recifei” CBS 110348 Nalanthamala diospyri T Nalanthamala guajavae 85 100 Nalanthamala vermoesenii Rubrinectria olivacea ‘Nectria’ rishbethii CBS 496.67 T Acremonium exiguum CBS 587.73 T 96 “Acremonium implicatum” CBS 397.70B exiguum 92 “Acremonium potronii” CBS 416.68 -clade 93 “Acremonium potronii” CBS 781.69 Nectriopsis squamulosa 1 100 Nectriopsis squamulosa 2 “Cephalosporium ballagii” CBS 134.33 T pseudozeylanicum 97 “Acremonium aff. curvulum” CBS 113275 77 Acremonium pseudozeylanicum CBS 560.73 T -clade Niesslia exilis 1 Acremonium nigrosclerotium CBS 154.72 T Niesslia exilis 2 D Fig. 2. (Continued). www.studiesinmycology.org 151 98 97 0.01 substitutions/site Fig. 2. (Continued). 152 Cordycipitaceae Clavicipitaceae Hypocreaceae 80 Cordyceps ramosipulvinata Melanospora brevirostris 100 83 Bertia moriformis Phaeoacremonium aleophilum Ophiocordyceps bispora Elaphocordyceps longisegmentis Haptocillium sinense Haptocillium zeosporum 99 Paecilomyces lilacinus 2 Paecilomyces lilacinus 1 82 Isaria takamizusanensis 98 Chaunopycnis alba 1 Chaunopycnis alba 2 Chaunopycnis pustulata Tolypocladium inflatum Verticillium leptobactrum CBS 109351 94 Simplicillium lanosoniveum CBS 321.72 92 78 Simplicillium lanosoniveum 98 Simplicillium obclavatum T Simplicillium lamellicola T Beauveria bassiana 74 Cordyceps militaris Pleurodesmospora coccorum Isaria javanica Syspastospora parasitica 91 Hyperdermium bertonii Lecanicillium antillanum Lecanicillium aranearum T Lecanicillium fusisporum T Lecanicillium psalliotae 1 Lecanicillium dimorphum T Lecanicillium aff. psalliotae 2 Lecanicillium lecanii Paecilomyces farinosus Lecanicillium attenuatum Epichloe typhina Pochonia bulbillosa 2 Pochonia bulbillosa 1 99 Pochonia gonioides Pochonia rubescens Metarhizium anisopliae var. frigidum Pochonia chlamydosporia var. catenulatum T 95 Verticillium epiphytum 2 Verticillium epiphytum 1 Torrubiella pruinosa 98 Torrubiella petchii Echinodothis tuberiformis Verticillium pseudohemipterigenum Ophionectria trichospora Torrubiella luteorostrata Acremonium camptosporum CBS 756.69 T Acremonium camptosporum CBS 835.91 camptosporum Acremonium camptosporum CBS 757.69 Acremonium camptosporum CBS 677.74 99 Acremonium camptosporum CBS 890.85 -clade Clypeosphaeria phillyreae Blistum tomentosum Neomunkia sydowii Hypocrea koningii Hypomyces chlorinigenus Sphaerostilbella aureonitens Sporophagomyces chrysostomus 81 Cladobotryum rubrobrunnescens Hypomyces subiculosus Mycogone rosea Verticillium incurvum Acremonium psammosporum CBS 590.63 T Ceratocystis fimbriata Ophiostoma piluliferum 2 Glomerella cingulata Ophiocordycipitaceae SuMMerbell et al. e acremonium phylogeny Another major hypocrealean Acremonium clade in Fig. 1A contains A. breve, A. radiatum, A. gamsii and, more distantly, with 96 % bootstrap support, A. curvulum. In Fig. 2C where phylogenetic signal is lower, A. curvulum loses its tight association with A. breve and its relatives and appears in unsupported juxtaposition with the genus Trichothecium and the corresponding teleomorph genus, Leucosphaerina. The clade containing Trichothecium roseum and Leucosphaerina indica (see taxonomic comments below) also contains two anamorph species that were long placed in different genera based on conidiogenesis, namely, Acremonium crotocinigenum and Spicellum roseum, here recombined into Trichothecium. The next clade in Fig. 1A is a loosely structured assemblage consisting of members of Acremonium subgenus Gliomastix, some of which are delineated below as members of a phylogenetically delineated genus Gliomastix, plus the teleomorphic genera Bionectria, linked to the well known penicillate hyphomycete anamorph genus Clonostachys (Schroers 2001), Hydropisphaera, and Roumegueriella. As Fig. 2B shows in more detail, the type species of the genus Gliomastix, originally named Gliomastix chartarum but currently called G. murorum, is in a relatively well supported clade (92 % bootstrap support) along with G. masseei, G. polychroma, and G. roseogrisa, three other species with melanised conidia that were placed in Acremonium subg. Gliomastix by Gams (1971). Related to Gliomastix are two clades of non-melanised Acremonium species placed in A. subg. Gliomastix, the A. persicinum clade, and A. pteridii clade. Smaller clades containing species in A. subg. Gliomastix such as A. biseptum, A. cerealis, A. luzulae, and A. rutilum (= A. roseum) are included in the large Gliomastix/Bionectria clade, which has 78 % bootstrap support. This clade includes additional teleomorphic fungi such as Heleococcum, Hydropisphaera, Nectriopsis, Ochronectria, Selinia, and Stephanonectria, along with the anamorph Sesquicillium microsporum. The Gliomastix/Bionectria clade, the sclerotigenum/Geosmithia clade, and other members of the Bionectriaceae form a weakly supported clade with 74 % bootstrap value as shown in Fig. 1A. Included in the sclerotigenum/Geosmithia clade is the penicillate anamorph genus Geosmithia sensu stricto and the ex-type isolates of Acremonium pinkertoniae and A. sclerotigenum as well as the cephalosporin-producer Acremonium chrysogenum and its close relative, the thermophilic A. lavum. It also includes non-type isolates identiied as A. blochii and A. egyptiacum. In the LSU-tree (Fig. 2A) the sclerotigenum/Geosmithia clade includes an extensive group of Acremonium species and cleistothecial Bionectriaceae with Acremonium-like anamorphs, namely, Emericellopsis, Hapsidospora, Mycoarachis, and Nigrosabulum. Among the anamorphic species in this group are most of the phylogenetically disparate isolates identiied as the type species of Acremonium, A. alternatum. One of these, CBS 407.66, is designated below as epitype of A. alternatum. Prominent subclades include the Acremonium sclerotigenum clade containing the ex-type isolates of A. sclerotigenum and A. sordidulum. Another major, well supported bionectriaceous subclade associated with the sclerotigenum clade is the Emericellopsis clade (Fig. 2A). It includes the type species of the synnematal hyphomycete genus Stilbella, S. imetaria (Seifert 1985) as well as the type of Stanjemonium (Gams et al. 1998) and the marine Acremonium tubakii sensu stricto and A. fuci (Zuccaro et al. 2004). Stilbella imetaria is closely related to the ex-type isolate of Acremonium salmoneum isolated from dung, also a typical habitat for S. imetaria (Seifert 1985). An adjacent weakly www.studiesinmycology.org supported clade includes Hapsidospora, Mycoarachis, and Nigrosabulum, and the two Acremonium species named for yellow pigmentation, A. chrysogenum and A. lavum. Although associated with A. chrysogenum and A. lavum in Fig. 1B, A. pinkertoniae and A. borodinense form a distinct clade in Fig. 2A along with an isolate included in the polyphyletic A. blochii (CBS 993.69), plus the cleistothecial teleomorphs Bulbithecium hyalosporum and Leucosphaerina arxii, both of which have unnamed Acremonium anamorphs. In Fig. 2A, the A. chrysogenum subclade appears to be distinct from the other clades containing A. sclerotigenum, Emericellopsis, and Geosmithia. The other clades within the overall sclerotigenum/Geosmithia clade include the A. fusidioides clade containing several acremonia forming similar conidial chains (A. cavaraeanum, A. fusidioides, A. hansfordii, A. hennebertii, one of the isolates labeled A. alternatum). A small A. brachypenium clade associated with the A. sclerotigenum clade includes A. brachypenium plus the ex-type strain of Cephalosporium purpurascens placed by Gams (1971) in A. persicinum. There is also an isolate of the polyphyletic, untypiied species A. potronii. Basal to these clades is another small clade that links an entomogenous isolate identiied as Verticillium insectorum with two isolates from human sources identiied as A. blochii; these conidial chain-forming isolates are sister to an isolate of the chain-forming entomogenous species Acremonium zeylanicum. The “A. blochii” isolate CBS 427.93, linked with a 99 % bootstrap value to A. pinkertoniae in Fig. 1B, is one of the two isolates associated with Acremonium zeylanicum in Fig. 2A. Adjacent and loosely linked to the bionectriaceous clades in Fig. 2B is a small clade in Fig. 2C containing the ex-type isolate of Acremonium incrustatum plus an isolate labeled A. potronii, and a sequence attributed to Linkosia fusiformis, although this sequence most likely represents a contaminant. Below the sclerotigenum/Geosmithia clade in Fig. 1A and above the Hypocreaceae in Fig. 2E fall clades representing the Clavicipitaceae sensu lato. These clades include the families Clavicipitaceae sensu stricto, Ophiocordycipitaceae, and Cordycipitaceae. Although many species in this group of three families have Acremonium-like anamorphic states, only two described Acremonium species are associated here. In Fig. 2E A. camptosporum sits basally in a clade adjacent to the Clavicipitaceae and is close to the poorly understood teleomorphic species Clypeosphaeria phillyreae, assuming the latter is correctly associated with the sequence attributed to it. Simplicillium obclavatum, originally described as Acremonium obclavatum, provides the only other clavicipitaceous species in Fig. 2 representing a named species of Acremonium. Below the Clavicipitaceaceae in Fig. 1B is a clade of ambiguous afinities containing Acremonium guillematii, A. minutisporum and A. vitellinum. This group also appears as two to three unafiliated clades in Fig. 2C. An insigniicant branch in Fig. 1B subtends Acremonium exiguum, A. psammosporum, and an isolate identiied as A. potronii. In Fig. 2D, just A. exiguum and the A. potronii entity remain associated while Acremonium psammosporum segregates into a basal hypocrealean clade of its own in Fig. 2E. The Nectriaceae is represented by Nectria cinnabarina in Fig. 1B along with the ex-type isolate of the tropical opportunistic pathogen of humans, Acremonium recifei. Fig. 2D shows A. recifei subtending multiple taxa with three non-type isolates splitting off as a separate clade. These clades have approximately the same status in the Nectriaceae as the genus Nalanthamala, including N. diospyri, the former Acremonium diospyri. Another nectriaceous Acremonium in Fig. 2D is A. tsugae, which is closely related 153 SuMMerbell et al. to Cylindrocarpon cylindroides. The broad morphotaxonomic concept of Acremonium berkeleyanum is polyphyletic consisting of isolates placed in the nectriaceous genus Cosmospora (Fig. 2D). Acremonium berkeleyanum sensu lato is represented in Fig. 2D by the newly recombined Cosmospora species, C. lavitskiae and C. khandalensis based on the ex-type isolates of Gliomastix lavitskiae and Cephalosporium khandalense (Gräfenhan et al. 2011). Another purported synonym of A. berkeleyanum, a Cadophora isolate received as A. butyri CBS 301.38, falls outside the Hypocreales (Fig. 1C). Basally in the Hypocreales in Fig. 1B, Acremonium roseolum appears in loose association with Stachybotrys species. In Fig. 2D, it appears in a clade along with the teleomorph Scopinella solani and three Acremonium inlatum isolates, including CBS 403.70, an atypical, catenate-conidial isolate identiied at CBS as A. atrogriseum. Nearby but statistically unlinked clades include Stachybotrys and allied fungi such as Peethambara spirostriata and Didymostilbe echinoibrosa (Castlebury et al. 2004). Acremonium nigrosclerotium represents an isolated Acremonium near the families Hypocreaceae and Niessliaceae (Fig. 1B). In Fig. 2D, A. nigrosclerotium is intercalated among two genotypes ascribed to N. exilis, and loosely associated (77 % bootstrap) with Acremonium pseudozeylanicum and the type culture of Cephalosporium ballagii, currently in synonymy with Acremonium charticola (Gams 1971). A distant outlier is Acremonium lichenicola at the bottom of Fig. 1C. This isolate, CBS 425.66, chosen to represent this species in lieu of ex-type material, blasts as a pezizalean fungus with afinities to another hyaline, phialidic fungus, Phialophora alba. A number of genera in addition to Acremonium were investigated for possible afinity with Acremonium clades as shown in Fig. 2. The sporodochial genus Sarcopodium was investigated and found to split into two groups (Fig. 2C, D). One isolate identiied as S. circinatum grouped with Sarcopodium circinosetiferum and S. vanillae in a widely separated clade along with Lanatonectria teleomorphs and a sequence identiied as Pseudonectria rousseliana (Fig. 2C). This clade appeared in LSU sequencing to be independently situated within the Hypocreales. Acremonium rhabdosporum appeared as a statistically unsupported, possible distant relative. The other two isolates of S. circinatum formed a clade near Myrothecium in the Stachybotrys/Peethambara clade (Fig. 2D). Also appearing in this clade was Parasarcopodium ceratocaryi, a monotypic genus recently described by Mel’nik et al. (2004). DIsCUssION The main morphotaxonomic groundwork for Acremonium as conceived in the late 20th century was laid by Gams (1971) in his monograph Cephalosporium-artige Schimmelpilze (Hyphomycetes). This monograph was radically more comprehensive than previous treatments of the species and was followed by several key adjunct studies, including but not limited to Gams & Lacey (1972), Gams (1975), and Ito et al. (2000). Gams’ studies were based on a meticulous morphological observation scheme that involved growing species on appropriate media, e.g., oatmeal agar, and then making camera lucida drawings that could be directly compared with subsequent isolates. The comparison was done by superimposing the virtual image of the new isolate directly over the camera lucida drawings of previous isolates drawn 154 at the same scale. This highly rigorous approach was necessary for a group of hyphomycetous fungi so morphologically simpliied as Acremonium. Gams (1971, 1975) also discovered a subtle character that allowed him to associate dark-conidial species, monographed by Dickinson (1968) as the genus Gliomastix, with numerous biologically related hyaline-conidial species. This character was "chondroid hyphae," which could be seen under the microscope as hyphae with wall thickenings, and which makes colonies somewhat resistant to being cut with a scalpel. The species Gams (1971) united using this character are, for the most part, grouped in the Gliomastix/Bionectria clade referred to earlier in this study. Despite the rigorous approach and the discovery of new, useful characters, a number of the morphotaxonomic species names ultimately were applied in the CBS collection to phylogenetically divergent organisms. Six distinct taxa from CBS investigated in this study were identiied as A. persicinum; three are now seen phylogenetically to fall within the Gliomastix clade and three sort elsewhere. These taxa are mostly directly visible as A. persicinum isolates in Fig. 2. Names without quotation marks are consistent with the type, while names in quotation marks sort into other phylogenetic groups. An exception is represented by CBS 149.62. This isolate, the ex-type of Cephalosporium purpurascens, was listed by Gams (1971) as a synonym of A. persicinum. Five taxa in Fig. 2 were labeled A. potronii in CBS, and four were called A. strictum. Within both A. potronii and A. strictum, as conceived morphologically, some isolates fall within A. sclerotigenum. The name A. alternatum was applied to four species, three of them visible in Fig. 2, plus isolates of A. sclerotigenum with catenulate conidia. “Acremonium blochii” was applied to three different species. Phylogenetic analysis compared to the morphological treatment of Acremonium Gams (1971, 1975) divided Acremonium into three major sections, Simplex, a name later updated as the type section Acremonium, Gliomastix, and Nectrioidea. Of these sections, only Gliomastix withstands phylogenetic scrutiny as a unit, albeit a loosely associated one. The type section Acremonium contained four widely phylogenetically scattered major clades (Fig. 2), speciically the A. sclerotigenum clade, Sarocladium clade, A. curvulum clade, and A. breve clade. As seen best in Fig. 1, the Sarocladium clade and the A. breve and A. curvulum clades comprise a distinct group that falls within the Hypocreales but outside any currently recognised family. Acremonium sclerotigenum falls into a distinct clade within the Bionectriaceae that also contains Emericellopsis and Geosmithia. This clade also includes about half the investigated CBS isolates identiied as the type species of Acremonium, A. alternatum, including CBS 407.66 as well as some isolates such as CBS 223.70 revealed as morphological variants of A. sclerotigenum. Despite the substantial phylogenetic distance between A. sclerotigenum and A. strictum, relatively glabrous, cylindrical-conidial isolates of A. sclerotigenum not producing sclerotia on special media (lupine stem agar according to Gams, 1971, later replaced at CBS by nettle stem agar) are essentially micromorphologically indistinguishable from A. strictum. Table 1 shows CBS 287.70 O as an A. sclerotigenum isolate identiied in CBS as A. strictum; ITS sequencing studies of additional strains (data not shown) have found two more such isolates, CBS 319.70 D and CBS 474.67. acremonium phylogeny The convergence among isolates of phylogenetically remote species is remarkable. An unknown proportion of the literature on A. strictum is based on studies of A. sclerotigenum. For example, in a study inluential in medical mycology, Novicki et al. (2003) labeled ITS-sequenced isolates of A. sclerotigenum in GenBank as "Acremonium strictum genogroup II." The complexity of A. sclerotigenum, not its earliest valid name, goes beyond the scope of this paper. Perdomo et al. (2010) have recently investigated the diversity of medically important isolates within this species. Besides the four clades mentioned above, Acremonium sect. Acremonium species also make up the non-synnematal anamorphs of the Emericellopsis clade, most of the A. fusidioides clade, and most of the small A. camptosporum, A. exiguum, A. minutisporum, A. pinkertoniae, and A. pseudozeylanicum clades. Gams (1975) accommodated A. byssoides, now known to belong in Simplicillium lanosoniveum (Zare & Gams 2001), in Acremonium sect. Acremonium, while commenting that it was suggestive of Verticillium sect. Prostrata, later recognised as Simplicillium (Zare & Gams 2001). He withheld A. byssoides from Verticillium because the colony margin was relatively lat and slightly fasciculate, rather than cottony. To some extent Acremonium sect. Acremonium was based on keying out all the relatively lat or fasciculate Acremoniumlike species together provided that they lacked the dark conidia or chondroid hyphae of Gliomastix. Acremonium sect. Nectrioidea as delineated by Gams (1971) included many Nectria sensu lato anamorphs. Some of these species are now placed in the genus Cosmospora by Gräfenhan et al. (2011). These include members of the A. berkeleyanum complex as well as A. arxii and A. cymosum. Acremonium falciforme in A. sect. Nectrioidea had already been recognised as a member of the Fusarium solani complex (Summerbell & Schroers 2002) and A. diospyri had been transferred into Nalanthamala along with other nectriaceous species (Schroers et al. 2005). Acremonium tsugae appears to be a microconidial Cylindrocarpon species. The Acremonium recifei complex still remains as an undisposed major group of nectriaceous Acremonium species originally included in A. sect. Nectrioidea. The placement of A. sect. Nectrioidea species A. alcalophilum, A. brunnescens, A. furcatum, A. nepalense, A. restrictum, and A. stromaticum in the Plectosphaerellaceae has already been shown by Zare et al. (2007). Acremonium apii also has been shown to belong to this family as a synonym of Verticillium alboatrum, and its ex-type strain, CBS 130.51, was used as the representative isolate of that species by Zare et al. (2007). Other anomalous elements of A. sect. Nectrioidea include A. crotocinigenum in the Trichothecium clade, A. radiatum in the phylogenetically isolated A. breve clade, A. biseptum in the A. cerealis clade near Gliomastix, A. salmoneum in the Emericellopsis clade near Stilbella imetaria, A. chrysogenum in a bionectriaceous clade containing cleistothecial teleomorphs such as Nigrosabulum, A. rutilum in a clade otherwise containing isolates identiied as A. persicinum, and a non-type A. hyalinulum isolate in another clade peripheral to Gliomastix. When Sarocladium zeae as A. zeae in A. sect. Nectrioidea was compared to the phylogenetically related S. kiliense as A. kiliense in A. sect. Acremonium by Gams (1971, p. 16), he noted that the latter species may sometimes also be strongly branched and thus resemble the former. The exigencies of dichotomous morphological keying tended to sort closely related species into widely separated Sections of the genus. The main heterogeneous element included in Gams’ (1971) original concept of sect. Gliomastix was the “Striatisporum series.” These were later distinguished as the separate genus Sagenomella (Gams 1978). Both Sagenomella and the recently described genus www.studiesinmycology.org Phialosimplex are members of the Eurotiales (Sigler et al. 2010). Another anomalous element in sect. Gliomastix, Acremonium atrogriseum, is here removed to the Cephalothecaceae. Other species included by Gams (1971, 1975) in A. sect. Gliomastix that can now be seen to be separated from the Gliomastix/Bionectria clade include “Cephalosporium purpurascens,” synonymised by Gams (1971) with A. persicinum as well as A. brachypenium, A. hennebertii, A. incrustatum, and A. inlatum. Species outside the Gliomastix/Bionectria clade that have well developed chondroid hyphae include A. hennebertii and A. incrustatum. TAxONOMy The main purpose of this study is to provide a phylogenetic overview of Acremonium plus distinctive LSU sequences to render the described species recognisable in molecular studies. In addition, some taxonomic changes are undertaken. What is Acremonium? The irst task at hand is to establish what Acremonium is. The lectotype species of Acremonium is A. alternatum as designated by Gams (1968). Gams (1968) studied and illustrated the type material used by Link (1809) in describing A. alternatum. This material consists of a thin fungal mycelium colonising a birch leaf. In choosing living cultures that best approximated this specimen, Gams (1968) listed four isolates. From among these, one is chosen with a dried culture to be designated here as the epitype with an ex-epitype culture. This is CBS 407.66, which groups with the extype isolate of Cephalosporium malorum, synonymised by Gams (1971) with A. charticola, as well as with A. sordidulum and A. charticola in the poorly deined A. sclerotigenum/Geosmithia clade. Use of the corresponding dried culture CBS H-20525 as an epitype specimen serves nomenclatural stability because the genus name Acremonium is then used to designate a large group of species currently accepted in Acremonium. Other candidate isolates included CBS 308.70 (called "Kultur 1127"), which died out and was replenished from its degenerated, nonsporulating subculture MUCL 8432, now also called CBS 114602. As a degenerated isolate, it makes poor potential epitype material. Another isolate mentioned by Gams (1968), CBS 406.66, is conspeciic with CBS 114602 and in good condition. Both isolates are included in a clade relatively distant from any other Acremonium group but deeply basal to the Sarocladium and A. breve clades, as seen in Fig. 1A. If Acremonium were epitypiied with one of these isolates, the generic name might be restricted to this single species. The inal isolate is CBS 223.70, an isolate that, despite its catenate conidia, is conspeciic with the type of A. sclerotigenum (100 % ITS sequence identity; GenBank AJ621772 for CBS 124.42 is essentially identical to A. sclerotigenum, U57674, CBS 223.70). Isolate CBS 223.70 strongly resembles pale greenish grey coloured, sclerotium-forming isolates identiied as A. egyptiacum (e.g., CBS 734.69), which are also conspeciic with A. sclerotigenum. It differs by not forming sclerotia. Catenate conidia may or may not be produced in this group and the greenish grey colonies produced by chain-forming isolates have explicitly been connected with A. egyptiacum, not A. alternatum. One other taxon that Gams (1971, 1975) consistently identiied as A. alternatum, a species in the A. fusidioides clade, is represented by CBS 831.97 155 SuMMerbell et al. and 381.70A. These isolates have the disadvantage of not having been explicitly compared with the type material. In addition, this clade is related to several clades with known teleomorphs, e.g., Emericellopsis and Nigrosabulum, and anamorphs, e.g., Stilbella and Geosmithia. In a revised nomenclatural system, it would root Acremonium as a broad unitary genus name encompassing the teleomorphs and complex anamorphs. Ultimately, it might epitypify Acremonium strictly as a genus name for the A. fusidioides clade. Acremonium alternatum Link : Fr., Mag. Ges. naturf. Fr. Berlin 3: 15. 1809 : Fries, Syst. Mycol. 3: 425. 1832. Holotype: Germany, Rostock, on leaf litter of Betula, collected by Ditmar, B-type specimen labeled in Link’s handwriting. Epitype designated here: Austria, Stangensteig near Innsbruck, ex Ustulina deusta, W. Gams, Dec. 1965, CBS-H 20525 dried culture of CBS 407.66, ex-epitype living culture CBS 407.66. Additional genera recognised here 1958). The distinction between G. murorum var. murorum having conidia in chains and G. murorum var. felina having conidia in mucoid heads does not appear to be supported by phylogenetic analysis. Gliomastix murorum var. felina isolates originally described as Graphium malorum (ex-type CBS 154.25) and Torula cephalosporioides (ex-type CBS 378.36) are molecularly conirmed as synonyms of G. murorum (Fig. 2B). Recently, Kiyuna et al. (2010) neotypiied Gliomastix felina (Marchal) Hammill, recombined as Acremonium felinum (Marchal) Kiyuna, An, Kigawa & Sugiy., with CBS 147.81. The sequences deposited in GenBank, e.g., AB540562, suggest that this isolate represents G. roseogrisea. The new combination is reduced to synonymy with that species below. 2. Gliomastix masseei (Sacc. ) Matsush., Icon. microfung. Matsush. lect. (Kobe): 76. 1975. Basionym: Trichosporium masseei Sacc., Syll. Fung. 22: 1356. 1913 [= Trichosporium aterrimum Massee, Bull. Misc. Inform. 1899: 167 non (Corda) Sacc. 1886] ≡ Acremonium masseei (Sacc.) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 83. 1971. Based on these analyses, three genera are represented in suficient detail and with high bootstrap support to be formally recognised here. In most cases, the genera and clades are not suficiently populated with their constituent members without analysis of additional sequences. For example, the Emericellopsis clade is missing 12 of its 13 species including two identiied as E. minima (Zuccaro et al. 2004) as well as one of its two Stanjemonium species. The name lacks an ex-type isolate. Although the isolate (CBS 794.69) sequenced is basal to the Gliomastix clade (Fig. 2B), it appears to be a suitable to serve as the basis for epitypiication. Gliomastix Epitype designated here: Italy, Turin, isolated from rabbit dung, A. Fontana, CBS H-8244, ex-epitype culture CBS 794.69. The core clade of Gliomastix including the type species is well delimited with a 92 % bootstrap value even in the very conservative LSU analysis. Although Gams (1971) placed this genus into Acremonium, several authors have recognised Gliomastix. Most notably, Matsushima (1975) placed Acremonium masseei and A. polychromum into Gliomastix and Lechat et al. (2010) linked G. fusigera with Hydropisphaera bambusicola. As circumscribed in this paper, the phylogenetically supported Gliomastix differs from previous morphological concepts by excluding several distantly related species such as Acremonium cerealis and A. inlatum. The closely related A. persicinum clade may also be included as suggested by Supplemental ig. 6E in Schoch et al. (2009) and discussed above. At the moment, we recognise only four species from the present study in Gliomastix. An additional species, published while the present manuscript was in preparation, Acremonium tumulicola (Kiyuna et al. 2010), should also be included in this concept of Gliomastix. The generic characters do not differ signiicantly from those summarised in the generic diagnosis of Dickinson (1968). 1. Type species. Gliomastix murorum (Corda) S. Hughes, Canad. J. Bot. 36: 769. 1958. Basionym: Torula murorum Corda, Icon. Fung. 2: 9. 1838. ≡ Sagrahamala murorum (Corda) Subram., Curr. Sci. 41: 49. 1972. ≡ Acremonium murorum (Corda) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 84. 1971. = Torula chartarum Corda, Icon. Fung. 2: 9. 1839. ≡ Gliomastix chartarum (Corda) Guég, Bull. Soc. Mycol. France 21: 240. 1905. For additional synonyms, see Gams (1971). The type species of Gliomastix, G. chartarum, is a synonym of G. murorum (Hughes 156 Holotype of Trichosporium masseei: India, Punjab, Changa Manga, on Morus indica, Jan. 1898, J. Gleadow, ex Herb. Massee, K; isotypes IMI 49,214 = IMI 87,346. 3. Gliomastix polychroma (J.F.H. Beyma) Matsush., Icon. microfung. Matsush. lect. (Kobe): 77. 1975. Basionym: Oospora polychroma J.F.H. Beyma, Verh. K. Ned. Akad. Wetensch., Sect. 2, 26 (2): 5. 1928. ≡ Sagrahamala polychroma (J.F.H. Beyma) Subram., Curr. Sci. 41: 49. 1972. ≡ Acremonium polychromum (J.F.H. Beyma) W. Gams, Cephalosporiumartige Schimmelpilze (Stuttgart): 81. 1971. Additional synonyms are given by Gams (1971). This clade includes the ex-type isolate of Oospora polychroma, basionym of G. polychroma, CBS 181.27 (Fig. 2B). Periconia tenuissima var. nigra is conirmed as a synonym via inclusion of its ex-type isolate CBS 151.26 (Fig. 2B). The status of the different isolate, CBS 617.94, from banana, requires further clariication. This isolate may be related to Acremonium musicola, a species not represented in CBS. 4. Gliomastix roseogrisea (S.B. Saksena) Summerbell, comb. nov. MycoBank MB519588. Basionym: Cephalosporium roseogriseum Mycologia 47: 895. 1956 [1955]. S.B. Saksena, ≡ Acremonium roseogriseum (S.B. Saksena) W. Gams [as ‘roseogriseum’], Cephalosporium-artige Schimmelpilze (Stuttgart): 87. 1971. = Acremonium felinum (Marchal) Kiyuna, An, Kigawa & Sugiy., Mycoscience 52: 13. 2010. Gliomastix roseogrisea, like G. murorum, has a variety of conidial forms including conidia in chains and conidia of various shapes in mucoid heads. This plasticity of form recalls the situation mentioned acremonium phylogeny above for A. sclerotigenum and may represent a relatively common situation in acremonioid species. As another example Gams (1971) lists “Gliomastix murorum var. felina pro parte in Dickinson in Mycol Pap. 115: 16, 1968” as an additional synonym of this taxon. As mentioned above in the discussion of the genus, Kiyuna et al. (2010) recently neotypiied Gliomastix felina (basionym Periconia felina Marchal, Bull. Soc. R. Bot. Belg. 34:141. 1895) with CBS 147.81, an isolate collected by Hammill (1981). This isolate is a typical G. roseogrisea, a taxon not studied by Kiyuna et al. (2010). 5. Gliomastix tumulicola (Kiyuna, An, Kigawa & Sugiy.) Summerbell, comb. nov. MycoBank MB519599. Basionym: Acremonium tumulicola Kiyuna, An, Kigawa & Sugiy., Mycoscience 52: 13. 2010. This newly described species is phylogenetically placed by its original authors (Kiyuna et al. 2010) in the Gliomastix clade and comparison of sequences conirms that placement. Although this information was received too late to include this species in our phylogenetic analyses, the species is placed in Gliomastix. Sarocladium The genus Sarocladium was described for two pinkish coloured fungal pathogens causing sheath blast of rice (Gams & Hawksworth 1976). The drawings in that paper and the photographs in Bills et al. (2004) show structures that overlap with those produced by the phylogenetically related A. kiliense, A. strictum, and A. zeae. As in Fusarium, plant pathogenic fungi that sporulate on above-ground plant parts are likely to produce upright, branching sporulating structures with mucoid conidia suggesting dispersal by insects that ly from plant to plant. Species with habitats where water lux or microarthropod movement may be important in dispersal, e.g., various Acremonia occurring in soil or Fusarium domesticum growing on cheese, may have simpliied conidiogenous structures. Bills et al. (2004) suggested that the generic placement of Acremonium kiliense and A. strictum should be re-examined in light of their close relationship with Sarocladium oryzae. The genus Sarocladium is delineated here to include several species previously recognised in Acremonium, as seen in Figs 1 and 2. In Fig. 2, where phylogenetic signal is relatively low, Sarocladium tepidly (84 % bootstrap) links to the A. bacillisporum clade. In Fig. 1, it links with a 99 % bootstrap value. Phylogenetic clustering algorithms often insert the A. bacillisporum clade between A. strictum and A. kiliense due to certain apo- or plesiomorphies shared with one or the other of these two members of the A. strictum clade (data not shown). On the other hand, the next most closely related clade in Fig. 1, the A. breve/A. curvulum clade, has ITS sequences with substantial sections that are dificult to align with those of the A. bacillisporum and A. strictum clades, indicating considerable evolutionary distance. The genus Sarocladium is emended here to include those species that belong to the A. strictum and A. bacillisporum clades. The generic name Sagrahamala is not a contender for this group because the type species is the unrelated Acremonium luzulae. In addition Acremonium luzulae is a species in need of epitypiication, because, as shown in the present study, more than one phylogenetic species is encompassed under the name. Sarocladium W. Gams & D. Hawksw., Kavaka 3: 57. 1976 [1975]. www.studiesinmycology.org Colonies on 2 % malt extract agar slimy-glabrous to moderately loccose to deeply dusty, sometimes ropy; with, in Gams’ terminology (Gams 1971), phalacrogenous, nematogenous, to plectonematogenous conidiation; growing 13–25 mm in 10 d at 20 °C, whitish to pinkish to salmonaceous or, when conidia are formed in chains, sometimes acquiring vivid conidial mass colouration such as ochraceous or greenish glaucous; reverse pale to pinkish orange to pale grey-brown, rarely greenish-blue. Conidiogenous apparatus ranging from adelophialides, solitary orthotropic phialides to conidiophore structures with one or a few branches, or with cymose branching or occasionally with one or two ranks of loosely structured verticils, sometimes with repeated branching extending to 90 μm long. Phialides subulate, aculeate to acerose, straight, slightly curved, or undulate, thin- and smooth-walled, 15–60(–75) μm long, tapering from a basal width of 1.2–2.5 μm, with minimal collarette; conidia borne in mucoid heads or dry chains, notably longer than broad, l/w mostly 2.2–7.0, cylindrical to fusiform to bacilliform, aseptate, smooth-walled, with rounded or tapered-truncate ends, 3.5–8(–14) × 0.5–2 μm. Chlamydospores present or absent, when present relatively thick-walled, smooth or slightly roughened, globose to ellipsoidal, intercalary or terminal, mostly solitary, occasionally in short chains, 4–8 μm. Internal transcribed spacer sequence mostly with distinctive CGGTCGCGCC motif in mid-ITS2 region. Several species of Sarocladium are noted for melanogenesis yielding ochre-brown to dark grey-brown colony reverse colours on Sabouraud agar: S. glaucum, S. kiliense, and S. zeae (Gams 1971). In the case of S. kiliense, this melanogenesis has the result that most mycetoma cases feature black "grains" or sclerotium-like balls of compacted fungal hyphae (Summerbell 2003); melanogenesis is a well known pathogenicity factor in fungal diseases of humans and animals (Gómez & Nosanchuk 2003). As recognised here Sarocladium yields a remarkable unity of species with elongated conidia and phialides. Several species including S. kiliense, S. oryzae, and S. strictum form adelophialides prominently, at least in some isolates; acremonioid species outside Sarocladium usually lack this character. The recognised species are given below. Acremonium implicatum may belong here, but the species lacks living ex-type or representative material. The “A. implicatum” isolate that grouped in Sarocladium, CBS 243.59, is noted by Gams (1971) as an authentic isolate of Fusidium terricola J.H. Mill., Giddens & A.A. Foster and this name could be used if A. implicatum sensu Gams is revealed as polyphyletic. The other “A. implicatum” isolate, CBS 397.70B, included in this study is not a Sarocladium; rather it is a member of the A. exiguum clade. 1. Type species. Sarocladium oryzae (Sawada) W. Gams & D. Hawksw., Kavaka 3: 58. 1976 [1975]. A description and synonymy are given by Gams & Hawksworth (1975). Bills et al. (2004) synonymised Sarocladium attenuatum with S. oryzae based on the reported identity of the ITS sequence of its ex-type isolate, CBS 399.73, with that of representative isolates of S. oryzae. We resequenced the ITS region of CBS 399.73 and obtained a sequence differing from Bills et al. (AY566995) by 6 base-pairs and 2 gaps. Some of the base pairs in our sequence appeared to be symplesiomorphies shared with A. kiliense or A. strictum but not S. oryzae, rather than random mutations or possible miscalls. Our resequencing of unequivocal S. oryzae isolates CBS 180.74 and CBS 361.75 yielded results consistent with those of Bills et al. (2004). The status of S. attenuatum thus requires further study. 157 SuMMerbell et al. 2. Sarocladium bacillisporum (Onions & Barron) Summerbell, comb. nov. MycoBank MB519589. Basionym: Paecilomyces bacillisporus Onions & G.L. Barron, Mycol. Pap. 107: 11. 1967. ≡ Acremonium bacillisporum (Onions & G.L. Barron) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 72. 1971. ≡ Sagrahamala bacillispora (Onions & G.L. Barron) Subram., Curr. Sci. 41: 49. 1972. This species was described by Gams (1971). It is easily confused with Verticillium leptobactrum, which can be relatively loccose and loosely structured although some isolates are very dense and slowgrowing (Gams, 1971). In addition colonies of S. bacillisporum at maturity have a pinkish colouration. 3. Sarocladium bactrocephalum (W. Gams) Summerbell, comb. nov. MycoBank MB519590. Basionym: Acremonium bactrocephalum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 44. 1971. As indicated by Gams (1971) this uncommon species is closely related to S. strictum, but is distinguished morphologically by its long, narrow conidia. It is molecularly distinguishable by LSU sequences. 4. Sarocladium glaucum (W. Gams) Summerbell, comb. nov. MycoBank MB519591. Basionym: Acremonium glaucum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 68. 1971. This species was described by Gams (1971). The ex-type culture CBS 796.69 indicates that this species belongs in Sarocladium. 5. Sarocladium kiliense (Grütz) Summerbell comb. nov. MycoBank MB519592. Basionym: Acremonium kiliense Grütz, Dermatol. Wochenschr. 80: 774. 1925. = Cephalosporium incoloratum Sukapure & Thirum., Sydowia 19: 171. 1966 [1965]. = Acremonium incoloratum (Sukapure & Thirum.) W. Gams, Cephalosporiumartige Schimmelpilze (Stuttgart): 50. 1971. Additional synonyms and a description of S. kiliense are given by Gams (1971) and Domsch et al. (2007); the species is also extensively illustrated by de Hoog et al. (2000). The ITS sequence of the ex-type strain of Acremonium incoloratum, CBS 146.62, is identical to that of the ex-type of S. kiliense, CBS 122.29 (data not shown). Though isolate CBS 146.62 is unusual in colour and lacks well differentiated chlamydospores that generally occur in S. kiliense, there is no phenetic difference profound enough to suggest that additional genes must be examined to be certain of their synonymy. The sequences deposited in GenBank by Novicki et al. (2003) for their “Acremonium strictum genogroup III” (ITS: AY138846; LSU: AY138484) are actually of S. kiliense. 6. Sarocladium ochraceum (Onions & Barron) Summerbell, comb. nov. MycoBank MB519593. Basionym: Paecilomyces ochraceus Onions & G.L. Barron, Mycol. Pap. 107: 15. 1967. ≡ Acremonium ochraceum (Onions & G.L. Barron) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 67. 1971. ≡ Sagrahamala ochracea (Onions & G.L. Barron) Subram. & Pushkaran, Kavaka 3: 89. 1975 [1976]. 158 This species was described by Gams (1971). We analysed the extype culture, CBS 428.67. 7. Sarocladium strictum (W. Gams) Summerbell, comb. nov. MycoBank MB519594. Basionym: Acremonium strictum W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 42. 1971. Descriptions of S. strictum are given by Gams (1971) and Domsch et al. (2007). The type isolate of S. strictum was conirmed in this genus (Fig. 2C). Of the three isolates illustrated by Gams (1971) under A. strictum, CBS 287.70 D, is conirmed by sequencing as S. strictum. The only isolate of Acremonium zonatum in this study, CBS 565.67, turned out to have an ITS sequence identical to that of S. strictum. This is one of three isolates examined by Gams (1971) as A. zonatum. He stated that another isolate, CBS 145.62, appeared to be A. kiliense, but that examination of herbarium material suggested that this species had been growing on the natural substrate mixed with the real A. zonatum and had been isolated accidentally. One herbarium specimen examined by Gams (1971) showed septate conidia, something not otherwise seen in Sarocladium, so there may indeed be a real A. zonatum. It is not clear if A. zonatum sensu Gams is a uniied concept or a designation of various acremonioid fungi forming leaf spots on tropical plants. In any case, the known connection of the genus Sarocladium with phytopathogenesis and endophytism as in S. zeae makes it plausible that species such as S. strictum and S. kiliense may play a role in plant disease. 8. Sarocladium zeae (W. Gams & D.R. Sumner) Summerbell, comb. nov. MycoBank MB519595. Basionym: Acremonium zeae W. Gams & D.R. Sumner, in Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 121. 1971. This economically important maize endophyte species its the description given by Gams (1971) as a fungus with felty to shaggy colonies. Two S. zeae isolates with more lattened colonies were accessed in CBS as A. strictum. Both CBS 646.75 and 226.84 were from maize and found to be producers of pyrrocidine metabolites as well as dihydroresorcylide, characteristic of S. zeae (Wicklow et al. 2008). Pyrrocidines are antagonistic to Aspergillus lavus and Fusarium verticillioides in maize inlorescences and are thus important in the ecology and economic signiicance of S. zeae. An additional A. strictum isolate, CBS 310.85, is also S. zeae as evidenced by pyrrocidine production, but has not yet been sequenced (Wicklow et al. 2008). Trichothecium A signiicant theme of the current volume is the pioneering of a new approach to dikaryomycete nomenclature: the unitary naming of genus-level clades based on the oldest valid generic name, whether originally anamorphic or teleomorphic in nature (see discussion in Gräfenhan et al. 2011). Because the irst named fungi were often species prominently in contact with humans and their environs and because the irst names usually were attached to the most frequently seen reproductive state, there is considerable wisdom to using the oldest name applied to either aspect of the holomorph in constructing a unitary nomenclature. The genus Trichothecium makes an excellent example, since the system used here preserves the best known species name in the group. A unitary system giving teleomorphs primacy acremonium phylogeny Fig. 3. A. Trichothecium roseum CBS 113334 showing retrogressive conidiation. B-D. conidiogenesis in “Trichothecium indicum”/ Leucosphaerina indica CBS 123.78 showing retrogressive development (B), phialidic development (B) and sympodial development (D). would replace the familiar "T. roseum" with a Leucosphaerina name. A system that retains primacy for morphology, which is the only reasonable basis for dual nomenclature in the molecular era, would divide the Trichothecium clade into four genera, as is the case today. One of those genera, Acremonium, would be quintessentially artiicial and almost completely divorced from evolutionary biological relationships. With increased emphasis on genomes, proteomes, and metabolomes, a focus on polyphyletic elements of microscopic shape seems counterproductive. Every new system of nomenclatural change will entail both fortunate and infelicitous changes and will receive some resistance in scientiic communities. A nomenclatural system based on phylogeny will be considerably more stable than any previous system. The interests of all would be best served if it bridged gracefully out of prephylogenetic taxonomy, preserving as many familiar elements as possible. Trichothecium roseum, a constant from 1809 to today, is one of those elements that is worthy of being preserved. The small, tightly uniied clade of Trichothecium includes isolates with three different anamorphic forms, currently classiied as Acremonium (phialoconidia), Spicellum (sympodial blastoconidia), and Trichothecium (retrogressive blastoconidia). The associated teleomorph, Leucosphaerina indica, produces anamorphic forms described as “Acremonium or Sporothrix” (Suh & Blackwell 1999). These morphs are illustrated by von Arx et al. (1978). The range of anamorphic forms produced by L. indica overlaps those produced by all the anamorphic species in the clade (Fig. 3). The four species studied here, Trichothecium roseum, Acremonium crotocinigenum, Leucosphaerina indica, and Spicellum roseum, have recently been associated with a ifth, newly described species, Spicellum ovalisporum. The dendrogram produced by Seifert et al. (2008) makes it clear that S. ovalisporum is related to S. roseum and is certainly a member of the Trichothecium clade. In parallel with the revision of the genus Microcera by Gräfenhan et al. (2011), this clade is redeined here as a genus with the oldest valid generic name, Trichothecium. As Fig. 2 shows, the second described Leucosphaerina species, L. arxii, is in the distant Acremonium pinkertoniae clade and is closely related to Bulbithecium hyalosporum. Malloch (1989) www.studiesinmycology.org commented that it differed from L. indica by lacking sheathing gel around the ascospores and by having an Acremonium anamorph. Trichothecium Link : Fr., Mag. Gesell. naturf. Freunde, Berlin 3: 18. 1809. = Spicellum Nicot & Roquebert, Revue Mycol., Paris 39: 272. 1976 [1975]. = Leucosphaerina Arx, Persoonia 13: 294. 1987. Older synonymy for the genus is given by Rifai & Cooke (1966). Colonies on malt extract agar 20–40 μm after 7 d at 24 °C, white to salmon orange or salmon pink (Methuen 6-7A2, 4-5A2-3), felty, loccose or lanose, sometimes appearing powdery with heavy conidiation. Ascomatal initials, if present, produced on aerial mycelium, irregularly coiled. Ascomata spherical or nearly so, nonostiolate, colourless or slightly pink, 150–300 μm; ascomatal wall persistent, nearly colourless, 10–13 μm thick, of indistinct hyphal cells; asci uniformly distributed in centrum, clavate to spherical, with thin, evanescent walls, 8-spored, 10–13 μm wide; ascospores ellipsoidal or reniform, with refractile walls and a 1–1.5 μm broad gelatinous sheath, smooth or inely striate, hyaline, yellow to pink en masse, without germ pore, 6–7 × 3–4 μm. Conidiogenous apparatus varying by species, featuring one or more of: conidiophores up to 125 μm long × 2–3.5 μm wide, septate, unbranched, with terminal phialides 10–65 μm long, producing unicellular, hyaline, smoothwalled phialoconidia, obovate, oblong or cylindrical 4.4–7.4 μm; or conidiophores up to 175 μm long, unbranched or uncommonly with one or more branches, retrogressive, shortening with production of each conidium, with each conidial base subsuming a portion of conidiophore apex; conidia 0–1-septate, ellipsoidal or ovate, with a decurved, abruptly narrowed basal hilum terminating in a distinct truncate end, 5–12 × 3–6.5 μm; or conidiophores ranging from unicellular conidiogenous cells to multicellular, multiply rebranched apparati extending indeinitely to beyond 200 μm long; terminal cells 9–37 μm long with a cylindrical basal part and a narrowing, apically extending conidiogenous rachis sympodially proliferating and producing oval to ellipsoidal to cylindrical or allantoid conidia 3.5–11 × 1.5–3.5 μm, with truncate bases. Chlamydospores absent 159 SuMMerbell et al. or present, when present mostly in intercalary chains, hyaline, smooth or inely warted, 5–8(–12) μm wide. Internal transcribed spacer sequence generally with distinct CACAAACCTCGCG motif in ITS2 region. The numerical position varies by species and isolate, cf. position 476 in GenBank record EU445372, ITS for Spicellum ovalisporum ex-type isolate DAOM 186447. Various taxa described as Trichothecium need to be investigated to determine their relationship to this phylogenetic genus. For example, Trichothecium luteum and T. parvum, not represented by living cultures, should be investigated, as should T. campaniforme and T. plasmoparae, which are represented by one isolate each in CBS. Trichothecium domesticum was recently redisposed as Fusarium domesticum (Bachmann et al. 2005). Of teleomorphs reported to have Trichothecium anamorphs, Heleococcum japonense is unrelated to the Trichothecium clade (Fig. 2; the sequence is erroneously listed as H. japonicum in GenBank); rather it is related to Gliomastix and Hydropisphaera. A Trichothecium state of Hypomyces subiculosus (syn. H. trichothecoides) was described, but Hypomyces, a member of the Hypocreaceae, is a remote relative of the Trichothecium clade within the Hypocreales (Fig. 2). 1. Type species. Trichothecium roseum (Pers.) Link, Mag. Gesell. naturf. Freunde, Berlin 3: 18. 1809. Synonymy is given in MycoBank record MB164181. 2. Trichothecium crotocinigenum (Schol-Schwarz) Summerbell, Seifert, & Schroers, comb. nov. MycoBank MB519596. with ITS sequence record EU445372 for the ex-type isolate of S. ovalisporum, DAOM 186447. Two more CBS isolates accessed as S. roseum, CBS 119.77 and CBS 146.78, also gave ITS sequences identical to EU445372. A recent partial ITS sequence made by K.A. Seifert for CBS 227.76 agreed with our sequence (data not shown). No one has thus been able to replicate the sequence given for S. roseum in AB019365 and we are uncertain of its signiicance, even though a similar sequence (GenBank AB019364) has been attributed to two other S. roseum isolates in the JCM collection by the same depositor, G. Okada. If the fallibilities of earlier sequencing chemistries are involved in these discrepancies, S. ovalisporum may be more closely related to T. sympodiale than is evident in the literature. Preliminary results have shown at least one substitution distinguishing the translation elongation factor α sequence of S. ovalisporum from that of T. sympodiale (Rehner, data not shown). Based on comparative morphology and habitat, the authors of S. ovalisporum are conident that their species is distinct, and thus the new combination is included here with their sanction. 5. Trichothecium sympodiale Summerbell, Seifert, & Schroers, nom. nov. MycoBank MB 519600. Basionym: Spicellum roseum Nicot & Roquebert, Revue Mycol., Paris 39: 272. 1976 [1975]. If recombined into Trichothecium, Spicellum roseum would result in a homonym of the type species, thus a new name is needed. Basionym: Cephalosporium crotocinigenum Schol-Schwarz, Trans. Brit. Mycol. Soc. 48: 53. 1965. Acremonium atrogriseum and Acremonium cf. alternatum CBs 109043 in the Cephalothecaceae: a study in comparative morphology vs. phylogeny As pointed out by Seifert et al. (2008, supplement), T. crotocinigenum has long been known to produce crotocin mycotoxins that are similar to the trichothecenes produced by T. roseum and T. sympodiale. The production of similar mycotoxins reinforces the argument for phylogenetic nomenclature such that scientiic names relect true relationships. Acremonium atrogriseum and an isolate identiied as Acremonium cf. alternatum CBS 109043 belong in the Cephalothecaceae (Fig. 1C). This isolate is a white coloured acremonioid fungus forming fusoid conidia in long chains. It also forms small, dark structures that may be aborted ascomatal initials. Sequencing of the ITS region (data not shown) reveals it to be a representative of Phialemonium obovatum. It is identical in all bases but one to the ITS sequence of ex-type strain CBS 279.76 (AB278187) and in all but two bases to another isolate of this species, CBS 116.74. Phialemonium obovatum was described as having conidia in slimy heads (Gams & McGinnis 1983). CBS 109043 shows that either mucoid heads or chains may be formed in this species, as in Acremonium persicinum, A. sclerotigenum, and Gliomastix murorum. Gams (1971) mentions an isolate of Sarocladium bacillisporum that tends to produce mucoid heads. Colonies producing conidia in chains often have a different look from their head-forming conspeciics; the mass colour of the chains may give the colony colours not found in the species descriptions, such as the chalk white colour of CBS 109043 in contrast to the normally pale greenish brown of P. obovatum or the greenish grey of A. sclerotigenum isolate 223.70, in contrast to the normal pale salmon pink of non-catenate A. sclerotigenum. Existing morphological keys and descriptions not just in Acremonium but in all the acremonioid fungi need to be cautiously and skeptically interpreted. At the very least, identiications for publication should be tested by sequencing. We hope that the LSU sequences in this paper will provide the foundation for a phylogenetically sound approach to the systematics and ecology of acremonioid fungi. ≡ Acremonium crotocinigenum (Schol-Schwarz) W. Cephalosporium-artige Schimmelpilze (Stuttgart): 112. 1971. Gams, 3. Trichothecium indicum (Arx, Mukerji & N. Singh) Summerbell, Seifert, & Schroers, comb. nov. MycoBank MB519597. Basionym: Leucosphaerina indica (Arx, Mukerji & N. Singh) Arx, Persoonia 13: 294. 1987. With phylogenetic hindsight, the photographs of this species’ anamorph in the original description by von Arx et al. (1978) can be seen to suggest Acremonium, Spicellum, and Trichothecium. 4. Trichothecium ovalisporum (Seifert & Rehner) Seifert & Rehner, comb. nov. MycoBank MB519598. Basionym: Spicellum ovalisporum Seifert & S.A. Rehner, Fungal Planet: no. 28. 2008. The relationship of the recently described Spicellum ovalisporum to T. sympodiale is not clear. The ex-type of T. sympodiale (CBS 227.76) was resequenced for the ITS region; the resulting sequence differed from the GenBank record AB019365 by 7 gaps and one C ↔ T transition. The sequence had 100 % identity 160 acremonium phylogeny ACKNOWleDGeMeNTs We greatly thank Arien van Iperen, Bert Gerrits-van den Ende, and Kasper Luijsterburg for essential technical support in this study, as well as Keith Seifert and Steve Rehner for scientiic contributions. Key work was done by co-op students Salvatore Lopes, Saghal Ahmed-Suleyman, Arwin van der Rhee, and Nienke Lancee as well as visiting Canadian student Jonathan Shapero. For sending type cultures, we thank Akira Nakagiri of the NITE Biological Resource Centre Fungi collection and Françoise Symoens of the BCCM-IHEM collection. The staff of the CBS Collection deserve special thanks for strain cultivation and additional work. The encouragement and mentorship of Walter Gams is highly appreciated, and we hope our partial resolution of the dilemmas posed by phylogenetic systematics in Acremonium will be recognised as complementary to his invaluable work. ReFeReNCes Arx JA von, Mukerji KG, Singh N (1978). 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