Phylogeny and nomenclature of the genus Talaromyces and taxa accommodated in Penicillium subgenus Biverticillium

Robert A Samson

Talaromyces is a monophyletic genus containing seven sections. The number of species in Talaromyces grows rapidly due to reliable and complete sequence data contributed from all over the world. In this study agricultural soil samples from Fujiang, Guangdong, Jiangxi, Shandong, Tibet and Zhejiang provinces of China were collected and analyzed for fungal diversity. Based on a polyphasic approach including phylogenetic analysis of partial ITS, BenA, CaM and RPB2 gene sequences, macro- and micro-morphological analyses, six of them could not be assigned to any described species, and one cannot be assigned to any known sections. Morphological characters as well as their phylogenetic relationship with other Talaromyces species are presented for these putative new species. Penicillium resedanum is combined in Talaromyces section Subinflati as T. resedanus.

Talaromyces verruculosus, T. aculeatus and T. apiculatus are the only Talaromyces species that produce conidiophores with ampulliform phialides, which taper into very thin necks and have rough walled, globose conidia. In this study, we introduce five new species with similar micromorphological features, but were found to display unique macro-morphological characters. Talaromyces australis (CBS 137102 T) is distinguished by its restricted growth on CYA at 25 and 37 C (16e24 mm; 9e13 mm) and red pigments produced on most media. Talaromyces kendrickii (CBS 136666 T) is distinguished by its inability to grow on CYA at 37 C. Talaromyces veerkampii (CBS 500.78 T) grows rapidly on MEA (38e42 mm) and colonies on YES has a bronze green reverse. Talaromyces fuscoviridis (CBS 193.69 T) colonies have dark green reverses on MEA and commonly produces red exudates on other media. Talaromyces stellenboschiensis (CBS 135665 T) grows faster on CYA at 25, 30 and 37 C (40e45 mm; 48e53 mm; 35e40 mm) than the others. Morphological findings were supported by both multigene phylogenetic analyses and the extrolites produced by these species.

Coprophilous fungi are saprotrophic organisms that show great diversity, mainly on herbivore dung. The physico-chemical characteristics of this peculiar substrate combined with the high level of fungal adaptation to different environmental conditions offer the perfect setting for discovering new taxa. This study focused on the species diversity of penicillium-like fungi isolated mainly from herbivore dung collected at different Spanish locations. From 130 samples, a total of 104 isolates were obtained, and 48 species were identified. Preliminary identifications were based on morphology and partial β-tubulin (tub2) gene sequences. Putative new taxa were characterized by a multi-gene sequencing analysis testing the tub2, the internal transcribed spacer rDNA (ITS), calmodulin (cmdA), and RNA polymerase II second largest subunit (rpb2) genes, and a detailed phenotypic study. Using this polyphasic approach and following the genealogical concordance phylogenetic species recognition (GCPSR...

Downloaded from orbit.dtu.dk on: Jun 07, 2020 Phylogeny and nomenclature of the genus Talaromyces and taxa accommodated in Penicillium subgenus Biverticillium Samson, R.A.; Yilmaz, N.; Houbraken, J.; Spierenburg, H.; Seifert, K.A.; Peterson, S.W.; Varga, J.; Frisvad, Jens Christian Published in: Studies in Mycology Link to article, DOI: 10.3114/sim.2011.70.04 Publication date: 2011 Document Version Publisher's PDF, also known as Version of record Link back to DTU Orbit Citation (APA): Samson, R. A., Yilmaz, N., Houbraken, J., Spierenburg, H., Seifert, K. A., Peterson, S. W., Varga, J., & Frisvad, J. C. (2011). Phylogeny and nomenclature of the genus Talaromyces and taxa accommodated in Penicillium subgenus Biverticillium. Studies in Mycology, 70, 159-184. https://doi.org/10.3114/sim.2011.70.04 General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights.  Users may download and print one copy of any publication from the public portal for the purpose of private study or research.  You may not further distribute the material or use it for any profit-making activity or commercial gain  You may freely distribute the URL identifying the publication in the public portal If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. available online at www.studiesinmycology.org doi:10.3114/sim.2011.70.04 StudieS in Mycology 70: 159–184. 2011. Phylogeny and nomenclature of the genus Talaromyces and taxa accommodated in Penicillium subgenus Biverticillium R.A. Samson1, N. Yilmaz1,6, J. Houbraken1,6, H. Spierenburg1, K.A. Seifert2, S.W. Peterson3, J. Varga4 and J.C. Frisvad5 1 CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; 2Biodiversity (Mycology), Eastern Cereal and Oilseed Research Centre, Agriculture & Agri-Food Canada, 960 Carling Ave., Ottawa, Ontario, K1A 0C6, Canada, 3Bacterial Foodborne Pathogens and Mycology Research Unit, National Center for Agricultural Utilization Research, 1815 N. University Street, Peoria, IL 61604, U.S.A., 4Department of Microbiology, Faculty of Science and Informatics, University of Szeged, H-6726 Szeged, Közép fasor 52, Hungary, 5Department of Systems Biology, Building 221, Technical University of Denmark, DK-2800, Kgs. Lyngby, Denmark; 6Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.. *Correspondence: R.A. Samson, r.samson@cbs.knaw.nl Abstract: The taxonomic history of anamorphic species attributed to Penicillium subgenus Biverticillium is reviewed, along with evidence supporting their relationship with teleomorphic species classified in Talaromyces. To supplement previous conclusions based on ITS, SSU and/or LSU sequencing that Talaromyces and subgenus Biverticillium comprise a monophyletic group that is distinct from Penicillium at the generic level, the phylogenetic relationships of these two groups with other genera of Trichocomaceae was further studied by sequencing a part of the RPB1 (RNA polymerase II largest subunit) gene. Talaromyces species and most species of Penicillium subgenus Biverticillium sensu Pitt reside in a monophyletic clade distant from species of other subgenera of Penicillium. For detailed phylogenetic analysis of species relationships, the ITS region (incl. 5.8S nrDNA) was sequenced for the available type strains and/or representative isolates of Talaromyces and related biverticillate anamorphic species. Extrolite profiles were compiled for all type strains and many supplementary cultures. All evidence supports our conclusions that Penicillium subgenus Biverticillium is distinct from other subgenera in Penicillium and should be taxonomically unified with the Talaromyces species that reside in the same clade. Following the concepts of nomenclatural priority and single name nomenclature, we transfer all accepted species of Penicillium subgenus Biverticillium to Talaromyces. A holomorphic generic diagnosis for the expanded concept of Talaromyces, including teleomorph and anamorph characters, is provided. A list of accepted Talaromyces names and newly combined Penicillium names is given. Species of biotechnological and medical importance, such as P. funiculosum and P. marneffei, are now combined in Talaromyces. Excluded species and taxa that need further taxonomic study are discussed. An appendix lists other generic names, usually considered synonyms of Penicillium sensu lato that were considered prior to our adoption of the name Talaromyces. Key words: anamorph, DNA phylogeny, single name nomenclature, teleomorph, Trichocomaceae. Taxonomic novelties: Taxonomic novelties: New species – Talaromyces apiculatus Samson, Yilmaz & Frisvad, sp. nov. New combinations and names –Talaromyces aculeatus (Raper & Fennell) Samson, Yilmaz, Frisvad & Seifert, T. albobiverticillius (H.-M. Hsieh, Y.-M. Ju & S.-Y. Hsieh) Samson, Yilmaz, Frisvad & Seifert, T. allahabadensis (B.S. Mehrotra & D. Kumar) Samson, Yilmaz & Frisvad, T. aurantiacus (J.H. Mill., Giddens & A.A. Foster) Samson, Yilmaz, & Frisvad, T. boninensis (Yaguchi & Udagawa) Samson, Yilmaz, & Frisvad, T. brunneus (Udagawa) Samson, Yilmaz & Frisvad, T. calidicanius (J.L. Chen) Samson, Yilmaz & Frisvad, T. cecidicola (Seifert, Hoekstra & Frisvad) Samson, Yilmaz, Frisvad & Seifert, T. coalescens (Quintan.) Samson, Yilmaz & Frisvad, T. dendriticus (Pitt) Samson, Yilmaz, Frisvad & Seifert, T. diversus (Raper & Fennell) Samson, Yilmaz & Frisvad, T. duclauxii (Delacr.) Samson, Yilmaz, Frisvad & Seifert, T. echinosporus (Nehira) Samson, Yilmaz & Frisvad, comb. nov. T. erythromellis (A.D. Hocking) Samson, Yilmaz, Frisvad & Seifert, T. funiculosus (Thom) Samson, Yilmaz, Frisvad & Seifert, T. islandicus (Sopp) Samson, Yilmaz, Frisvad & Seifert, T. loliensis (Pitt) Samson, Yilmaz & Frisvad, T. marneffei (Segretain, Capponi & Sureau ) Samson, Yilmaz, Frisvad & Seifert, T. minioluteus (Dierckx) Samson, Yilmaz, Frisvad & Seifert, T. palmae (Samson, Stolk & Frisvad) Samson, Yilmaz, Frisvad & Seifert, T. panamensis (Samson, Stolk & Frisvad) Samson, Yilmaz, Frisvad & Seifert, T. paucisporus (Yaguchi, Someya & Udagawa) Samson & Houbraken T. phialosporus (Udagawa) Samson, Yilmaz & Frisvad, T. piceus (Raper & Fennell) Samson, Yilmaz, Frisvad & Seifert, T. pinophilus (Hedgcock) Samson, Yilmaz, Frisvad & Seifert, T. pittii (Quintan.) Samson, Yilmaz, Frisvad & Seifert, T. primulinus (Pitt) Samson, Yilmaz & Frisvad, T. proteolyticus (Kamyschko) Samson, Yilmaz & Frisvad, T. pseudostromaticus (Hodges, G.M. Warner, Rogerson) Samson, Yilmaz, Frisvad & Seifert, T. purpurogenus (Stoll) Samson, Yilmaz, Frisvad & Seifert, T. rademirici (Quintan.) Samson, Yilmaz & Frisvad, T. radicus (A.D. Hocking & Whitelaw) Samson, Yilmaz, Frisvad & Seifert, T. ramulosus (Visagie & K. Jacobs) Samson, Yilmaz, Frisvad & Seifert, T. rubicundus (J.H. Mill., Giddens & A.A. Foster) Samson, Yilmaz, Frisvad & Seifert, T. rugulosus (Thom) Samson, Yilmaz, Frisvad & Seifert, T. sabulosus (Pitt & A.D. Hocking) Samson, Yilmaz & Frisvad, T. siamensis (Manoch & C. Ramírez) Samson, Yilmaz & Frisvad, T. sublevisporus (Yaguchi & Udagawa) Samson, Yilmaz & Frisvad, T. variabilis (Sopp) Samson, Yilmaz, Frisvad & Seifert, T. varians (G. Sm.) Samson, Yilmaz & Frisvad, T. verruculosus (Peyronel) Samson, Yilmaz, Frisvad & Seifert, T. viridulus Samson, Yilmaz & Frisvad. INTRODUCTION The modern concept of Penicillium (referred to in this paper as Penicillium sensu lato), was derived from the pioneering monographic revisions of Thom (1930), Raper & Thom (1949), and formalised by the recognition of four subgenera, Aspergilloides, Furcatum, Penicillium and Biverticillium by Pitt (1980). Over the past decade, the realisation has grown that Penicillium subgenus Biverticillium is phylogenetically distinct from other subgenera of Penicillium and that this distinctiveness should be reflected in its formal taxonomy. Because of their usually symmetrical, biverticillate conidiophores, the group has been recognised since Wehmer (1914) segregated them in an informal subdivision of Penicillium that he called "Verticillatae". The delineation, species composition and taxonomic rank of this group were modified in subsequent monographs by Thom (1930), Raper & Thom (1949), Pitt (1980), and Ramírez (1982), culminating in the widespread recognition of subgenus Biverticillium and the use of this name in many taxonomic and phylogenetic studies. Malloch (1985), based on a consideration of morphological and ecological factors, and anamorph-teleomorph connections, may have been the first to speculate that subgenus Biverticillium should be removed from Penicillium as a separate genus. The teleomorph genera historically associated with Penicillium sensu lato are Talaromyces and Eupenicillium (in single name nomenclature, the latter is now considered a synonym of Copyright 2011 CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 159 SaMSon eT al. Penicillium sensu stricto, see Houbraken & Samson 2011). The teleomorphs of these two groups produce distinctive ascomata. In Talaromyces, the soft ascomatal walls are comprised of multiple layers of interwoven hyphae and the ascomata mature quickly, usually within a few weeks in agar culture. In Penicillium sensu stricto, the sclerotium-like ascomata have rigid walls of thick-walled, isodiametric cells and the ascomatal maturity can take months and often ascospores do not form at all. Furthermore, in Talaromyces the ascus initials sometimes have morphologically distinguishable gametangia and the mature asci are produced in chains (Stolk & Samson 1972), while the ascomatal initials in Penicillium sensu stricto are irregularly interwoven, loosely branched hyphae masses (Emmons 1935), and the mature asci are single. Raper & Thom (1949) already recognised that there was considerable evidence that Penicillium subgenus Biverticillium constituted a natural and homogenous group. A comparison of the anamorphs of these two teleomorph types reveals a correlation with phialide shape, with anamorphs of Talaromyces (until now classified in Penicillium subgenus Biverticillium) having narrower phialides that are aculeate or lanceolate, and anamorphs in Penicillium sensu stricto having broader, ampulliform or flask-shaped phialides. One consequence of the differences in phialide shape is that the symmetrical nature of the conidiophores of species allied with Talaromyces tends to be emphasised, because in general the phialides are more densely packed. The colonies of subgenus Biverticillium can often be distinguished from those of Penicillium sensu stricto by the naked eye. They often have darker green conidia, more or less yellow pigmented and encrusted aerial hyphae, and colony reverses in yellow, orange or red to purplish red shades. Once DNA-based studies of fungal phylogeny began, it quickly became apparent that the differences between Penicillium sensu stricto and Talaromyces were more than a matter of degree, and that there might be a significant problem with the generic concept of Penicillium sensu lato. Penicillium sensu stricto and Talaromyces occur as distinct clades within Trichocomaceae, which could be considered subfamilies (LoBuglio et al., 1993, LoBuglio & Taylor 1993). Using small subunit nuclear ribosomal DNA sequences (18S), Berbee et al. (1995) showed that Penicillium is polyphyletic if subgenus Biverticillium is included, a conclusion reconfirmed in one of the first reviews of the impact of molecular phylogenetics on Ascomycete taxonomy (Sugiyama 1998) using an analysis of 18S rDNA sequences. Removal of subgenus Biverticillium transforms Penicillium sensu stricto into a monophyletic group. This dichotomy between Penicillium sensu stricto and Talaromyces was shown repeatedly in studies employing nuclear ribosomal RNA genes, for example by Peterson (2000), who analysed a combination of the nuclear ribosomal internal transcribed spacer regions (ITS) and large subunit ribosomal DNA (28S) sequences (Ogawa et al. 1997, Ogawa & Sugiyama 2000), and by Wang & Zhuang (2007) in a phylogeny based on calmodulin sequences. The results of these analyses are all confirmed in the multigene phylogenetic analyses presented elsewhere in this volume by Houbraken & Samson (2011), using genes selected for their ability to accurately reflect molecular phylogeny. As indicated by Houbraken & Samson (2011), when other genera assigned to Trichocomaceae are included in phylogenetic analyses, the division between subgenus Biverticillium and Penicillium sensu stricto becomes even clearer. In that study, intervening genera include Aspergillus, Paecilomyces sensu stricto (with Byssochlamys as a synonym), and several small and less well-known genera such as Thermoascus, Penicilliopsis, Thermomyces and the recently described Rasamsonia (Houbraken et al. 2011). 160 In a molecularly defined, phylogenetically accurate taxonomic system, maintaining subgenus Biverticillium in Penicillium sensu stricto is untenable. However, almost every aspect of the biology, biochemistry, and physiology of these two groups emphasises their fundamental distinctiveness, although sometimes with limited taxon sampling. For example, Pitt (1980) emphasised the distinctiveness of subgenus Biverticillium by using a low wateractivity medium, G25N (which includes 25 % glycerol) in his standard plating regime. Strains assigned to this subgenus grow slowly on this medium, less than 10 mm diam at 25 °C in 7 d, whereas species of the other subgenera are more xerophilic and grow faster. Cell-wall components seem to differ significantly. Leal & Bernabé (1998) reported on the complex glucomannogalactan components of the water soluble polysaccharide fraction of several species of Trichocomaceae, suggesting that a characteristic heteropolysaccharide composed of 4 galactose: 1 mannose: 1 glucose was unique to species of subgenus Biverticillium. Species of Penicillium sensu stricto species were characterised by the presence of a β-(1-5)(1-6)-galactofuran polysaccharide in the same fraction. Cell wall components as reflected by their exoantigens were screened in about 50 species of Penicillium sensu lato using an ELISA reaction to antibodies raised to P. digitatum (subgenus Penicillium). These antibodies reacted well with all the species of subgenera Furcatum, Penicillium and Aspergilloides, but did not react with the four species of subgenus Biverticillium tested (P. funiculosum, P. islandicum, P. rubrum, and P. tardum) (Notermans et al. 1998). Kuraishi et al. (1991) first noted that the pattern of ubiquinones in Penicillium sensu lato and showed a distinct pattern in subgenus Biverticillium. Paterson (1998) examined 335 strains and 118 species of Penicillium sensu lato and determined that the Q9 ubiqinone type was predominant in the species of Penicillium sensu stricto. In contrast, species of Talaromyces, Trichocoma and subgenus Biverticillium had different versions of the Q10 ubiquinone type. Exceptions to these patterns can be explained by the small number of species whose classification in, or elimination from, subgenus Biverticillium has been uncertain or controversial. Frisvad et al. (1990a) provided an overview of the extrolites of Talaromyces species, and demonstrated the occurrence of characteristic extrolites such as mitorubins, bisanthaquinones such as rugulosin and skyrin, vermicellin, vermistatin, vermiculine, duclauxin and glauconic acid. None of these compounds were found in cultures of Penicillium sensu stricto (Frisvad et al. 1990b). The soon to be published International Code of Nomenclature for Algae, Fungi and Plants removes the primacy of teleomorphover anamorph-typified names, leaving both kinds of names competing equally for priority (Norvell 2011). Because of these changes, we apply the principle of ‘one fungus - one name’ and in the nomenclatural revision, priority is given to the oldest genus and species name irrespective of whether they were originally described for teleomorphs or anamorphs (Hawksworth et al. 2011). In this respect, Penicillium returns to the single named, but pleomorphic, nomenclatural and taxonomic system used by many of the founders of its taxonomy, and actively promoted by the Peoria school (Thom 1930, Raper & Thom 1949). Talaromyces, now also defined as a pleomorphic genus, is adopted for the anamorphic species formerly included in Penicillium subgenus Biverticillium. In this study, the phylogenetic relationships of species of subgenus Biverticillium and other members of the Trichocomaceae were studied by sequencing a part of the RPB1 (RNA polymerase II largest subunit) gene. Furthermore, we discuss the taxonomy and nomenclature of species of this expanded concept of Talaromyces, based on phylogenetic, phenotypic and extrolite data. For detailed PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium phylogenetic analysis below genus level, the ITS regions (including the 5.8S nrDNA) of ex-type strains and/or representatives were sequenced. As discussed below, this paper is not meant as a monographic treatment, because many complexes have not yet been studied comprehensively. MATERIALS AND METHODS Sources of cultures The fungi examined include type strains or representatives of all available species of Talaromyces and Biverticillium. The strains are maintained in the CBS-KNAW Fungal Biodiversity Centre (CBS) culture collection and an overview of strains used for phylogenetic analysis is shown in Table 1. In a few cases, the ex-type strain was unavailable and sequence data present in GenBank were used. Morphology and physiology Cultures were grown for 7 d on Czapek agar, Czapek yeast autolysate agar (CYA), oatmeal agar (OA) and/or malt extract agar (MEA) plates at 25 °C or, if required, another temperature. Medium compositions follow Samson et al. (2010). Cultures were grown for up to 3 wk for ascomata production. Extrolite analysis Nearly all species described in the genera Penicillium sensu lato (including those formerly classified in Eupenicillium), Penicillium subgenus Biverticillium, Talaromyces, Aspergillus and its many associated teleomorphic genera, and Paecilomyces (including those formerly or still classified in the associated teleomorph genus Byssochlamys) were analysed qualitatively for their profiles of secondary metabolites as determined by HPLC with diode array detection. Many strains of each species were examined, whenever available, but in some cases only the ex-type culture was available. Cultures were inoculated on the media CYA, MEA (Blakeslee formula, using Difco malt extract), YES agar (Samson et al. 2010, Difco yeast extract) and OA. All cultures were analysed chemically using three agar plugs from a 7 d old culture grown at 25 °C (Smedsgaard 1997). Different methods were used for HPLC analysis, but the methods were essentially based on Frisvad & Thrane (1987, 1993). Since 1997, the method for Nielsen & Smedsgaard (2003) was used and after 2010 the UPLC method of Nielsen et al. (2011) was applied. Metabolites were identified via their diode-array based UV-VIS spectra and in some cases by their mass spectra, and by comparison to authenticated standards (Nielsen et al. 2011). For the extrolites analyses, the biosynthetic families of the sampled genera were compared using UPGMA cluster analysis (NTSYS version 2.11). All metabolites were classified according to biosynthetic families; for example the viridicatin biosynthetic family consists of cyclopenol, cyclopenin, cyclopeptin, dehydrocyclopeptin, viridicatin, viridicatol and 3-methoxyviridicatin (Turner & Aldridge 1983). This family was scored as one character in the cluster analysis. The exometabolites were also combined into biosynthetic families and tabulated as such. For example, many species of Talaromyces and Penicillium subgenus Biverticillium produce the azaphilones mitorubrin, mitorubrinal, mitorubrinol, mitorubrinol acetate, mitorubrinic acid, funicone, deoxyfunicone, actofunicone, www.studiesinmycology.org 3-O-methylfunicone, kasanosin A and B, diazaphilonic acid, and wortmin; they are here collectively called the mitorubrins, while the related metabolites vermistatins and penicidones are called vermistatins (see Šturdíková et al. 2000, Nicoletti et al. 2009, Osmanova et al. 2010). Some chlorinated azaphilones such as helicusins (Yoshida et al. 1995) and luteusins (Fujimoto et al. 1990, Yoshida et al. 1996a, b) are epimers of the sclerotiorins from P. sclerotiorum, and are treated as two families, albeit closely related to the mitorubrins. DNA extraction, amplification and sequencing Isolates used for molecular studies were grown on MEA for 7–14 d at the required temperature prior to DNA extraction. DNA was extracted from the cells using the UltraClean™ Microbial DNA Kit (MoBio Laboratories), following the protocols of the manufacturer. A part of the RPB1 gene was amplified to study the phylogenetic relationships among Penicillium and other related genera. This fragment was amplified using the primer pair RPB1-F1843 5’-ATTTYGAYGGTGAYGARATGAAC-3’ and RPB1-R3096 5’-GRACRGTDCCRTCATAYTTRACC-3’ (Houbraken & Samson 2011). Primer RPB1-F1843 corresponds with position 1490–1512 of GenBank no. XM_002146871 (P. marneffei, ATCC 18224) and RPB1-R3096 corresponds with position 2610–2633. An addition primer, RPB1-R2623 5’-GCRTTGTTSARATCCTTMARRCTC-3’ was occasionally used as an internal primer for sequencing (Houbraken & Samson 2011). The ITS regions were sequenced to study the relationship among Talaromyces and the related biverticillate anamorphic species. Fragments containing the ITS region were amplified using primers V9G (de Hoog & Gerrits van den Ende 1998) and LS266 (Masclaux et al. 1995). Sequencing reactions were performed with the Big Dye Terminator Cycle Sequencing Ready Reaction Kit v. 3.1 (Applied Biosystems) and carried out for both strands to ensure consistency of the consensus sequence. Data analyses For the DNA sequence analyses, alignments were performed using the software Muscle as implemented in the MEGA5 programme (Tamura et al. 2011). The RAxML (randomised accelerated maximum likelihood) software (v. 7.2.8, Stamatakis et al. 2008) was used for the Maximum Likelihood (ML) analysis. The robustness of trees in the ML analyses was evaluated by 100 bootstrap replications. The phylogram based on RPB1 sequences is rooted with Coccidioides immitis (strain RS; full genome strain), and Trichocoma paradoxa (CBS 788.83) is used as an outgroup in the ITS analysis. RESULTS Phylogenetic generic delimitation of Talaromyces and biverticillate anamorphic species The phylogenetic relationships of Talaromyces and species of Penicillium subgenus Biverticillium among other related genera were studied using partial RPB1 sequences. One-hundred fifty-six strains were included in this analysis. The length of the alignment was 496 characters (exon data only, no introns observed) and 323 of those characters were variable. The proportion of gaps and 161 SaMSon eT al. Table 1. Strains used in phylogenetic analysis of Talaromyces. Name Collection no. Origin RPB1 ITS “Aphanoascus cinnabarinus” CBS 267.72 = ATCC 26215 Soil, Japan JN121625 JN899376 Aspergillus aculeatus CBS 172.66T = ATCC 16872 = IMI 211388 Tropical soil JN121590 Aspergillus clavatoflavus CBS 473.65NT = ATCC 16866 = IMI 124937 Rain forest soil,Tulley, Queensland, Australia JN121686 Aspergillus flavus NRRL 3357 = CBS 128202 = ATCC 200026 Peanut cotyledons, USA Unpublished Aspergillus fumigatus Af293 Patient with invasive aspergillosis Nierman et al. (2005) Aspergillus niger CBS 513.88 Derived from NRRL 3122 and currently used as enzyme production strain Pel et al. (2007) Aspergillus ochraceoroseus CBS 101887 = ATCC 42001 = IBT 14580 Soil, Tai National Forest, Ivory Coast JN121557 Aspergillus ochraceus CBS 108.08NT = ATCC 1008 = CBS 547.65 = IMI 016247 = IMI 016247iii = IMI 016247iv = NRRL 1642 = NRRL 398 Unknown source JN121562 Aspergillus penicillioides CBS 130294 Indoor environment, Germany JN121578 Aspergillus robustus CBS 649.93T = CBS 428.77 = IBT 14305 Surface soil from thorn-forest, near Mombasa, Kenya JN121711 Aspergillus sparsus CBS 139.61NT = ATCC 16851 = IMI 019394 = IMI 019394ii = MUCL 31314 = NRRL 1933 Soil, Costa Rica JN121586 Aspergillus steynii CBS 112812T = IBT 23096 Dried arabica green coffee bean, on parchment, internal infection, Chamumdeshuran Estata, Karnataka, district Giris, India JN121569 Aspergillus sydowii CBS 264.81 Grains and milling fractions, Triticum aestivum, India JN121624 Aspergillus versicolor CBS 245.65 = ATCC 11730 = ATCC 16020 = IMI 045554 = IMI 045554ii = IMI 045554iii = IMI 045554iv = MUCL 19008 Cellophane, Indiana, USA JN121614 Aspergillus zonatus CBS 506.65NT = ATCC 16867 = IMI 124936 Forest soil, Province of Linon, Fortuna, Costa Rica JN121691 Byssochlamys nivea CBS 100.11T = ATCC 22260 Unknown source JN121511 Byssochlamys spectabilis CBS 101075 = ATCC 90900 = FRR 5219 Heat processed fruit beverage, Tokyo, Japan JN121554 Byssochlamys verrucosa CBS 605.74T = ATCC 34163 Nesting material of Leipoa ocellata (Malleefowl), Pulletop Nature Reserve, New South Wales, Australia JN680311 Chrysosporium inops CBS 132.31T = IMI 096729 = UAMH 802 Skin of man, Italy JN121584 Coccidioides immitis Strain “RS” Vaccine strain - origin unknown Sharpton et al. (2009) Emericella nidulans FGSC A4 (= ATCC 38163 = CBS 112.46) Unknown source Galagan et al. (2005) Eurotium herbariorum CBS 516.65NT = ATCC 16469 = IMI 211383 = NRRL 116 Unpainted board, Washington, USA JN121693 Geosmithia viridis CBS 252.87T = FRR 1863 = IMI 288716 Soil, bank of creek flowing into Little River, New South Wales JN680284 Hamigera avellanea CBS 295.48T = ATCC 10414 = IMI 040230 = NRRL 1938 Soil, San Antonio, Texas, USA JN121632 Hamigera striata CBS 377.48NT = ATCC 10501 = IMI 039741 = NRRL 717 Canned blueberries, USA JN121665 Monascus purpureus CBS 109.07T = ATCC 16365 = ATCC 16426 = IMI 210765 = NRRL 1596 Fermented rice grain, ‘ang-quac’ (purple coloured rice), Kagok-Tegal, imported from China, Prov. Quouan-toung, Java, Indonesia JN121563 Paecilomyces aerugineus CBS 350.66T = IMI 105412 Debris of Glyceria maxima, Attenborough, Notts., UK JN121657 JN899388 Paecilomyces pascuus CBS 253.87T = FRR 1925 Pasture grass, Otara, New Zealand JN899292 JN899321 162 T GenBank Accession number JN899314 PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Table 1. (Continued). Name Collection no. Origin GenBank Accession number Penicilliopsis clavariiformis CBS 761.68 = CSIR 1135 Unknown source, Pretoria, South Africa Penicillium aculeatum CBS 100105 = CBS 289.48 = ATCC 10409 = IMI 040588 = NRRL 2129 = NRRL A-1474 Textile, USA JN899389 CBS 289.48NT = ATCC 10409 = IMI 040588 = NRRL 2129 = NRRL A-1474 Textile, USA JN899378 Penicillium aculeatum var. apiculatum CBS 312.59T = ATCC 18315 = FRR 635 = IMI 068239 Soil, Japan JN680293 JN899375 Penicillium allahabadense CBS 453.93T = ATCC 15067 = CBS 304.63 Soil of cultivated field, pH 6.9, Allahabad, India JN680309 JN899345 Penicillium arenicola CBS 220.66T = ATCC 18321 = ATCC 18330 = IMI 117658 = NRRL 3392 Soil from pine forest, Kiev, Ukraine JN121601 Penicillium aurantiacum CBS 314.59T = ATCC 13216 = IMI 099722 = NRRL 3398 Soil, Georgia JN899380 Penicillium aureocephalum CBS 102801T Quercus ruber, Gerona, Selva de Mar, Catalanıa, Spain JN899392 Penicillium brunneum CBS 227.60T = ATCC 18229 = FRR 646 = IFO 6438 = IHEM 3907 = IMI 078259 = MUCL 31318 Milled rice imported into Japan, Thailand JN680281 JN899365 Penicillium calidicanium CBS 112002T Soil, Nantou County, Taiwan JN899305 JN899319 Penicillium canescens CBS 300.48NT = ATCC 10419 = IMI 028260 = MUCL 29169 = NRRL 910 Soil, England JN121636 Penicillium catenatum CBS 352.67T = ATCC 18543 = IMI 136241 Desert soil, Upington, Cape Province, South Africa JN121659 Penicillium cinnamopurpureum CBS 490.66 = ATCC 18337 = IMI 114483 Cultivated soil, South Africa JN121690 Penicillium citrinum CBS 139.45T = ATCC 1109 = IMI 091961 = MUCL 29781 = NRRL 1841 Unknown source JN121585 Penicillium coalescens CBS 103.83T Soil under Pinus sp., near Vulladolid, Spain RPB1 ITS JN121716 JN899366 Penicillium concavorugulosum CBS 898.73 = ATCC 20202 Unknown substrate, Japan JN899304 JN899390 Penicillium crateriforme CBS 184.27T = FRR 1057 = IMI 094165 = LSHB P164 = MUCL 29224 = NRRL 1057 Soil, Luisiana JN680270 JN899373 Penicillium dendriticum CBS 660.80T = IMI 216897 Leaf litter of Eucalyptus pauciflora, Kosciusko National Park, New South Wales, Australia JN121714 JN899339 Penicillium diversum CBS 320.48T = ATCC 10437 = DSM 2212 = IMI 040579 = IMI 040579ii = NRRL 2121 Leather, USA JN680297 JN899341 Penicillium duclauxii CBS 322.48T = ATCC 10439 = IMI 040044 = MUCL 28672 = MUCL 29094 = MUCL 29212 = NRRL 1030 Canvas, France JN121643 JN899342 Penicillium echinosporum CBS 293.62T = ATCC 18319 = DSM 2230 = FRR 3411 = IMI 080450 = IMI 101214 Wood pulp, Surrey, Kenley, UK Penicillium erythromellis CBS 644.80T = FRR 1868 = IMI 216899 Soil from creek bank, Little River, New South Wales, Australia JN680315 Penicillium euglaucum CBS 323.71NT Soil, Argentina JN121644 Penicillium expansum CBS 325.48 = ATCC 7861 = IBT 5101 = IMI 039761= MUCL 29192 = NRRL 976 Fruit of Malus sylvestris, USA JN121645 Penicillium fellutanum CBS 229.81NT = ATCC 10443 = CBS 326.48 = FRR 746 = IFO 5761 = IMI 039734 = IMI 039734iii = NRRL 746 Unknown source, USA JN121605 Penicillium funiculosum CBS 272.86NT = IMI 193019 Lagenaria vulgaris, India JN680288 Penicillium glabrum CBS 125543 = IBT 22658 = IMI 91944 Unknown source JN121717 www.studiesinmycology.org T NT JN899363 JN899383 JN899377 163 SaMSon eT al. Table 1. (Continued). Name Collection no. Origin GenBank Accession number Penicillium herquei CBS 336.48T = ATCC 10118 = FRR 1040 = IMI 028809 = MUCL 29213 = NRRL 1040 Leaf, France Penicillium ilerdanum CBS 168.81T = IJFM 5596 = IMI 253793 Air, Madrid, Spain Penicillium isariiforme CBS 247.56T = ATCC 18425 = IMI 060371 = MUCL 31191 = MUCL 31323 = NRRL 2638 Woodland soil, Zaire JN121616 Penicillium islandicum CBS 338.48NT = ATCC 10127 = IMI 040042 = MUCL 31324 = NRRL 1036 Unknown source, Cape Town, South Africa JN121648 Penicillium janthinellum CBS 340.48NT = ATCC 10455 = IMI 040238 = NRRL 2016 Soil, Nicaragua JN131650 Penicillium javanicum CBS 341.48T = ATCC 9099 = IMI 039733 = MUCL 29099 = NRRL 707 Root of Camellia sinensis, Indonesia, Java JN121651 Penicillium kewense CBS 344.61T = ATCC 18240 = IMI 086561= MUCL 2685 = NRRL 3332 Culture contaminant of mineral oil CMI 1959; Kew, Surrey, UK JN121654 Penicillium korosum CBS 762.68T Rhizosphere, India Penicillium lapidosum CBS 343.48 = ATCC 10462 = IMI 039743 = NRRL 718 Canned blueberry, Washington, USA JN121653 Penicillium liani CBS 225.66T = ATCC 18325 = ATCC 18331 = IMI 098480 = NRRL 3380 = VKM F-301 Soil, China JN680280 JN899395 Penicillium loliense CBS 643.80T = ATCC 52252 = FRR 1798 = IMI 216901 = MUCL 31325 Lolium, Palmerston North, New Zealand JN680314 JN899379 Penicillium marneffei CBS 388.87T = ATCC 18224= CBS 334.59 = IMI 068794ii = IMI 068794iii Rhizomys sinensis (bamboo rat), Vietnam JN899298 JN899344 Penicillium minioluteum CBS 642.68T = IMI 089377 = MUCL 28666 Unknown source JN121709 JN899346 Penicillium mirabile CBS 624.72T = CCRC 31665 = FRR 1959 = IMI 167383 = MUCL 31206 Forest soil, Crimea, Ukraine JN680312 JN899322 Penicillium namylowskii CBS 353.48T = ATCC 11127 = IMI 040033 = MUCL 29226 = NRRL 1070 Soil under Pinus sp., Puszceza Bialowieska, square “652”, Poland JN121660 Penicillium oblatum CBS 258.87T = FRR 2234 Spoiled baby food, Sydney, New South Wales, Australia JN680285 Penicillium ochrosalmoneum CBS 489.66 = ATCC 18338 = IMI 116248ii Cornmeal, South Africa JN121689 Penicillium osmophilum CBS 462.72T = IBT 14679 Agricultural soil, Wageningen, Netherlands JN121683 Penicillium palmae CBS 442.88 = IMI 343640 Seed, Wageningen, Netherlands JN680308 JN899396 Penicillium panamense CBS 128.89T = IMI 297546 Soil, Barro Colorado Island, Panama JN899291 JN899362 Penicillium phialosporum CBS 233.60T = ATCC 18481 = FRR 203 = IMI 078256 Milled Californian rice, California, USA JN680282 JN899340 Penicillium piceum CBS 361.48T = ATCC 10519 = IMI 040038 = NRRL 1051 Unknown source Penicillium pinophilum CBS 631.66NT = ATCC 36839 = CECT 2809 = DSM 1944 = IAM 7013 =IMI 114933 PVC, Centre d’Études du Bouchet, M. Magnoux, France JN680313 JN899382 Penicillium pittii CBS 139.84T = IMI 327871 Clay soil, under poplar trees, bank of Duero River, Valladolid, Spain JN680274 JN899325 Penicillium primulinum CBS 321.48T = ATCC 10438 = CBS 439.88 = FRR 1074 = IMI 040031 = MUCL 31321 = MUCL 31330 = NRRL 1074 USA JN680298 JN899317 Penicillium proteolyticum CBS 303.67T = ATCC 18326 = NRRL Granite soil, Ukraine 3378 JN680292 JN899387 Penicillium pseudostromaticum CBS 470.70T = ATCC 18919 = FRR 2039 JN899300 JN899371 RPB1 164 T T Feather, near Itasca State Park, Hubbard Co., Minnesota, USA ITS JN121647 JN899311 JN899318 JN899347 JN899364 JN899370 PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Table 1. (Continued). Name Collection no. Origin RPB1 ITS Penicillium purpurogenum CBS 286.36T = IMI 091926 Unknown source JN680271 JN899372 Penicillium purpurogenum var. rubisclerotium CBS 274.95 Sculpture, castle Troja, Prague, Czech Republic JN899295 JN899316 CBS 270.35T = ATCC 4713 = ATCC 52244 = FRR 1064 = IBT 4302 = MUCL 29225 = NRRL 1064 = NRRL 1142 Zea mays, Castle Rock, Virginia, USA JN680287 JN899381 Penicillium rademirici CBS 140.84T = CECT 2771 = IMI 282406 = IMI 327870 Air under willow tree, bank of river Duero, Herrera, Valladolid, Spain JN899386 Penicillium radicum CBS 100489T = FRR 4718 Root of seedling of Triticum aestivum, Wagga Wagga, New South Wales, Australia JN899324 Penicillium rotundum CBS 369.48T = ATCC 10493 = IMI 040589 = NRRL 2107 Wood, Chiriqui Prov., Panama JN899353 Penicillium rubicundum CBS 342.59T = ATCC 13217 = IMI 099723 = NRRL 3400 Soil, Georgia, USA "Penicillium rubrum" CBS 196.88 = FRR1714 Unknown source JN680278 JN899312 CBS 206.89 = IFO 6580 Japan JN680279 JN899313 CBS 263.93 Bronchoalveolair lavage of immunecompetent female patient with pneumonia by Nocardia JN680286 JN899315 Penicillium rugulosum CBS 371.48T = ATCC 10128 = IMI 040041 = MUCL 31201 = NRRL 1045 Tuber (Solanum tuberosum), Connecticut, USA JN680302 JN899374 Penicillium sabulosum CBS 261.87T = FRR 2743 Spoiled pasteurized fruit juice, New South Wales, Sydney, Australia JN899294 Penicillium samsonii CBS 137.84T = CECT 2772 = IMI 282404 = IMI 327872 Fruit, damaged by insect, Valladolid, Spain JN680273 Penicillium shearii CBS 290.48T = ATCC 10410 = IMI 039739 = IMI 039739iv = NRRL 715 Soil, Tela, Honduras JN121631 Penicillium siamense CBS 475.88T = IMI 323204 Forest soil, Lampang, Thurn District, Ban Daen Tham, Thailand Penicillium simplicissimum CBS 372.48NT = ATCC 10495 = IMI 039816 Flannel bag, Cape, South Africa Penicillium stipitatum CBS 375.48T = ATCC 10500 = NRRL Rotting wood, Louisiana, USA 1006 = IMI 39805 JN680303 Penicillium stolkiae CBS 315.67T = IMI 136210 = ATCC 18546 Peaty forest soil, Eastern Transvaal, South-Africa JN680295 Penicillium tardum CBS 258.37T = NRRL 2116 Unknown source JN899293 CBS 378.48T = ATCC 10503 = IMI 040034 = NRRL 1073 Dead twig, France JN899297 Penicillium tularense CBS 430.69T = ATCC 22056 = IMI 148394 Soil, under Pinus ponderosa and Quercus kelloggii, Tulare Co., Pine Flat, California, USA JN121681 Penicillium variabile CBS 385.48NT = ATCC 10508= IMI 040040 = NRRL 1048 Cocos fibre, Johannesburg, South Africa JN680304 JN899343 Penicillium varians CBS 386.48T = ATCC 10509 = IMI 040586 = NRRL 2096 Cotton yarn, UK JN680305 JN899368 Penicillium verruculosum CBS 388.48NT = ATCC 10513= DSM 2263= IMI 040039 = NRRL 1050 Soil, Texas, USA Penicillium victoriae CBS 274.36T = IMI 058412 = MUCL 9651 Dried leaf, Tobaheide, Sumatra JN680289 Penicillium viridicatum CBS 390.48NT = ATCC 10515= IBT 23041 = IMI 039758 = IMI 039758ii = NRRL 963 Air, District of Columbia, Washington D.C., USA JN121668 Phialosimplex caninus CBS128032T = UAMH 10337 Bone marrow aspirate ex canine, San Antonio, Texas, USA JN121587 Phialosimplex chlamydosporus CBS 109945T = FMR 7371 = IMI 387422 Disseminated infection in a dog JN121566 Phialosimplex sclerotialis CBS 366.77T = IAM 14794 Fodder of ray-grass and lucerne, France JN121661 Rasamsonia eburnea CBS 100538 = IBT 17519 Soil, Taipei, Taiwan JN680325 www.studiesinmycology.org T GenBank Accession number JN680301 JN899384 JN899369 JN899385 JN121662 JN899348 JN899367 JN899393 165 SaMSon eT al. Table 1. (Continued). Name Collection no. Origin GenBank Accession number Rasamsonia argillacea CBS 101.69T = IMI 156096 = IBT 31199 Mine tip with a very high surface temperature; Staffordshire, UK JN121556 Rasamsonia byssochlamydoides CBS 413.71T = IBT 11604 Dry soil under Douglas fir, Oregon, USA JN121675 Rasamsonia emersonii CBS 393.64 = DTO 48I1 = IBT 21695 = ATCC 16479 = IMI 116815 = IMI 116815ii Compost, Italy JN121670 Sagenoma viride CBS 114.72T ATCC 22467 = NRRL 5575 Soil, Australia JN121571 Sagenomella bohemica CBS 545.86T = CCF 2330 = IAM 14789 Peloids for balneological purposes, Frantiskovy Lázne Spa, West Bohemia, Czech Republic JN121699 Sagenomella diversispora CBS 398.69 Forest soil under Populus tremuloides, Petawawa, Ontario, Canada JN121673 CBS 399.69 = MUCL 15012 Forest soil under Thuja occidentalis, Aberfoyle, Ontario, Canada JN121674 Sagenomella griseoviridis CBS 426.67 T = ATCC 18505 = IMI 113160 Unknown source JN121677 Sagenomella humicola CBS 427.67T = ATCC 18506 = IMI 113166 Forest soil under Thuja occidentalis, Ontario, Canada JN121678 Sagenomella striatispora CBS 429.67T = ATCC 18510 = IMI 113163 Soil, Guelph, Ontario, Canada JN121679 Sagenomella verticillata CBS 415.78A Gymnosperm forest soil, Sweden JN680307 Sclerocleista ornata CBS 124.53 = ATCC 16921 = IMI 055295 = MUCL 15643 = NRRL 2256 Soil in oak forest, Dane Co., Madison, Wisconsin, USA JN121581 Talaromyces assiutensis CBS 118440 Soil, Fes, Morocco CBS 147.78T Soil, amended with crushed buffalo hoofs and incubated for 5 months at 35 oC, Egypt JN680275 JN899323 Talaromyces austrocalifornicus CBS 644.95T = IBT 17522 Soil, campus Univ. South California, Los Angelos, USA JN680316 JN899357 Talaromyces bacillisporus CBS 296.48T = ATCC 10126 = IMI 040045 = NRRL 1025 Begonia leaf, New York City, New York, USA JN121634 JN899329 Talaromyces barcinensis CBS 649.95T = IBT 17518 Soil, Barcelona, Spain JN680318 JN899349 Talaromyces brevicompactus CBS 102661T = AS 3.4676 Moulded vegetables, Prov. Sechuan, Wolong, China JN680326 Talaromyces convolutus CBS 100537T = IBT 14989 Soil, Kathmandu, Nepal JN121553 JN899330 Talaromyces cyanescens CBS 114900 = FMR 8388 Tortosa, Catalina, Spain Talaromyces derxii CBS 412.89T = NHL 2981 Cultivated soil, Okayama Prefecture, Kurashiki City, Higashitomii, Japan JN680306 JN899327 CBS 413.89T = NHL 2982 Cultivated soil, Okayama Prefecture, Kurashiki City, Higashitomii, Japan JN899299 JN899326 Talaromyces emodensis CBS 100536T = IBT 14990 Soil, Kathmandu, Nepal JN121552 JN899337 Talaromyces flavus CBS 310.38 = IMI 197477 = NRRL 2098 Unknown substrate, New Zealand JN121639 JN899360 Talaromyces galapagensis CBS 751.74T = IFO 31796 Shaded soil under Maytenus obovata, Isla Santa Cruz, Galapagos Islands, Ecuador JN680321 JN899358 Talaromyces gossypii CBS 645.80T = FRR 1966 = IMI 198365 Gossypium, India JN680317 JN899334 Talaromyces helicus var. boninensis CBS 650.95T = IBT 17516 Lawn soil, Kominato, Chichijima, Ogasawaramura, Tokyo-to, Japan JN680319 JN899356 Talaromyces helicus var. helicus CBS 335.48T = ATCC 10451 = DSM 3705 = IMI 040593 = NRRL 2106 Soil, Sweden JN680300 JN899359 Talaromyces helicus var. major CBS 652.66T = IMI 100914 Swamp soil, near Attenborough, Nottingham, UK JN680320 JN899335 Talaromyces indigoticus CBS 100534T = IBT 17590 Soil, Nagasaki-ken, Minamikushiyama-mura, Japan JN680323 JN899331 Talaromyces intermedius CBS 152.65T = BDUN 267 = IFO 31752 = IMI 100874 Alluvial pasture and swamp soil, Attenborough, Nottingham, England JN680276 JN899332 RPB1 166 T NT NT ITS JN899400 JN899320 JN899391 PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Table 1. (Continued). Name Collection no. Origin GenBank Accession number Talaromyces leycettanus CBS 398.68T = ATCC 22469 = IMI 178525 Coal spoil tip soil, Leycett, Staffordshire, England, UK JN121672 Talaromyces luteus CBS 348.51NT = IMI 089305 Soil, UK JN121656 Talaromyces macrosporus CBS 317.63 = FRR 404 = IMI 197478 Apple juice, Stellenbosch, South Africa JN680296 JN899333 Talaromyces mimosinus CBS 659.80T = FRR 1875 = IMI 223991 Soil from creek bank, Nattai River, New South Wales, Australia JN899302 JN899338 Talaromyces muroii CBS 756.96T = PF 1153 Soil, Hualien County, Chingpu, Taiwan JN680322 JN899351 Talaromyces ocotl CBS 102855 Heat-treated soil from forest of Pinus hartwegii, Veracruz, Mexico JN680327 Talaromyces ohiensis CBS 127.64T Soil treated with cyanamide, Germany JN680272 JN899355 Talaromyces purpureus CBS 475.71T = ATCC 24069 = ATCC 52513 = FRR 1731 = IMI 181546 Soil, near Esterel, France JN121687 JN899328 Talaromyces subinflatus CBS 652.95T = IBT 17520 Copse soil, Hahajima, Ogasawara-mura, Tokyo-to, Japan JN899301 JN899397 Talaromyces tardifaciens CBS 250.94T Unknown source JN680283 JN599361 Talaromyces thermophilus CBS 236.58T = ATCC 10518 = IMI 048593 = NRRL 2155 Parthenium argentatum, decaying plant; California, USA JN121611 Talaromyces trachyspermus CBS 373.48T = ATCC 10497 = IMI 040043 = NRRL 1028 Unknown source, USA JN121664 JN899354 Talaromyces ucrainicus CBS 162.67T = ATCC 22344 = FRR 3462 Unknown source JN680277 JN899394 Talaromyces udagawae CBS 579.72T = FRR 1727 = IMI 197482 Soil, Misugimura, Japan JN680310 JN899350 Talaromyces unicus CBS 100535T = CCRC 32703 = IBT 18385 Soil, Chiayi County, Funlu, Taiwan JN680324 JN899336 Talaromyces wortmanii CBS 391.48T = ATCC 10517 = IMI 040047 = NRRL 1017 Unknown source JN121669 JN899352 Thermoascus aurantiacus CBS 396.78 Sawdust, in lumber yard, Toronto, Ontario, Canada JN121671 CBS 891.70 = IMI 173037 Wood, Firenze, Italy JN121719 Thermoascus crustaceus CBS 181.67 = ATCC 16462 = IMI 126333 Parthenium argentatum, decaying plant; Salinas, California, USA JN121591 Thermoascus thermophilus CBS 528.71NT = IMI 123298 = NRRL 5208 Wood and bark of Pinus, Sweden JN121697 Thermomyces lanuginosus CBS 218.34 = MUCL 8338 Fruit shell of Theobroma cacao JN121599 CBS 224.63 = MUCL 8337 Mushroom compost; Gossau-Zürich Switzerland JN121602 CBS 288.54 = MUCL 8340 Stomach of bovine foetus, Netherlands JN680291 CBS 103.73 Unknown source, Japan JN121558 CBS 247.57 = MUCL 39666 = IBT 31159 Unknown source, Hachijô, Japan JN121617 CBS 788.83 Rotting stump of cut down tree, Myojoji Temple near Hakui Noto Park, Ishikawa Pref., Japan JN121718 CBS 512.65NT = ATCC 16919 = IMI 075885 = NRRL 4376 Jungle soil, Berakas-Muara, Brunei JN121692 RPB1 Trichocoma paradoxa Warcupiella spinulosa T T T completely undetermined characters in the alignment was 0.60 %. Figure 1 shows that members of the subgenus Biverticillium and Talaromyces are accommodated in a well-supported (97 % bs), monophyletic clade (= Talaromyces s. str.) and that species of the Penicillium subgenera Aspergilloides, Furcatum and Penicillium form an independent, well-supported clade (Penicillium s. str.). The majority of described Talaromyces species belong to Talaromyces s. str., but some species are dispersed in other clades, including Talaromyces ocotl, T. luteus, T. thermophilus, T. www.studiesinmycology.org ITS JN899398 eburneus, T. emersonii, T. byssochlamydoides, T. spectabilis, T. brevicompactus, T. striatus and T. leycettanus. Talaromyces ocotl is in a well-supported clade with the type species of Sagenomella, S. diversispora, and other Sagenomella species. The former T. emersonii, T. eburneus and T. byssochlamydoides form a clade recently recognised and described as the genus Rasamsonia (Houbraken et al. 2011). Talaromyces thermophilus is also excluded from Talaromyces s. str. and is closely related to the type species of Thermomyces, Therm. lanuginosus. Basal to Therm. lanuginosus 167 SaMSon eT al. 100 CBS 196.88 “Penicillium rubrum” CBS 342.59T Penicillium rubicundum CBS 100534T Talaromyces indigoticus CBS 312.59T Penicillium aculeatum var. apiculatum CBS 274.36T Penicillium victoriae CBS 751.74T Talaromyces galapagensis 100 CBS 413 413.89 89T Talaromyces T l d derxiiii 92 CBS 412.89T Talaromyces derxii T CBS 274.36 Penicillium victoriae CBS 225.66T Penicillium liani 100 CBS 321.48T Penicillium primulinum CBS 320.48T Penicillium diversum CBS 252.87T Geosmithia viridis CBS 272.86NT Penicillium funiculosum 78 CBS 317.63T Talaromyces macrosporus CBS 334.59T Penicillium marneffei 100 CBS 388.87T Penicillium marneffei 96 CBS 112002T Penicillium calidicanium CBS 322 322.48 48T Penicillium duclauxii 79 CBS 310.38NT Talaromyces flavus CBS 128.89T Penicillium panamense CBS 114.72T Sagenoma viride CBS152.65T Talaromyces intermedius 96 CBS 545.86T Sagenomella bohemica 100 CBS 350.66T Paecilomyces aerugineus 100 CBS 267.72 “Aphanoascus cinnabarinus” CBS 756.96T Talaromyces muroii T CBS 649.95 Talaromyces barcinensis 96 CBS 335.48T Talaromyces helicus var. helicus 72 98 CBS 652.66T Talaromyces helicus var. major 100 CBS 650.95T Talaromyces helicus var. boninensis CBS 386 386.48 48T Penicillium P i illi varians i 100 CBS 184.27T Penicillium crateriforme T CBS 286.36 Penicillium purpurogenum CBS 375.48 Talaromyces stipitatus T 100 CBS 373.48T Talaromyces trachyspermus CBS 147.78 Talaromyces assiutensis 98 CBS 645.80T Talaromyces gossypii CBS 127.64T Talaromyces ohiensis CBS 162.67T Talaromyces ucrainicus 98 100 CBS 642.68T Penicillium minioluteum 99 CBS 270.35T Penicillium purpurogenum var. rubisclerotiorum CBS 137.84T Penicillium samsonii 99 94 CBS 579.72T Talaromyces udagawae 100 CBS 624 624.72 72T Penicillium P i illi mirabile i bil 99 CBS 100537T Talaromyces convolutus CBS 644.95T Talaromyces austrocalifornicus 99 CBS 644.80T Penicillium erythromellis CBS 206.89 “Penicillium rubrum” CBS 660.80T Penicillium dendriticum 100 CBS 253.87T Paecilomyces pascuus 98 CBS 258.87T Penicillium oblatum CBS 470.70T Penicillium pseudostromaticum 98 100 CBS 139.84T Penicillium pittii CBS 475.71T Talaromyces purpureus NT Penicillium islandicum 80 CBS 338.48 T 95 CBS 643.80 T Penicillium loliense CBS 227.60 Penicillium brunneum 90 CBS 453.93T Penicillium allahabadense CBS 385.48NT Penicillium variabile 100 CBS 391.48T Talaromyces wortmanii 98 CBS 898.73T Penicillium concavorugulosum CBS 250.94T Talaromyces tardifaciens CBS 233.60T Penicillium phialosporum 95 CBS 371.48T Penicillium rugulosum CBS 303.67T Penicillium proteolyticum 100 CBS 100535T Talaromyces unicus CBS 100536T Talaromyces emodensis CBS 659.80T Talaromyces mimosinus CBS 296.48T Talaromyces y bacillisporus p 85 CBS 442.88T Penicillium palmae CBS 652.95T Talaromyces subinflatus 91 97 CBS 258.37T Penicillium tardum CBS 274.95 Penicillium purpurogenum var. rubisclerotiorum 100 CBS 263.93 “Penicillium rubrum” 93 CBS 631.66NT Penicillium pinophilum 97 Clade 1 Talaromyces Clade 2A Clade 2B Fig. 1. Best-scoring Maximum Likelihood tree calculated using RAxML, based on partial RPB1 sequences showing the relationships among members of Talaromyces and Penicillium subgenus Biverticillium and related genera. The bootstrap support percentages of the maximum likelihood (ML) analysis are presented at the nodes. Bootstrap support values less than 70 % are not shown and branches with bootstrap support values > 70 % are thickened. The bar indicates the number of substitutions per site. The tree is rooted with Coccidioides immitis (strain RS). and T. thermophilus is Talaromyces luteus. This species is on a separate branch and no other closely related species were found in our analysis. The uniqueness of the species is supported by the production of large amounts of the prenylated diketopiperaziners talathermophilins A and B, not found in any other species (Chu et al. 2010). The phylogenetic position of T. leycettanus is not convincingly defined. This species is positioned near Warcupiella 168 spinulosa and Hamigera striata (= Talaromyces striatus), but bootstrap support is lacking. Talaromyces brevistipitatus occurs on a well-supported branch with H. avellanea. Comparison of ITS and calmodulin sequences shows that this species is closely related to NRRL 2108, an undescribed, phylogenetically distinct Hamigera species (ITS 100 % bs, calmodulin 99 % bs) (Peterson et al. 2010). The majority of members of subgenus Biverticillium sensu PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium 97 94 76 CBS 426.67T Sagenomella griseoviridis CBS 102855T Talaromyces ocotl 88 CBS 398.69 Sagenomella diversispora CBS 399.69 Sagenomella diversispora 100 CBS 429.67T Sagenomella striatispora 100 CBS 427.67T Sagenomella humicola CBS 415.78A 415 78AT Sagenomella S ll verticillata ti ill t CBS 288.54 Thermomyces lanuginosus 100 CBS 224.63 Thermomyces lanuginosus 100 CBS 218.34 Thermomyces lanuginosus 97 CBS 236.58T Talaromyces thermophilus 92 NT CBS 348.51 Talaromyces luteus CBS 788.83 Trichocoma paradoxa 100 CBS 247.57 Trichocoma paradoxa CBS 103.73 Trichocoma paradoxa 98 CBS 101.69T Rasamsonia argillacea 73 CBS 100538T Rasamsonia eburnea 90 CBS 393.64 393 64T Rasamsonia emersonii CBS 413.71T Rasamsonia byssochlamydoides CBS 101075T Byssochlamys spectabilis CBS 100.11T Byssochlamys nivea CBS 336.48T Penicillium herquei CBS 125543NT Penicillium glabrum CBS 229.81NT Penicillium fellutanum CBS 343.48T Penicillium lapidosum 96 T CBS 353.48 Penicillium namyslowskii CBS 261.87T Penicillium sabulosum CBS 352.67T Penicillium catenatum 100 CBS 340.48 340 48NT Penicillium janthinellum 100 CBS 341.48T Penicillium javanicum CBS 372.48NT Penicillium simplicissium CBS 323.71NT Penicillium euglaucum 92 CBS 290.48T Penicillium shearii CBS 139.45T Penicillium citrinum CBS 315.67T Penicillium stolkiae 78 CBS 247.56T Penicillium isariiforme CBS 489.66T Penicillium ochrosalmoneum CBS 490.66T Penicillium cinnamopurpureum 99 CBS 390.48NT Penicillium viridicatum CBS 325.48 Penicillium expansum CBS 378.48T Penicillium tardum 98 CBS 344.61T Penicillium kewense 100 CBS 462.72T Penicillium osmophilum 100 CBS 430.69T Penicillium tularense CBS 300.48NT Penicillium canescens 86 CBS 506.65T Aspergillus zonatus CBS 761.68 Penicilliopsis clavariiformis CBS 473.65T Aspergillus clavatoflavus 70 CBS 220.66T Penicillium arenicola CBS 124.53T Sclerocleista ornata 100 CBS 109945T Phialosimplex p chlamydosporus y p 93 CBS 128032T Phialosimplex caninus T CBS 366.77 Phialosimplex sclerotialis Af293 Aspergillus fumigatus NRRL 3357 Aspergillus flavus 81 DTO 11C3 Aspergillus penicillioides CBS 516.65NT Eurotium herbariorum CBS 513.88 Aspergillus niger CBS 172.66 Aspergillus aculeatus 94 CBS 264.81 Aspergillus sydowii 77 CBS 245.65 Aspergillus versicolor CBS 112.46 Emericella nidulans 91 CBS 139.61 139 61 Aspergillus A ill sparsus CBS 101887 Aspergillus ochraceoroseus 99 CBS 108.08NT Aspergillus ochraceus CBS 112812T Aspergillus steynii CBS 649.93T Aspergillus robustus CBS 512.65T Warcupiella spinulosa CBS 398.68T Talaromyces leycettanus CBS 377.48T Hamigera striata 100 CBS 102661T Talaromyces brevicompactus CBS 295.48T Hamigera avellanea 86 CBS 132.31T Chrysosporium inops CBS 109.07 109 07T Monascus M purpureus 90 CBS 181.67T Thermoascus crustaceus 100 NT CBS 528.71 Thermoascus thermophilus 98 CBS 605.74T Byssochlamys verrucosa 100 CBS 891.70 Thermoascus aurantiacus CBS 396.78 Thermoascus aurantiacus Strain ‘RS’ Coccidioides immitis 95 0.1 Sagenomella Thermomyces Trichocoma Rasamsonia Paecilomyces / Byssochlamys Penicillium s. str. Penicilliopsis Miscellaneous Phialosimplex Aspergillus s. str. Hamigera, Warcupiella Monascus Thermoascus Fig. 1. (Continued). Pitt (1980) are phylogenetically placed within Talaromyces s. str., with P. isariiforme as the only exception. This species belongs to Penicillium s. str. and is closely related to P. ochrosalmoneum. This relationship was also confirmed by extrolite data (see below). Figure 1 indicates that the following species phylogenetically belong in Talaromyces: Aphanoascus cinnabarinus (CBS 267.72), www.studiesinmycology.org Sagenomella bohemica (CBS 545.86T), Paecilomyces aerugineus (CBS 350.66T), Geosmithia viridis (CBS 252.87T) and Sagenoma viride (CBS 114.72T). The former three strains are on a wellsupported sister clade basal to Talaromyces muroii CBS 756.96. 169 SaMSon eT al. Species delimitation and synonymies within Talaromyces The ITS analysis (Fig. 2) was used in this study to provide a preliminary circumscription of the species belonging to the Talaromyces clade. Ninety-seven strains were included in the ITS analysis. The used primer pair V9G and LS266 also amplifies a part of the 18S and 28S rDNA; however, for analysis, only the span including the ITS regions and 5.8S rDNA was used. The length of the alignment was 483 characters and 221 characters were variable. Most bootstrap support values in the ITS analysis are low, less than 70 %. Only a few branches are supported with values higher than 70 %. The majority of Talaromyces species are on a branch with 96 % bootstrap support (clade 1, Fig. 2). This clade is also present in the RPB1 analysis (100 % bs). Another large clade was present in the ITS phylogram and this clade is supported with 96 % boostrap (clade 2). This clade can be divided in two subclades (2A and 2B), both present in the RPB1 analysis; however, the relationship among these subclades is not supported statistically. Talaromyces dendriticus, T. oblatus, and Paecilomyces pascuus are in the same lineage and the former two species share the same ITS sequence. Talaromyces assiutensis and T. gossypii also have similar ITS sequences and are phenotypically similar (Frisvad et al. 1990a). Extrolite analysis In general, Talaromyces species produce many biosynthetic families of polyketides and meroterpenoids, but rather few families of nonribosomal peptides and terpenes. By examining HPLCDAD results from all described species of Penicillium, Aspergillus and their teleomorphs, and by searching the literature for families of exometabolites produced by these fungi, it is obvious that Talaromyces species have unique and specific extrolites (Table 2). Figure 3 shows the common exometabolite families in Talaromyces/ Biverticillium, Penicillium, Aspergillus and other genera. Aspergillus and Penicillium share 91 biosynthetic families, but shares more of these with other fungal genera than with Talaromyces. A few exometabolites are shared among Talaromyces, Penicillium and Aspergillus including alternariols, asperphenamate, botryodiploidin, dehydrocarolic acid, emodins, geodins, gregatins, herqueinone, 3-hydroxyphtalic acid, italinic acid, lichexanthones, mellein, monordens, pinselin, rugulosuvines, rugulovasines, secalonic acids and zeorins. Most of these metabolites have relatively simple structures, and many occur in other genera less related phylogenetically to any of the penicilloid and aspergilloid genera. Considering the large number of shared exometabolite biosynthetic families in common between Penicillium and Aspergillus, Talaromyces is clearly different, which corresponds with all other data for these genera. Among the few extrolites shared by Penicillum, Aspergillus and Talaromyces are the ergochromes, secalonic acid D & F. These anthraquinone derived metabolites are found in P. isariiforme, P. chrysogenum, Aspergillus aculeatinus, P. dendriticum and P. pseudostromaticum (Samson et al. 1989, Frisvad & Samson 2004, Houbraken et al. 2011). It is also possible that there are optical antipodes of these compounds produced in these genera, as was found in Aspergillus versicolor ((+) versicolamide)) and A. sclerotiorum ((-)-versicolamide) (Williams 2011). If this is so, it may indicate that the extrolites of Talaromyces and Penicillium / 170 Aspergillus may also differ in stereochemical aspects. Another example of shared yet different extrolites is the azaphilones, which are common in species of Talaromyces and related biverticillate anamorphic species (Frisvad et al. 1990a, Nicoletti et al. 2009, Osmanova et al. 2010), but could not be found in Aspergillus and Penicillium sensu stricto. When similar compounds were found in Talaromyces, stereoisomers of the compounds were found in Aspergillus and Penicillium. For example, while sclerotiorins occur in P. sclerotiorum, the epimers are found in Talaromyces helicus and T. luteus (Yoshida et al. 1995, 1996a, b). Austdiol was isolated from Aspergillus pseudoustus (Vleggaar et al. 1974, Samson et al. 2011), but 7-epi-austdiol from a Talaromyces species (Liu et al. 2010). Misidentifications of strains can make these comparisons difficult, but the overwhelming majority of extrolites found in Talaromyces are not found in Aspergillus or Penicillium. Although vermistatins, penisimplisins, penisimplicissins were reported from Penicillium simplicissimum (Komai et al. 2005), the producing strain was misidentified and actually represents a species of Talaromyces. The opposite has also happened, and metabolites attributed to a species of subgenus Biverticillium are later found to be produced by species of Penicillium sensu stricto. Penicillium verruculosum was reported to produce verruculogen, hence the name (Cole et al. 1972, Cole & Kirksey 1973), but the strain was later reidentified as P. brasilianum (Frisvad 1989). Penicillium isariiforme (Samson et al. 1989) and P. ochrosalmoneum (Wicklow & Cole 1984) both produce large amounts of citreoviridin, supporting their close relationship indicated by the phylogenetic analyses, as noted above (Fig. 1). DISCUSSION The symmetrical, biverticillate penicillus was used as a defining character by Wehmer (1914), and Thom (1915a, b). Wehmer (1914) proposed to call this group the Verticillata, while Thom (1915a) referred to it as the Penicillium luteum-purpurogenum group. Biourge (1923) was the first who named this group as the subgenus Biverticillium, but included species such as P. citrinum (as P. aurifluum), P. atramentosum etc., which are no longer regarded as members of this subgenus (Houbraken et al. 2010). The characteristic lanceolate or acerose phialides was used as a more definitive morphological character of subgenus Biverticillium and related Talaromyces anamorphs (Raper & Thom 1949), because biverticillate branched conidiophores with flaskshaped phialides are mainly found in unrelated species such as P. citrinum. Although the lanceolate phialides occur in most species of subgenus Biverticillium, some species, e.g. P. rugulosum, have phialides that are not slender and have an apical portion tapering into a long acuminate point. Thom (1930) treated some of the Penicillia in his BiverticillateSymmetrica group and distinguished four sections: Ascogena, Coremigena, Luteo-virida (Funiculosa and Luteo-purpurogena) and Miscellanea. Later Raper & Thom (1949) subdivided the group into the P. luteum series, P. duclauxii series, P. funiculosum series, P. purpurogenum series, P. rugulosum series and P. herquei series. This grouping is inconsistent with our phylogenetic analysis of the biverticillate group. The classification proposed by Pitt (1980) is more in concordance with the phylogenetic and taxonomic treatment proposed here, although he included a few species in Penicillium subgenus Biverticillium, namely P. isariiforme, P. clavigerum and PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium 96 76 0.1 96 CBS 762.68T – P. korosum CBS 631.66NT – P. pinophilum PF1150T – Erythrogymnotheca paucispora* CBS 225.66T – P. liani CBS 289.48 – P. aculeatum CBS 312.59T – P. aculeatum var. apiculatum CBS 751.74T – T. galapagensis CBS 152.65T – T. intermedius CBS 100105 – P. aculeatum CBS 102801T – P. aureocephalum CBS 274.36T – P. victoriae CBS 272.86NT – P. funiculosum CBS 274.95 – P. purpurogenum var. rubisclerotiorum 97 CBS 263 263.93 93 – “P P. rubrum rubrum” CBS 388.48NT – P. verruculosum CBS 196.88 – “P. rubrum” CBS 475.88T – P. siamense CBS 310.38NT – T. flavus CBS 317.63T – T. macrosporus 99 CBS 286.36T – P. purpurogenum CBS 184.27T – P. crateriforme CBS 375.48T – T. stipitatus CBS 413.89T – T. derxii CBS 412.89T – T. derxii CBS 342.59T – P. rubicundum CBS 114900 – T. cyanescens CBS 100534T – T. indigoticus CBS 314.59T – P. aurantiacum AF 2857821 – Sagenoma viride viride* PF 1203T – T. euchlorocarpius* T CBS 128.89 – P. panamense CBS 293.62T – P. echinosporum CBS 388.87T – P. marneffei CBS 322.48T – P. duclauxii CBS 386.48T – P. varians 75 CBS 649.95T – T. barcinensis 77 CBS 652.66T – T. helicus var. major 98 CBS 335.48T – T. helicus var. helicus CBS 650.95T – T. helicus var. boninensis CBS 545.86T – Sagonema bohemica 79 CBS 350.66T – Paecilomyces aerugineus 72 NHL 2917 – T. ryukyuensis* CBS 267.72 – “Aphanoascus cinnabarinus” CBS C S 756.96 56 96T – T. muroii uo CBS 252.87T – Geosmithia viridis CBS 112002T – P. calidicanium 91 CBS 320.48T – P. diversum CBS 321.48T – P. primulinum CBS 660.80T – P. dendriticum CBS 258.87T – P. oblatum 89 CBS 253.87T – Paecilomyces pascuus 100 CBS 103.83T – P. coalescens 99 97 DAOM 233329T – P. cecidicola* KAS 2792 – P. ramulosum* 94 CBS 470.70T – P. pseudostromaticum T CBS 139.84 – P. pittii CBS 642.68T – P. minioluteum 97 CBS 270.35T – P. purpurogenum var. rubisclerotium 96 CBS 137.84 137 84T – P. P samsonii 100 CBS 206.89 – “Penicillium rubrum” CBS 644.80T – P. erythromellis 90 CBS 624.72T – P. mirabile CBS 579.72T – T. udagawae T 99 CBS 645.80 T – T. gossypii CBS 373.48 – T. trachyspermus 99 CBS 147.78T – T. assiutensis CBS 118440 – T. assiutensis 96 100 CBS 127.64T – T. ohiensis CBS 162.67T – T. ucrainicus 100 CBS 644.95T – T. austrocalifornicus CBS 100537T – T. convolutus CBS 140.84T – P. rademirici CBS 475.71T – T. purpureus 98 CBS 168 168.81 81T – P. P ilerdanum CBS 361.48T – P. piceum CBS 371.48T – P. rugulosum CBS 227.60T – P. brunneum CBS 338.48NT – P. islandicum CBS 898.73T – P. concavorugulosum CBS 391.48T – T. wortmanii 90 AB 176638 – T. sublevisporus* CBS 385.48NT – P. variabile 98 CBS 369.48T – T. rotundus CBS 233.60T – P. phialosporum CBS 643.80T – P. loliense CBS 250.94T – T. tardifaciens CBS 100489T – P. radicum CBS 453.93T – P. allahabadense CBS 100535T – T. unicus 91 CBS 100536T – T. emodensis 92 AB176620 – T. hachijoensis* 84 CBS 303.67T – P. proteolyticum 72 CBS 659.80T – T. mimosinus CBS 296.48T – T. bacillisporus CBS 652.95T – T. subinflatus CBS 442.88T – P. palmae CBS 788.83 – Trichocoma paradoxa Clade 1 Clade 2A Clade 2B Fig. 2. Best-scoring Maximum Likelihood tree calculated using MEGA 5.0 based on ITS sequences showing the relationship among members of the Talaromyces and members of Penicillium subgenus Biverticillium. The bootstrap support percentages of the maximum likelihood (ML) analysis are presented at the nodes. Bootstrap support values less than 70 % are not shown and branches with bootstrap support values > 75 % are thickened. The bar indicates the number of substitutions per site. The tree is rooted with Trichocoma paradoxa (CBS 788.83). T. = Talaromyces; P. = Penicillium. Strains indicated with * are ITS sequencing obtained from GenBank. www.studiesinmycology.org 171 SaMSon eT al. Table 2. Secondary metabolite (exometabolite) biosynthetic families known from Talaromyces and Penicillium subgenus Biverticillium. (P) means also found in Penicillium and its teleomorphic state Eupenicillium, (A) means also found in species of Aspergillus. (Others) means also found in other fungi outside Penicillium, Aspergillus, Talaromyces and related genera. Secondary metabolite (exometabolite) biosynthetic families AF-110 5-Hydroxymethylfurfural Purpurogenones Alternariols * (P and others) Hydromethylmaltol Rasfonin Anthglutin 4-Hydroxy-4,5-dicarboxy pentadecanoic acid (T. spiculisporus) Rubratoxins Apiculides (incl. NG-011’s * (others)) 7-Hydroxy-2,5-dimethylchromane Rugulosins (& flavoskyrin) * (others) AS-186-G 3-Hydroxymethyl-6,8-dimethoxycoumarin Rugulotrosins Asperphenamates & asperglaucid * (A, P) 3-Hydroxyphthalic acid * (P) Rugulosuvine * (P) Atrovenetinon methyl acetal (P. verruculosum) Islandic acids Rugulovasines * (P) Epi-Austdiols (7-epiaustdiol & 8-O-methylepiaustdiol) (the stereoisomer austdiol found in Aspergillus) (+)-Isocitric acid + Decylcitric acid (T. spiculisporus) Secalonic acids * (A, P, others) Austins * (A, P) Italinic acids * (P) Speciferone* (others) BE-24811 Juglones Spiculisporic acids (= minioluteic acids) BE-31405’s Lichexanthone * (others) SQ 30957 Berkeleyamides Luteusins Stemphyperylenole Botryodipoidin * (P & others) Maculosin * (others) Stipitatic acids Chrodinanine A Mellein * (A) Talaperoxides Cordyanhydrides Methyl-4-carboxy-5-hydroxyphthalaldehydrate Talaroconvolutins Cyclochlorotines & islanditoxin 3-Methyl-6-hydroxy-8-methoxy-3,4-dihydroisocoumarins Talaroderxine Dehydrocarolic acids * (A, P) Miniolutelides, berkeleydione, berkeleytriones, berkeleyacetals, dhilirolides Talaroflavones Diethylphthalate (Artefact?) Mitorubrins & kasanosins & funicones Talaromycins 5,6-Dihydro-3,5-dihydroxy-6-hydroxymethyl-2H-pyran-2-one Monascins & monascorubramin Talarotoxins 4,6-Dihydroxy-5-methylphthalide Monordens * (A, others) TAN-931 (2E,2E’,7S,7’E)-4,9-Dioxo-7-(4’,9’-dioxo-2’,7’decadienoyloxy)2-decanoic acid NG-061 Thailandolides Diversonols NK-374200 Trachyspermic acids Duclauxins OF-4949’s Trachyspic acid Emodins * (A, P, others) Penicilliopsin * (others) Triacetic lactone Erythroskyrins Penisimplicins (-)-2,3,4-Trihydroxy-butanamide Flavomannin Penisimplicissins Vermicellins Funiculosic acids Penitrinic acid & penitricins Vermiculins Funiculosin Pevalic acid Vermilutins Geodins * (A, P) PF-1092A Vermistatins & penicidones Glauconic acids Pinselic acid Vertoskyrin Gregatins and penicilliols * (A, P) Pinselin * (A, others) Wortmannilactones Helicusins Purpactins ( = penicillides = vermixocins) Wortmannins * (others) Herqueinones* (P) Purpuride Xanthoradones Zeorins * (A, others) P. vulpinum (as P. claviforme) that are now classified in Penicillium sensu stricto. The same conclusion was shown by the early molecular results of LoBuglio & Taylor (1993), and subsequently supported by the physiological, morphological and extrolite characters reviewed in the Introduction, and generated during this study. In general, Penicillium sensu stricto and Aspergillus share many more features with each other than they do with Talaromyces. This includes micro- and macro-morphology, good growth on low water activity media, and the many shared exometabolite families. Talaromyces produces a series of metabolites that are apparently unique to this genus (J.C. Frisvad unpubl. data). The characteristic yellow and red colony and mycelial colours in Talaromyces are often caused by accumulation of mitorubrins and other azaphilones 172 and unique anthraquinones and mitorubrins that are not found in Aspergillus and Penicillium. Some azaphilones are found in Penicillium sclerotiorum and Penicillium hirayamae, but only their optical antipodes are found in Talaromyces. Penicillium and Talaromyces species excluded from the revised Talaromyces genus Figure 1 shows that a number of species described in the genus should be excluded from Talaromyces s. str. Phylogenetically, T. ocotl CBS 102855T belongs to Sagenomella, as also suggested using phenotypic characters (Heredia et al. 2001). The anamorph of this species was not formally named, described only as Hylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Common exometabolite families inPTalaromyces, Penicillium, Aspergillus and other genera Aspergillus p g 338 61 1 3 Penicillium P i illi 173 23 4 TTalaromyces l 85 3 2 43 1 Other genera > 1000 Fig. 3. Common exometabolite families in Talaromyces/Biverticillium, Penicillium, Aspergillus and other genera. Sagenomella sp., and thus the new combination Sagenomella ocotl is proposed in the taxonomy section below. Our analysis confirms the distinctiveness of the recently described genus Rasamsonia erected for thermotolerant or thermophilic species with distinctly rough-walled conidiphore stipes, olive-brown conidia, and ascomata, if present, with a scanty hyphal covering. Talaromyces eburneus, T. emersonii, T. byssochlamydoides were assigned to this genus, together with the anamorphic species originally described as Geosmithia argillacea and G. cylindrospora (Houbraken et al. 2011). Talaromyces thermophilus is the only member of Talaromyces section Thermophila (Stolk & Samson 1972). LoBuglio et al. (1993) already noted that this species is the most divergent Talaromyces species, occupying a basal position to the major Talaromyces clade. Houbraken et al. (2011) showed that this species is closely related to Thermomyces lanuginosus and our partial RPB1 sequence data confirm this relationship (Fig. 1). We did not examine type material of Talaromyces thermocitrinus (as ‘thermocitrinum’) and the conclusion of Mouchacca (2007), who tentatively placed this species in synonymy with T. thermophilus, is not followed here. Talaromyces luteus is further basal to T. thermophilus and Therm. lanuginosus and this species might represent a distinct genus. For the present, T. thermophilus and T. luteus will be retained in Talaromyces. More research is needed to confirm whether the assignment of these species to Thermomyces is warranted. Udagawa & Suzuki (1994) described Talaromyces spectabilis with a Paecilomyces anamorph. Houbraken et al. (2008) transferred this species to Byssochlamys and showed that it is the teleomorph of Paec. variotii. In a single name system, Paec. variotii is the oldest genus and species name for this taxon, and thus the correct name for the holomorph. Talaromyces brevicompactus, T. striatus (= Hamigera striata) and T. leycettanus are distant from Talaromyces s. str. and phylogenetically more closely related to Penicillium s. str. and Aspergillus. Figure 1 shows that H. striata and T. leycettanus are closely related. Further phylogenetic support for this relationship was presented in the studies of Ogawa & Sugiyama (2000) and Houbraken & Samson (2011). These two species are phylogenetically distant from www.studiesinmycology.org Talaromyces s. str. and more closely related to Hamigera. Peterson et al. (2010) delimited Hamigera phylogenetically but stated that T. leycettanus and H. striata do not belong to this genus, and followed Benjamin’s (1955) placement of H. striata in Talaromyces. In this study, we retain H. striata and T. leycettanus in Hamigera and Talaromyces, respectively. A thorough study on Hamigera and related genera is needed to clarify the correct placement of these species. Kong (1999) described Talaromyces brevicompactus, stating that this species is closely related to Hamigera avellanea (as Talaromyces avellaneus). The anamorph of this species was described in Merimbla, thus confirming the relationship with Hamigera. Sequence comparisons of this species showed that it is similar to NRRL 2108, a phylogenetically undescribed Hamigera species (J. Houbraken, unpubl. data, Peterson et al. 2010). We wait with combining this species in Hamigera until a more data and strains become available. Species described in other genera but phylogenetically within Talaromyces Phylogenetic analysis shows that “Aphanoascus cinnabarinus”, Sagenomella bohemica, Paecilomyces aerugineus, Geosmithia viridis and Sagenoma viride belong to Talaromyces. The genus Sagenoma is typified with S. viride, and therefore this genus can be considered as a synonym of Talaromyces. Our data support the conclusions of von Arx (1987), who correctly transferred this species in Talaromyces, and this is reflected in the taxonomy section below. Houbraken & Samson (2011) discussed the confusion over Aphanoascus cinnabarinus, which has persisted since the description of the genus Aphanoascus by Zukal (1890). Most authors follow Apinis (1968) and consider the genus Aphanoascus to be typified by A. fulvescens Zukal. In addition, the neotypification of A. cinnabarinus by Udagawa & Takada (1973) was incorrect, because their neotype strain had a Paecilomyces anamorph, whereas Zukal’s original description and illustrations clearly showed a Chrysosporium-like anamorph (Stolk & Samson 1983). Based on morphological features, Stolk & Samson (1983) indicated that Chromocleista cinnabarina (as A. cinnabarinus sensu Udagawa & Takada) belongs to the Eurotiales and suggested that this species is intermediate between Thermoascus and Talaromyces. Our phylogenetic study, and that of Houbraken & Samson (2011), clarified that C. cinnabarina belongs to Talaromyces s. str. The taxonomic position of Chromocleista cinnabarina (as A. cinnabarinus sensu Udagawa & Takada) will be discussed in a forthcoming paper. Paecilomyces aerugineus was proposed by Samson (1974) for Spicaria silvatica Oudemans sensu Apinis. This species resembles the anamorph of A. cinnabarinus sensu Udagawa & Takada and a more detailed study is necessary to clarify this relationship. TAxONOMY Penicillium itself has a long list of generic synonyms (see Seifert et al. 2011) that must be considered for the species formerly included in subgenus Biverticillium. These synonyms of Penicillium are discussed in the Appendix to this paper. As it turns out, none of these are appropriate for subgenus Biverticillium, leaving the comparatively young Talaromyces as the oldest well-known generic name as the new home for the anamorphic species of subgenus Biverticillium. 173 SaMSon eT al. Yaguchi et al. (1994a) introduced Erythrogymnotheca for the single species E. paucispora. No specimens of E. paucispora were studied; however, examination of the available ITS data on GenBank and the original description shows that this species belongs in Talaromyces. As a consequence, Erythrogymnotheca is synonymised with Talaromyces. Comparison of an ITS sequence of E. paucispora (AB176603) shows that it is related to P. korosum, P. pinophilum and P. liani in Talaromyces (Fig. 2). The original description suggests that Talaromyces and Erythrogymnotheca differ in ascus characteristics and ascospore morphology. However, these genera also share characters. The ascomatal initials of E. paucispora approximate those of Talaromyces flavus and other species of Talaromyces. Furthermore, E. paucispora produces a loose hyphaI yellow- or red-pigmented ascomata similar to those of other Talaromyces species and the main ubiquinone systems are Q-10 and Q-10 (H2), also indicating a relationship with Talaromyces (Paterson 1998, Yaguchi et al. 1994a). Matsushima (2001) described Paratalaromyces from soil collected in Taiwan, distinguishing it by a distinct textura epidermoidea layer in the ascomatal wall, and the presence of spinulose marginal hyphae. We have not seen the type but the description of Paratalaromyces lenticularis is similar to that of Talaromyces unicus (Tzean et al. 1992). We consider the genus a synonym here. Visagie & Seifert (unpubl. data) report on the generic name Lasioderma Mont., typified by L. flavo-virens Durieu & Mont., which is conspecific with Penicillium aureocephalum Munt.-Cvetk., Hoyo & Gómez-Bolea. The name Lasioderma is widely used as an insect genus, and a formal proposal for the conservation of Talaromyces against this older name is being prepared. Talaromyces C.R. Benj., Mycologia 47: 681. 1955. = Penicillium Link subgenus Biverticillium Dierckx apud Biourge Cellule 33: 31. 1923. = Penicillium subg. Biverticillata-Symmetrica Thom, The Penicillia: 158. 1930. = Sagenoma Stolk & G.F. Orr, Mycologia 66: 676. 1974. = Erythrogymnotheca Yaguchi & Udagawa, Mycoscience 35: 219. 1994. = Paratalaromyces Matsush., Matsush. Mycol. Mem. 10: 111 (2003) [2001]. Ascomata cleistothecial, usually with a distinctly hyphal exterior wall, often yellow, occasionally white, creamish, pinkish or reddish. Asci 8-spored, globose to ellipsoidal, ascus initials sometimes with morphologically distinguishable gametangia, mature asci produced in chains. Ascospores one-celled, rarely smooth-walled, but often with surface ornamentation and wings, hyaline to yellow, in strains producing abundant red pigment occasionally red. Conidiophores comprising smooth or rough-walled elements, with long hyaline stipes, generally terminating in a single whorl of 3–10 metulae, appearing symmetrical in face view (in some species with a single subterminal lateral branch that afterwards repeats the branching pattern of the main axis, but then with the whole conidiophore appearing asymmetrical), each metula with a terminal whorl of phialides. Conidiogenous cells phialidic, aculeate or acerose, rarely ampulliform, periclinal thickening usually visible in the conidiogenous aperture, with or without a cylindrical collarette. Conidia aseptate, green in mass, in basipetal connected chains, usually ellipsoidal to fusiform. Type species: Talaromyces vermiculatus (P.A. Dang.) C.R. Benj., Mycologia 47: 684. 1955. The name Talaromyces was introduced by Benjamin (1955), and the type species is T. vermiculatus (P.A. Dang.) C.R. Benj. One of 174 the authors (RAS) personally visited several herbaria in Paris to locate holotype or other original material of Penicillium vermiculatum P.A. Dang. Dangeard (1907) described and illustrated both the anamorph and teleomorph under this name, but his material could not be located. To repair the shortcoming of the typification of Talaromyces, the lectotype for P. vermiculatum is here designated as Plate XVIII in Dangeard (1907, available at the Biodiversity Heritage Library, www.biodiversitylibrary.org). It was selected from among the plates XVI−XX because it includes the most detailed drawings of the anamorph, but also includes elements of the teleomorph. Herb. IMI 197477 is here designated as the epitype of Penicillium vermiculatum P.A. Dang. This specimen, which is also the holotype of Penicillium dangeardii J. Pitt, the seldom-used name for the anamorph of T. flavus, is derived from the equivalent cultures CBS 310.38, IMI 19447, and NRRL 2098. The latter strain was considered typical of P. vermiculatum by Raper & Thom (1949), the last major treatment to use this Penicillium name as a distinct species. List of species The following list includes previously accepted species of Talaromyces and proposals to transfer the species of Penicillium subgenus Biverticillium to Talaromyces. Our phylogenetic studies demonstrate that several taxa represent complexes of morphologically cryptic phylogenetic species, requiring further study. For example, we analysed members of the Penicillium purpurogenum complex (including P. purpurogenum, P. rubrum, P. crateriforme, P. sanguineum) and found that several species group could be distinguished by sequencing certain genes (N. Yilmaz, unpubl. data) and had distinct macromorphological features and unique extrolite profiles. The full phylogenetic diversity of the P. purpurogenum species complex requires more investigation, and a more detailed account will be published elsewhere. ACCEPTED SPECIES IN TALAROMYCES Talaromyces aculeatus (Raper & Fennell) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560639. Basionym: Penicillium aculeatum Raper & Fennell, Mycologia 40: 535. 1948. Talaromyces albobiverticillius (H.-M. Hsieh, Y.-M. Ju & S.-Y. Hsieh) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560683. Basionym: Penicillium albobiverticillium H.-M. Hsieh, Y.-M. Ju & S.Y. Hsieh, Fung. Sci. 25: 26. 2010. Talaromyces allahabadensis (B.S. Mehrotra & D. Kumar) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560640. Basionym: Penicillium allahabadense B.S. Mehrotra & D. Kumar, Canad. J. Bot. 40: 1399. 1962. Talaromyces apiculatus Samson, Yilmaz & Frisvad, sp. nov. MycoBank MB560641. = Penicillium aculeatum var. apiculatum Abe, S., 1956, J. Gen. Appl. Microbiol., Tokyo 2: 124. 1956 (nom. inval., Art. 36). PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Penicillio aculeato simile, sed conidiis apiculatis distinguitur. Typus: Japan from soil (CBS H-20755 Holotype, culture ex-type CBS 312.59) Note: Species similar to Penicillium aculeatum but differing by apiculate conidia. Talaromyces assiutensis Samson & Abdel-Fattah, Persoonia 9: 501. 1978. Anamorphic synonym: Penicillium assiutense Samson & Abdel Fattah (simultaneously published, identical holotype). Talaromyces aurantiacus (J.H. Mill., Giddens & A.A. Foster) Samson, Yilmaz, & Frisvad, comb. nov. MycoBank MB560642. Basionym: Penicillium aurantiacum J.H. Mill., Giddens & A.A. Foster, Mycologia 49: 797. 1957. Talaromyces austrocalifornicus Yaguchi & Udagawa Trans. Mycol. Soc. Japan 34: 245. 1993. Anamorphic synonym: Penicillium austrocalifornicum Yaguchi & Udagawa (simultaneously published, identical holotype). Talaromyces bacillisporus (Swift) C. R. Benj., Mycologia 47: 682. 1955. ≡ Penicillium bacillisporum Swift, Bull. Torrey Bot. Club 59: 221, 1932. Talaromyces boninensis (Yaguchi & Udagawa) Samson, Yilmaz, & Frisvad, comb. nov. MycoBank MB560643. Basionym: Talaromyces helicus var. boninensis Yaguchi & Udagawa, Transactions Mycological Society Japan 33: 511. 1992. Talaromyces brunneus (Udagawa) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560644. Basionym: Penicillium brunneum Udagawa, J. Agric. Sci. (Tokyo) Nogyo Daigaku 5: 16. 1959. Talaromyces calidicanius (J.L. Chen) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560645. Basionym: Penicillium calidicanium J.L. Chen, Mycologia 94(5): 870. 2002. Talaromyces cecidicola (Seifert, Hoekstra & Frisvad) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560646. Talaromyces dendriticus (Pitt) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560648. Basionym: Penicillium dendriticum Pitt, The Genus Penicillium: 413. 1980. Talaromyces derxii Takada & Udagawa, Mycotaxon 31: 418. 1988. Anamorphic synonym: Penicillium derxii Takata & Udagawa (simultaneously published, identical holotype). Talaromyces diversus (Raper & Fennell) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560649. Basionym: Penicillium diversum Raper & Fennell, Mycologia 40: 539. 1948. Talaromyces duclauxii (Delacr.) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560650. Basionym: Penicillium duclauxii Delacr., Bull. Soc. Mycol. France 7: 107. 1891. Talaromyces echinosporus (Nehira) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560651. Basionym: Penicillium echinosporum Nehira, J. Ferment. Technol., Osaka 11: 861. 1933. Note: Penicillium asperosporum G. Smith, Trans. Brit. Mycol. Soc. 48: 275. 1965. (= Penicillium echinosporum G. Sm., Trans. Brit. Mycol. Soc. 45: 387. 1962 (non Nehira in J. Ferment. Technol. 11: 849. 1933) belongs in Penicillium section Aspergilloides (Houbraken & Samson 2011). Talaromyces emodensis Udagawa, Mycotaxon 48: 146. 1993. Anamorphic synonym: Penicillium emodense (simultaneously published, identical holotype). Udagawa Talaromyces erythromellis (A.D. Hocking) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560652. Basionym: Penicillium erythromellis A.D. Hocking apud Pitt, The Genus Penicillium: 459. 1980. Talaromyces euchlorocarpius Yaguchi, Someya & Udagawa, Mycoscience 40: 133. 1999. Anamorphic synonym: Penicillium euchlorocarpium Yaguchi, Someya & Udagawa (simultaneously published, identical holotype). Basionym: Penicillium cecidicola Seifert, Hoekstra & Frisvad, Stud. Mycol. 50: 520. 2004. Note: We have not seen the type, but the description and the ITS sequences available in GenBank (AB176617) show that this is a distinct species of Talaromyces. Talaromyces coalescens (Quintan.) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560647. Talaromyces flavo-virens (Durieu & Mont.) Visagie, Llimona & Seifert, ined. Talaromyces convolutus Udawaga, Mycotaxon 48: 141. 1993. Note: A manuscript on this species and its relationship to Penicillium aureocephalum Munt.-Cvetk., Hoyo & Gómez-Bolea is being prepared for publication in Mycotaxon. Basionym: Penicillium coalescens Quintan., Mycopathol. 84: 115. 1984. Anamorphic synonym: Penicillium convolutum (simultaneously published, identical holotype). Udagawa Talaromyces flavus (Klöcker) Stolk & Samson, Stud. Mycol. 2: 10. 1972. Anamorphic synonym: Penicillium dangeardii Pitt, The Genus Penicillium: 472. 1980. www.studiesinmycology.org 175 SaMSon eT al. Talaromyces funiculosus (Thom) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560653. Talaromyces minioluteus (Dierckx) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560657. Basionym: Penicillium funiculosum Thom, Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 69. 1910 Basionym: Penicillium minioluteum Dierckx, Ann. Soc. Sci. Bruxelles 25: 87. 1901. Talaromyces galapagensis Samson & Mahoney, Trans. Brit. Mycol. Soc. 69: 158. 1977. Talaromyces muroii Yaguchi, Someya & Udagawa, Mycoscience 35: 252. 1994. Anamorphic synonym: Penicillium galapagense Samson & Mahoney (simultaneously published, identical holotype). Talaromyces hachijoensis Yaguchi, Someya & Udagawa, Mycoscience 37: 157. 1996. Note: We have not seen the type but the description and the ITS sequences available in GenBank (AB176620) show that this is a distinct species of Talaromyces. It is unusual in the genus for its apparent lack of an anamorph. Talaromyces helicus (Raper & Fennell) C.R. Benj., Mycologia 47: 684. 1955. ≡ Penicillium helicum Raper & Fennell, Mycologia 40: 515. 1948. Talaromyces indigoticus Takada & Udagawa, Mycotaxon 46: 129. 1993. Anamorphic synonym: Penicillium indigoticum Takada & Udagawa (simultaneously published, identical holotype). Note: This species is unusual in Talaromyces because of its lack of a known anamorph. Talaromyces palmae (Samson, Stolk & Frisvad) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560658. Basionym: Penicillium palmae Samson, Stolk & Frisvad, Stud. Mycol. 31: 135. 1989. Talaromyces panamensis (Samson, Stolk & Frisvad) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560659. Basionym: Penicillium panamense Samson, Stolk & Frisvad, Stud. Mycol. 31: 136. 1989. Talaromyces paucisporus (Yaguchi, Someya & Udagawa) Samson & Houbraken, comb.nov. MycoBank MB560684. Basionym: Erythrogymnotheca paucispora Yaguchi, Someya & Udagawa, Mycoscience 35: 219. 1994. Talaromyces intermedius (Apinis) Stolk & Samson, Stud. Mycol. 2: 21. 1972. Talaromyces phialosporus (Udagawa) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560660. Talaromyces islandicus (Sopp) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560654. Talaromyces piceus (Raper & Fennell) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560661. Talaromyces loliensis (Pitt) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560655. Talaromyces pinophilus (Hedgcock) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560662. Talaromyces macrosporus (Stolk & Samson) Frisvad, Samson & Stolk, Ant. van Leeuwenhoek J. Microbiol. Serol. 57: 186. 1990. Talaromyces pittii (Quintan.) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560663. Anamorphic synonym: Penicillium intermedium Stolk & Samson, Stud. Mycol. 2: 21. 1972. Basionym: Penicillium islandicum Sopp, Skr. Vidensk.-Selsk. Christiania, Math.-Naturvidensk. Kl. 11: 161. 1912. Basionym: Penicillium loliense Pitt, The Genus Penicillium: 450. 1980 Anamorphic synonym: Penicillium macrosporum Frisvad, Filt., Samson & Stolk. nom. illegit. Art. 53 (non P. macrosporum Berk. & Broome 1882). Talaromyces marneffei (Segretain, Capponi & Sureau) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560656. Basionym: Penicillium marneffei Segretain, Capponi & Sureau apud Segretain, Bull. Soc. Mycol. France 75: 416. 1959 [1960]. Talaromyces mimosinus A.D. Hocking apud Pitt, The Genus Penicillium: 507. 1980. Anamorphic synonym: Penicillium mimosinum A. D. Hocking (simultaneously published, identical holotype). 176 Basionym: Penicillium phialosporum Udagawa, J. Agric. Sci. (Tokyo) Nogyo Daigaku 5: 11. 1959. Basionym: Penicillium piceum Raper & Fennell, Mycologia 40: 533. 1948. Basionym: Penicillium pinophilum Hedgcock apud Thom, Bull. Bur. Anim. Ind. US Dept. Agric. 118: 37. 1910. Basionym: Penicillium pittii Quintan., Mycopathol. 91: 69. 1985. Talaromyces primulinus (Pitt) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560664. Basionym: Penicillium primulinum Pitt, The Genus Penicillium: 455. 1980. Talaromyces proteolyticus (Kamyschko) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560665. Basionym: Penicillium proteolyticum Kamyschko, Nov. Sist. niz. Rast. Sist. niz. Rast. 14: 227. 1961. Talaromyces pseudostromaticus (Hodges, G.M. Warner, Rogerson) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560666. PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Basionym: Penicillium pseudostromaticum Hodges, G.M. Warner & Rogerson, Mycologia 62: 1106. 1970. Talaromyces purpureus (E. Müll. & Pacha-Aue) Stolk & Samson, Stud. Mycol. 2: 57. 1972. Anamorphic synonym: Penicillium purpureum Stolk & Samson, Stud. Mycol. 2: 57. 1972. Talaromyces purpurogenus (Stoll) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560667. Basionym: Penicillium purpurogenum Stoll, Beitr. Charakt. Penicillium-Arten: 32. 1904. Talaromyces stipitatus (Thom) C.R. Benj., Mycologia 47: 684. 1955. ≡ Penicillium stipitatum Thom, Mycologia 27: 138. 1935. Talaromyces sublevisporus (Yaguchi & Udagawa) Samson, Yilmaz & Frisvad, comb. et stat. nov. MycoBank MB560675. Basionym: Talaromyces wortmannii var. sublevisporus Yaguchi & Udagawa, Mycoscience 35: 63. 1994. Note: We have not examined the ex-type of this species but from the ITS data (GenBank AB176638), this seems to be a separate species. Talaromyces rademirici (Quintan.) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560668. Talaromyces tardifaciens Udagawa, Mycotaxon 48: 150. 1993. Basionym: Penicillium rademirici Quintan., Mycopathol. 91: 69. 1985. Anamorphic synonym: Penicillium tardifaciens (simultaneously published, identical holotype). Talaromyces radicus (A.D. Hocking & Whitelaw) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560669. Talaromyces trachyspermus (Shear) Stolk & Samson, Stud. Mycol. 2: 32. 1972. Udagawa Basionym: Penicillium radicum A.D. Hocking & Whitelaw, Mycol. Res. 102: 802. 1998. Anamorphic synonym: Penicillium spiculisporum Leman, Mycologia 12: 268. 1920 Talaromyces ramulosus (Visagie & K. Jacobs) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560670. Talaromyces ucrainicus Udagawa, in Stolk & Samson, Stud. Mycol. 2: 34. 1972. Basionym: Penicillium ramulosum Visagie & K. Jacobs, Mycologia 101: 890. 2009. Anamorphic synonym: Penicillium ucrainicum Mycologia 56: 59. 1964. Talaromyces rotundus (Raper & Fennell) C.R. Benj., Mycologia 47: 683. 1955. Talaromyces udagawae Stolk & Samson, Stud. Mycol. 2: 36. 1972. ≡ Penicillium rotundum Raper & Fennell, Mycologia 40: 518. 1948. Talaromyces ryukyuensis (S. Ueda & Udagawa) Arx, Persoonia 13: 282. 1987. ≡ Sagenoma ryukyuense S. Ueda & Udagawa, Mycotaxon 20: 499. 1984. Note: We have not seen the type but the description and the ITS sequences available in GenBank (AB176628) show that this is a distinct species of Talaromyces. Talaromyces rubicundus (J.H. Mill., Giddens & A.A. Foster) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560671. Basionym: Penicillium rubicundum J.H. Mill., Giddens & A.A. Foster, Mycologia 49: 797. 1957. Talaromyces rugulosus (Thom) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560672. Basionym: Penicillium rugulosum Thom, Bull. Bur. Anim. Ind. US Dept. Agric. 118: 60. 1910. Talaromyces sabulosus (Pitt & A.D. Hocking) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560673. Basionym: Penicillium sabulosum Pitt & A. D. Hocking, Mycologia 77: 818. 1985. Talaromyces siamensis (Manoch & C. Ramírez) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560674. Basionym: Penicillium siamense Manoch & C. Ramírez, Mycopathol. 101: 32. 1988. www.studiesinmycology.org Panasenko, Anamorphic synonym: Penicillium udagawae Stolk & Samson (simultaneously published, identical holotype). Talaromyces unicus Tzean, J.L. Chen & Shiu, Mycologia 84: 739. 1992. Anamorphic synonym: Penicillium unicum Tzean, J.L. Chen & Shiu (simultaneously published, identical holotype). Talaromyces variabilis (Sopp) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560676. Basionym: Penicillium variabile Sopp, Skr. Vidensk.-Selsk. Christiania, Math.-Naturvidensk. Kl. 11: 169. 1912. Talaromyces varians (G. Sm.) Samson, Yilmaz & Frisvad, comb. nov. MycoBank MB560677. Basionym: Penicillium varians G. Sm., Trans. Brit. Mycol. Soc. 18: 89. 1933. Talaromyces verruculosus (Peyronel) Samson, Yilmaz, Frisvad & Seifert, comb. nov. MycoBank MB560678. Basionym: Penicillium verruculosum Peyronel, Germi Atmosf. Fung. Micel.: 22. 1913. Talaromyces viridis (Stolk & G.F. Orr) von Arx, Persoonia 13(3): 2821. 1987 ≡ Sagenoma viride Stolk & G.F. Orr, Mycologia 66: 677. 1974. Talaromyces viridulus Samson, Yilmaz & Frisvad, nom. nov. MycoBank MB560679. Basionym: Geosmithia viridis Pitt & A.D. Hocking, Mycologia 77: 822. 1985 = P. viride (Pitt & A.D. Hocking) Frisvad, Samson & 177 SaMSon eT al. Stolk, Persoonia 14: 229. 1990, nom. illegit. Art. 53 (non Fres. 1851 nec Rivera 1873 nec Sopp 1912 nec (Matr.) Biourge 1923). Non Talaromyces viridis (Stolk & G.F. Orr) Arx. Talaromyces wortmannii (Klöcker) C.R. Benjamin, Mycologia 47: 683. 1955 ≡ Penicillium wortmannii Klöcker, Compt-Rend. Trav. Carlsberg Lab. 6: 100. 1903. ExCLUDED SPECIES AND TAxA, WHICH NEED FURTHER TAxONOMIC STUDY Penicillium concavorugulosum S. Abe, J. Gen. Appl. Microbiol, Tokyo 2: 127. 1956 (nom. inval. Art. 36). Note: This species was invalidly described, but our ITS data (Fig. 2) show that it is related to T. wortmanii. Further study is required but extrolite data indicate that this species is unique (J.C. Frisvad, unpublished data). Penicillium crateriforme J.C. Gilman & E.V. Abbott, Iowa State Coll. J. Sc. 1: 293. 1927. Note: Our ITS data (Fig. 2) show that this species is a synonym of P. purpurogenum. Penicillium ilerdanum C. Ramírez, A.T. Martínez & Berer., Mycopathol. 72: 32. 1980. Note: Frisvad et al. (1990b) considered this species synononymous with Penicillium piceum Raper & Fennell, which is confirmed by our ITS data (Fig. 2) Penicillium mirabile Beliakova & Milko, Mikol. Fitopatol. 6: 145. 1972. Note: The ex-type culture is in poor condition and although our ITS data (Fig. 2) indicate that is a distinct species, it should be further investigated. Penicillium oblatum Pitt & A.D. Hocking, Mycologia 77: 810. 1985. Note: In our ITS phylogeny (Fig. 2), this species is close to Paecilomyces pascuus and Penicillium dendriticum and needs further study. Penicillium pascuum (Pitt & A.D. Hocking) Frisvad, Samson & Stolk, Persoonia 14: 229. 1990 ≡ Paecilomyces pascuus Pitt & A. D. Hocking, Mycologia 77: 822. 1985. Note: See on the position of this species under P. oblatum above. Penicillium rubrum Stoll, Beitr. Charakt. Penicillium-Arten: 35. 1904. Note: Although the name is well-known, the taxonomic position of the taxon remains doubtful because no type material has been located. A possible solution would be lectotypification from Stoll’s illustrations, followed by epitypification to become a usable name. Penicillium purpurogenum var. rubrisclerotium Thom, Mycologia 7: 137. 1915. Note: Our ITS data (Fig. 2) indicate that this species is synonymous with P. minioluteum. Penicillium isariiforme Stolk & J.A. Mey., Trans. Brit. Mycol. Soc. 40: 187. 1957. Penicillium samsonii Quintan., Mycopathol. 91: 69. 1985. Note: According to Houbraken & Samson (2011), this species, included in subgenus Biverticillium by Pitt (1980), is correctly classified in Penicillium sensu lato. Penicillium tardum Thom, The Penicillia: 485. 1930. Penicillium korosum J.N. Rai, Wadhwani & J.P. Tewari, Ant. van Leeuwenhoek 35: 430. 1969. Note: This species requires further investigation, but our ITS sequence (Fig. 2) indicates that it is similar to P. pinophilum. Penicillium krugeri C. Ramírez, Mycopathol. 110: 23. 1990. = Talaromyces minioluteus (Dierckx) Samson, Yilmaz, Frisvad & Seifert (see above). Note: Raper & Thom (1949) pointed out that there is confusion about the type culture and the status of this species will be subject of further studies. Penicillium victoriae Szilv., Archiv. Hydrobiol. 14, Suppl. 6: 535. 1936 = Penicillium janthinellum Biourge, Cellule 33: 258. 1923 (Pitt, 1980). Note: We have been unable to examine authentic material, and the correct classification of this species is uncertain. Note: Pitt (1980) synonymised this species under Penicillium janthinellum, but our studies showed that it clearly belongs in Talaromyces. Because there is only one strain, the exact identity of this fungus requires further study. Penicillium lignorum Stolk, Ant. van Leeuwenhoek 35: 264. 1969. Talaromyces barcinensis Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 15. 1993. Note: A preliminary phylogenetic analysis indicates that this species does not belong to Talaromyces and might represent a new genus (J. Houbraken, unpubl. data). 178 Anamorphic synonym: Penicillium barcinense Yaguchi & Udagawa (simultaneously published, identical holotype). Note: Our ITS sequence data show that this species is close to Talaromyces helicus and further study should determine its correct taxonomic position. PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium Talaromyces brevicompactus Kong, Mycosystema 18: 9. 1999. Anamorphic synonym: Merimbla brevicompacta Kong, Mycosystema 18: 9. 1999 (simultaneously published, identical holotype). Note: Fig. 1 shows that this species belongs in Hamigera. Comparison of partial β-tubulin and calmodulin sequences of the ex-type strain of T. brevicompactus with recent published data shows that this species represents a distinct species (J. Houbraken, unpubl. data). The new combination in Hamigera will be made elsewhere. Talaromyces byssochlamydoides Stolk & Samson, Stud. Mycol. 2: 45. 1972. Anamorphic synonym: Paecilomyces byssochlamydoides Stolk & Samson (simultaneously published, same holotype). = Rasamsonia byssochlamydoides (Stolk & Samson) Houbraken & Frisvad, Ant. van Leeuwenhoek, in press. Talaromyces eburneus Yaguchi, Someya & Udagawa, Mycoscience 35: 249. 1994. Anamorphic synonym: Geosmithia eburnea Yaguchi, Someya & Udagawa (simultaneously described, holotype identical) ≡ Rasamsonia eburnea (Yaguchi, Someya & Udagawa) Houbraken & Frisvad, Ant. van Leeuwenhoek, in press. Talaromyces emersonii Stolk, Ant. van Leeuwenhoek 31: 262. 1965. Anamorphic synonym: Penicillium emersonii Stolk (simultaneously described, holotype identical), Ant. van Leeuwenhoek 31: 262. 1965. = Rasamsonia emersonii (Stolk) Houbraken & Frisvad, Ant. van Leeuwenhoek, in press. Talaromyces gossypii Pitt, The Genus Penicillium: 500. 1980 = T. assiutensis, Samson & Abdel-Fattah, Persoonia 9: 501. 1978 (fide Frisvad et al. 1990a). Talaromyces lagunensis Udagawa, Uchiy. & Kamiya, Mycoscience 35: 403. 1994. Anamorphic synonym: Penicillium lagunense Udagawa, Uchiy. & Kamiya (simultaneously published, identical holotype). Note: We have been unable to examine authentic material, and the correct classification of this species is uncertain. Talaromyces leycettanus H.C. Evans & Stolk, Trans. Brit. Mycol. Soc. 56: 45. 1971. Anamorphic synonym: Penicillium leycettanus H.C. Evans & Stolk (simultaneously published, identical holotype) ≡ Paecilomyces leycettanus (H.C. Evans & Stolk) Stolk, Samson & H.C. Evans, Persoonia 6: 342. 1971. Note: Houbraken & Samson (2011) showed that this species is phylogenetically unrelated to Talaromyces and close to Hamigera. Its taxonomic position requires further investigation. Talaromyces luteus (Zukal) C.R. Benj., Mycologia 47: 681. 1955. ≡ Penicillium luteum Zukal, Sitzungsber Kaiserl. Akad. Wiss. MathNaturwiss. C1., Abt. 1, 98: 561. 1890. www.studiesinmycology.org Note: Although the phenotype of this species resembles species of Talaromyces, our molecular analysis shows that it is phylogenetically unique and basal to T. thermophilus. Talaromyces malagensis (Thüm.) Stalpers & Samson 1984, in Stalpers, Stud. Mycol. 24: 69. 1984. Note: Stolk & Samson (1972) considered Sporotrichum malagense a dubious synonym of T. udagawae, based on their failure to find ascospores and conidia in the type material (herb. W). Later, Stalpers (1984) studied material preserved in herb. BR which is authentic and labelled as “type”. It agrees with Thümen’s original diagnosis and contains both fertile Talaromyces cleistothecia and a sporulating biverticillate anamorph. Therefore, the new combination to Talaromyces was proposed. The species resembles T. udagawae or T. luteus, but in the absence of a living culture we cannot determine its precise taxonomic identity. Talaromyces ocotl Bills & Heredia, Mycologia 90: 533. 1998. Note: Figure 1 shows that this species belongs to Sagenomella and the new combination is proposed here: Sagenomella ocotl (Bills & Heredia) Samson, Houbraken & Frisvad, comb. nov. MycoBank MB560681. Basionym: Talaromyces ocotl Bills & Heredia, Mycologia 93: 533. 1998. Talaromyces ohiensis Pitt, The Genus Penicillium: 502. 1980. Anamorphic synonym: Penicillium ohiense L. H. Huang & J. A. Schmitt, Ohio J. Sci. 75: 78. 1975. Note: Pitt (1980) considered this species to be related to T. luteus, but our ITS data clearly show that is synonymous with T. ucrainicus. Talaromyces panasenkoi Pitt, The Genus Penicillium: 482. 1980. Anamorphic synonym: Penicillium panasenkoi Pitt (simultaneously published, identical holotype). Note: Pitt (1980) proposed T. panasenkoi as a new species for the invalidly published P. ucraininum Panasenko; however, Stolk & Samson (1972) had already proposed Talaromyces ucrainicus Udagawa for this taxon. T. panasenkoi Pitt is therefore a synonym of T. ucrainicus. Talaromyces retardatus Udagawa, Kamiya & Kaori Osada, Trans. Mycol. Soc. Japan 34: 9. 1993. Anamorphic synonym: Penicillium retardatum Udagawa, Kamiya & Kaori Osada (simultaneously published, identical holotype). Note: No strain was available for examination and the status of this species is thus unknown. Talaromyces spectabilis Udagawa & Suzuki, Mycotaxon 50: 82. 1994. = Byssochlamys spectabilis (Udagawa & Suzuki) Houbraken & Samson, Appl. Environ. Microbiol. 74: 1618. 2008. = Paecilomyces variotii Bainier Bull. Soc. mycol. Fr. 23: 27. 1907. 179 SaMSon eT al. Note: The oldest generic and species name for this species is P. variotii, which becomes the correct name for the holomorph. Coremium Link : Fr., Mag. Ges. naturf. Freunde, Berlin 3: 19. 1809 : Fries, Syst. mycol. 1: xlviii, 1821. Talaromyces striatus (Raper & Fennell) C.R. Benj., Mycologia 47: 682. 1955 Type species: C. glaucum Link 1809. = Hamigera striata (Raper & Fennell) Stolk & Samson, Persoonia 6: 347. 1971. Talaromyces thermocitrinus Subrahm. & Gopalkr., Ind. Bot. Reporter 35: 35. 1984 (as ‘T. thermocitrinum’). Note: We have not seen the type, but judging from the substrate (dust on books), and the mention of yellow cleistothecia, it is possible that this species is a Eurotium species, a typical contaminant of books and other material in archives. However, its reported thermophily is different from known species of the mesophilic Eurotium species. Talaromyces thermophilus Stolk, Ant. van Leeuwenhoek 31: 268. 1965. Basionym: Penicillium dupontii Griffon & Maubl., Bull. Trimmest. Soc. mycol. Fr. 27: 73. 1911. Note: Figure 1 shows that this species is related to Thermomyces lanuginosus, and should be transferred to Thermomyces (Houbraken et al. 2011, Houbraken & Samson 2011). ACKNOWLEDGEMENTS We are appreciative of discussions with Dave Malloch, and contributions by Ellen Hoekstra in the early years of this project. Cobus Visagie (South Africa) and Xavier Llimona (Spain) allowed us to access unpublished data on P. aureocephalum and the genus Lasioderma. We are also grateful for nomenclatural advice received from Scott Redhead and Uwe Braun. We also thank Sung-Yuan Hsieh for providing the ITS sequence and the culture of Penicillium albobiverticillium. APPENDIx: OTHER POSSIBLE GENERIC NAMES As noted above in the Taxonomy section, in order to adopt Talaromyces as the generic name for the former Penicillium subgenus Biverticillium, older genera considered synonyms of Penicillium sensu lato had to be considered. These are treated below. Aspergillopsis Sopp, Vid.-Selsk. Skr. I. Math.-naturv. Kl. 11: 201. 1912. (Taf. xx, Fig. 149, Taf. xxiii, Fig. 31). Type species: A. fumosus Sopp 1912. Note: This generic name is illegitimate (Art. 53), being a later homonym of Aspergillopsis Speg. 1910. Pitt (1980) considered Sopp’s genus a tentative synonym of Merimbla Pitt. Citromyces Wehmer, Ber. dt. Bot. Ges. 11: 338. 1893. Type species: C. pfefferianus Wehmer 1893 = Penicillium glabrum (Wehmer) Westling 1911, fide Pitt 1980. Note: Wehmer’s genus was considered a synonym of Penicillium by many authors, including Raper & Thom (1949) and Pitt (1980), with C. pfefferianus considered a probable synonym of P. glabrum (subgenus Aspergillioides) by Pitt (1980). Therefore, the genus remains a synonym of Penicillium sensu stricto. 180 Note: This genus was described in the same publication as Penicillium. Raper & Thom (1949) and Seifert & Samson (1985) both considered the type species to be a synonym of the type species of Penicillium, P. expansum Link 1809. Therefore, Coremium remains a synonym of Penicillium sensu stricto. Eladia G. Sm., Trans. Br. mycol. Soc. 44: 47. 1961. Type species: Eladia saccula (Dale) G. Sm. 1961 = Penicillium sacculum Dale 1926. Note: This genus was considered a synonym of Penicillium by Stolk & Samson (1985), but was considered distinct by Pitt (1980), and von Arx (1981). In the multigene phylogenetic study by Houbraken & Samson (2011), Eladia is clearly included in Penicillium sensu stricto and that synonymy is accepted here. Floccaria Grev., Scott. Crypt. Fl., Vol. 6, Pl. 301. 1828. Type species: F. glauca Grev. 1828. Note: There is no known extant type according to Seifert & Samson (1985), who searched for it in K and E. The illustration shows a synnematous fungus that could well be P. expansum, but there are no microscopic details. Therefore, this name can be discounted as a possible generic name for the species formerly ascribed to subgenus Biverticillium. Geosmithia Pitt , Can. J. Bot. 57: 2021. 1980. Type species: Geosmithia lavendula (Raper & Fennell) Pitt 1980 = Penicillium lavendulum Raper & Fennell 1948. Note: Although von Arx (1981) considered Geosmithia a synonym of Penicillium, it is polyphyletic as presently circumscribed. Using SSU sequences, Ogawa et al. (1997) showed that G. lavendula, and a second common species G. putterilli, belong to the Bionectriaceae, Hypocreales. Similar results were obtained using ITS sequences by Kolařík et al. (2004), using LSU sequences by Schroers et al. (2005) and then multigene phylogenies by Kolařík & Kirkendall (2010). Despite this, some anamorphs attributed to Geosmithia have been described recently in Talaromyces (e.g. Yaguchi et al. 2005). Because the type species is not associated with the same order as Penicillium, Geosmithia need not be considered as a possible home for species of subgenus Biverticillium, but neither should it be considered a synonym of Penicillium. Hormodendrum Bonord., Handbuch allg. Mykol.: 76. 1851. Type species: Amphitrichum olivaceum Corda 1837 = Hormodendrum olivaceum (Corda) Bonord. 1851, lectotype selected by Clements & Shear 1931. Note: Hormodendron has variously been treated as a synonym of Penicillium by von Arx (1974) and de Hoog & Hermanides-Nijhoff (1977) but more often as a synonym of Cladosporium Link, following PHylogeny and noMenclature of tHe genuS Talaromyces and taxa accoMModated in Penicillium SubgenuS BiverTicillium the study of the type specimen by Hughes (1958). There is no reason to consider this name further as a synonym of Penicillium or as a possible receptacle for the species of subgenus Biverticillium. Merimbla Pitt, Can. J. Bot. 57: 2394. 1980. Type species: M. ingelheimensis (F.H. Beyma) Pitt 1980 = Penicillium ingelheimense F.H. Beyma 1942. Note: Merimbla was considered a possible synonym of Penicillium by von Arx (1981), but this has not generally been accepted. Merimbla ingelheimensis was considered the anamorph of Hamigera avellanea by Stolk & Samson (1971), but is now known to be a closely related but phylogenetically distinct species (Peterson et al. 2010). The Hamigera clade is phylogenetically distinct from subgenus Biverticillium in the multigene analyses of Peterson et al. (2010) and Houbraken & Samson (2011). In a single name system, we consider Merimbla a synonym of the older genus Hamigera. Monilia Fr., Syst. mycol. 3: 409. 1832. Type species: M. caespitosa (L. : Fr. ) Fr. 1832 / Mucor caespitosus L. 1753. Note: Donk (1963) suggested that M. caespitosa might be a species of Penicillium based on the protologue. However, this generic name was formally rejected to conserve usage of Monilia Bonorden for the well-known genus of fruit pathogens. Therefore, it is unavailable as a possible generic name for species included in subgenus Biverticillium. Rhodocephalus Corda, Ic. Fung. 1: 21. 1837 (Tab. vi, Fig. 282). Type species: R. candidus Corda 1837 Type species: Penicillium leucocephalum Rabenh. 1844. Note: Corda (1837) illustrated and described his species as having aseptate stipes, a branched, asymmetrical penicillate head, with long chains of ameroconidia. Rabenhorst (1844) renamed the species in Penicillium, changing the epithet, a conclusion followed by Lindau (1907). Thom (1930) and Raper & Thom (1949) disagreed, stating that the illustration in the protologue has branched conidial chains that would exclude the fungus from Penicillium. This a debatable conclusion, because the chains are simply overlapping in the illustration and there is no clear indication of branching. Pitt (1980) evidently did not examine the protologue when he suggested a synonymy with Aspergillus candidus. Hughes (1958) did not report on the type, and according to Holubová (in litt. to Seifert, 1991), there is no material of Rhodocephalus in the Corda herbarium (PRM). The asymmetrical conidiophores illustrated by Corda discount this as a possible genus for species of subgenus Biverticillium, but its exact identity is unknown. Torulomyces Delitsch, Systematik der Schimmelpilze: 91. 1943 (Taf. 30, Figs 232–235). Type species: T. lagena Delitsch 1943 = Monocillium lagena (Delitsch) Hashmi, W.B. Kendr. & Morgan-Jones 1972 = Penicillium lagena (Delitsch) Stolk & Samson 1983. Moniliger Letell., Fig. Champ., Pl. 668. 1839. Figs 3, 4. Note: Torulomyces was included as a synonym of Penicillium sensu stricto in the phylogenetic study of Houbraken & Samson (2011). Type species: not designated, two original species. Yunnania H.-Z. Kong, Mycotaxon 69: 320. 1998. Note: According to Seifert et al. (2011), Letellier included two species, with illustrations clearly representing Aspergillus. The synonymy of Moniliger with Penicillium proposed by Kirk et al. (2008) thus seems unlikely, and the genus is better listed as a synonym of Aspergillus. Type species: Y. penicillata H.-Z. Kong 1998. Penicillium Link : Fr., Mag. Ges. naturf. Freunde, Berlin 3: 16. 1809. : Fries, Syst. mycol. 3: 406.1832. Type species: P. expansum Link 1809, fide Thom 1910. Note: With this revision, and that of Houbraken & Samson (2011), Penicillium is now used exclusively for the nominal Clade including P. expansum, and species in the now synonymous genus Eupenicillium F. Ludw. 1892 (Houbraken & Samson 2011). Pritzeliella Henn., Hedwigia Beibl. 42: 88. 1903. Type species: P. caerulea Henn. 1903. Note: Clements & Shear (1931) suggested that Pritzeliella should be considered a synonym of Penicillium without further commenting on the identity of its type species. Seifert & Samson (1985) examined the holotype of P. caerulea and considered it a synonym of Penicillium coprophilum (subgenus Penicillium). 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