Acremonium phylogenetic overview and revision of ... - CBS - KNAW

available online at www.studiesinmycology.org StudieS in Mycology 68: 139–162. 2011. 

doi:10.3114/sim.2011.68.06 

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 1, 2 

and J.A. Scott 

1 Sporometrics, Inc. 219 Dufferin Street, Suite 20C, Toronto, Ont., Canada M6K 1Y9; 2 Dalla Lana School of Public Health, University of Toronto, 223 College St., Toronto ON 

Canada M5T 1R4; 3 CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; 4 Department of Botany, The Natural History Museum, Cromwell 

Road, SW7 5BD London, United Kingdom; 5 Agricultural Institute of Slovenia, Hacquetova 17, 1000 Ljubljana, Slovenia, Mycology Unit, Medical School; 6 IISPV, 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 identification. 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 five 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 filamentous 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, typified 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. 

Copyright 2011 CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

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 defined, 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 specifically 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 

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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 identification-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 

classification and reliable species identification. 

MATeRIAls AND MeThODs 

Two separate sets of data matrices were assembled (Table 1). 

The first 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 first 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 five 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 fimbriata, 

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 

Medium (Raper & Raper 1972). The LSU region of ribosomal 

DNA (rDNA) was amplified with primers V9G (de Hoog & Gerrits 

van den Ende 1998) and LR5 (Vilgalys & Hester 1990). The SSU 

region was amplified 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 (final elongation) followed by chilling to 4 °C. The PCR 

products were purified with a GFX purification 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 

scientific names. 

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.

www.studiesinmycology.org 

Acremoniella lutzi T 

Petriella setifera 

Doratomyces stemonitis Microascales (1) 

93 Microascus longirostris 

Microascus trigonosporus 

Aniptodera chesapeakensis 

99 

Halosphaeria appendiculata 

80 

96 

99 

Lignincola laevis 

Nimbospora effusa 

Nohea umiumi Halosphaeriales 

95 

Corollospora maritima 

Varicosporina ramulosa 

Ceriosporopsis halima 

Graphium penicillioides Microascales (2) 

Ambrosiella xylebori 

Ceratocystis fimbriata 

Microascales (3) 

92 

89 

Acremonium alcalophilum CBS 114.92 T 

Acremonium restrictum CBS 178.40 T 

Acremonium antarcticum CBS 987.87 

Acremonium stromaticum CBS 863.73 T 

75 

Acremonium furcatum CBS 122.42 T 

Verticillium alboatrum (Acremonium apii T) CBS 130.51 

Verticillium dahliae 

Acremonium nepalense CBS 971.72 T 

Plectosphaerella cucumerina 

96 

70 

Acremonium cucurbitacearum CBS 683.88 

Acremonium brunnescens CBS 559.73 T 

Glomerella cingulata 

“Acremonium alternatum” CBS 406.66 

92 Sarocladium bacillisporum CBS 425.67 T 

“Acremonium implicatum” CBS 243.59 

bacillisporum 

-clade 

96 99 Sarocladium bactrocephalum CBS 749.69 T 

Sarocladium strictum CBS 346.70 T 

Sarocladium kiliense CBS 146.62 

95 

Sarocladium kiliense CBS 122.29 T 

strictum 

-clade 

99 Sarocladium zeae CBS 801.69 T 

Acremonium breve CBS 150.62 T 

Acremonium radiatum CBS 142.62 T breve/curvulum 

96 

Acremonium gamsii CBS 726.71 T 

Acremonium curvulum CBS 430.66 T 

-clade 

98 

“Acremonium blochii” CBS 424.93 

Acremonium persicinum CBS 310.59 T 

persicinum 

-clade 

99 

“Acremonium hyalinulum” CBS 271.36 

Roumegueriella rufula 

75 

Hydropisphaera erubescens 1 

99 

85 Acremonium spinosum CBS 136.33 T 

Acremonium rutilum CBS 396.66 T 

Bionectria ochroleuca 2 

99 “Acremonium blochii” CBS 427.93 

97 Acremonium pinkertoniae CBS 157.70 T 

74 Acremonium chrysogenum CBS 144.62 T chrysogenum 

Acremonium flavum CBS 596.70 T 

Acremonium sclerotigenum CBS 124.42 T 

Geosmithia lavendula 

99 Geosmithia putterillii 

“Acremonium egyptiacum” CBS 303.64 

-clade 

acremonium phylogeny 

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. 

Gliomastix/Bionectria clade 

Sarocladium clade 

sclerotigenum/Geosmithia clade 

Hypocreales 

Plectosphaerellaceae 

Glomerellales 

A 

141


Fig. 1. (Continued). 

142 

71 

96 

Simplicillium lanosoniveum CBS 321.72 

Cordyceps cardinalis 

Acremonium camptosporum CBS 756.69 T 

Paecilomyces lilacinus 2 

Elaphocordyceps capitata 

Elaphocordyceps ophioglossoides 

Balansia henningsiana 

Epichloe typhina 

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 

Acremonium recifei CBS 137.35 T 

Nectria cinnabarina 

Clavicipitaceae 

Acremonium roseolum CBS 289.62 

Stachybotrys chartarum 

Stachybotrys subsimplex 

Myrothecium roridum 1 

Peethambara spirostriata 

Pseudonectria rousseliana 

Acremonium nigrosclerotium CBS 154.72 T 

96 Hypocrea americana 

Hypocrea lutea 

Sphaerostilbella berkeleyana 

Hypocreaceae 

Niesslia exilis 

Nectria haematococca 

Chaetosphaerella phaeostroma 

96 

Melanospora tiffanii 

Melanospora zamiae 

Lindra thalassiae 

Lulworthia grandispora 

“Acremonium alabamense” CBS 456.75 

Cercophora terricola 

Apiosordaria verruculosa 

97 Cercophora striata 

Podospora decipiens 

Bombardia bombarda 

Lasiosphaeria ovina 

71 

Cercophora newfieldiana 

Gelasinospora tetrasperma 

99 

Neurospora crassa 

Immersiella immersa 

Lasiosphaeria hispida 

Podospora fibrinocaudata 

Strattonia carbonaria 

84 

98 

Camarops amorpha 

Camarops microspora 

Camarops tubulina 

Boliniales 

94 Camarops ustulinoides 

89 97 

Chaetosphaeria ovoidea 

Menispora tortuosa 

Melanochaeta hemipsila 

74 Lasiosphaeriella nitida 

Linocarpon appendiculatum 

T 

Cordycipitaceae 

Ophiocordycipitaceae 

T 

Stachybotrys/ 

Peethambara 

-clade 

Lulworthiales 

Sordariales 

Chaetosphaeriales 

Hypocreales 

B



99 “Acremonium cf. alternatum” CBS 109043 

Acremonium atrogriseum CBS 544.79 

75 


80 Acremonium atrogriseum CBS 774.97 




Acremonium atrogriseum CBS 604.67 T 

88 


Cephalotheca sulfurea 

Acremonium thermophilum CBS 734.71 T 

Albertiniella polyporicola 

Coniochaeta ostrea 

99 

98 Coniochaeta savoryi Coniochaetales 

70 Coniochaeta discoidea 

Annulatascus triseptatus 

89 

91 Fragosphaeria purpurea 

97 

Ophiostoma piliferum 1 

Ophiostoma stenoceras 

Papulosa amerospora 

Apiognomonia errabunda 

92 Plagiostoma euphorbiae 

Cryptodiaporthe aesculi 

99 Gnomonia gnomon 

Cryptosporella hypodermia 

95 Melanconis alni 

Melanconis stilbostoma 

81 

89 99 

Melanconis marginalis 

97 

Chromendothia citrina 

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 

Graphostroma platystoma 

Xylariales 

77 

Xylaria acuta 

90 Xylaria hypoxylon 

Seynesia erumpens 

Leotia lubrica 

Microglossum rufum 

Pseudeurotium zonatum 

Chalara aurea 

Acremonium butyri (?) CBS 301.38 T 

Acremonium lichenicola CBS 425.66 T 

Botryotinia fuckeliana 

0.01 substitutions/site 

Cephalothecaceae 

Ophiostomatales 

Diaporthales 

Leotiomycetes (outgroup) 


C 

143


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 confirmed name 

are listed under the name under which they are currently held in the CBS collection. 

Currently assigned species name Collection numbers nuclsU nucssU Notes 

Acremoniella lutzi T CBS 103.48 hQ231971 – Ex-type of Acremoniella lutzi 

Acremonium acutatum T CBS 682.71 hQ231965 

Acremonium alabamense CBS 456.75 hQ231972 – 

Acremonium alcalophilum T CBS 114.92 hQ231973 – Ex-type of Acremonium alcalophilum 

Acremonium alternatum T CBS 407.66 hQ231988 

“Acremonium alternatum” CBS 381.70A hQ231986 

CBS 406.66 hQ231987 hQ232178 

CBS 831.97 hQ231989 

CBS 114602 hQ231990 

“Acremonium cf. alternatum” CBS 109043 hQ231974 – 

Acremonium antarcticum CBS 987.87 hQ231975 – 

Acremonium atrogriseum T CBS 604.67 hQ231981 – Ex-type of Phaeoscopulariopsis atrogrisea 

CBS 252.68 hQ231977 – 







Acremonium biseptum T CBS 750.69 hQ231998 Ex-type of Acremonium biseptum 

“Acremonium blochii” CBS 324.33 hQ231999 




Acremonium borodinense T CBS 101148 hQ232003 Ex-type of Acremonium borodinense 

Acremonium brachypeniumT CBS 866.73 hQ232004 

Acremonium breve T CBS 150.62 hQ232005 hQ232183 Ex-type of Cephalosporium roseum var. breve 

Acremonium brunnescens T CBS 559.73 hQ231966 hQ232184 Ex-type of Acremonium brunnescens 

Acremonium butyri T CBS 301.38 hQ231967 – Ex-type of Tilachlidium butyri; synonym of Cadophora malorum 

Acremonium camptosporum T CBS 756.69 hQ232008 hQ232186 Ex-type of Acremonium camptosporum 





Acremonium cavaraeanum CBS 758.69 hQ232012 

Acremonium cerealis CBS 207.65 hQ232013 Ex-type of Gliomastix guttuliformis 


Acremonium chrysogenum T CBS 144.62 hQ232017 hQ232187 Ex-type of Cephalosporium chrysogenum 

Acremonium cucurbitacearum CBS 683.88 hQ231968 – Previously identified as Acremonium strictum 

Acremonium curvulum T CBS 430.66 hQ232026 hQ232188 Ex-type of Acremonium curvulum 





CBS 384.70A hQ232023 

CBS 384.70C hQ232024 





“Acremonium aff. curvulum” CBS 100551 hQ232031 


Acremonium egyptiacum CBS 303.64 hQ232034 hQ232189 

Acremonium exiguum T CBS 587.73 hQ232035 hQ232190 Ex-type of Acremonium exiguum 

Acremonium exuviarum T UAMH 9995 hQ232036 

Acremonium flavum T CBS 596.70 hQ232037 hQ232191 Ex-type of Acremonium flavum 

Acremonium fuci UAMH 6508 hQ232038 

Acremonium furcatum T CBS 122.42 AY378154 hQ232192 Ex-type of Acremonium furcatum 

144

Table 1. (Continued). 




Acremonium fusidioides T CBS 840.68 hQ232039 Ex-type of Paecilomyces fusidioides 

Acremonium gamsii T CBS 726.71 hQ232040 hQ232193 Ex-type of Acremonium gamsii 

Acremonium guillematii T CBS 766.69 hQ232042 hQ232194 Ex-type of Acremonium guillematii 

Acremonium hansfordii CBS 390.73 hQ232043 

Acremonium hennebertii T CBS 768.69 hQ232044 Ex-type of Acremonium hennebertii 

Acremonium hyalinulum CBS 271.36 hQ232045 hQ232195 

“Acremonium implicatum” CBS 243.59 hQ232046 hQ232196 Authentic strain of Fusarium terricola 

CBS 397.70B hQ232047 

Acremonium incrustatumT CBS 159.70 hQ232049 

Acremonium inflatum T CBS 212.69 hQ232050 Ex-type of Gliomastix inflata 


CBS 403.70 hQ231991 In CBS as Acremonium atrogriseum 

Acremonium lichenicola T CBS 425.66 hQ231969 – Ex-type of Acremonium lichenicola 

Acremonium longisporum CBS 113.69 hQ232057 

“Acremonium luzulae” CBS 495.67 hQ232058 


Acremonium minutisporum T CBS 147.62 hQ232061 hQ232199 Ex-type of Cephalosporium minutisporum 

det 267B hQ232062 

Acremonium nepalense T CBS 971.72 hQ231970 – Ex-type of Acremonium nepalense 

Acremonium nigrosclerotium T CBS 154.72 hQ232069 hQ232200 Ex-type of Acremonium nigrosclerotium 

Acremonium persicinum T CBS 310.59 hQ232077 hQ232201 Ex-type of Paecilomyces persicinus 



CBS 295.70M hQ232076 



CBS 378.70D hQ232081 

CBS 378.70 E hQ232082 





“Acremonium persicinum” CBS 378.70C hQ232080 


“Acremonium aff. persicinum” CBS 203.73 hQ232073 


“Acremonium cf. persicinum” CBS 102877 hQ232087 

Acremonium pinkertoniae T CBS 157.70 hQ232089 hQ232202 Ex-type of Acremonium pinkertoniae 

“Acremonium potronii” CBS 189.70 hQ232094 

CBS 379.70F hQ232096 




Acremonium psammosporum T CBS 590.63 hQ232100 hQ232204 Ex-type of Acremonium psammosporum 

Acremonium pseudozeylanicum T CBS 560.73 hQ232101 Ex-type of Acremonium pseudozeylanicum 

Acremonium pteridii T CBS 782.69 hQ232102 Ex-type of Acremonium pteridii 


Acremonium radiatum T CBS 142.62 hQ232104 hQ232205 Ex-type of Cephalosporium acremonium var. radiatum 

Acremonium recifei T CBS 137.35 hQ232106 hQ232206 Ex-type of Cephalosporium recifei 











CBS 400.85 hQ232025 In CBS as Acremonium curvulum 

CBS 505.94 hQ232027 In CBS as Acremonium cf. curvulum 

“Acremonium recifei” CBS 485.77 hQ232113 


145





Acremonium restrictum T CBS 178.40 hQ232119 – Ex-type of Verticillium dahliae f. restrictum 

Acremonium rhabdosporum T CBS 438.66 hQ232120 Ex-type of Acremonium rhabdosporum 

Acremonium roseolum T CBS 289.62 hQ232123 hQ232207 Ex-type of Paecilomyces roseolus 

Acremonium rutilum T CBS 396.66 hQ232124 hQ232208 Ex-type of Acremonium rutilum 

Acremonium salmoneum T CBS 721.71 hQ232125 Ex-type of Acremonium salmoneum 

Acremonium sclerotigenum T CBS 124.42 hQ232126 hQ232209 Ex-type of Cephalosporium sclerotigenum 






OMH F1648.97 hQ232132 

OMH F2365.97 hQ232133 

OMH F2969.97 hQ232134 

OMH F3691.97 hQ232135 

CBS 287.70O hQ232140 In CBS as Acremonium strictum 

CBS 379.70D hQ232095 In CBS as Acremonium potronii 

CBS 223.70 hQ231985 In CBS as Acremonium alternatum 

Acremonium sordidulum T CBS 385.73 hQ232136 Ex-type of Acremonium sordidulum 

Acremonium sp. CBS 314.72 hQ232156 

Acremonium spinosum T CBS 136.33 hQ232137 hQ232210 Ex-type of Cephalosporium spinosum 

“Acremonium strictum” CBS 106.23 hQ232138 


Acremonium stromaticum T CBS 863.73 hQ232143 – Ex-type of Acremonium stromaticum 

Acremonium tectonae T CBS 725.87 hQ232144 Ex-type of Acremonium tectonae 

Acremonium thermophilum T CBS 734.71 hQ232145 – Ex-type of Acremonium thermophilum 

Acremonium tsugae T CBS 788.69 hQ232146 Ex-type of Acremonium tsugae 

Acremonium tubakii T CBS 790.69 hQ232148 Ex-type of Acremonium tubakii 

“Acremonium tubakii” CBS 824.69 hQ232149 

Acremonium verruculosum T CBS 989.69 hQ232150 Ex-type of Acremonium verruculosum 

Acremonium vitellinum T CBS 792.69 hQ232151 hQ232212 Ex-type of Acremonium vitellinum 

Acremonium zeylanicum CBS 746.73 hQ232154 

Acremonium zonatum CBS 565.67 hQ232155 

“Cephalosporium acremonium var. cereum” T CBS 140.62 hQ232147 Ex-type of Cephalosporium acremonium var. cereum. In CBS as 

Acremonium tubakii 

“Cephalosporium acremonium var. CBS 141.62 hQ232053 Ex-type of Cephalosporium acremonium var. funiculosum. In CBS 

funiculosum” T 

as Acremonium kiliense 

“Cephalosporium ballagii” T CBS 134.33 hQ232016 Ex-type of Cephalosporium ballagii. In CBS as Acremonium 

charticola 

“Cephalosporium malorum” T CBS 117.25 hQ232015 Ex-type of Cephalosporium malorum. In CBS as Acremonium charticola 

“Cephalosporium purpurascens” T CBS 149.62 hQ232071 Ex-type of Cephalosporium purpurascens. In CBS as Acremonium 

persicinum 

Cosmospora khandalensis T CBS 356.65 hQ231996 Ex-type of Cephalosporium khandalense. In CBS as Acremonium 

berkeleyanum 

Cosmospora lavitskiaeT CBS 530.68 hQ231997 Ex-type of Gliomastix lavitskiae. In CBS as Acremonium berkeleyanum 

Gliomastix masseei CBS 794.69 hQ232060 In CBS as Acremonium masseei 

Gliomastix murorum CBS 154.25 hQ232063 Ex-type of Graphium malorum. In CBS as Acremonium murorum 

var. felina 

CBS 195.70 hQ232064 In CBS as Acremonium murorum var. felina 

CBS 119.67 hQ232066 In CBS as Acremonium murorum var. murorum 

CBS 157.72 hQ232067 In CBS as Acremonium murorum var. murorum 

CBS 378.36 hQ232068 Ex-type of Torula cephalosporioides. In CBS as Acremonium 

murorum var. murorum 

Gliomastix polychroma T CBS 181.27 hQ232091 Ex-type of Oospora polychroma. In CBS as Acremonium polychromum 

CBS 151.26 hQ232090 Ex-type of Periconia tenuissima var. nigra. In CBS as Acremonium 

polychromum 

CBS 617.94 hQ232093 In CBS as Acremonium polychromum 

Gliomastix roseogrisea T CBS 134.56 hQ232121 Ex-type of Cephalosporium roseogriseum. In CBS as Acremonium 

roseogriseum 

CBS 279.79 hQ232122 In CBS as Acremonium roseogriseum 

CBS 213.69 hQ232092 In CBS as Acremonium polychromum 

CCFC 226570 AY283559 Identified as Acremonium murorum var. felina 

CBS 211.69 hQ232065 In CBS as Acremonium murorum var. felina 

Lanatonectria flavolanata CBS 230.31 hQ232157 

146





Lanatonectria flocculenta CBS 113461 hQ232158 

Leucosphaerina arxii T CBS 737.84 hQ232159 Ex-type of Leucosphaerina arxii 

Nalanthamala diospyri T CBS 560.89 hQ232160 Ex-type of Cephalosporium diospyri = Acremonium diospyri 

Nectria rishbethii T CBS 496.67 hQ232162 Ex-type of Nectria rishbethii 

Neocosmospora endophytica AR 2674 U17411 Anamorph is Acremonium fungicola 

Paecilomyces lilacinus CBS 101068 hQ232163 hQ232214 Atypical monophialidic isolate, Acremonium+E402-like 

Pochonia bulbillosa CBS 102853 hQ232164 Atypical isolate 

Sarcopodium circinatum CBS 376.81 hQ232167 



Sarcopodium circinosetiferum CBS 100251 hQ232170 




Sarcopodium vanillae CBS 100582 hQ232174 

Sarocladium attenuatum T CBS 399.73 hQ232165 Ex-type of Sarocladium attenuatum 

Sarocladium bacillisporum T CBS 425.67 hQ231992 hQ232179 Ex-type of Acremonium bacillisporum 

Sarocladium bactrocephalum T CBS 749.69 hQ231994 hQ232180 Ex-type of Acremonium bactrocephalum 

NRRL 20583 hQ231995 

Sarocladium glaucum T CBS 796.69 hQ232041 Ex-type of Acremonium glaucum 

Sarocladium kiliense T CBS 122.29 hQ232052 hQ232198 Ex-type of Acremonium kiliense 

CBS 146.62 hQ232048 hQ232197 Ex-type of Cephalosporium incoloratum. In CBS as Acremonium 

incoloratum 

CBS 155.61 hQ232054 Ex-type of Cephalosporium incarnatum. In CBS as Acremonium 

kiliense 

CBS 156.61 hQ232055 Ex-type of Cephalosporium incarnatum var. macrospora. In CBS as 

Acremonium kiliense 

CBS 157.61 hQ232056 Ex-type of Cephalosporium infestans.In CBS as Acremonium kiliense 

Sarocladium ochraceum T CBS 428.67 hQ232070 Ex-type of Paecilomyces ochraceus. In CBS as Acremonium ochraceum 

Sarocladium oryzae CBS 180.74 hQ232166 

Sarocladium strictum T CBS 346.70 hQ232141 hQ232211 Ex-type of Acremonium strictum 

“Sarocladium cf. strictum” JY03-006 hQ232142 

Sarocladium zeae T CBS 801.69 hQ232152 hQ232213 Ex-type of Acremonium zeae 

KAS 965 hQ232153 

Simplicillium lanosoniveum CBS 321.72 hQ232006 hQ232185 Ex-type of Acremonium byssoides 

Simplicillium obclavatum T CBS 311.74 hQ232175 Ex-type of Acremonium obclavatum 

Trichothecium crotocinigenum T CBS 129.64 hQ232018 Ex-type of Acremonium crotocinigenum 

“Trichothecium 

indicaT 

indicum”/ Leucosphaerina CBS 123.78 AF096194 Ex-type of ‘Leucosphaera’ indica 

Trichothecium roseum DAOM 208997 U69891 

Trichothecium sympodiale ATCC 36477 U69889 In CBS as Spicellum roseum 

Verticillium alboatrum CBS 130.51 hQ231976 – Ex-type of Cephalosporium apii, in CBS as Acremonium apii 

Verticillium insectorum CBS 101239 hQ248107 

Verticillium leptobactrum CBS 109351 hQ231993 In CBS as Acremonium cf. bacillisporum 

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 

conspecific 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 

identified 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" 

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


148 

85 

94 

75 Acremonium acutatum CBS 682.71 T 

“Cephalosporium acremonium var. cereum” CBS 140.62 T 

Acremonium sclerotigenum CBS 223.70 

Acremonium sclerotigenum CBS 379.70D 

Acremonium sclerotigenum OMH F2365.97 




Mycopepon smithii 

Pseudonectria sp. 

Acremonium sclerotigenum CBS 100816 

Acremonium sclerotigenum CBS 287.70O 




89 Acremonium sclerotigenum CBS 124.42 T 


Acremonium alternatum CBS 407.66 T 

“Cephalosporium malorum” CBS 117.25 T 

Acremonium sordidulum CBS 385.73 T 

Acremonium brachypenium CBS 866.73 T 

“Cephalosporium purpurascens” CBS 149.62 T 

83 


Acremonium exuviarum UAMH 9995 T 

“Acremonium tubakii” CBS 824.69 

79 

Emericellopsis terricola 

Acremonium fuci UAMH 6508 

“Acremonium potronii” CBS 379.70F 

99 

Acremonium salmoneum CBS 721.71 T 

93 Stilbella fimetaria 

Acremonium tubakii CBS 790.69 T 

Stanjemonium grisellum 

Acremonium chrysogenum CBS 144.62 T 

90 Acremonium flavum CBS 596.70 T 

71 Hapsidospora irregularis 

74 

Nigrosabulum globosum 

Mycoarachis inversa 

Geosmithia lavendula 

Geosmithia putterillii 


96 

99 Acremonium borodinense CBS 101148 T 

Acremonium pinkertoniae CBS 157.70 T 

Bulbithecium hyalosporum 

Leucosphaerina arxii T 

pinkertoniae 

-clade 

“Acremonium cf. persicinum” CBS 102877 

85 “Acremonium alternatum” CBS 381.70A 

“Acremonium alternatum” CBS 831.97 

“Acremonium cavaraeanum” CBS 758.69 fusidioides 

Acremonium hennebertii CBS 768.69 T 

Acremonium fusidioides CBS 840.68 T 

“Acremonium hansfordii” CBS 390.73 

“Acremonium egyptiacum” CBS 303.64 

-clade 

96 

99 “Acremonium blochii” CBS 427.93 


Verticillium insectorum 

“Acremonium zeylanicum” CBS 746.73 

Stilbocrea macrostoma 

sclerotigenum 

-clade 

brachypenium 

-clade 

Emericellopsis 

-clade 

chrysogenum 

-clade 

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. 

sclerotigenum/Geosmithia-clade 

A



86 

78 

85 

Acremonium biseptum CBS 750.69 

Acremonium cerealis CBS 207.65 

Acremonium cerealis CBS 215.69 

93 

cerealis 

-clade 


Acremonium persicinum CBS 378.70E 

Acremonium persicinum CBS 378.70D 

Acremonium persicinum CBS 378.70A 

Acremonium persicinum CBS 102349 

Acremonium persicinum CBS 169.65 

Acremonium persicinum CBS 295.70M 

Acremonium persicinum CBS 295.70A 


Acremonium persicinum CBS 101694 


91 


Acremonium verruculosum CBS 989.69 T 

Acremonium persicinum CBS 310.59 T 



Gliomastix masseei CBS 794.69 T 

Gliomastix murorum CBS 157.72 



80 

92 



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 

72 Roumegueriella rufula 

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 

100 Acremonium spinosum CBS 136.33 T 

Acremonium pteridii CBS 782.69 T 

Ochronectria calami 

93 Bionectria grammicospora 

Bionectria ochroleuca 2 

72 Nectria sesquicillii 

Bionectria pityrodes 

Stephanonectria keithii 

99 

Nectriopsis sporangiicola 

93 Nectriopsis violacea 

Sesquicillium microsporum 

Kallichroma glabrum 

Tilachlidium brachiatum 

T 

Gliomastix clade 

pteridii 

-clade 

persicinum 

-clade 


Gliomastix/Bionectria-clade 

B 

149


93 

Acremonium incrustatum CBS 159.70 T 


Linkosia fusiformis 

Acremonium guillematii CBS 766.69 T 

“Acremonium persicinum” CBS 110646 

Acremonium sp. CBS 314.72 

Acremonium vitellinum CBS 792.69 T minutisporum 

100 Acremonium minutisporum CBS 147.62 T 

Acremonium minutisporum det 267B 

-clade 

100 “Acremonium alternatum” CBS 406.66 

“Acremonium alternatum” CBS 114602 

99 Sarocladium bacillisporum CBS 425.67 T 

77 “Acremonium implicatum” CBS 243.59 

Sarocladium glaucum CBS 796.69 T 

bacillisporum 

-clade 

94 Sarocladium bactrocephalum CBS 749.69 T 

Sarocladium bactrocephalum NRRL 20583 

84 Sarocladium cf. strictum JY03-006 

83 Sarocladium strictum CBS 346.70 T 

84 “Acremonium zonatum” CBS 565.67 

100 Sarocladium attenuatum T 

Sarocladium oryzae 

94 Sarocladium kiliense CBS 146.62 

Sarocladium kiliense CBS 122.29 T 




Sarocladium zeae CBS 801.69 T 

98 Sarocladium zeae KAS 965 

Sarocladium ochraceum CBS 428.67 T 

strictum 

-clade 

Sarocladium 

-clade 

100 

Acremonium breve CBS 150.62 T 

76 Acremonium radiatum CBS 142.62 T 

91 “Acremonium strictum” CBS 147.49 

“Cephalosporium acremonium var. funiculosum” CBS 141.62 

Acremonium gamsii CBS 726.71 T 

T 

100 

83 Trichothecium crotocinigenum CBS 129.64 T 

“Trichothecium indicum”/Leucosphaerina indica T 

Trichothecium sympodiale 

73 Trichothecium roseum 

Trichothecium 

-clade 

74 

75 

Acremonium curvulum CBS 430.66 T 

Acremonium curvulum CBS 229.75 

Acremonium curvulum CBS 384.70A 


100 Acremonium curvulum CBS 761.69 



curvulum 

-clade 

95 

100 

Acremonium curvulum CBS 110514 

Acremonium curvulum CBS 384.70C 



98 

Valetoniellopsis laxa 

Acremonium rhabdosporum CBS 438.66 T 

Lanatonectria flavolanata 1 

Lanatonectria flavolanata 2 

Sarcopodium vanillae 

94 Sarcopodium circinatum 2 

Sarcopodium circinosetiferum 3 Lanatonectria 

82 

84 

Sarcopodium circinosetiferum 4 

Lanatonectria flocculenta 



Pseudonectria rousseliana 

-clade 


150 

breve 

-clade 

C



100 Acremonium inflatum CBS 403.70 



100 

“Acremonium luzulae” CBS 579.73 

Scopinella solani 

Acremonium roseolum CBS 289.62 

Peethambara sundara 

Albosynnema elegans 

Didymostilbe echinofibrosa 

Peethambara spirostriata 

Sarcopodium circinatum 3 

Sarcopodium circinatum 1 

Myrothecium cinctum 

Myrothecium inundatum 



Myrothecium leucotrichum 

Myrothecium verrucaria 

Parasarcopodium ceratocaryi 

Stachybotrys chartarum 

Chaetosphaeria aterrima 

96 Didymostilbe matsushimae 

Melanopsamma pomiformis 

80 

Cosmospora lavitskiae CBS 530.68 

Cosmospora episphaeria 

“Acremonium aff. curvulum” CBS 100551 

Cosmospora vilior 

Cosmospora khandalensis CBS 356.65 

Acremonium recifei CBS 400.85 













“Acremonium strictum” CBS 106.23 

Fusarium dimerum 

Fusarium domesticum 

Fusarium oxysporum 1 

Fusarium oxysporum 2 

Fusarium verticillioides 

Persiciospora africana 

Nectria rigidiuscula 

Fusarium falciforme 

Fusarium solani 

Fusarium lichenicola 

Neocosmospora endophytica 

Acremonium tsugae CBS 788.69 

Cylindrocarpon cylindroides 

Heliscus lugdunensis 

Calonectria morganii 

Nectria radicicola 

Leuconectria clusiae 

Dematiocladium celtidis 

Nectria cinnabarina 

“Acremonium recifei” CBS 485.77 

“Acremonium recifei” CBS 482.78 

“Acremonium recifei” CBS 110348 

Nalanthamala diospyri 

Nalanthamala guajavae 

Nalanthamala vermoesenii 

Rubrinectria olivacea 

‘Nectria’ rishbethii CBS 496.67 

Acremonium exiguum CBS 587.73 

“Acremonium implicatum” CBS 397.70B 



Nectriopsis squamulosa 1 

Nectriopsis squamulosa 2 

“Cephalosporium ballagii” CBS 134.33 

“Acremonium aff. curvulum” CBS 113275 

Acremonium pseudozeylanicum CBS 560.73 

Niesslia exilis 1 

Acremonium nigrosclerotium CBS 154.72 

Niesslia exilis 2 

76 

T 

T 

88 

98 

86 

78 

T 

86 

T 

88 

T 

89 

81 

100 

91 

99 

87 

70 

89 

T 

90 

T 

85 100 

96 

T 

T 

exiguum 

92 

93 

-clade 

100 

97 

T 

77 

T 

T 

inflatum 

-clade 

Stachybotrys/Peethambara 

-clade 

Cosmospora 

Fusarium 

-clade 

pseudozeylanicum 

-clade 

Nectriaceae/recifei-clade 


D 

151



152 

97 

98 

80 

0.01 substitutions/site 

Cordyceps ramosipulvinata 

100 

Melanospora brevirostris 

83 

Bertia moriformis 

Phaeoacremonium aleophilum 

Ophiocordyceps bispora 

Elaphocordyceps longisegmentis 

Haptocillium sinense 

Haptocillium zeosporum 

99 Paecilomyces lilacinus 2 

Paecilomyces lilacinus 1 

82 Isaria takamizusanensis 

98 

99 

Chaunopycnis alba 1 

Chaunopycnis alba 2 

Chaunopycnis pustulata 

Tolypocladium inflatum 

Verticillium leptobactrum CBS 109351 

92 94 Simplicillium lanosoniveum CBS 321.72 

78 Simplicillium lanosoniveum 

98 Simplicillium obclavatum T 

91 

74 

Simplicillium lamellicola 

Beauveria bassiana 

Cordyceps militaris 

Pleurodesmospora coccorum 

Isaria javanica 

Syspastospora parasitica 

Hyperdermium bertonii 

Lecanicillium antillanum 

Lecanicillium aranearum 

Lecanicillium fusisporum 

Lecanicillium psalliotae 1 

Lecanicillium dimorphum 

Lecanicillium aff. psalliotae 2 

Lecanicillium lecanii 

Paecilomyces farinosus 

Lecanicillium attenuatum 

Epichloe typhina 

Pochonia bulbillosa 2 

Pochonia bulbillosa 1 

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 



99 Acremonium camptosporum CBS 890.85 

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 

T 

T 

T 

T 

camptosporum 

-clade 

Ophiocordycipitaceae 

Cordycipitaceae 

Clavicipitaceae 

Hypocreaceae 

e

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. flavum. It also includes non-type 

isolates identified 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 identified 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. fimetaria (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 fimetaria is closely related to the ex-type 

isolate of Acremonium salmoneum isolated from dung, also a 

typical habitat for S. fimetaria (Seifert 1985). An adjacent weakly 



supported clade includes Hapsidospora, Mycoarachis, and 

Nigrosabulum, and the two Acremonium species named for yellow 

pigmentation, A. chrysogenum and A. flavum. Although associated 

with A. chrysogenum and A. flavum 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, untypified species A. potronii. Basal to these clades 

is another small clade that links an entomogenous isolate identified 

as Verticillium insectorum with two isolates from human sources 

identified 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 

affinities containing Acremonium guillematii, A. minutisporum and 

A. vitellinum. This group also appears as two to three unaffiliated 

clades in Fig. 2C. An insignificant branch in Fig. 1B subtends 

Acremonium exiguum, A. psammosporum, and an isolate identified 

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


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 inflatum isolates, including CBS 

403.70, an atypical, catenate-conidial isolate identified at CBS as 

A. atrogriseum. Nearby but statistically unlinked clades include 

Stachybotrys and allied fungi such as Peethambara spirostriata 

and Didymostilbe echinofibrosa (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 affinities 

to another hyaline, phialidic fungus, Phialophora alba. 

A number of genera in addition to Acremonium were 

investigated for possible affinity 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 identified 

as S. circinatum grouped with Sarcopodium circinosetiferum and 

S. vanillae in a widely separated clade along with Lanatonectria 

teleomorphs and a sequence identified 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 20 th 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 simplified 

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 identified 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), specifically 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 

identified 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 identified 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.

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 influential 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 flat and slightly fasciculate, rather 

than cottony. To some extent Acremonium sect. Acremonium was 

based on keying out all the relatively flat 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 fimetaria, A. chrysogenum in a bionectriaceous 

clade containing cleistothecial teleomorphs such as Nigrosabulum, 

A. rutilum in a clade otherwise containing isolates identified 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 



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. inflatum. 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 first 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 defined 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 conspecific 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 epitypified 

with one of these isolates, the generic name might be restricted 

to this single species. The final isolate is CBS 223.70, an isolate 

that, despite its catenate conidia, is conspecific 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 identified as A. 

egyptiacum (e.g., CBS 734.69), which are also conspecific 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 identified as A. alternatum, a 

species in the A. fusidioides clade, is represented by CBS 831.97 

155


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 

Based on these analyses, three genera are represented in sufficient 

detail and with high bootstrap support to be formally recognised here. 

In most cases, the genera and clades are not sufficiently populated 

with their constituent members without analysis of additional 

sequences. For example, the Emericellopsis clade is missing 12 of 

its 13 species including two identified as E. minima (Zuccaro et al. 

2004) as well as one of its two Stanjemonium species. 

Gliomastix 

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. inflatum. 

The closely related A. persicinum clade may also be included as 

suggested by Supplemental fig. 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 significantly 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 

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 confirmed 

as synonyms of G. murorum (Fig. 2B). Recently, Kiyuna et al. 

(2010) neotypified 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 


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 epitypification. 

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. 

Epitype designated here: Italy, Turin, isolated from rabbit dung, A. 

Fontana, CBS H-8244, ex-epitype culture CBS 794.69. 

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 


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 confirmed 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 clarification. 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 S.B. Saksena, 

Mycologia 47: 895. 1956 [1955]. 

≡ 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

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 neotypified 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 confirms 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 fly from plant to plant. Species with habitats where water 

flux or microarthropod movement may be important in dispersal, 

e.g., various Acremonia occurring in soil or Fusarium domesticum 

growing on cheese, may have simplified 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 difficult 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 epitypification, 

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]. 



Colonies on 2 % malt extract agar slimy-glabrous to moderately 

floccose 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


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, 


≡ 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 floccose 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, 


Basionym: Acremonium bactrocephalum W. Gams, 


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 


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 


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, 


Basionym: Paecilomyces ochraceus Onions & G.L. Barron, Mycol. 

Pap. 107: 15. 1967. 

≡ Acremonium ochraceum (Onions & G.L. Barron) W. Gams, 


≡ 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 


Descriptions of S. strictum are given by Gams (1971) and Domsch 

et al. (2007). The type isolate of S. strictum was confirmed in this 

genus (Fig. 2C). Of the three isolates illustrated by Gams (1971) 

under A. strictum, CBS 287.70 D, is confirmed 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 unified 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, 


Basionym: Acremonium zeae W. Gams & D.R. Sumner, in Gams, 


This economically important maize endophyte species fits the 

description given by Gams (1971) as a fungus with felty to shaggy 

colonies. Two S. zeae isolates with more flattened 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 flavus 

and Fusarium verticillioides in maize inflorescences and are thus 

important in the ecology and economic significance 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 significant 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 first named fungi were often 

species prominently in contact with humans and their environs 

and because the first 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



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 artificial 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 scientific 

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 unified clade of Trichothecium includes 

isolates with three different anamorphic forms, currently classified 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 fifth, 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 redefined 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) 

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, 

floccose 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 finely 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 indefinitely 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


or present, when present mostly in intercalary chains, hyaline, 

smooth or finely 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. 

Basionym: Cephalosporium crotocinigenum Schol-Schwarz, Trans. 

Brit. Mycol. Soc. 48: 53. 1965. 

≡ Acremonium crotocinigenum (Schol-Schwarz) W. Gams, 


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 scientific names reflect true 

relationships. 

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 

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 significance, 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 

confident 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. 

Acremonium atrogriseum and Acremonium cf. 

alternatum CBs 109043 in the Cephalothecaceae: a 

study in comparative morphology vs. phylogeny 

Acremonium atrogriseum and an isolate identified 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 conspecifics; 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, identifications 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.

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 scientific 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). Leucosphaera, a new genus of the 

Pseudeurotiaceae. Persoonia 10: 141–143. 

Bachmann HP, Bobst C, Bütikofer U, Casey MG, Dalla Torre M, Fröhlich-Wyder MT, 

Fürst M (2005). Occurrence and significance of Fusarium domesticum alias 

Anticollanti on smear-ripened cheeses. LWT - Food Science and Technology 

38: 399–407. 

Bills GF, Platas G, Gams W (2004). Conspecificity of the cerulenin and helvolic 

acid producing ‘Cephalosporium caerulens’, and the hypocrealean fungus 

Sarocladium oryzae Mycological Research 108: 1291–1300. 

Castlebury LA, Rossman AY, Sung GH, Hyten AS, Spatafora JW (2004). Multigene 

phylogeny reveals new lineage for Stachybotrys chartarum, the indoor air 

fungus. Mycological Research 108: 864–872. 

Dickinson CH (1968). Gliomastix Guéguen. Mycological Papers 115: 1–26. 

Domsch KH, Gams W, Anderson T-H (2007) Compendium of soil fungi. IHW-Verl., 

Eching, Germany. 

Gams W (1968). Typisierung der Gattung Acremonium. Nova Hedwigia 16: 141–145. 

Gams W (1971). Cephalosporium-artige Schimmelpilze (Hyphomycetes). 1–262. 

G. Fischer, Stuttgart. 

Gams W (1975). Cephalosporium-like Hyphomycetes: some tropical species. 

Transactions of the British Mycological Society 64: 389–404. 

Gams W (1978). Connected and disconnected chains of phialoconidia and 

Sagenomella gen. nov. segregated from Acremonium. Persoonia 10: 97–112. 

Gams W, Hawksworth DL (1976) [‘1975’]. The identity of Acrocylindrium oryzae 

Sawada and a similar fungus causing sheath rot of rice. Kavaka 3: 57–61. 

Gams W, Lacey J (1972). Cephalosporium-like Hyphomycetes. Two species of 

Acremonium from heated substrates. Transactions of the British Mycological 

Society 59: 519–522. 

Gams W, McGinnis MR (1983). Phialemonium, a new anamorph genus intermediate 

between Phialophora and Acremonium. Mycologia 75: 977–987. 

Gams W, O’Donnell K, Schroers H-J, Christensen M (1998). Generic classification 

of some more hyphomycetes with solitary conidia borne on phialides. Canadian 

Journal of Botany 76: 1570–1583. 

Gargas A, Taylor JW (1992). Polymerase chain reaction (PCR) primers for amplifying 

and sequencing 18S rDNA from lichenized fungi. Mycologia 84: 589–592. 

Glenn AE, Bacon CW, Price R, Hanlin RT (1996). Molecular phylogeny of 

Acremonium and its taxonomic implications. Mycologia 88: 369–383. 

Gómez BL, Nosanchuk JD (2003). Melanin and fungi. Current Opinion in Infectious 

Disease 16: 91–96. 

Gräfenhan T, Schroers H-J, Nirenberg HI, Seifert KA (2011). An overview of the 

taxonomy, phylogeny, and typification of nectriaceous fungi in Cosmospora, 

Acremonium, Fusarium, Stilbella, and Volutella. Studies in Mycology 68: 79– 

113 (this issue). 

Hammill TM (1981). On Gliomastix murorum and G. felina. Mycologia 73: 229–237 

Hoog GS de, Guarro J, Gené J, Figueras MJ (2000). Atlas of Clinical Fungi. 2 nd ed. 

Utrecht, Netherlands: Centraalbureau voor Schimmelcultures. 

Hoog GS de, Gerrits van den Ende AHG (1998). Molecular diagnostics of clinical 

strains of filamentous basidiomycetes. Mycoses 41: 183–189. 

Hughes SJ (1958). Revisiones hyphomycetum aliquot cum appendice de nominibus 

rejiciendis. Canadian Journal of Botany 36: 727–836. 

Ito T, Okane I, Nakagiri A, Gams W (2000). Two species of Acremonium section 

Acremonium: A. borodinense sp. nov. and A. cavaraeanum rediscovered. 

Mycological Research 104: 77–80. 

Kiyuna T, An K-D, Kigawa R, Sano C, Miura S, Sugiyama J (2010). Molecular 

assessment of fungi in ‘‘black spots" that deface murals in the Takamatsuzuka 

and Kitora Tumuli in Japan: Acremonium sect. Gliomastix including Acremonium 

tumulicola sp. nov. and Acremonium felinum comb. nov. Mycoscience 52: 1–17. 

Lechat C, Farr DF, Hirooka Y, Minnis AM, Rossman AY (2010). A new species of 

Hydropisphaera, H. bambusicola, is the sexual state of Gliomastix fusigera. 

Mycotaxon 111: 95–102. 



Liang ZQ, HanYF, Chu HL, Fox RTV (2009). Studies on the genus Paecilomyces 

in China V. Taifanglania gen. nov. for some monophialidic species. Fungal 

Diversity 34: 69–77. 

Link H (1809). Observationes in ordines plantarum naturals. Gesellschaft 

Naturforschender Freunde zu Berlin Magazin 3: 1–42. 

Lutzoni F, Wagner P, Reeb V, Zoller S (2000). Integrating ambiguously aligned 

regions of DNA sequences in phylogenetic analyses without violating positional 

homology. Systematic Biology 49: 628–651 

Maddison WP, Maddison DR (2003). MacClade: analysis of phylogeny and character 

evolution. V. 4.06. Sinauer, Sunderland, Massachusetts. 

Malloch D (1989). An undescribed species of Leucosphaerina. Studies in Mycology 

31: 107–111. 

Mason-Gamer R, Kellogg E (1996). Testing for phylogenetic conflict among 

molecular datasets in the tribe Triticeae (Graminae). Systematic Biology 45: 

524–545. 

Matsushima T (1975). Icones Microfungorum a Matsushima lectorum. Published by 

the author, Kobe, Japan, 209 pp. 

Mel’nik V, Lee S, Groenewald J Z, Crous PW (2004). New hyphomycetes from 

Restionaceae in fynbos: Parasarcopodium ceratocaryi gen. et sp. nov., and 

Rhexodenticula elegiae sp. nov. Mycological Progress 3: 19–28. 

Morgan-Jones G, Gams W (1982). Notes on Hyphomycetes. XLI. An endophyte of 

Festuca arundinacea and the anamorph of Epichloe typhina, new taxa in one 

of two new sections of Acremonium. Mycotaxon 15: 311–318. 

Novicki TJ, LaFe K, Bui L, Bui U, Geise R, Marr K, Cookson BT (2003). Genetic 

diversity among clinical isolates of Acremonium strictum determined during an 

investigation of a fatal mycosis. Journal of Clinical Microbiology 41: 2623–2628. 

O’Donnell K (1993). Fusarium and its near relatives. In: The fungal holomorph: 

mitotic, meiotic and pleomorphic speciation in fungal systematics. (Reynolds 

R, Taylor JW, eds.), CBA International, Wallingford, United Kingdom: 225–233. 

Perdomo H, Sutton DA, García D, Fothergill AW, Cano J, Gené J, Summerbell 

RC, Rinaldi MG, Guarro J (2010). Spectrum of clinically relevant Acremonium 

species in the United States. Journal of Clinical Microbiology. doi:10.1128/ 

JCM.00793-10 

Raper JR, Raper CA (1972). Genetic analysis of the life cycle of Agaricus bisporus. 

Mycologia 64:1088–1117. 

Reeb V, Roux C, Lutzoni F (2004). Contribution of RPB2 to multilocus phylogenetic 

studies of the euascomycetes (Pezizomycotina, Fungi) with special emphasis 

on the lichen-forming Acarosporaceae and evolution of polyspory. Molecular 

Phylogenetics and Evolution 32: 1036–1060. 

Rifai MA, Cooke RC (1966). Studies on some didymosporous genera of nematodetrapping 

Hyphomycetes. Transactions of the British Mycological Society 49: 

147 – 168. 

Rossman AY, Samuels GJ, Rogerson CT, Lowen R (1999). Genera of Bionectriaceae, 

Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in 

Mycology 42: 1–248. 

Schoch CL, Sung G-H, López-Giráldez F, Townsend JP, Miadlikowska J, Hofstetter 

V, Robbertse B, Matheny PB, Kauff F, Wang Z, Guiedan C, Andrie RM, Trippe K, 

Ciufetti LM, Wynns A, Fraker E, Hodkinson BP, Bonito G, Yahr R, Groenewald 

JZ, Arzanlou M, de Hoog GS, Crous PW, Hewitt D, Pfister DH, Peterson K, 

Gryzenhout M, Wingfield MJ, Aptroot A, Suh S-O, Blackwell M, Hillis DM, 

Griffith GW, Castlebury LA, Rossman AY, Lumbsch HT, Lücking R, Büdel B, 

Rauhut A, Diederich P, Ertz D, Geiser DM, Hosaka K, Inderbitzin P, Kohlmeyer 

J, Volkmann-Kohlmeyer B, Mostert L, O’Donnell K, Sipman H, Rogers JD, 

Shoemaker RA, Sugiyama J, Summerbell RC, Untereiner W, Johnston P, 

Stenroos S, Zuccaro A, Dyer PS, Crittenden PD, Cole MS, Hansen K, Trappe 

JM, Lutzoni F, Spatafora JW (2009) The Ascomycota tree of life: a phylumwide 

phylogeny clarifies the origin and evolution of fundamental reproductive and 

ecological traits. Systematic Biology 58: 224–239. 

Schroers H-J (2000). Generic delimitation of Bionectria (Bionectriaceae, 

Hypocreales) based on holomorph characters and rDNA sequences. Studies 

in Mycology 45: 63–82. 

Schroers H-J (2001). A monograph of Bionectria (Ascomycota, Hypocreales, 

Bionectriaceae) and its Clonostachys anamorphs. Studies in Mycology 46: 

1–214. 

Schroers H-J, Geldenhuis MM, Wingfield MJ, Schoeman MH, Yen YF, Shen WC, 

Wingfield BD (2005). Classification of the guava wilt fungus Myxosporium 

psidii, the palm pathogen Gliocladium vermoesenii and the persimmon wilt 

fungus Acremonium diospyri in Nalanthamala. Mycologia 97: 375–395. 

Seifert KA (1985). A monograph of Stilbella and allied hyphomycetes. Studies in 

Mycology 27: 1–235. 

Seifert KA, Rehner SA, Sugita T, Okada G (2008). Spicellum ovalisporum Seifert & 

Rehner, sp. nov. Fungal Planet 28: 1–4. 

Sigler L, Sutton DA, Gibas CF, Summerbell RC, Noel RK, Iwen PC (2010) 

Phialosimplex, a new anamorphic genus associated with infections in dogs 

and having phylogenetic affinity to the Trichocomaceae. Medical Mycology 48: 

335–345. 

161


Stamatakis A, Hoover P, Rougemont J (2008). A rapid bootstrap algorithm for the 

RAxML web-servers. Systematic Biology 75: 758–771. 

Stamatakis A, Ludwig T, Meier H (2005). RAxML-III: A fast program for maximum 

likelihood-based inference of large phylogenetic trees. Bioinformatics 21: 

456–463. 

Suh S-O, Blackwell M (1999). Molecular phylogeny of the cleistothecial fungi placed 

in Cephalothecaceae and Pseudeurotiaceae Mycologia 91: 836–848. 

Summerbell RC (2003). Aspergillus, Fusarium, Sporothrix, Piedraia and their 

relatives. In: Pathogenic Fungi in Humans and Animals (Howard DH, ed) 

Marcel Dekker Press, New York: 237–498. 

Summerbell RC, Schroers H-J (2002). Analysis of phylogenetic relationship of 

Cylindrocarpon lichenicola and Acremonium falciforme to the Fusarium solani 

species complex and a review of similarities in the spectrum of opportunistic 

infections caused by these fungi. Journal of Clinical Microbiology 40: 2866– 

2875. 

Sung GH, Hywell-Jones NL, Sung JM, Luangsa-ard JJ, Shrestha B, Spatafora JW 

(2007). Phylogenetic classification of Cordyceps and the clavicipitaceous fungi. 

Studies in Mycology 57: 5–59. 

Vilgalys R, Hester M (1990). Rapid genetic identification and mapping of 

enzymatically amplified ribosomal DNA from several Cryptococcus species. 

Journal of Bacteriology 172: 4238–4246. 

162 

White TJ, Bruns TD, Lee SB, Taylor JW (1990). Amplification and direct sequencing 

of fungal ribosomal RNA genes for phylogenetics. In: PCR protocols: A guide to 

methods and applications. (Innis MA, Gelfand DA, Sninsky JJ, White TJ, eds.), 

Academic Press, San Diego, California, USA: 315–322. 

Wicklow DT, Poling SM, Summerbell RC (2008). Occurrence of pyrrocidine and 

dihydroresorcylide production among Acremonium zeae populations from 

maize grown in different regions. Canadian Journal of Plant Pathology 30: 

425–433. 

Zare R, Gams W (2001). A revision of Verticillium section Prostrata. IV. The genera 

Lecanicillium and Simplicillium gen. nov. Nova Hedwigia 73: 1–50. 

Zare R, Gams W, Starink-Willemse M, Summerbell RC (2007). Gibellulopsis, a 

suitable genus for Verticillium nigrescens, and Musicillium, a new genus for V. 

theobromae. Nova Hedwigia 85: 463–489. 

Zuccaro A, Summerbell RC, Gams W, Schroers H-J, Mitchell JI (2004). A new 

Acremonium species associated with Fucus spp., and its affinity with a 

phylogenetically distinct marine Emericellopsis clade. Studies in Mycology 50: 

283–297.

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