Polyphasic taxonomy of the genus Talaromyces

Jens Frisvad

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.

available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 78: 175–341. Polyphasic taxonomy of the genus Talaromyces N. Yilmaz1,2, C.M. Visagie1, J. Houbraken1, J.C. Frisvad3*, and R.A. Samson1* 1 CBS-KNAW Fungal Biodiversity Centre, 3508 AD, Utrecht, The Netherlands; 2Department of Biology, Utrecht University, Utrecht, The Netherlands; 3Center for Microbial Biotechnology, Department of Systems Biology, Building 221, Technical University of Denmark, DK-2800, Kgs. Lyngby, Denmark *Correspondence: R.A. Samson, r.samson@cbs.knaw.nl; J.C. Frisvad, jcf@bio.dtu.dk Abstract: The genus Talaromyces was described by Benjamin in 1955 as a sexual state of Penicillium that produces soft walled ascomata covered with interwoven hyphae. Phylogenetic information revealed that Penicillium subgenus Biverticillium and Talaromyces form a monophyletic clade distinct from the other Penicillium subgenera. Subsequently, in combination with the recent adoption of the one fungus one name concept, Penicillium subgenus Biverticillium was transferred to Talaromyces. At the time, the new combinations were made based only on phylogenetic information. As such, the aim of this study was to provide a monograph on Talaromyces applying a polyphasic species concept, including morphological, molecular and physiological characters. Based on an ITS, BenA and RPB2 multigene phylogeny, we propose a new sectional classiﬁcation for the genus, placing the 88 accepted species into seven sections, named sections Bacillispori, Helici, Islandici, Purpurei, Subinﬂati, Talaromyces and Trachyspermi. We provide morphological descriptions for each of these species, as well as notes on their identiﬁcation using morphology and DNA sequences. For molecular identiﬁcation, BenA is proposed as a secondary molecular marker to the accepted ITS barcode for fungi. Key words: Ascomycetes, Trichocomaceae, Ascospores, Single name nomenclature, Talaromyces marneffei, Food spoilage, Biotechnology. Taxonomic novelties: New sections: Talaromyces section Helici Yilmaz, Frisvad & Samson, Talaromyces section Bacillispori Yilmaz, Frisvad & Samson, Talaromyces section Subinﬂati Yilmaz, Frisvad & Samson; New combinations: Talaromyces section Islandici (Pitt) Yilmaz, Frisvad & Samson, Talaromyces aerugineus (Samson) Yilmaz, Frisvad & Samson, Talaromyces bohemicus (Fassat. & Peckova) Yilmaz, Frisvad & Samson, Talaromyces liani (Kamyschko) Yilmaz, Frisvad & Samson, Talaromyces cinnabarinus (S.C. Jong & E.E. Davis) Yilmaz, Samson & Frisvad. Published online 22 September 2014; http://dx.doi.org/10.1016/j.simyco.2014.08.001. Hard copy: June 2014. Studies in Mycology INTRODUCTION Benjamin (1955) introduced the genus Talaromyces for teleomorphic Penicillium species with T. vermiculatus (P.A. Dang.) C.R. Benj. (= T. ﬂavus (Klöcker) Stolk & Samson) as generic type. The genus was characterised by soft ascocarps with a cleistothecial wall of interwoven hyphae and typically yellow ascomata, with ovate to globose asci containing mostly spiny ascospores (Benjamin 1955). Stolk & Samson (1971) introduced Hamigera for Talaromyces species that produce single asci, limiting Talaromyces to species producing asci in chains (Stolk & Samson 1972). Although Pitt (1980) considered Hamigera synonymous with Talaromyces, Houbraken & Samson (2011) showed that it is a distinct genus closely related to Warcupiella. Thermophilic Talaromyces species were shown to be distinct from Talaromyces (Houbraken & Samson 2011) and were classiﬁed in the new genus Rasamsonia and Thermomyces (Houbraken et al. 2012, 2014). Talaromyces used to be associated with the anamorph genera Geosmithia, Merimbla, Paecilomyces and Penicillium (Pitt et al. 2000). Merimbla was shown to belong to a monophyletic clade with Hamigera, while both Geosmithia and Paecilomyces were shown to be polyphyletic (Houbraken & Samson 2011), with G. lavendula (generic type) belonging in the Hypocreales and P. variotii (generic type) monophyletic with Byssochlamys in the Thermoascaceae. The remaining Geosmithia species belong in Penicillium, Talaromyces and Rasamsonia (Houbraken & Samson 2011), while Paecilomyces aerugineus belongs in Talaromyces (Samson et al. 2011). A number of species described as Talaromyces were shown to not belong in the genus. Three thermophilic species, T. byssochlamydoides, T. eburneus and T. emersonii, belong in the genus Rasamsonia and were renamed as R. byssochlamydoides, R. eburnea and R. emersonii (Houbraken et al. 2012). Talaromyces thermophilus is a biotechnologically important species and belongs to Thermomyces, with Houbraken et al. (2014) introducing the new combination Thermomyces dupontii for the species. Houbraken et al. (2012) showed that T. leycettanus belongs in the Hamigera/Warcupiella clade, and this species exact position within the clade should be further investigated. Talaromyces luteus morphologically resembles T. udagawae based on their ascospores having transverse to spiral ridges (Stolk & Samson 1972). Phylogenetically, however, T. luteus is basal to the Thermomyces dupontii and T. lanuginosus clade, and most probably represents a distinct genus (Houbraken et al. 2014). It was well documented that subgenus Biverticillium resulted in Penicillium being polyphyletic (Frisvad et al. 1990a, b, LoBuglio et al. 1993, Berbee et al. 1995, Ogawa et al. 1997, Ogawa & Sugiyama 2000, Peterson 2000, Heredia et al. 2001, Seifert et al. 2004, Wang & Zhuang 2007). Houbraken & Samson (2011), using a four-gene phylogeny, showed that Penicillium subgenus Biverticillium and Talaromyces species form a monophyletic clade, with Samson et al. (2011) recombining these Penicillia into Talaromyces in support of singlename nomenclature (McNeill et al. 2012). Talaromyces contains species that are medically important. Talaromyces marneffei is an emerging fungal pathogen causing a fatal mycosis in especially immunocompromised individuals Peer review under responsibility of CBS-KNAW Fungal Biodiversity Centre. Copyright © 2014, CBS-KNAW Fungal Biodiversity Centre. Production and hosting by ELSEVIER B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/ licenses/by-nc-nd/3.0/). 175 YILMAZ ET AL. from East Asian countries such as China, Taiwan, Thailand and Vietnam (Deng et al. 1988, Supparatpinyo et al. 1994, Chiang et al. 1998, Hien et al. 2001), even though infections in HIV negative individuals have been reported (Kwan et al. 1997, Saadiah et al. 1999). Talaromyces marneffei is the only known dimorphic species in the genus, producing ﬁlamentous growth at 25 C and a yeast phase at 37 C (Andrianopoulos 2002). Because T. marneffei used to be classiﬁed in Penicillium, its associated disease is unfortunately still referred to as penicilliosis. However, true Penicillium species causing human infections are very rare. Other Talaromyces species have also been reported to be medically important. For example, T. indigoticus was isolated from skin and nail lesions from a male who was affected by onychomycosis in Western Panama (Weisenborn et al. 2010), T. piceus caused both fungaemia (Horre et al. 2001) and rib osteomyelitis in an X-linked chronic granulomatous disease (X-CGD) patient (Santos et al. 2006), T. radicus caused a fatal infection in a German shepherd dog (de Vos et al. 2009), T. helicus caused granulomatous lymphadenitis in a labrador retriever (Tomlinson et al. 2011), and T. amestolkiae and T. stollii were isolated from the lungs and sputum of immunocompromised patients (Yilmaz et al. 2012). Talaromyces is also of importance in the food industry. Heat resistant ascospores are produced in T. macrosporus, T. ﬂavus, T. bacillisporus, T. helicus, T. stipitatus, T. trachyspermus and T. wortmannii, and cause spoilage of pasteurised juices and other fruit based products (Pitt & Hocking 1997, Dijksterhuis 2007). Some species also produce mycotoxins in food products. Rubratoxin is a potential hepatotoxin (Engelhardt & Carlton 1991) and is produced by T. purpurogenus (Yilmaz et al. 2012). A teenaged boy needed an immediate liver transplant after drinking rhubarb-wine, which was contaminated with T. purpurogenus and contained a high concentration of rubratoxin (Sigler et al. 1996, Richer et al. 1997). Talaromyces islandicus is one of the most destructive and harmful fungi affecting rice in storage, causing the yellowing of rice (Saito et al. 1971, Sakai et al. 2005, Oh et al. 2008) and also produces mycotoxins such as cyclochlorotine, islanditoxin, erythroskyrine and luteoskyrin, which are hepatotoxic agents and carcinogenic (Uraguchi et al. 1961, 1972, Uraguchi 1962, Ueno & Ishikawa 1969, Bouhet et al. 1976, Stark et al. 1978). Talaromyces islandicus, T. radicus, T. rugulosus and T. wortmannii produce rugulosin and skyrin. Rugulosin is a bis-anthraquinoid pigment (Breen et al. 1955) with a speciﬁc antibacterial effect against Staphylococcus aureus (Yamazaki et al. 2010a, b, c) and was also shown to be moderately active against Pythium (Ueno et al. 1980). Studies have shown that rugulosin could also be a hepatocarcinogen (Ueno et al. 1980). Even though they have been classiﬁed as mycotoxins, extrolites such as erythroskyrin have also been reported to be antitumour agents (Kenkyusho 1983). Species in Talaromyces generally produce yellow, orange and red pigments in the mycelium or as diffusing pigments. Azaphilone polyketide pigments like the mitorubrins (mitorubrin, mitorubrinol, mitorubrinol acetate and mitorubrinic acid) (Büchi et al. 1965) and the Monascus red pigments (N-glutaryl monascorubramin, N-glutarylrubropunctamin, monascorubramine, monascin, PP-R and others (Mapari et al. 2009)) are responsible for the red pigments and are produced in different ratios and amounts between different isolates and species (Samson et al. 1989, Frisvad et al. 1990a, b, van Reenen-Hoekstra et al. 1990, Samson et al. 2011). Talaromyces purpurogenus, T. marneffei, T. albobiverticillius, T. minioluteus and T. atroroseus produce large volumes of 176 mitorubrins (Yilmaz et al. 2012, Frisvad et al. 2013). However, only T. atroroseus consistently produces the azaphilone biosynthetic families mitorubrins and Monascus pigments without any mycotoxins and can thus be used for the biotechnological production of these pigments (Frisvad et al. 2013). Talaromyces ﬂavus is one of the most important fungal antagonists used as a bio-control agent of soil-borne pathogens such as Verticillium dahliae, V. albo-atrum, Rhizoctonia solani and Sclerotinia sclerotiorum (Marois et al. 1984, Punja 2001, Brunner et al. 2005, Gohel et al. 2006). It was shown to suppress Verticillium wilt of tomato (Naraghi et al. 2010b), aubergine (Fahima & Henis 1997), potato (Naraghi et al. 2010b, Naraghi et al. 2012), cotton (Naraghi et al. 2012) and green house cucumber (Naraghi et al. 2010a). Their ability to produce enzymes and soluble pigments make Talaromyces an important genus for biotechnological purposes. Important enzyme producers include T. rugulosus (β-rutinosidase and phosphatase; Reyes et al. 1999, Narikawa et al. 2000), T. pinophilus (endoglucanase, cellulase; Pol et al. 2012), T. funiculosus (cellulase; Maeda et al. 2013) and T. cellulolyticus (Acremonium cellulolyticus = T. pinophilus), which is reported as an important cellulose-degrading fungus used for biomass degradation (Fujii et al. 2013, Houbraken et al. 2014). Samson et al. (2011) redeﬁned Talaromyces by combining Penicillium subgenus Biverticillium into Talaromyces using sequence data from the ITS and RPB1 loci. The aim of this study was to follow on from Samson et al. (2011) and provide a taxonomic treatment of the genus Talaromyces using a polyphasic approach. We classify the 88 accepted species in seven sections based on a multigene phylogeny of the ITS, β-tubulin (BenA) and RPB2 gene regions. We provide morphological descriptions using macro- and micromorphological characters and provide tables and notes to be used for morphological identiﬁcations. We also provide a nomenclatural list of all accepted species. For molecular identiﬁcation, GenBank accession numbers to ITS barcodes and BenA sequences for ex-type and reference cultures are given in this list, with BenA proposed as an alternative identiﬁcation marker for Talaromyces species. MATERIAL AND METHODS Strains Strains used in this study (summarised in Table 1) were obtained from the public collection of the CBS-KNAW Fungal Biodiversity Centre, Utrecht, the Netherlands; the working collection of the Department of Applied and Industrial Mycology (DTO), housed at CBS-KNAW; the Department of Systems Biology, DTU, Lyngby, Denmark (IBT); and the United States Department of Agriculture, Agricultural Research Service (USDA-ARS), Peoria, United States (NRRL). Morphological analysis Macroscopic characters were studied on different media and growth conditions. Cultures were inoculated in three point fashion onto Czapek yeast extract agar (CYA), CYA supplemented with 5 % NaCl (CYAS), creatine sucrose agar (CREA), dichloran 18 % glycerol agar (DG18), oatmeal agar (OA), malt extract agar (Oxoid; MEA) and yeast extract sucrose agar (YES) (Samson et POLYPHASIC TAXONOMY OF TALAROMYCES Table 1. Strains used in phylogenetic analysis of the genus Talaromyces. Species name CBS no. Other collections Substrate and origin Talaromyces aculeatus CBS 136673 DTO 277-E3 = IBT 14255 Weathering wood stakes, Palmerston North, New Zealand CBS 282.92 IBT 132626 Soil in secondary forest, Brazil CBS 289.48T ATCC 10409 = IMI 040588 = NRRL 2129 = NRRL A-1474 IBT 14843 = IBT 36624 Textile, USA Nut, South Africa CBS 290.65 CBS 563.92 Stem of Dicymbe Altsonii, French Guiana Talaromyces aerugineus CBS 350.66T BDUN 276 = IMI 105412 Debris, United Kingdom Talaromyces albobiverticillius CBS 133440T Decaying leaves of a broad-leaved tree, Taiwan CBS 137397 DTO 166-E5 = BCRC 34774 = IBT 31667 DTO 166-E6 = BCRC 34775 = IBT 31668 DTO 245-E3 House dust, Mexico CBS 137399 DTO 267-H6 House dust, Thailand CBS 133441 Talaromyces allahabadensis Talaromyces amestolkiae CBS 178.81 ATCC 48474 = FRR 3579 = IMI 253805 Type of P. zacinthae, Crepis zacintha, Alicante, Spain CBS 441.89 DTO 247-D5 Seed groud, Denmark CBS 453.93T CBS 304.63 = ATCC 15067 = NRRL 3397 = FRR 3397 = IBT 3926 = IBT 10824 DTO 179-F5 Cultivated soil, Allahabad, India DTO 179-E4 House dust, South Africa DTO 179-F1 House dust, South Africa CBS 132696T DTO 179-F6 KACC 46611 = DTO 303-E2 CBS 101366 CBS 312.59T CBS 548.73 Talaromyces assiutensis House dust, South Africa House dust, South Africa T Talaromyces angelicus Talaromyces apiculatus Decaying leaves of a broad-leaved tree, Taiwan Dried roots of Angelica gigas, Pyeongchang, Korea Soil, Hong Kong, China ATCC 18315 = FRR 635 = IMI 068239 = IFO 5728 = IBT 10894 = IBT 14261 IBT 5037 Soil, Japan Soil, Suriname CBS 116554 Pasteurised canned strawberries, the Netherlands CBS 118440 Soil, Fes, Marocco CBS 147.78 T Soil, Egypt CBS 645.80 FRR 1966 = IFO 31750 = IMI 198365 Type of T. gossypii, Gossypium, India Talaromyces atricola CBS 255.31T Unknown Talaromyces atroroseus CBS 133442T NRRL 1052 = FRR 1052 = Thom 4640.439 = ATCC 52257 IBT 32470 = DTO 178-A4 House dust, South Africa DTO 267-I1 House dust, Thailand DTO 270-D5 House dust, Mexico DTO 270-D6 House dust, Mexico ATCC 13216 = IMI 099722 = NRRL 3398 IBT 17522 Soil, Georgia T Talaromyces aurantiacus CBS 314.59 Talaromyces austrocalifornicus CBS 644.95T Talaromyces bacillisporus CBS 102389 Sludge of anaerobic pasteurised organic household waste, Sweden CBS 110774 Rye bread, the Netherlands CBS 116927 CBS 296.48T Talaromyces bohemicus CBS 545.86T Talaromyces boninensis CBS 650.95 T Talaromyces brunneus CBS 227.60T Talaromyces calidicanius CBS 112002T Talaromyces cecidicola CBS 101419T Talaromyces chloroloma Soil, California, USA Soil, the Netherlands ATCC 10126 = IMI 040045 = NRRL 1025 CCF 2330 = IAM 14789 Leaf, New York, USA Peloids for balneological purposes, Czech Republic IBT 17516 Lawn soil, Japan ATCC 18229 = FRR 646 = IFO 6438 = IHEM 3907 = IMI 078259 = MUCL 31318 Milled rice imported into Japan, Thailand Soil, Nantou County, Taiwan DAOM 233329 DAOM 241016 Cynipid insect galls on Quercus paciﬁca twigs, Oregon, USA T Fynbos soil, Western Cape, South Africa (continued on next page) www.studiesinmycology.org 177 YILMAZ ET AL. Table 1. (Continued). Species name CBS no. Talaromyces chloroloma Talaromyces cinnabarinus CBS 267.72T CBS 357.72 Talaromyces cnidii Other collections Substrate and origin DTO 180-F4 Fynbos soil, South Africa DTO 182-A5 Air sample, Malmesbury, South Africa ATCC 26215 = NHL 2673 Soil, Japan NHL 2674 Soil, Japan DTO 269-H8 House dust, Thailand DTO 270-A4 House dust, Thailand DTO 270-A8 House dust, Thailand DTO 270-B7 House dust, Thailand KACC 46617T = DTO 303-E1 Dried roots of Cnidium ofﬁcinale, Jecheon, Korea IBT 13019 = DTO 189-A5 Chicken feed (Unga), Nairobi, Kenya NRRL 58644 Air, Maryland, USA T Talaromyces coalescens CBS 103.83 Talaromyces columbinus CBS 137393 Soil under Pinus sp., Spain NRRL 58811 T Air, Loisiana, USA NRRL 62680 Corn grits, Illinois, USA Talaromyces convolutus CBS 100537T IBT 14989 = SUM 3018 Soil, Kathmandu, Nepal Talaromyces dendriticus CBS 660.80T IMI 216897 Eucalyptus pauciﬂora leaf litter, New South Wales, Australia DAOM 226674 Doryanthes excelsa spathes, Mangrove Mountain, New South Wales, Australia DAOM 233861 Unindentiﬁed insect gall on Eucalyptus leaf, Kalnura, New South Wales, Australia DTO 183-G3 Mite, Struisbaai, South Africa NHL 2981 Cultivated soil, Japan ATCC 10437 = DSM 2212 = IMI 040579 = IMI 040579ii = NRRL 2121 DTO 133-A7 Leather, USA House dust, Thailand DTO 133-E4 House dust, Thailand DTO 133-I6 Lotus tea, produced in Vietnam, imported to the Netherlands DTO 244-E6 House dust, New Zealand ATCC 10439 = IMI 040044 = MUCL 28672 = MUCL 29094 = MUCL 29212 = NRRL 1030 IBT 14990 Canvas, France FRR 1868 = IMI 216899 Soil from creek bank, New South Wales PF 1203T = DTO 176I3 = DTO 176I4 Soil, Yokohama, Japan IBT 27044 Dead leaves of Quercus ilex, Parque del Retiro, Madrid, Spain DAOM 236381 Leaves of Quercus suber, port de la Selva, Girona, Spain DAOM 236382 Leaves of Quercus suber, Selva de Mar, Girona, Spain DAOM 236383 Leaves of Quercus suber, Barraca d'en Rabert, Paau, Girona, Spain DAOM 236384 Leaves of Quercus suber, Xovar, Alt Palacia, Valencia Talaromyces derxii Talaromyces diversus CBS 412.89T CBS 320.48 T T Talaromyces duclauxii CBS 322.48 Talaromyces emodensis CBS 100536T Talaromyces erythromellis CBS 644.80 T Talaromyces euchlorocarpius Talaromyces ﬂavovirens Talaromyces ﬂavus T CBS 102801 T CBS 310.38 IMI 197477 = NRRL 2098 CBS 437.62 Talaromyces funiculosus CBS 171.91 NRRL 1035 Unknown CBS 272.86T IMI 193019 Lagenaria vulgaris, India CBS 883.70 Unknown, Java CBS 884.70 Unknown, Java CBS 751.74T Talaromyces hachijoensis Talaromyces helicus 178 Unknown, New Zealand Compost, Bonn, Germany CBS 885.71 Talaromyces galapagensis Soil, Kathmandu, Nepal CBS 134.67 Air, Java, Jakarta IFO 31796 Shaded soil under Maytenus obovata, Galapagos Islands, Isla Santa Cruz, Ecuador IFM 53624T = PF 1174 Soil, Hachijojima, Japan Green house soil under Lycopersicon esculentum, Wageningen, the Netherlands POLYPHASIC TAXONOMY OF TALAROMYCES Table 1. (Continued). Species name Talaromyces helicus CBS no. CBS 335.48 T Other collections Substrate and origin ATCC 10451 = DSM 3705 = IMI 040593 = NRRL 2106 Soil, Sweden Saline soil, Vallee de la Seille, France CBS 550.72A T Talaromyces indigoticus CBS 100534 IBT 17590 = SUM 3010 Soil, Japan Talaromyces intermedius CBS 152.65T BDUN 267 = IFO 31752 = IMI 100874 Allauvial pasture and swamp soil, Nottingham, England CBS 117284 DTO 002-C7 Wheat ﬂour, the Netherlands CBS 338.48T Unknown, Cape Town, South Africa CBS 394.50 ATCC 10127 = IMI 040042 = MUCL 31324 = NRRL 1036 DTO 093-B9 CBS 165.81 ATCC 42240; IMI 253796 Talaromyces islandicus Talaromyces liani CBS 118434 Soil in orchid garden, Sanur, Bali, Indonesia CBS 118885 CBS 225.66T Talaromyces loliensis Talaromyces macrosporus Soil of pepper ﬁeld, DaeJeon, Korea ATCC 18325 = ATCC 18331 = IMI 098480 = NRRL 3380 = VKM F-301 DTO 058-F2 Heat tretaed corn kernels, the Netherlands CBS 172.91 DTO 105-E9 Soil, New Zealand CBS 643.80T ATCC 52252 = FRR 1798 = IMI 216901 = MUCL 31325 NRRL 2101 Rye grass (Lolium), New Zealand CBS 117.72 CBS 131.87 FRR 404 = IMI 197478 CBS 353.72 DTO 105-C4 Apple juice, Stellenbosch, South Africa Pine apple concentrate, the Netherlands Unknown CBS 108.89 Human (male), China CBS 119456 Male blood, Thailand CBS 122.89 Male AIDS patient after travel to Indonesia CBS 135.94 Haemoculture, Nonthaburi, Thailand CBS 388.87T CBS 549.77 Talaromyces mimosinus CBS 659.80 Talaromyces minioluteus CBS 137.84 T CBS 270.35 CBS 642.68T ATCC 18224 = CBS 334.59 = IMI 068794ii = IMI 068794iii ATCC 24100 Man spleen, unknown FRR 1875 = IMI 223991 Soil from creek bank, New South Wales Bamboo rat (Rhizomys sinensis), Vietnam Fruit, damaged by insect, Valladolid, Spain ATCC 4713 = ATCC 52244 = FRR 1064 = IBT 4302 = NRRL 1064 = NRRL 1142 IMI 089377 = MUCL 28666 = NRRL 1714 Zea mays, Virginia, USA Unknown CBS 261.55 Clematis, Boskoop, the Netherlands CBS 283.58 Jute potato bag, treated with copper oxide ammonia, unknown CBS 284.58 Unknown, the Netherlands CBS 351.61 CBS 756.96T Chicken crop, the Netherlands PF 1153 Soil, Taiwan DTO 180-B4 House dust, South Africa CBS 889.96 Talaromyces oumae-annae Cotton fabric, USA Tentage, New Guinea DTO 077-C5 Talaromyces muroii Soil, China Faecal pellet of grasshopper, Malaysia CBS 317.63T Talaromyces marneffei Kapok ﬁbre, unkown Type of P. aurantioﬂammiferum, spice mixture used in sausage making industry, Spain CBS 138207 Dung of sheep, Papua New Guinea CBS 138208T DTO 269-E8 House dust, South Africa Talaromyces palmae CBS 442.88 IMI 343640 Chrysalidocarpus lutescens seed, Wageningen, the Netherlands Talaromyces panamensis CBS 128.89T IMI 297546 Soil, Barro Colorado Island, Panama T PF 1150 = IFM 53616 Soil, Aso-machi, Japan CBS 116872 DTO 247-E1 Production plant, the Netherlands CBS 132063 DTO 191-C5 Straw used in horse stable, the Netherlands CBS 137363 DTO 058-D1 Pectin, unknown CBS 137377 DTO 178-F3 House dust, South Africa Talaromyces paucisporus Talaromyces piceus (continued on next page) www.studiesinmycology.org 179 YILMAZ ET AL. Table 1. (Continued). Species name CBS no. T Talaromyces piceus CBS 361.48 Talaromyces pinophilus CBS 101709 Other collections Substrate and origin ATCC 10519 = IMI 040038 = NRRL 1051 Unknown Soil, Japan CBS 173.91 Unknown, USA CBS 235.94 ATCC 11797 = NRRL 3647 = NRRL A1616 = NRRL A-622 CBS 269.73 Unknown, Germany CBS 440.89 CBS 631.66 Unknown, USA Zea mays, India T ATCC 36839 = CECT 2809 = DSM 1944 = IAM 7013 = IMI 114933 CBS 762.68 PVC, France Type of Penicillium korosum, Rhizosphere, India DTO 183-I6 T Protea repens infructescense, Struisbaai, South Africa IMI 327871 Clay soil under poplar trees, Spain Unknown, USA CBS 303.67T ATCC 10438 = CBS 439.88 = FRR 1074 = IMI 040031 = MUCL 31321 = MUCL 31330 = NRRL 1074 ATCC 18326 = NRRL 3378 Granite soil, Ukraine CBS 470.70T ATCC 18919 = FRR 2039 Feather of Hylocichla fuscescens, Minnesota, USA DAOM 241017T = DTO 180-E7 Fynbos soil, Malmesbury, South Africa DTO 180-E9 Fynbos soil, Malmesbury, South Africa DTO 180-F1 Fynbos soil, Malmesbury, South Africa ATCC 24069 = ATCC 52513 = FRR 1731 = IMI 181546 NRRL 1059 = ATCC 10064 = IBT 10612 = IBT 3560 = DTO 049-F7 IMI 136128 = IBT 3658 = IBT 3560 = DTO 189-A1 IMI 094165 = FRR 1057 = NRRL 1057 Soil, France Talaromyces pittii CBS 139.84 Talaromyces primulinus CBS 321.48T Talaromyces proteolyticus Talaromyces pseudostromaticus Talaromyces ptychoconidium Talaromyces purpureus CBS 475.71T Talaromyces purpurogenus CBS 122434 CBS 132707 CBS 184.27 T Unknown Moulded ﬁeld corn, Wisconsin, USA Soil, Lousiana, USA IMI 091926 = Thom 17 Parasitic on a culture of Aspergillus oryzae, Japan Talaromyces rademirici CBS 140.84T CECT 2771 = IMI 282406 = IMI 327870 Air under willow tree, Valladolid, Spain Talaromyces radicus CBS 100488 DTO 037-F6 Wheat root, New South Wales CBS 100489T FRR 4718 = IBT 14379 Root seadling, New South Wales CBS 100490 DTO 037-F8 Wheat root, New South Wales CBS 137382 DTO 181-D5 Fynbos soil, South Africa DTO 181-D4 Fynbos soil, South Africa CBS 286.36 DTO 181-D7 Fynbos soil, South Africa T DAOM 241660 = DTO 184-B8 Soil, Malmesbury, South Africa DTO 181-E3 Mite, Stellenbosch, South Africa DTO 181-F6 Protea repens infructescense, Stellenbosch, South Africa DTO 182-A3 Protea repens infructescense, Stellenbosch, South Africa DTO 182-A6 Air, Malmesbury, South Africa DTO 183-A7 Protea repens infructescense, Malmesbury, South Africa Cardboard, Norway CBS 196.88 ATCC 10493 = IMI 040589 = NRRL 2107 DTO 193-H6 = IBT 10703 = CBS 113137 FRR 1714 = IBT 3951 Unknown CBS 237.93 ACC 828-81 Unknown CBS 370.48 Currency paper, Washington, USA Talaromyces ramulosus T Talaromyces rotundus CBS 369.48 Talaromyces ruber CBS 132704T Air craft fuel tank, United Kingdom Talaromyces rubicundus CBS 342.59T Talaromyces rugulosus CBS 137366 ATCC 10520 = IMI 00036 = NRRL 1062 = IBT 4431 = IBT 3927 ATCC 13217 = IMI 099723 = NRRL 3400 DTO 061-E8 CBS 344.51 IFO 6016 Type of P. echinosporum Nehira, unknown, Japan ATCC 10128 = IMI 040041 = MUCL 31201 = NRRL 1045 Roating potato tubers (Solanum tuberosum), USA CBS 371.48 180 T Soil, Georgia Type of P. chrysitis, air sample, beer producing factory, Kaulille, Belgium POLYPHASIC TAXONOMY OF TALAROMYCES Table 1. (Continued). Species name Other collections Substrate and origin Talaromyces rugulosus NRRL 1053 = FRR 1053 = IMI 028259 Unknown Type of P. tardum & P. elongatum, decaying twigs, France Talaromyces ryukyuensis NRRL 1073 = FRR 1073 = IMI 040034 = ATCC 10503 = Thom 4640.444 NHL 2917T = DTO 176-I6 Soil, Naha, Japan CBS 138204T DTO 245-H1 House dust, Mexico CBS 138205 DTO 245-H2 House dust, Mexico CBS 138206 DTO 245-H3 House dust, Mexico CBS 233.60 NRRL203 = IMI 78256 = FRR 203 = ATCC 18481 = IFO 6437 NRRL 1129 = FRR 1129 Type of T. phialosporus, milled Californian rice, Japan Talaromyces sayulitensis Talaromyces scorteus CBS no. CBS 340.34T CBS 499.75 CBS 500.75 Talaromyces siamensis Talaromyces solicola Military equipment, Japan Unknown, Nigeria Unknown, Sierra Leone CBS 475.88T T CBS 133445 DTO 270-A6 House dust, Thailand IMI 323204 Forest soil Thailand DTO 269-I3 House dust, Thailand DAOM 241015 = CV 2800 = DTO 180D4 Soil, Malmesbury, South Africa CBS 133446 Soil, Malmesbury, South Africa Talaromyces stipitatus CBS 375.48T ATCC 10500 = NRRL 1006 = IMI 39805 Decaying wood, Louisiana, USA Talaromyces stollii CBS 169.91 NRRL 1033 Unknown substrate, South Africa CBS 265.93 Bronchoalveolar lavage of patient after lung transplantation (subclinical), France CBS 408.93T AIDS patient, the Netherlands CBS 581.94 Unknown CBS 624.93 Ananas camosus cultivar, Martinique T Talaromyces subinﬂatus CBS 652.95 Talaromyces tardifaciens CBS 250.94T T IBT 17520 Copse soil, Japan SUM 3017 = IBT 14986 Paddy soil, Bhaktapur, Nepal KUFC 3399 Soil, Thailand Talaromyces thailandensis CBS 133147 Talaromyces trachyspermus CBS 116556 Pasteurised canned strawberries, Germany CBS 118437 Soil, Marocco CBS 118438 Soil, Marocco CBS 133146T ATCC 10497 = IMI 040043 = NRRL 1028 KUFC 3383 Soil, Trat, Thailand CBS 137400 DTO 270-F5 House dust, Mexico CBS 137401 NRRL1013 = FRR 1013 Carbonated beverage, Washington D.C., USA CBS 162.67T ATCC 22344 = FRR 3462 = NHL 6086 Unknown CBS 583.72A IFO 8395 Soil, Japan CBS 583.72C IFO 8891 Soil, Japan FRR 1727 = IMI 197482 Soil, Misugimura, Japan CCRC 32703 = IBT 18385 = FRR 4436 Soil, Taiwan ATCC 10509 = IMI 040586 = NRRL 2096 IBT 5047 Cotton yarn, England CBS 373.48 Talaromyces tratensis Talaromyces ucrainicus T CBS 127.64 Type of T. ohiensis, soil treated with cyanamide, Germany T Talaromyces udagawae CBS 579.72 Talaromyces unicus CBS 100535T Talaromyces varians CBS 386.48 Talaromyces verruculosus CBS 254.56 Unknown, USA T CBS 388.48T ATCC 10513 = DSM 2263 = IMI 040039 = NRRL 1050 DTO 129-H4 Unknown, Yangambi, Zaire Soil, Texas, USA House dust, Thailand DTO 129-H5 House dust, Thailand Talaromyces viridis CBS 114.72T ATCC 22467 = NRRL 5575 Soil, Australia Talaromyces viridulus CBS 252.87T FRR 1863 = IMI 288716 Soil from bank of creek ﬂoading into Little river, New South Wales (continued on next page) www.studiesinmycology.org 181 YILMAZ ET AL. Table 1. (Continued). Species name Talaromyces wortmannii CBS no. CBS 137376 PF 1130 Type of T. sublevisporus, soil, Japan DTO 108-A4 Unknown CBS 385.48 IMI 40040 = NRRL 1048 = FRR 1048 = ATCC 10508 = IFO 6111 Type of T. variabilis, coconut matting, Johannesburg, South Africa CBS 391.48T NRRL 1017 = IMI 040047 = FRR 1017 = ATCC 10517 = IFO 7738 = Thom 4733.126.1 = IBT 4838 ATCC 20201 = IFO 4683 = BCRC 31677 NRRL 2125 = FRR 2125 = DTO 278-E7 Soil, Denmark Weathering canvas, Panama CBS 138209 DTO 268-E5 House dust, Micronesia CBS 138210T DTO 268-E7 House dust, Micronesia al. 2010), using 90 mm Petri dishes. Plates were incubated for 7 d at 25 C in darkness. Additional CYA plates were incubated at 30 and 37 C, while additional MEA plates were incubated at 30 C. After 7 d of incubation, colony diameters were recorded. The degree of sporulation, obverse and reverse colony colours and the production of soluble pigments were noted. Colony colour codes used in descriptions refer to Kornerup & Wanscher (1967). For ascoma production, OA, MEA and hay infusion agar (HAY) plates were incubated for up to four weeks. Microscope preparations were made from 1 to 2 wk old colonies grown on MEA, with OA and HAY used when sporulation was absent on MEA. OA was used for observing ascomata, asci and ascospores. Lactid acid (60 %) was used as mounting ﬂuid. Characters were captured using a Zeiss SteReo Discovery.V20 dissecting microscope and Zeiss AX10 Imager.A2 compound microscope. The microscopes are equipped with AxioCam MRc5 cameras and uses AxioVs40 v. 4.8.2.0 software. Microscopic measurements were made using Nikon NIS-elements D v. 4.0. Photo plates were prepared in Adobe® Photoshop® CS6. DNA extraction, PCR ampliﬁcation and sequencing DNA extractions were made from 1 to 2 wk old colonies grown on MEA using the Ultraclean™ Microbial DNA isolation Kit (MoBio, Solana Beach, USA) with extracts stored at −20 C. PCR reactions were prepared as described in Houbraken & Samson (2011). Generally, a standard thermal cycle was used, which ran 35 cycles and had a 55 C annealing temperature. The Internal Transcribed Spacer region (ITS) was ampliﬁed using primer pair V9G (de Hoog & Gerrits van den Ende 1998) and LS266 (Masclaux et al. 1995). For the DNA-dependent RNA polymerase II second largest subunit (RPB2), primer-pair RPB2-5F and RPB2-7Cr (Liu et al. 1999) was used with a step-up PCR that started with 5 cycles and annealing temperature of 48 C, followed by 5 cycles at 50 C and a ﬁnal 25 cycles at 52 C. β-tubulin (BenA) was generally ampliﬁed with primer pair Bt2a and Bt2b (Glass & Donaldson 1995), while primer pair T10 and Bt2b (Glass & Donaldson 1995) was used with a 52 C annealing temperature for section Islandici species. Sequencing reactions were set up using the BigDye Terminator v. 3.1 Cycle Sequencing Kit (Applied Biosystems, CA) with the same primer sets used for PCR ampliﬁcation. Sequences were determined on an ABI PRISM 3730xl genetic analyser (Applied Biosystems, 182 Substrate and origin CBS 319.63 CBS 895.73 Talaromyces yelensis Other collections Unkown, Japan California, USA). Sequence contigs were assembled using Seqman Pro v. 9.0.4 (DNAstar Inc.) and newly generated sequences deposited into GenBank. Accession numbers are included in the phylogenetic trees, with accession numbers of ex-type sequences provided in the accepted species list. Data analysis All data sets were aligned using Muscle software included with the MEGA v. 5.2.2 software package (Tamura et al. 2011). Aligned data sets were analysed using Maximum likelihood (ML). The best model for ML was selected based on the Akaike Information Criterion (AIC), which was calculated in MEGA. ML analysis was done by calculating an initial tree using BioNJ and the subsequent Heuristic search done with the Nearest-Neighbour-Interchange (NNI) option. Support in nodes was calculated using a bootstrap analysis of 1 000 replicates. For the multigene phylogeny, the aligned ITS, BenA and RPB2 data sets were concatenated in SeaView (Gouy et al. 2010) and analysed using both ML and Bayesian inference (BI). The most suitable model for BI was selected based on AIC, calculated in MrModeltest v. 2.3 (Nylander et al. 2004). The analysis was run in MrBayes v. 3.2.1 (Huelsenbeck & Ronquist 2001) with two sets of four chains (one cold, three heated), until an average deviation for split frequencies reached 0.01. The sample frequency was set at 100, with 25 % of trees removed as burnin. In the presented phylogenies, thickened branches indicate supported branches with bootstrap support above 80 % and or a posterior probability above 0.95. Trees were visually prepared and edited using Adobe® Illustrator® CS6. Extrolites Extrolites were extracted from fungal strains grown on CYA and YES. In some cases, extractions were made from strains also grown on MEA and OA at 25 C for 7 d. Three agar plugs of each medium were extracted as described in Nielsen et al. (2011) and Houbraken et al. (2012). The extracts were analysed by high performance liquid chromatography with diodearray detection (HPLC-DAD) (Frisvad & Thrane 1987). Analyses made after 2011, used UHPLC-DAD (Houbraken et al. 2012). Comparing retention time, retention index and UV spectra measured at 200–600 nm, the detected eluted compounds were identiﬁed. The UV spectra were compared to a database of UV spectra (Nielsen et al. 2011) and data from POLYPHASIC literature. Authentic standards of the different extrolites were available for most of the Talaromyces compounds (Klitgaard et al. 2014). RESULTS & DISCUSSION Phylogeny of Talaromyces Previous studies showed that Penicillium was a polyphyletic genus with the asexual Penicillium subgenus Biverticillium and the sexual Talaromyces forming a monophyletic clade distinct from Penicillium sensu stricto (LoBuglio et al. 1993, Berbee et al. 1995, Ogawa et al. 1997, Ogawa & Sugiyama 2000, Peterson 2000, Heredia et al. 2001, Wang & Zhuang 2007, Houbraken & Samson 2011, Samson et al. 2011). The move to single name nomenclature in the International Code of Nomenclature for algae, fungi and plants (McNeill et al. 2012) allows phylogenetically related species to be treated under the same generic name regardless of its sexual or asexual morphs. As such, Samson et al. (2011) recombined Penicillium subgenus Biverticillium species into Talaromyces. This means that Talaromyces sensu stricto now contains species, which can reproduce sexually (ascomata) and/or asexually (conidiophores). The generic description of Talaromyces is amended below. In this study, we used sequence data of the ITS, BenA and RPB2 gene regions for deﬁning relationships within Talaromyces (Fig. 1). The aligned concatenated data set, which includes a subset of species classiﬁed in Talaromyces, had a total length of 1 368 bp (ITS, 494 bp; BenA, 336 bp; RPB2, 538 bp). The Kimura 2-parameter (K2) model with gamma distributed (+G) and invariant sites (+I) was the most suitable model for ML, while the general time reversible (GTR) model (+G+I) was most suitable for BI. Trichocoma paradoxa was selected as out-group. Topologies for ML and BI trees were identical and thus, the ML tree was used for illustrating the phylogeny, with bootstrap support (bs) above 80 % and/or posterior probabilities (pp) above 0.95 indicated above thickened branches. Based on this, and the ITS and BenA phylogenies shown below, we divided the 88 accepted Talaromyces species into seven clades and propose these as a new sectional classiﬁcation for the genus. This is discussed below. Talaromyces sensu stricto also now contains species, previously classiﬁed in different genera, for example T. cinnabarinus is introduced for Aphanoascus cinnabarinus and T. bohemicus for Sagenomella bohemica, both phylogenetically belonging in Talaromyces. Some species do in fact produce both the sexual and asexual forms. An example is T. wortmannii, which together with T. sublevisporus, T. variabilis and Penicillium concavorugulosum, forms a coherent clade (based on ITS, BenA, CaM and RPB2 sequence data) under Genealogical Concordance Phylogenetic Species Recognition (GCPSR; Taylor et al. 2010) and as a result are synonymised with T. wortmannii (data will be published elsewhere). This was also conﬁrmed by our morphological studies, where conidiophores of T. sublevisporus and T. wortmannii (known for their teleomorphic states) are identical to that of P. concavorugulosum and T. variabilis. Additionally, extrolite data also supported this. Sexual Talaromyces species are generally homothallic; however, there are two species, which are heterothallic. Talaromyces derxii was the ﬁrst Talaromyces species to be described as heterothallic (Takada & Udagawa 1988). Recent studies have www.studiesinmycology.org TAXONOMY OF TALAROMYCES shown that species, which were previously thought to be asexual, could in fact produce ascomata. One of these heterothallic species is T. ﬂavovirens (= P. aureocephalum) which produces ascomata only on damaged Oak leaf-litter in Catalonia, Spain, but never under laboratory conditions (Visagie et al. 2012). Other species shown to contain both the MAT-1 and MAT-2 loci in populations include T. funiculosus, T. marneffei and T. pinophilus, but a sexual state has not been induced in any of these (Woo et al. 2006, Lopez-Villavicencio et al. 2010, Henk et al. 2012). Mating types in Talaromyces will be the focus of future studies. The genus Lasioderma was described by Montagne (1845) with L. ﬂavovirens as type. Visagie et al. (2012) showed that P. aerocephalum is a synonym of L. ﬂavovirens and that it belongs in the Talaromyces clade. This created a problem where Lasioderma represented an older name than the more widely used Talaromyces and resulted in Seifert et al. (2012) proposing the conservation of Talaromyces over Lasioderma. Generic diagnosis The concept of Talaromyces was adapted in Samson et al. (2011) to accommodate the transfer of Penicillium subgenus Biveriticillium. Here we further adapt the concept based on species transferred to Talaromyces below in the taxonomy section, as well as new characters observed for species. Talaromyces C.R. Benj., Mycologia 47: 681. 1955. = Lasioderma Mont. In Ann. Sci. Nat., Bot., ser. 3, 4: 364. 1845, nom. rej. prop. = 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). Typus: T. vermiculatus (P.A. Dang.) C.R. Benj. (= Talaromyces ﬂavus (Klöcker) Stolk & Samson) Colonies on CYA commonly produce yellow or red reverse and/ or soluble pigments, on CYAS no growth to very restricted growth, sometimes determinate or indeterminate synnemata produced. Conidiophores having smooth or rough-walled elements, characteristically symmetrically biverticillate, with a minor proportion having subterminal branches (in some species with a single subterminal lateral branch that afterwards repeats the branching pattern of the main axis), some species monoverticillate or with solitary phialides. Stipes usually hyaline, terminating in a whorl of metulae of 3–10, appearing symmetrical in face view. Conidiogenous cells phialidic, approximately equal length to metulae, typically acerose, rarely ﬂaskshaped. Conidia aseptate, green en masse, in basipetal chains, usually ellipsoidal to fusiform, rarely globose to subglobose or ovoidal. Ascomata cleistothecial when produced, usually with a distinctly hyphal exterior soft wall, often yellow, occasionally white, creamish, pinkish, orange, reddish or green. Asci produced in chains, containing eight ascospores, rarely two. Ascospores one-celled, ellipsoidal to globose, rarely smoothwalled, but often with spines and/or less commonly ridges, hyaline to yellow, occasionally red. 183 YILMAZ ET AL. T. macrosporus CBS 317.63T 0.98/93 T. funiculosus CBS 272.86T T. rubicundus CBS 342.59T T. pinophilus CBS 631.66T T. apiculatus CBS 312.59T T. aculeatus CBS 289.48T T. siamensis CBS 475.88T */* T. cnidii KACC 46617T T. primulinus CBS 321.48T T. verruculosus CBS 388.48T T. viridulus CBS 252.87T 0.96/- T. flavus CBS 310.38T T. marneffei CBS 388.87T T. duclauxii CBS 322.48T */98 */99 T. calidicanius CBS 112002T T. stollii CBS 408.93T */* */* T. ruber CBS 132704T T. amestolkiae CBS 132696T T. panamensis CBS 128.89T */- T. viridis CBS 114.72T 0.97/- T. euchlorocarpius DTO 176-I3T */- T. thailandensis CBS 133147T */* T. derxii CBS 412.89T T. stipitatus CBS 375.48T */* sect. Talaromyces T. purpurogenus CBS 286.36T */88 T. bohemicus CBS 545.86 */* T T. aerugineus CBS 350.66T */* T. cinnabarinus CBS 267.72T */- T. varians CBS 386.48T */* T. boninensis CBS 650.95T 0.95/- */* */* T. helicus CBS 652.66 T. helicus CBS 335.48T */* T. helicus CBS 649.95 sect. Helici T. rademirici CBS 140.84T T. purpureus CBS 475.71T */* T. ptychoconidium DAOM 241017T */* */95 */96 T. pittii CBS 139.84T T. pseudostromaticus CBS 470.70T T. dendriticus CBS 660.80T */* T. chloroloma DAOM 241016T */* T. ramulosus DAOM 241660T 0.97/91 0.99/88 T. coalescens CBS 103.83T T. cecidicola CBS 101419T sect. Purpurei Fig. 1. Combined phylogeny of the ITS, BenA and RPB2 gene regions of species from Talaromyces. Trichocoma paradoxa was chosen as out-group. Support in nodes is indicated above thick branches and is represented by posterior probabilities (BI analysis) of 0.95 and higher and/or bootstrap values (ML analysis) of 80 % and higher. Full support (1.00/100 %) is indicated with an asterisk (*); support lower than 0.95/80 is indicated with a dash (–). T = ex type. 184 POLYPHASIC */* */84 TAXONOMY OF TALAROMYCES T. trachyspermus CBS 373.48T T. assiutensis CBS 147.78T T. ucrainicus CBS 162.67T */* T. atroroseus CBS 133442T T. minioluteus CBS 642.68T */* T. diversus CBS 320.48T T. convolutus CBS 100537T */* T. solicola DAOM 241015T */* T. erythromellis CBS 644.80T */- T. albobiverticillius CBS 133440T */* T. emodensis CBS 100536T */99 sect. Trachyspermi T. proteolyticus CBS 303.67T 0.97/89 T. mimosinus CBS 659.80T sect. Bacillispori sect. Subinflati T. bacillisporus CBS 296.48T T. palmae CBS 442.88T */* T. subinflatus CBS 652.95T */* */96 T. columbinus NRRL 58811T T. piceus CBS 361.48T T. rotundus CBS 369.48T T. tratensis CBS 133146T */*/* T. scorteus CBS 340.34T -/88 */* */- T. atricola CBS 255.31T T. rugulosus CBS 371.48T T. tardifaciens CBS 250.94T */* */* T. allahabadensis CBS 453.93T T. radicus CBS 100489T T. wortmannii CBS 391.48T 0.96/92 T. loliensis CBS 643.80T -/90 */- T. islandicus CBS 338.48T sect. Islandici T. brunneus CBS 227.60T Trichocoma paradoxa 0.1 Fig. 1. (Continued). Sectional classiﬁcation Subgeneric classiﬁcations have traditionally been used for the taxonomy of Talaromyces and Penicillium. Stolk & Samson (1972) divided Talaromyces into four sections based on differences in their asexual states (Table 2). Pitt (1980) accepted this sectional classiﬁcation and introduced different series for these sections. He did this also for Penicillium subgenus Biverticillium. At the time, classiﬁcations were generally based only on phenotypic characters. However, Yaguchi et al. (1996) used the ubiquinone systems for infrageneric classiﬁcation of Talaromyces. Ubiquinone Q-10(H2) was found in the majority of Talaromyces species; however, they found a mixture of ubiquinone Q-10(H2) and ubiquinone Q-10(H4) in T. trachyspermus and related taxa. Therefore Yaguchi et al. (1996) introduced Talaromyces section Trachyspermi [as ‘trachyspermus’]. In modern taxonomies, phylogenies often do not support classiﬁcations based on criteria such as morphology. Thus, similar to Houbraken & Samson (2011), we use a multigene phylogeny (Fig. 1) for deﬁning a new sectional classiﬁcation of Talaromyces. Species were resolved into seven distinct clades and we propose www.studiesinmycology.org sections Talaromyces, Helici, Purpurei, Trachyspermi, Bacillispori, Subinﬂati and Islandici for these clades. These are discussed below. For each section, we include ITS and BenA phylogenies containing each species classiﬁed in the section, in order to show the use of these genes for sequence based identiﬁcation. The results of this are summarised in the phylogenetic species recognition section below. Talaromyces section Talaromyces. Figs 1, 2. = Penicillium subgenus Biverticillium section Simplicium series Miniolutea Pitt, The genus Penicillium: 419. 1980. = Penicillium subgenus Biverticillium section Coremigenum series Duclauxii Raper & Thom ex Pitt, The genus Penicillium: 404. 1980. = Talaromyces section Talaromyces series Flavi Pitt, The genus Penicillium: 471. 1980. Typus: Talaromyces ﬂavus (Klöcker) Stolk & Samson, Stud. Mycol. 2: 10. 1972. Stolk & Samson (1972) introduced Talaromyces section Talaromyces for species that produce yellow ascomata, which can occasionally be white, creamish, pinkish or reddish, and 185 YILMAZ ET AL. Table 2. Overview of sectional classiﬁcation in different studies of Talaromyces. Stolk & Samson (1972) Pitt (1980) Current study Section Type species Section/Series Type species Section Type species Talaromyces sect. Emersonii Talaromyces sect. Purpurea Talaromyces sect. Talaromyces Talaromyces sect. Thermophila T. emersonii Penicillium subgenus Biverticillium sect. Coremigenum ser. Dendritica Penicillium subgenus Biverticillium sect. Coremigenum ser. Duclauxii Talaromyces sect. Purpureus ser. Purpurei P. dendriticum sect. Bacillispori T. bacillisporus P. duclauxii sect. Helici T. helicus T. purpureus sect. Islandici T. islandicus Penicillium sect. Simplicium ser. Islandica P. islandicum sect. Purpurei T. purpureus Penicillium sect. Simplicium ser. Miniolutea P. minioluteum sect. Subinﬂati T. subinﬂatus Talaromyces sect. Talaromyces ser. Flavi T. ﬂavus sect. Talaromyces T. ﬂavus Talaromyces sect. Talaromyces ser. Lutei T. luteus sect. Trachyspermi T. trachyspermus T. purpureus T. ﬂavus ( =T. vermiculatus) T. thermophilus Talaromyces sect. Talaromyces ser. Trachyspermi T. trachyspermus Talaromyces sect. Thermophilus ser. Thermophili T. thermophilus have yellow ascospores. Conidiophores are usually of the biverticillate-symmetrical type, with some species that have reduced conidiophores with solitary phialides. Phialides are usually acerose, with a minor proportion of species having wider bases (Stolk & Samson 1972). Pitt (1980) used T. minioluteus (= P. minioluteum) for typifying Penicillium subgenus Biverticillium section Simplicium series Miniolutea Pitt. However, he used an incorrect strain for his description of T. minioluteus (CBS 196.88), which was correctly identiﬁed as T. ruber (Yilmaz et al. 2012) and belongs in Talaromyces section Talaromyces. As a result, the series Miniolutea Pitt is synonymised with Talaromyces section Talaromyces. Talaromyces duclauxii (type of Penicillium subgenus Biverticillium section Coremigenum series Duclauxii Pitt) is also resolved in section Talaromyces (Table 2). Section Talaromyces species are commonly isolated from soil, indoor environments, humans with penicilliosis and food products. Common species include T. ﬂavus, T. funiculosus, T. macrosporus, T. marneffei, T. pinophilus and T. purpurogenus. Talaromyces purpurogenus produces rubratoxin, which is a well-known hepa-carcinogenic toxin (Burnside et al. 1957, Kihara et al. 2001). Talaromyces marneffei is the only thermal dimorphic Talaromyces species and is a pathogen of especially HIV patients mostly diagnosed in East Asia (Deng et al. 1988, Supparatpinyo et al. 1994, Chiang et al. 1998, Hien et al. 2001). Talaromyces pinophilus and T. funiculosus are important enzyme producers (Rao et al. 1983, Wood & McCrae 1986, Rando et al. 1997, Sukhacheva et al. 2004). Talaromyces cellulolyticus was recently introduced for Acremonium cellulolyticus (strains Y-94), an important cellulosedegrading species (Fujii et al. 2013). However, based on their sequence data (ITS: AB474749 and BenA: AB773823) we do not consider this species distinct from T. pinophilus and synonymise it with the latter. Delmas et al. (2014) and Lafond et al. (2014) have incorrectly linked the unpublished name T. versatilis (IMI 378536) as a “basionym” of Penicillium funiculosum (≡ T. funiculosus). Even though T. versatilis has not been validly published, the BenA sequence for IMI 378536 (KC992272) is distinct from all accepted Talaromyces species and is closely related to T. angelicus. Accepted species: Talaromyces aculeatus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 174. 2011. [MB560639]. — Herb.: IMI 040588. Ex-type: CBS 289.48 = ATCC 10409 = IMI 040588 = NRRL 2129 = NRRL A-1474. ITS barcode: KF741995. 186 (Alternative markers; BenA = KF741929; CaM = KF741975; RPB1 = n.a.; RPB2 = KM023271). Talaromyces amestolkiae Yilmaz et al., Persoonia 29: 48. 2012. [MB801358]. — Herb.: CBS H-21050. Ex-type: CBS 132696 = DTO 179-F5. ITS barcode: JX315660. (Alternative markers; BenA = JX315623; CaM = KF741937; RPB1 = JX315679; RPB2 = JX315698). Talaromyces angelicus S.H. Yu, T.-J. An & H. Sang, J. Microbiol. 51: 707. 2013. [MB804807]. — Herb.: KACC 46611. Ex-type: KACC 46611. ITS barcode: KF183638. (Alternative markers; BenA = KF183640; CaM = KJ885259; RPB1 = n.a.; RPB2 = n.a.). Talaromyces apiculatus Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 174. 2011. [MB560641]. — Herb.: CBS H-20755. Ex-type: CBS 312.59 = ATCC 18315 = FRR 635 = IMI 068239. ITS barcode: JN899375. (Alternative markers; BenA = KF741916; CaM = KF741950; RPB1 = JN680293; RPB2 = KM023287). Talaromyces aurantiacus (J.H. Mill., Giddens & A.A. Foster) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. [MB560642]. — Herb.: No. 1736 (A.A. Foster). Ex-type: CBS 314.59 = ATCC 13216 = IMI 099722 = NRRL 3398. ITS barcode: JN899380. (Alternative markers; BenA = KF741917; CaM = KF741951; RPB1 = KC202951; RPB2 = n.a.). Talaromyces calidicanius (J.L. Chen) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. [MB560645]. — Herb.: CFC-7 (isotype TNM F12246). Extype: CBS 112002. ITS barcode: JN899319. (Alternative markers; BenA = HQ156944; CaM = KF741934; RPB1 = JN899305; RPB2 = KM023311). Talaromyces cnidii S.H. Yu, T.-J. An & H. Sang, J. Microbiol. 51: 707. 2013. [MB804809]. — Herb.: KACC 46617. Ex-type: KACC 46617. ITS barcode: KF183639. (Alternative markers; BenA = KF183641; CaM = KJ885266; RPB1 = n.a.; RPB2 = KM023299). Talaromyces derxii Takada & Udagawa, Mycotaxon 31: 418. 1988. [MB133755]. — Herb.: NHL 2980. Ex-type: CBS 412.89 = NHL 2981. ITS barcode: JN899327. (Alternative markers; BenA = JX494305; CaM = KF741959; RPB1 = JN680306; RPB2 = KM023282). Talaromyces duclauxii (Delacr.) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560650]. — Herb.: IMI 24312. Ex-type: CBS 322.48 = ATCC 10439 = IMI 040044 = MUCL 28672 = MUCL 29094 = MUCL 29212 = NRRL 1030. ITS barcode: JN899342. (Alternative markers; BenA = JX091384; CaM = KF741955; RPB1 = JN121643; RPB2 = JN121491). Talaromyces euchlorocarpius Yaguchi, Somaya & Udagawa, Mycoscience 40: 133. 1999. [MB460481]. — Herb.: PF 1203. Ex-type: PF 1203 = DTO 176I3 = DTO 176-I4. ITS barcode: AB176617. (Alternative markers; BenA = KJ865733; CaM = KJ885271; RPB1 = n.a.; RPB2 = KM023303). Talaromyces ﬂavovirens (Durieu & Mont.) Visagie, Llimona & Seifert, Mycotaxon 122: 404. 2012. [MB800438]. — Herb.: Lectotype PC 0088796; Epitype BCC 473 = BCN 473. Ex-type: CBS 102801 = IBT 27044. ITS barcode: JN899392. (Alternative markers; BenA = JX091376; CaM = KF741933; RPB1 = n.a.; RPB2 = n.a.). Talaromyces ﬂavus (Klöcker) Stolk & Samson, Stud. Mycol. 2: 10. 1972. [MB324416]. — Herb.: CBS H-7820. Ex-type: CBS 310.38 = IMI 197477 = NRRL 2098. ITS barcode: JN899360. (Alternative markers; BenA = JX494302; CaM = KF741949; RPB1 = JN121639; RPB2 = JF417426). POLYPHASIC 94 KM066186 T. marneffei CBS 119456 KJ775217 T. cnidii DTO 269-H8 KM066187 T. marneffei CBS 108.89 KJ775224 T. cnidii DTO 270-B7 KM066185 T. marneffei CBS 549.77 KJ775223 T. cnidii DTO 270-A8 KM066184 T. marneffei CBS 135.94 KF183641 T. cnidii KACC 46617T KM066183 T. marneffei CBS 122.89 KJ775219 T. siamensis DTO 269-I3 94 JN899314 T. viridulus CBS 252.87T JX091379 T. siamensis CBS 475.88T JN899319 T. calidicanius CBS 112002T JX091377 T. flavovirens DAOM 236383 AF285782 T. viridis CBS 114.72T JX091374 T. flavovirens DAOM 236382 99 AB176617 T. euchlorocarpius DTO 176-I3T JX091373 T. flavovirens DAOM 236381 JN899317 T. primulinus CBS 321.48T JX091375 T. flavovirens DAOM 236384 JN899362 T. panamensis CBS 128.89T JX091376 T. flavovirens CBS 102801T KJ775710 T. oumae-annae CBS 138207 KF741910 T. apiculatus CBS 101366 96 KJ775720 T. oumae-annae CBS 138208T KF741916 T. apiculatus CBS 312.59T JX315674 T. stollii CBS 408.93T KF741919 T. apiculatus CBS 548.73 JX315676 T. stollii CBS 624.93 98 KF741915 T. aculeatus CBS 290.65 82 JX315670 T. stollii CBS 265.93 KF741927 T. aculeatus CBS 136673 JX315664 T. stollii CBS 169.91 KF741929 T. aculeatus CBS 289.48T 98 JX315675 T. stollii CBS 581.94 KF741914 T. aculeatus CBS 282.92 KJ775706 T. amestolkiae DTO 179-E4 KF741920 T. aculeatus CBS 563.92 JX315660 T. amestolkiae CBS 132696T KF183640 T. angelicus KACC 46611T KJ775707 T. amestolkiae DTO 179-F1 KJ775207 T. sayulitensis CBS 138205 99 KJ775708 T. amestolkiae DTO 179-F6 96 KJ775208 T. sayulitensis CBS 138206 KJ775206 T. sayulitensis CBS 138204T JX315677 T. ruber CBS 868.96 JX315662 T. ruber CBS 132704T 83 KM066140 T. liani DTO 058-F2 JX091380 T. liani CBS 225.66T JX315666 T. ruber CBS 196.88 92 JX315673 T. ruber CBS 370.48 KM066139 T. liani CBS 118434 JX315667 T. ruber CBS 237.93 KM066138 T. liani CBS 118885 JX898041 T. thailandensis CBS 133147T JX494301 T. pinophilus CBS 762.68 KM066191 T. macrosporus CBS 131.87 JX091621 T. pinophilus DTO 183-I6 KM066192 T. macrosporus DTO 077-C5 KM066141 T. pinophilus CBS 173.91 KM066190 T. macrosporus DTO 105-C4 KM066142 T. pinophilus CBS 101709 JN899333 T. macrosporus CBS 317.63T KM066143 T. pinophilus CBS 440.89 KM066188 T. macrosporus CBS 117.72 85 JN899377 T. funiculosus CBS 272.86T KM066146 T. macrosporus DTO 105-C4 KM066194 T. funiculosus CBS 885.71 KM066147 T. macrosporus CBS 131.87 KM066195 T. funiculosus CBS 884.70 KM066148 T. macrosporus CBS 117.72 95 JX091382 T. macrosporus CBS 317.63T KM066196 T. funiculosus CBS 883.70 KJ775731 T. cnidii DTO 270-B7 KM066149 T. macrosporus CBS 353.72 KJ775726 T. siamensis DTO 269-I3 KM066150 T. macrosporus DTO 077-C5 JX494305 T. primulinus CBS 321.48T KJ775730 T. cnidii DTO 270-A8 88 KM066144 T. pinophilus CBS 269.73 KM066145 T. pinophilus CBS 235.94 KM066193 T. funiculosus CBS 171.91 93 JX091381 T. pinophilus CBS 631.66T 84 KM066189 T. macrosporus CBS 353.72 84 KJ775724 T. cnidii DTO 269-H8 93 KJ775203 T. oumae-annae CBS 138207 94 KF183639 T. cnidii KACC 46617T KJ775213 T. oumae-annae CBS 138208T JX091385 T. viridulus CBS 252.87T KJ775729 T. cnidii DTO 270-A4 JN899385 T. siamensis CBS 475.88T KF741928 T. verruculosus CBS 388.48T 98 JN899327 T. derxii CBS 412.89T KJ775191 T. verruculosus DTO 129-H4 JN899342 T. duclauxii CBS 322.48T KJ775192 T. verruculosus DTO 129-H5 99 JN899380 T. aurantiacus CBS 314.59T KF741913 T. verruculosus CBS 254.56 JN899348 T. stipitatus CBS 375.48T KF741917 T. aurantiacus CBS 314.59T JN899372 T. purpurogenus CBS 286.36T 89 99 JX494305 T. derxii CBS 412.89T JX315663 T. purpurogenus CBS 122434 KM066151 T. muroii CBS 283.58 JX315665 T. purpurogenus CBS 184.27 KM066152 T. muroii CBS 284.58 JX315661 T. purpurogenus CBS 132707 99 JN899332 T. intermedius CBS 152.65T KF741977 T. apiculatus CBS 101366 KJ865727 T. muroii CBS 756.96T KM066155 T. muroii CBS 351.61 90 KF741985 T. apiculatus CBS 548.73 KM066154 T. muroii CBS 889.96 KM066153 T. muroii CBS 261.55 JN899375 T. apiculatus CBS 312.59T JX013914 T. flavovirens DAOM 236383 JX013915 T. flavovirens DAOM 236384 JX091388 T. galapagensis CBS 751.74T JX494309 T. rubicundus CBS 342.59T 95 JX494308 T. indigoticus CBS 100534T JX315633 T. stollii CBS 408.93T JX013913 T. flavovirens DAOM 236382 JX013912 T. flavovirens DAOM 236381 JX315632 T. stollii CBS 581.94 JN899392 T. flavovirens CBS 102801T 100 KF183638 T. angelicus KACC 46611T KM066200 T. muroii CBS 261.55 JX315636 T. stollii CBS 624.93 KJ775201 T. amestolkiae DTO 179-F6 98 KM066198 T. muroii CBS 351.61 100 KM066197 T. muroii CBS 283.58 KM066201 T. muroii CBS 889.96 KJ775200 T. amestolkiae DTO 179-F1 JX315629 T. ruber CBS 132704T KF741980 T. verruculosus CBS 254.56 KJ775699 T. verruculosus DTO 129-H5 KF741994 T. verruculosus CBS 388.48T JN899360 T. flavus CBS 310.38T KM066202 T. flavus CBS 437.62 JX315623 T. amestolkiae CBS 132696T KJ775199 T. amestolkiae DTO 179-E4 82 JN899351 T. muroii CBS 756.96T KJ775698 T. verruculosus DTO 129-H4 JX315635 T. stollii CBS 265.93 JX315634 T. stollii CBS 169.91 KM066199 T. muroii CBS 284.58 92 TALAROMYCES KJ775222 T. cnidii DTO 270-A4 87 JN899344 T. marneffei CBS 388.87T TAXONOMY OF JX315627 T. ruber CBS 196.88 100 JX315628 T. ruber CBS 237.93 JX315630 T. ruber CBS 370.48 JX315631 T. ruber CBS 868.96 JX494294 T. thailandensis CBS 133147T JX091387 T. intermedius CBS 152.65T Fig. 2. ML phylogenies of ITS (left) and BenA (right) for species classiﬁed in Talaromyces sect. Talaromyces. Talaromyces dendriticus was chosen as out-group. Support in nodes is indicated above thick branches and is represented by bootstrap values of 80 % and higher. T = ex type. Model selected: Tamura 3-parameter (T92) +G+I for ITS, K2+G for BenA, alignment 537 (ITS) and 458 (BenA) bp. www.studiesinmycology.org 187 YILMAZ ET AL. KJ865733 T. euchlorocarpius DTO 176-I3 99 HQ156944 T. calidicanius CBS 112002 JX091384 T. duclauxii CBS 322.48 99 JX494302 T. flavus CBS 310.38 KM066156 T. flavus CBS 437.62 JX091389 T. marneffei CBS 388.87 KM066157 T. marneffei CBS 108.89 99 KM066158 T. marneffei CBS 135.94 KM066159 T. marneffei CBS 549.77 KM066160 T. marneffei CBS 119456 KM066161 T. marneffei CBS 122.89 HQ156948 T. panamensis CBS 128.89 JX494310 T. viridis CBS 114.72 88 84 98 KM066162 T. funiculosus CBS 171.91 JX091383 T. funiculosus CBS 272.86 KM066163 T. funiculosus CBS 883.70 KM066164 T. funiculosus CBS 884.70 KM066165 T. funiculosus CBS 885.71 KM111288 T. stipitatus CBS 375.48 JX315640 T. purpurogenus CBS 122434 90 100 JX315638 T. purpurogenus CBS 132707 JX315637 T. purpurogenus CBS 184.27 JX315639 T. purpurogenus CBS 286.36 JX091391 T. dendriticus CBS 660.80 CBS 660.80T Fig. 2. (Continued). Talaromyces funiculosus (Thom) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560653]. — Herb.: IMI 193019. Ex-type: CBS 272.86 = IMI 193019. ITS barcode: JN899377. (Alternative markers; BenA = JX091383; CaM = KF741945; RPB1 = JN680288; RPB2 = KM023293). Talaromyces galapagensis Samson & Mahoney, Trans. Brit. Mycol. Soc. 69: 158. 1977. [MB324417]. — Herb.: CBS H-7489. Ex-type: CBS 751.74 = IFO 31796. ITS barcode: JN899358. (Alternative markers; BenA = JX091388; CaM = KF741966; RPB1 = JN680321; RPB2 = n.a.). Talaromyces indigoticus Takada & Udagawa, Mycotaxon 46: 129. 1993. [MB359290]. — Herb.: CBM SUM-3010. Ex-type: CBS 100534 = IBT 17590. ITS barcode: JN899331. (Alternative markers; BenA = JX494308; CaM = KF741931; RPB1 = JN680323; RPB2 = n.a.). Talaromyces intermedius (Apinis) Stolk & Samson, Stud. Mycol. 2: 21. 1972. [MB324418]. — Herb.: CBS H-7828. Ex-type: CBS 152.65 = BDUN 267 = IFO 31752 = IMI 100874. ITS barcode: JN899332. (Alternative markers; BenA = JX091387; CaM = KJ885290; RPB1 = JN680276; RPB2 = n.a.). Talaromyces liani (Kamyschko) Yilmaz, Frisvad & Samson, (this study). [MB809555]. — Herb.: unknown. Ex-type: CBS 225.66 = ATCC 18325 = ATCC 18331 = IMI 098480 = NRRL 3380 = VKM F-301. ITS barcode: JN899395. (Alternative markers; BenA = JX091380; CaM = KJ885257; RPB1 = JN680280; RPB2 = n.a.). Talaromyces macrosporus (Stolk & Samson) Frisvad, Samson & Stolk, Antonie van Leeuwenhoek 57: 186. 1990. [MB126704]. — Herb.: CBS H-7822. Extype: CBS 317.63 = FRR 404 = IMI 197478. ITS barcode: JN899333. (Alternative markers; BenA = JX091382; CaM = KF741952; RPB1 = JN680296; RPB2 = KM023292). Talaromyces marneffei (Segretain, Capponi & Sureau) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560656]. — Herb.: IMI 68794iii. Ex-type: CBS 388.87 = ATCC 18224 = CBS 334.59 = IMI 068794ii = IMI 068794iii. ITS barcode: JN899344. (Alternative markers; BenA = JX091389; CaM = KF741958; RPB1 = JN899298; RPB2 = KM023283). Talaromyces muroii Yaguchi, Someya & Udagawa, Mycoscience 35: 252. 1994. [MB362930]. — Herb.: CBM PF-1153. Ex-type: CBS 756.96 = PF 1153. ITS barcode: JN899351. (Alternative markers; BenA = KJ865727; CaM = KJ885274; RPB1 = n.a.; RPB2 = n.a.). Talaromyces oumae-annae Visagie et al., Stud. Mycol. 78: 130. 2014. [MB809187]. — Herb.: CBS H-21797. Ex-type: CBS 138208 = DTO 269-E8. ITS barcode: KJ775720. (Alternative markers; BenA = KJ775213; CaM = KJ775425; RPB1 = n.a.; RPB2 = n.a.). Talaromyces panamensis (Samson, Stolk & Frisvad) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560659]. — Herb.: CBS 128.89. Ex-type: CBS 128.89 = IMI 297546. ITS barcode: JN899362. (Alternative markers; BenA = HQ156948; CaM = KF741936; RPB1 = JN899291; RPB2 = KM023284). 188 Talaromyces paucisporus (Yaguchi, Someya & Udagawa) Samson & Houbraken, Stud. Mycol. 71: 176. 2011. [MB560684]. — Herb.: PF 1150. Ex-type: PF 1150 = IFM 53616. ITS barcode: AB176603. (Alternative markers; BenA = n.a.; CaM = n.a.; RPB1 = n.a.; RPB2 = n.a.). Talaromyces pinophilus (Hedgc.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560662]. — Herb.: IMI 114933. Ex-type: CBS 631.66 = ATCC 36839 = CECT 2809 = DSM 1944 = IAM 7013 = IMI 114933. ITS barcode: JN899382. (Alternative markers; BenA = JX091381; CaM = KF741964; RPB1 = JN680313; RPB2 = KM023291). Talaromyces primulinus (Pitt) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011. [MB560664]. — Herb.: IMI 040031. Ex-type: CBS 321.48 = ATCC 10438 = CBS 439.88 = FRR 1074 = IMI 040031 = MUCL 31321 = MUCL 31330 = NRRL 1074. ITS barcode: JN899317. (Alternative markers; BenA = JX494305; CaM = KF741954; RPB1 = JN680298; RPB2 = KM023294). Talaromyces purpurogenus (Stoll) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560667]. — Herb.: IMI 091926. Ex-type: CBS 286.36 = IMI 091926. ITS barcode: JN899372. (Alternative markers; BenA = JX315639; CaM = KF741947; RPB1 = JN680271; RPB2 = JX315709). Talaromyces ruber (Stoll) Yilmaz et al., Persoonia 29: 48. 2012. [MB801360]. — Herb.: CBS H-21052. Ex-type: CBS 132704 = DTO 193-H6 = IBT 10703 = CBS 113137. ITS barcode: JX315662. (Alternative markers; BenA = JX315629; CaM = KF741938; RPB1 = JX315681; RPB2 = JX315700). Talaromyces rubicundus (J.H. Mill., Giddens & A.A. Foster) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560671]. — Herb.: No. 2531 (A.A. Foster). Ex-type: CBS 342.59 = ATCC 13217 = IMI 099723 = NRRL 3400. ITS barcode: JN899384. (Alternative markers; BenA = JX494309; CaM = KF741956; RPB1 = JN680301; RPB2 = KM023296). Talaromyces sayulitensis Visagie et al., Stud. Mycol. 78: 132. 2014. [MB809188]. — Herb.: CBS H-21798. Ex-type: CBS 138204 = DTO 245-H1. ITS barcode: KJ775713. (Alternative markers; BenA = KJ775206; CaM = KJ775422; RPB1 = n.a.; RPB2 = n.a.). Talaromyces siamensis (Manoch & C. Ramírez) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. [MB560674]. — Herb.: CBS 475.88. Ex-type: CBS 475.88 = IMI 323204. ITS barcode: JN899385. (Alternative markers; BenA = JX091379; CaM = KF741960; RPB1 = n.a.; RPB2 = KM023279). Talaromyces stipitatus (Thom) C.R. Benj., Mycologia 47: 684. 1955. [MB306722]. — Herb.: CBS H-7835. Ex-type: CBS 375.48 = ATCC 10500 = NRRL 1006 = IMI 39805. ITS barcode: JN899348. (Alternative markers; BenA = KM111288; CaM = KF741957; RPB1 = JN680303; RPB2 = KM022380). Talaromyces stollii Yilmaz et al., Persoonia 29: 52. 2012. [MB801359]. — Herb.: CBS H-21053. Ex-type: CBS 408.93. ITS barcode: JX315674. POLYPHASIC TAXONOMY OF TALAROMYCES KM066176 T. helicus CBS 134.67 Fig. 3. ML phylogenies of ITS (left) and BenA (right) for species classiﬁed in Talaromyces sect. Helici. Talaromyces ucrainicus was chosen as out-group. Support in nodes is indicated above thick branches and is represented by bootstrap values of 80 % and higher. T = ex type. Model selected: T92+G for ITS, K2+I for BenA, alignment 463 (ITS) and 510 (BenA) bp. (Alternative markers; BenA = JX315633; CaM = JX315646; RPB1 = JX315693; RPB2 = JX315712). Talaromyces thailandensis Manoch, Dethoup & Yilmaz, Mycoscience 54: 339. 2013. [MB801737]. — Herb.: CBS H-21075. Ex-type: CBS 133147 = KUFC 3399. ITS barcode: JX898041. (Alternative markers; BenA = JX494294; CaM = KF741940; RPB1 = JX898043; RPB2 = KM023307). Talaromyces verruculosus (Peyronel) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560678]. — Herb.: IMI 040039. Ex-type: CBS 388.48 = ATCC 10513 = DSM 2263 = IMI 040039 = NRRL 1050. ITS barcode: KF741994. (Alternative markers; BenA = KF741928; CaM = KF741944; RPB1 = n.a.; RPB2 = KM023306). Talaromyces viridis (Stolk & G.F. Orr) Arx, Persoonia 13: 2821. 1987. [MB132097]. — Herb.: CBS H-7732 (isotype), CBS H-7733 (isotype), CBS H-7734 (isotype). Ex-type: CBS 114.72 = ATCC 22467 = NRRL 5575. ITS barcode: AF285782. (Alternative markers; BenA = JX494310; CaM = KF741935; RPB1 = JN121571; RPB2 = JN121430). Talaromyces viridulus Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. [MB560679]. — Herb.: FRR 1863. Ex-type: CBS 252.87 = FRR 1863 = IMI 288716. ITS barcode: JN899314. (Alternative markers; BenA = JX091385; CaM = KF741943; RPB1 = JN680284; RPB2 = JF417422). Talaromyces section Helici Yilmaz, Frisvad & Samson, sect. nov. MycoBank MB809558. Figs 1, 3. Typus: Talaromyces helicus (Raper & Fennel) C.R. Benj., Mycologia 47: 684. 1955. Description: Conidiophores biverticillate, occasionally consisting of solitary phialides, with stipes generally pigmented, CYA reverse yellowish brown or dark green, species generally grow at 37 C and do not produce acid on CREA, while some do not grow on this medium. Some species produce yellow to deep red or green ascomata. Talaromyces section Helici is basal to section Talaromyces and includes a main clade containing T. helicus, T. boninensis and T. varians, which produce conidiophores with pigmented stipes, and the second clade containing T. cinnabarinus (≡ Paecilomyces cinnabarinus), T. aerugineus (≡ Paecilomyces aerugineus), T. bohemicus (≡ Sagenomella bohemica) and T. ryukyuensis (≡ Sagenoma ryukyuense) (Fig. 3). Accepted species: Talaromyces aerugineus (Samson) Yilmaz, Frisvad & Samson, (this study). [MB809553]. — Herb.: CBS H-7448. Ex-type: CBS 350.66 = BDUN 276 = IMI 105412. ITS barcode: AY753346. (Alternative markers; BenA = KJ865736; CaM = KJ885285; RPB1 = n.a.; RPB2 = JN121502). Talaromyces bohemicus (Fassat. & Peckova) Yilmaz, Frisvad & Samson, (this study). [MB809554]. — Herb.: unknown. Ex-type: CBS 545.86 = CCF 2330 = IAM 14789. ITS barcode: JN899400. (Alternative markers; www.studiesinmycology.org BenA = KJ865719; CaM = KJ885286; RPB1 = JN121699; RPB2 = JN121532). Talaromyces boninensis (Yaguchi & Udagawa) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. [MB560643]. — Herb.: CBM PF-1103. Ex-type: CBS 650.95 = IBT 17516. ITS barcode: JN899356. (Alternative markers; BenA = KJ865721; CaM = KJ885263; RPB1 = JN680319; RPB2 = KM023276). Talaromyces cinnabarinus (S.C. Jong & E.E. Davis) Yilmaz, Samson & Frisvad, (this study). [MB809557]. — Herb.: CBS H-6686. Ex-type: CBS 267.72 = ATCC 26215 = NHL 2673. ITS barcode: JN899376. (Alternative markers; BenA = AY753377; CaM = KJ885256; RPB1 = n.a.; RPB2 = JN121477). Talaromyces helicus (Raper & Fennel) C.R. Benj., Mycologia 47: 684. 1955. [MB306715]. — Herb.: IMI 040593. Ex-type: CBS 335.48 = ATCC 10451 = DSM 3705 = IMI 040593 = NRRL 2106. ITS barcode: JN899359. (Alternative markers; BenA = KJ865725; CaM = KJ885289; RPB1 = JN680300; RPB2 = KM023273). Talaromyces ryukyuensis (S. Ueda & Udagawa) Arx, Persoonia 13: 282. 1987. [MB132096]. — Herb.: unknown. Ex-type: NHL 2917 = DTO 176-I6. ITS barcode: AB176628. (Alternative markers; BenA = n.a.; CaM = n.a.; RPB1 = n.a.; RPB2 = n.a.). Talaromyces varians (G. Sm.) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. [MB560677]. — Herb.: IMI 040586. Ex-type: CBS 386.48 = ATCC 10509 = IMI 040586 = NRRL 2096. ITS barcode: JN899368. (Alternative markers; BenA = KJ865731; CaM = KJ885284; RPB1 = JN680305; RPB2 = KM023274). Talaromyces section Purpurei Stolk & Samson [as ‘Purpurea’], Stud. Mycol. 2: 56. 1972. MycoBank MB39627. Figs 1, 4. = Penicillium subgenus Biverticillium sect. Coremigenum ser. Dendritica Pitt, The genus Penicillium: 413. 1980. Typus: Talaromyces purpureus (E. Müll. & Pacha-Aue) Stolk & Samson, Stud. Mycol. 2: 57. 1972. Description: Conidiophores biverticillate often with additional subterminal branches. Synnemata usually produced after two to three weeks of incubation. Colonies in general grow rapidly on CYA and MEA. Talaromyces section Purpurei includes species that generally produce synnemata after two to three weeks of incubation, the exceptions being T. rademirici, T. purpureus and T. ptychoconidium (Fig. 4). Pitt (1980) used synnemata production as a deﬁning character for his section Coremigenum (type species Talaromyces dendriticus ≡ P. dendriticum), which we synonomise here with section Purpurei. In general, this group of species produces biverticillate conidiophores that have additional subterminal branches. Talaromyces purpureus is the exception, which 189 YILMAZ ET AL. JX091495 T. ramulosus DTO 181-F6 JX091629 T. ramulosus DTO 181-F6 JX091493 T. ramulosus DTO 183-A7 JX091632 T. ramulosus DTO 183-A7 JX091494 T. ramulosus DTO 181-E3 JX091630 T. ramulosus DTO 182-A3 T EU795706 T. ramulosus DAOM 241660 JX091496 T. ramulosus DTO 182-A3 JX091497 T. ramulosus DTO 182-A6 FJ160273 T. chloroloma DTO 180-F4 JX091485 T. chloroloma DTO 182-A5 FJ160273 T. chloroloma DAOM 241016T AY787844 T. cecidicola CBS 101419T JN899366 T. coalescens CBS 103.83T JN899325 T. pittii CBS 139.84T JN899371 T. pseudostromaticus CBS 470.70T FJ753290 T. ramulosus DAOM 241660T JX091626 T. ramulosus DTO 181-E3 JX091631 T. ramulosus DTO 182-A6 FJ753295 T. cecidicola CBS 101419T JX091390 T. coalescens CBS 103.83T JX091597 T. chloroloma DTO 182-A5 GU385737 T. chloroloma DTO 180-F4 GU385736 T. chloroloma DAOM 241016T KJ865728 T. pittii CBS 139.84T HQ156950 T. pseudostromaticus CBS 470.70T JN899339 T. dendriticus CBS 660.80T JX091391 T. dendriticus CBS 660.80T AY787842 T. dendriticus DAOM 226674 FJ753293 T. dendriticus DAOM 226674 AY787843 T. dendriticus DAOM 233861 JX091486 T. dendriticus DTO 183-G3 JN899328 T. purpureus CBS 475.71T FJ753294 T. dendriticus DAOM 233861 JX091619 T. dendriticus DTO 183-G3 GU385739 T. purpureus CBS 475.71T JN899386 T. rademirici CBS 140.84T GU385735 T. ptychoconidium DTO 180-F1 GQ414762 T. ptychoconidium DTO 180-F1 GU385734 T. ptychoconidium DTO 180-E9 FJ160267 T. ptychoconidium DTO 180-E9 GU385733 T. ptychoconidium DAOM 241017T FJ160266 T. ptychoconidium DAOM 241017T JN899319 T. calidicanius CBS 112002T KJ865734 T. rademirici CBS 140.84T HQ156944 T. calidicanius CBS 112002T Fig. 4. ML phylogenies of ITS (left) and BenA (right) for species classiﬁed in Talaromyces sect. Purpurei. Talaromyces calidicanius was chosen as out-group. Support in nodes is indicated above thick branches and is represented by bootstrap values of 80 % and higher. T = ex type. Model selected: T92+G for ITS, K2+G for BenA, alignment 521 (ITS) and 376 (BenA) bp. produces conidiophores that can be monoverticillate or have solitary phialides. A number of species seems to be associated with very speciﬁc habitats. Talaromyces cecidicola was isolated from wasp insect galls (Seifert et al. 2004), T. dendriticus is commonly found in the presence of Eucalyptus (Pitt 1980, Seifert et al. 2004), while T. chloroloma and T. ramulosus are very common in Protea repens infructescences and other substrates like apples grown in the Fynbos region of the Western Cape, South Africa (Visagie et al. 2009, Van der Walt et al. 2010, Visagie & Jacobs 2012). Accepted species: Talaromyces cecidicola (Seifert, Hoekstra & Frisvad) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560646]. — Herb.: DAOM 233329. Ex-type: CBS 101419 = DAOM 233329. ITS barcode: AY787844. (Alternative markers; BenA = FJ753295; CaM = KJ885287; RPB1 = n.a.; RPB2 = KM023309). Talaromyces chloroloma Visagie & K. Jacobs, Persoonia 28: 18. 2012. [MB564326]. — Herb.: PREM 60033. Ex-type: DAOM 241016 = CV 2802. ITS barcode: FJ160273. (Alternative markers; BenA = GU385736; CaM = KJ885265; RPB1 = n.a.; RPB2 = KM023304). Talaromyces coalescens (Quintan.) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. [MB560647]. — Herb.: CBS 103.83. Ex-type: CBS 103.83. ITS barcode: JN899366. (Alternative markers; BenA = JX091390; CaM = KJ885267; RPB1 = n.a.; RPB2 = KM023277). Talaromyces dendriticus (Pitt) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560648]. — Herb.: IMI 216897. Ex-type: CBS 660.80 = IMI 216897. ITS barcode: JN899339. (Alternative markers; BenA = JX091391; CaM = KF741965; RPB1 = JN121714; RPB2 = KM023286). Talaromyces pittii (Quintan.) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560663]. — Herb.: CBS 139.84. Ex-type: CBS 139.84 = IMI 327871. ITS barcode: JN899325. (Alternative markers; BenA = KJ865728; CaM = KJ885275; RPB1 = JN680274; RPB2 = KM023297). Talaromyces pseudostromaticus (Hodges, G.M. Warner & Rogerson) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560666]. — Herb.: Warner 18 (NY). Ex-type: CBS 470.70 = ATCC 18919 = FRR 2039. ITS barcode: JN899371. (Alternative markers; BenA = HQ156950; CaM = KJ885277; RPB1 = JN899300; RPB2 = KM023298). Talaromyces ptychoconidium Visagie & K. Jacobs, Persoonia 28: 18. 2012. [MB564327]. — Herb.: PREM 60041. Ex-type: DAOM 241017 = CV 2808 = DTO 180-E7. ITS barcode: FJ160266. (Alternative markers; BenA = GU385733; CaM = JX140701; RPB1 = n.a.; RPB2 = KM023278). 190 Talaromyces purpureus (E. Müll. & Pacha-Aue) Stolk & Samson, Stud. Mycol. 2: 57. 1972. [MB324420]. — Herb.: CBS H-7832 (isotype). Ex-type: CBS 475.71 = ATCC 24069 = ATCC 52513 = FRR 1731 = IMI 181546. ITS barcode: JN899328. (Alternative markers; BenA = GU385739; CaM = KJ885292; RPB1 = JN121687; RPB2 = JN121522). Talaromyces rademirici (Quintan.) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. [MB560668]. — Herb.: CBS 140.84. Ex-type: CBS 140.84 = CECT 2771 = IMI 282406 = IMI 327870. ITS barcode: JN899386. (Alternative markers; BenA = KJ865734; CaM = n.a.; RPB1 = n.a.; RPB2 = KM023302). Talaromyces ramulosus (Visagie & K. Jacobs) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560670]. — Herb.: PREM 59947. Ex-type: DAOM 241660 = CV 2837 = DTO 184-B8. ITS barcode: EU795706. (Alternative markers; BenA = FJ753290; CaM = JX140711; RPB1 = n.a.; RPB2 = KM023281). Talaromyces section Trachyspermi Yaguchi & Udagawa [as ‘trachyspermus’], Mycoscience 37: 57. 1996. MycoBank MB538978. Figs 1, 5. Typus: Talaromyces trachyspermus (Shear) Stolk & Samson, Stud. Mycol. 2: 32. 1972. Description: Restricted growth on CYA, YES and DG18, growing slightly faster on MEA, no growth to poor growth on CREA. Some species produce abundant red pigments. Conidiophores are generally biverticillate and some species produce creamish white or yellow ascomata. Talaromyces section Trachyspermi includes species that generally grow restrictedly on CYA, YES and DG18, and slightly faster on MEA. This group of species also grows poorly on CREA and some species produce colonies with abundant red pigments. Conidiophores are generally biverticillate and ascomata, when produced, have a creamish white or yellow colour. Talaromyces atroroseus, T. minioluteus and T. albobiverticillius are biotechnologically important, with their pigments used as colourants in the food industry (Frisvad et al. 2013). POLYPHASIC KM066166 T. trachyspermus CBS 118438 TAXONOMY OF TALAROMYCES KJ865720 T. assiutensis CBS 147.78T KM066167 T. assiutensis CBS 116554 KF114802 T. assiutensis CBS 645.80 KM066168 T. assiutensis CBS 118440 KM066124 T. assiutensis CBS 116554 KM066169 T. trachyspermus CBS 118437 KM066125 T. assiutensis CBS 118440 KM066170 T. trachyspermus CBS 116556 KF114803 T. trachyspermus CBS 373.48T JN899323 T. assiutensis CBS 147.78T KM066128 T. trachyspermus CBS 118438 JN899334 T. assiutensis CBS 645.80 KM066126 T. trachyspermus CBS 116556 JN899354 T. trachyspermus CBS 373.48T KM066127 T. trachyspermus CBS 118437 JN899394 T. ucrainicus CBS 162.67T KF114796 T. udagawae CBS 579.72T KM066173 T. ucrainicus CBS 127.64 KF114799 T. minioluteus CBS 642.68T KM066174 T. ucrainicus CBS 583.72A KF114798 T. minioluteus CBS 137.84 KM066175 T. ucrainicus CBS 583.72C KM066129 T. minioluteus CBS 270.35 KJ775734 T. atroroseus DTO 270-D5 KM066131 T. ucrainicus CBS 583.72C KJ775735 T. atroroseus DTO 270-D6 KF114773 T. ucrainicus CBS 127.64 T KF114747 T. atroroseus CBS 133442 KF114771 T. ucrainicus CBS 162.67T KJ775716 T. atroroseus DTO 267-I1 KM066130 T. ucrainicus CBS 583.72A T JN899350 T. udagawae CBS 579.72 KF114789 T. atroroseus CBS 133442T JN899346 T. minioluteus CBS 642.68T KJ775227 T. atroroseus DTO 270-D5 KM066171 T. minioluteus CBS 137.84 KJ775228 T. atroroseus DTO 270-D6 KM066172 T. minioluteus CBS 270.35 KJ775209 T. atroroseus DTO 267-I1 FJ160264 T. solicola CBS 133445T KJ775205 T. diversus DTO 244-E6 KF114730 T. solicola CBS 133446 KJ865723 T. diversus CBS 320.48T HQ605705 T. albobiverticillius CBS 133440T KJ775195 T. diversus DTO 133-E4 KF114733 T. albobiverticillius CBS 133441 KJ775193 T. diversus DTO 131-I6 JN899383 T. erythromellis CBS 644.80T KJ775700 T. diversus DTO 131-I6 KJ775194 T. diversus DTO 133-A7 KJ865732 T. austrocalifornicus CBS 644.95T KJ775701 T. diversus DTO 133-A7 KF114773 T. convolutus CBS 100537T KJ865740 T. diversus CBS 320.48T GU385731 T. solicola CBS 133445T KJ775702 T. diversus DTO 133-E4 KF114775 T. solicola CBS 133446 KJ775712 T. diversus DTO 244-E6 HQ156945 T. erythromellis CBS 644.80T T JN899357 T. austrocalifornicus CBS 644.95 JN899330 T. convolutus CBS 100537T T JN899372 T. purpurogenus CBS 286.36 KF114778 T. albobiverticillius CBS 133440T KF114777 T. albobiverticillius CBS 133441 JX315639 T. purpurogenus CBS 286.36T Fig. 5. ML phylogenies of ITS (left) and BenA (right) for species classiﬁed in Talaromyces sect. Trachyspermi. Talaromyces purpurogenus was chosen as out-group. Support in nodes is indicated above thick branches and is represented by bootstrap values of 80 % and higher. T = ex type. Model selected: T92+G+I for ITS, K2+G for BenA, alignment 495 (ITS) and 376 (BenA) bp. Accepted species: Talaromyces albobiverticillius (H.-M. Hsieh, Y.M. Ju & S.Y. Hsieh) Samson et al., Stud. Mycol. 71: 174. 2011. [MB560683]. — Herb.: BCRC 34774. Extype: CBS 133440 = DTO 166-E5 = YMJ 1292. ITS barcode: HQ605705. (Alternative markers; BenA = KF114778; CaM = KJ885258; RPB1 = n.a.; RPB2 = KM023310). Talaromyces assiutensis Samson & Abdel-Fattah, Persoonia 9: 501. 1978. [MB324414]. — Herb.: CBS 147.78. Ex-type: CBS 147.78. ITS barcode: JN899323. (Alternative markers; BenA = KJ865720; CaM = KJ885260; RPB1 = JN680275; RPB2 = KM023305). Talaromyces atroroseus Yilmaz et al., PLoS ONE 8: e84102-page 8. 2013. [MB804901]. — Herb.: CBS H-21790. Ex-type: CBS 133442 = IBT 32470 = DTO 178-A4. ITS barcode: KF114747. (Alternative markers; BenA = KF114789; CaM = KJ775418; RPB1 = n.a.; RPB2 = KM023288). Talaromyces austrocalifornicus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 245. 1993. [MB361182]. — Herb.: CBM-PF 1117. Ex-type: CBS 644.95 = IBT 17522. ITS barcode: JN899357. (Alternative markers; BenA = KJ865732; CaM = KJ885261; RPB1 = JN680316; RPB2 = n.a.). Talaromyces convolutus Udawaga, Mycotaxon 48: 141. 1993. [MB360474]. — Herb.: CBM SUM-3018. Ex-type: CBS 100537 = IBT 14989. ITS barcode: JN899330. (Alternative markers; BenA = KF114773; CaM = n.a.; RPB1 = JN121553; RPB2 = JN121414). Talaromyces diversus (Raper & Fennell) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. [MB560649]. — Herb.: IMI 040579. Ex-type: CBS 320.48 = ATCC 10437 = DSM 2212 = IMI 040579 = IMI 040579ii = NRRL 2121. ITS barcode: KJ865740. (Alternative markers; BenA = KJ865723; CaM = KJ885268; RPB1 = JN680297; RPB2 = KM023285). Talaromyces erythromellis (A.D. Hocking) Samson et al., Stud. Mycol. 71: 175. 2011. [MB560652]. — Herb.: IMI 216899. Ex-type: CBS 644.80 = FRR 1868 = IMI 216899. ITS barcode: JN899383. (Alternative markers; BenA = HQ156945; CaM = KJ885270; RPB1 = JN680315; RPB2 = KM023290). Talaromyces minioluteus (Dierckx) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560657]. — Herb.: CBS 642.68. Ex-type: CBS 642.68 = IMI 089377 = MUCL 28666. ITS barcode: JN899346. (Alternative markers; BenA = KF114799; CaM = KJ885273; RPB1 = JN121709; RPB2 = JF417443). Talaromyces solicola Visagie & K. Jacobs, Persoonia 28: 20. 2012. [MB564328]. — Herb.: PREM 60037. Ex-type: DAOM 241015 = CV 2800 = DTO 180-D4. ITS barcode: FJ160264. (Alternative markers; BenA = GU385731; CaM = KJ885279; RPB1 = n.a.; RPB2 = KM023295). www.studiesinmycology.org Talaromyces trachyspermus (Shear) Stolk & Samson, Stud. Mycol. 2: 32. 1972. [MB324421]. — Herb.: IMI 040043. Ex-type: CBS 373.48 = ATCC 10497 = IMI 040043 = NRRL 1028. ITS barcode: JN899354. (Alternative markers; BenA = KF114803; CaM = KJ885281; RPB1 = JN121664; RPB2 = JF417432). Talaromyces ucrainicus (Panas.) Udagawa, Trans. Mycol. Soc. Japan 7: 94. 1966. [MB449587]. — Herb.: unknown. Ex-type: CBS 162.67 = ATCC 22344 = FRR 3462 = NHL 6086. ITS barcode: JN899394. (Alternative markers; BenA = KF114771; CaM = KJ885282; RPB1 = JN680277; RPB2 = KM023289). Talaromyces udagawae Stolk & Samson, Stud. Mycol. 2: 36. 1972. [MB324424]. — Herb.: CBS H-7841. Ex-type: CBS 579.72 = FRR 1727 = IMI 197482. ITS barcode: JN899350. (Alternative markers; BenA = KF114796; CaM = n.a.; RPB1 = JN680310; RPB2 = n.a.). Talaromyces section Bacillispori Yilmaz, Frisvad & Samson, sect. nov. MycoBank MB809642. Figs 1, 6. Typus: Talaromyces bacillisporus (Swift) C.R. Benj., [as ‘bacillosporus’], Mycologia 47: 682. 1955. Description: Restricted growth on CYA, DG18, YES and CREA. Some species produce creamish white to yellow ascomata. Conidiophores mono- to biverticillate. The section is introduced for T. bacillisporus and its closely related species. Species typically produce creamish white to yellow ascomata and generally grow restrictedly on CYA, DG18, YES and CREA, with the exception of T. proteolyticus, which grows relatively well and lacks a sexual state. Accepted species: Talaromyces bacillisporus (Swift) C.R. Benj. [as ‘bacillosporus’], Mycologia 47: 682. 1955. [MB118745]. — Herb.: CBS H-7813 (Isotype). Ex-type: CBS 296.48 = ATCC 10126 = IMI 040045 = NRRL 1025. ITS barcode: KM066182. (Alternative markers; BenA = AY753368; CaM = KJ885262; RPB1 = JN121634; RPB2 = JF417425). Talaromyces emodensis Udagawa, Mycotaxon 48: 146. 1993. [MB360476]. — Herb.: CBM SUM-3025. Ex-type: CBS 100536 = IBT 14990. ITS barcode: 191 YILMAZ ET AL. JN899336 T. unicus CBS 100535T AB176620 T. hachijoensis IFM53624T KM066135 T. bacillisporus CBS 102389 JN899387 T. proteolyticus CBS 303.67T KM066136 T. bacillisporus CBS 110774 JN899337 T. emodensis CBS 100536T AY753368 T. bacillisporus CBS 296.48T JN899338 T. mimosinus CBS 659.80T KM066137 T. bacillisporus CBS 116927 KM066179 T. bacillisporus CBS 102389 KJ865726 T. mimosinus CBS 659.80T KM066180 T. bacillisporus CBS 110774 KJ865724 T. emodensis CBS 100536T KM066181 T. bacillisporus CBS 116927 KJ865735 T. unicus CBS 100535T KM066182 T. bacillisporus CBS 296.48T KJ865729 T. proteolyticus CBS 303.67T JN899374 T. rugulosus CBS 371.48T KF984575 T. rugulosus CBS 371.48T Fig. 6. ML phylogenies of ITS (left) and BenA (right) for species classiﬁed in Talaromyces sect. Bacillispori. Talaromyces rugulosus was chosen as out-group. Support in nodes is indicated above thick branches and is represented by bootstrap values of 80 % and higher. T = ex type. Model selected: T92+G for ITS, K2+G for BenA, alignment 457 (ITS) and 469 (BenA) bp. JN899337. (Alternative markers; BenA = KJ865724; CaM = KJ885269; RPB1 = JN121552; RPB2 = JN121552). Talaromyces hachijoensis Yaguchi, Someya & Udagawa, Mycoscience 37: 157. 1996. [MB416016]. — Herb.: PF 1174. Ex-type: PF 1174 = IFM 53624. ITS barcode: AB176620. (Alternative markers; BenA = n.a.; CaM = n.a.; RPB1 = n.a.; RPB2 = n.a.). Talaromyces mimosinus A.D. Hocking, The genus Penicillium: 507. 1980. [MB116382]. — Herb.: IMI 223991. Ex-type: CBS 659.80 = FRR 1875 = IMI 223991. ITS barcode: JN899338. (Alternative markers; BenA = KJ865726; CaM = KJ885272; RPB1 = JN899302; RPB2 = n.a.). Talaromyces proteolyticus (Kamyschko) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011. [MB560665]. — Herb.: CBS 303.67. Ex-type: CBS 303.67 = ATCC 18326 = NRRL 3378. ITS barcode: JN899387. (Alternative markers; BenA = KJ865729; CaM = KJ885276; RPB1 = JN680292; RPB2 = KM023301). Talaromyces unicus Tzean, J.L. Chen & S.H. Shiu, Mycologia 84: 739. 1992. [MB360172]. — Herb.: PPH 16; Holotype Nat. Taiwan Univ. Ex-type: CBS 100535 = CCRC 32703 = IBT 18385. ITS barcode: JN899336. (Alternative markers; BenA = KJ865735; CaM = KJ885283; RPB1 = JN680324; RPB2 = n.a.). Description: Restricted growth on general media. Colonies have prominent yellow mycelia. Most species grow at 37 C and some species produce yellow ascomata. The section is introduced for Penicillium sect. Simplicium ser. Islandica Pitt. Species typically produce restrictedly growing colonies and have prominent yellow mycelia. A four-gene phylogeny, morphology and extrolite data revealed that T. variabilis, P. concavorugulosum and T. sublevisporus are synonymous with T. wortmannii (Yilmaz et al. unpubl.). Section Islandici includes species, which are biotechnologically (e.g. T. variabilis and T. rugulosus) and medically important (e.g. T. piceus, T. columbinus and T. radicus) (Barthomeuf et al. 1991, Petruccioli et al. 1999, Horre et al. 2001, de Vos et al. 2009, Peterson & Jurjevic 2013). Accepted species: Talaromyces section Subinﬂati Yilmaz, Frisvad & Samson, sect. nov. MycoBank MB809564. Figs 1, 7. Typus: Talaromyces subinﬂatus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 249. 1993. Description: Poor growth on CREA and DG18. Sometimes indeterminate synnemata produced at colony margin after one to two weeks. The section is introduced for T. subinﬂatus and T. palmae. Morphologically, T. subinﬂatus and T. palmae do not resemble each other, although both grow poorly on CREA and DG18. Talaromyces palmae produces indeterminate synnemata after one to two weeks, while T. subinﬂatus grows poorly on all media except MEA and OA. Accepted species: Talaromyces palmae (Samson, Stolk & Frisvad) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560658]. — Herb.: CBS 442.88. Ex-type: CBS 442.88 = IMI 343640. ITS barcode: JN899396. (Alternative markers; BenA = HQ156947; CaM = KJ885291; RPB1 = JN680308; RPB2 = KM023300). Talaromyces subinﬂatus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 249. 1993. [MB361184]. — Herb.: CBM PF-1113. Ex-type: CBS 652.95 = IBT 17520. ITS barcode: JN899397. (Alternative markers; BenA = KJ865737; CaM = KJ885280; RPB1 = JN680301; RPB2 = KM023308). Talaromyces section Islandici (Pitt) Yilmaz, Frisvad & Samson, comb. et stat. nov. MycoBank MB809565. Figs 1, 7. Basionym: Penicillium sect. Simplicium ser. Islandica Pitt, The genus Pencillium: 445. 1980. Typus: Talaromyces islandicus (Sopp) Samson et al., Stud. Mycol. 71: 176. 2011. 192 Talaromyces allahabadensis (B.S. Mehrotra & D. Kumar) Samson et al., Stud. Mycol. 71: 174. 2011. [MB560640]. — Herb.: University of Allahabad P-26. Ex-type: CBS 453.93 = ATCC 15067 = CBS 304.63. ITS barcode: KF984873. (Alternative markers; BenA = KF984614; CaM = KF984768; RPB1 = JN680309; RPB2 = KF985006). Talaromyces atricola S.W. Peterson & Jurjevic, PLoS ONE 8: e78084-page 8. 2013. [MB804733]. — Herb.: unknown. Ex-type: CBS 255.31 = NRRL 1052 = FRR 1052 = Thom 4640.439 = ATCC 52257. ITS barcode: KF984859. (Alternative markers; BenA = KF984566; CaM = KF984719; RPB1 = n.a.; RPB2 = KF984948). Talaromyces brunneus (Udagawa) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. [MB560644]. — Herb.: NHL 6054. Ex-type: CBS 227.60 = ATCC 18229 = FRR 646 = IFO 6438 = IHEM 3907 = IMI 078259 = MUCL 31318. ITS barcode: JN899365. (Alternative markers; BenA = KJ865722; CaM = KJ885264; RPB1 = JN680281; RPB2 = KM023272). Talaromyces columbinus S.W. Peterson & Jurjevic, PLoS ONE 8: e78084-page 6. 2013. [MB804732]. — Herb.: BPI 892668. Ex-type: NRRL 58811. ITS barcode: KJ865739. (Alternative markers; BenA = KF196843; CaM = KJ885288; RPB1 = n.a.; RPB2 = KM023270). Talaromyces islandicus (Sopp) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560654]. — Herb.: IMI 040042. Ex-type: CBS 338.48 = ATCC 10127 = IMI 040042 = MUCL 31324 = NRRL 1036. ITS barcode: KF984885. (Alternative markers; BenA = KF984655; CaM = KF984780; RPB1 = JN121648; RPB2 = KF985018). Talaromyces loliensis (Pitt) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011. [MB560655]. — Herb.: IMI 216901. Ex-type: CBS 643.80 = ATCC 52252 = FRR 1798 = IMI 216901 = MUCL 31325. ITS barcode: KF984888. (Alternative markers; BenA = KF984658; CaM = KF984783; RPB1 = JN680314; RPB2 = KF985021). Talaromyces piceus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 176. 2011. [MB560661]. — Herb.: IMI 040038. Ex-type: CBS 361.48 = ATCC 10519 = IMI 040038 = NRRL 1051. ITS barcode: KF984792. (Alternative markers; BenA = KF984668; CaM = KF984680; RPB1 = n.a.; RPB2 = KF984899). Talaromyces radicus (A.D. Hocking & Whitelaw) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560669]. — Herb.: DAR 72374. Ex-type: CBS 100489 = FRR 4718. ITS barcode: KF984878. (Alternative markers; BenA = KF984599; CaM = KF984773; RPB1 = n.a.; RPB2 = KF985013). Talaromyces rotundus (Raper & Fennell) C.R. Benj., Mycologia 47: 683. 1955. [MB306719]. — Herb.: IMI 040589. Ex-type: CBS 369.48 = ATCC 10493 = IMI POLYPHASIC TAXONOMY OF TALAROMYCES KF984875 T. radicus CBS 137382 KF984599 T. radicus CBS 100489T KF984880 T. radicus DTO 181-D4 KF984600 T. radicus CBS 100490 KF984879 T. radicus CBS 100490 KF984598 T. radicus CBS 100488 KF984878 T. radicus CBS 100489T KF984602 T. radicus CBS 137382 KF984877 T. radicus CBS 100488 KF984601 T. radicus DTO 181-D4 KF984881 T. radicus DTO 181-D7 KF984603 T. radicus DTO 181-D7 KF984864 T. allahabadensis CBS 137397 KF984605 T. allahabadensis CBS 137397 KF984866 T. allahabadensis CBS 137399 KF984607 T. allahabadensis CBS 137399 KF984863 T. allahabadensis CBS 178.81 KF984614 T. allahabadensis CBS 453.93T KF984872 T. allahabadensis CBS 441.89 KF984612 T. allahabadensis CBS 178.81 KF984873 T. allahabadensis CBS 453.93T KF984613 T. allahabadensis CBS 441.89 JN899365 T. brunneus CBS 227.60T KF984651 T. wortmannii CBS 319.63 KF984887 T. loliensis CBS 172.91 KF984648 T. wortmannii CBS 391.48T KF984888 T. loliensis CBS 643.80T 98 KF984882 T. islandicus CBS 117284 KF984626 T. wortmannii CBS 895.73 KF984883 T. islandicus CBS 165.81 KF984635 T. wortmannii NRRL 2125 KF984885 T. islandicus CBS 338.48T KF984886 T. islandicus CBS 394.50 KF984632 T. wortmannii CBS 137376 KF984657 T. loliensis CBS 172.91 97 JN899361 T. tardifaciens CBS 250.94T KF984828 T. wortmannii CBS 319.63 KF196853 T. wortmannii CBS 385.48 KF984658 T. loliensis CBS 643.80T KJ865722 T. brunneus CBS 227.60T KF984829 T. wortmannii CBS 391.48T KF984652 T. islandicus CBS 117284 KF196915 T. wortmannii CBS 385.48 KF984653 T. islandicus CBS 165.81 KF984797 T. wortmannii NRRL 2125 KF984655 T. islandicus CBS 338.48T KF984800 T. wortmannii CBS 137376 KF984656 T. islandicus CBS 394.50 KF984811 T. wortmannii CBS 895.73 KF984577 T. rugulosus NRRL 1053 JN899353 T. rotundus CBS 369.48T KF984574 T. rugulosus CBS 344.51 KF984895 T. scorteus CBS 233.60 99 KF984896 T. scorteus CBS 500.75 KF984894 T. scorteus CBS 499.75 KF984893 T. scorteus DTO 270-A6 KF984892 T. scorteus CBS 340.34T KF984572 T. rugulosus CBS 137366 KF984575 T. rugulosus CBS 371.48T KF984579 T. rugulosus NRRL 1073 KF984566 T. atricola CBS 255.31T KF984561 T. scorteus DTO 270-A6 KJ775717 T. yelensis CBS 138209T KF984562 T. scorteus CBS 233.60 KJ775719 T. yelensis CBS 138210 KF984564 T. scorteus CBS 500.75 KF984891 T. tratensis CBS 133146T KF984563 T. scorteus CBS 499.75 KF984889 T. tratensis CBS 137400 KF984565 T. scorteus CBS 340.34T KF984890 T. tratensis CBS 137401 KJ865730 T. rotundus CBS 369.48T KF984889 T. tratensis DTO 270-F5 KF984859 T. atricola CBS 255.31T KF984832 T. rugulosus NRRL 1073 KJ775210 T. yelensis CBS 138209T KJ775212 T. yelensis CBS 138210 KF984559 T. tratensis CBS 133146T KF984858 T. rugulosus CBS 344.51 KF984557 T. tratensis CBS 137400 KF984850 T. rugulosus CBS 137366 KF984558 T. tratensis CBS 137401 KF984834 T. rugulosus CBS 371.48T KF984557 T. tratensis DTO 270-F5 KF984848 T. rugulosus NRRL 1053 KC202954 T. tardifaciens CBS 250.94T KF196899 T. columbinus NRRL 58644 KF196843 T. columbinus NRRL 58811T KJ865739 T. columbinus NRRL 58811T KF196844 T. columbinus NRRL 62680 KF196901 T. columbinus NRRL 62680 KF196842 T. columbinus NRRL 58644 KF984794 T. columbinus CBS 137393 KF984659 T. columbinus CBS 137393 KF984788 T. piceus CBS 116872 KF984660 T. piceus CBS 116872 KF984789 T. piceus CBS 132063 KF984664 T. piceus CBS 137363 KF984787 T. piceus CBS 137363 KF984665 T. piceus CBS 132063 KF984784 T. piceus CBS 137377 KF984661 T. piceus CBS 137377 T KF984668 T. piceus CBS 361.48T KF984792 T. piceus CBS 361.48 JN899396 T. palmae CBS 442.88T JN899397 T. subinflatus CBS 652.95T HQ156947 T. palmae CBS 442.88T KJ865737 T. subinflatus CBS 652.95T Fig. 7. ML phylogenies of ITS (left) and BenA (right) for species classiﬁed in Talaromyces sect. Islandici. Talaromyces palmae and T. subinﬂatus (Talaromyces sect. Subinﬂati) were chosen as out-group. Support in nodes is indicated above thick branches and is represented by bootstrap values of 80 % and higher. T = ex type. Model selected: GTR+G for ITS, K2+G for BenA, alignment 584 (ITS) and 398 (BenA) bp. 040589 = NRRL 2107. ITS barcode: JN899353. (Alternative markers; BenA = KJ865730; CaM = KJ885278; RPB1 = n.a.; RPB2 = KM023275). Talaromyces rugulosus (Thom) Samson et al., Stud. Mycol. 71: 177. 2011. [MB560672]. — Herb.: IMI 040041. Ex-type: CBS 371.48 = ATCC 10128 = IMI 040041 = MUCL 31201 = NRRL 1045. ITS barcode: KF984834. (Alternative markers; BenA = KF984575; CaM = KF984702; RPB1 = JN680302; RPB2 = KF984925). Talaromyces scorteus (Nakazawa, Takeda & Suematsu) S.W. Peterson & Jurjevic PLoS ONE 8: e78084-page 8. 2013. [MB492647]. — Herb.: unknown. Ex-type: CBS 340.34 = NRRL 1129 = FRR 1129. ITS barcode: KF984892. (Alternative markers; BenA = KF984565; CaM = KF984684; RPB1 = n.a.; RPB2 = KF984916). Talaromyces tardifaciens Udagawa, Mycotaxon 48: 150. 1993. [MB360478]. — Herb.: CBM SUM 3017. Ex-type: CBS 250.94. ITS barcode: JN899361. (Alternative markers; BenA = KC202954; CaM = KF984682; RPB1 = JN680283; RPB2 = KF984908). Talaromyces tratensis Manoch, Dethoup & Yilmaz, Mycoscience 54: 337. 2013. [MB801738]. — Herb.: CBS H-21074. Ex-type: CBS 133146 = KUFC 3383. www.studiesinmycology.org ITS barcode: KF984891. (Alternative markers; BenA = KF984559; CaM = KF984690; RPB1 = JX898042; RPB2 = KF984911). Talaromyces wortmannii (Klöcker) C.R. Benj., Mycologia 47: 683. 1955. [MB344294]. — Herb.: IMI 040047. Ex-type: CBS 391.48 = ATCC 10517 = IMI 040047 = NRRL 1017. ITS barcode: KF984829. (Alternative markers; BenA = KF984648; CaM = KF984756; RPB1 = JN121669; RPB2 = KF984977). Talaromyces yelensis Visagie et al., Stud. Mycol. 78: 134. 2014. [MB809189]. — Herb.: CBS H-21799. Ex-type: CBS 138209 = DTO 268-E5. ITS barcode: KJ775717. (Alternative markers; BenA = KJ775210; CaM = n.a.; RPB1 = n.a.; RPB2 = n.a.). Phylogenetic species recognition The ITS rDNA region was recently accepted as the ofﬁcial DNA barcode for fungi (Schoch et al. 2012). It was chosen because of the universal primer sets available for PCR ampliﬁcation and is 193 YILMAZ ET AL. the most widely sequenced gene for fungi. The barcode does have its limitations and is often not sufﬁciently variable for making a species identiﬁcation. For example, in section Talaromyces (Fig. 2), T. siamensis and T. cnidii have identical ITS sequences, while there is lack of support in most branches of the phylogeny. Another example is T. assiutensis and T. trachyspermus, classiﬁed in section Trachyspermi (Fig. 5), which also have identical ITS sequences. Overall ITS does, however, perform very well for species recognition in the genus, even though it should be used cautiously since variability is low in a few clades. Previous studies on Penicillium and Talaromyces showed that BenA sequences have more variation compared to ITS (Seifert & Louis-Seize 2000, Samson et al. 2004, Visagie et al. 2009, Visagie & Jacobs 2012, Yilmaz et al. 2012, Frisvad et al. 2013). In our study, BenA easily distinguished between all species, also allowing for some infraspecies variation. The ITS and BenA phylogenies (Figs 2–7) cover the variation accepted in each species. Peterson & Jurjevic (2013) mentioned problems with the ampliﬁcation of BenA paralogs in section Islandici using primer pairs Bt2a and Bt2b or Bt2f and T22. We had similar results with these primers, with gel-electrophoresis showing one band after ampliﬁcation, but subsequent sequences that have mixed electropherograms. As a result, we suggest primer pair T10 and Bt2b (Glass & Donaldson 1995), at annealing temperatures of 50 or 52 C, for the ampliﬁcation and sequencing of BenA in section Islandici. Apart from ITS and BenA, several other genes have been used for phylogenetic comparisons in Talaromyces, including calmodulin, RPB1 and RPB2 (Yilmaz et al. 2012, Frisvad et al. 2013). These genes work well for species recognition, but ampliﬁcation of these genes are often challenging. For example, RPB1 and RPB2 ampliﬁcation is difﬁcult across all sections of Talaromyces, while ampliﬁcation of calmodulin often is difﬁcult in section Trachyspermi using primer pairs CMD5 and CMD6 or CF1 and CF4. As a result, we propose the use of BenA as the secondary molecular marker for species identiﬁcations in Talaromyces. Morphological species recognition In aid of morphological species identiﬁcation, tables (Table 3–9) are provided summarising the most important characters. Tables 3 & 4 contains all Talaromyces species and summarises general macro- (Table 3) and micromorphological characters (Table 4). Additional tables are provided for ascomata producers (Table 5), synnemata producers (Table 6), red soluble pigment producers (Table 7) and species that produce globose rough-walled conidia (Table 8). In Table 9 growth rates on CYA at 37 C are given. Table 3 provides a summary of the most important macromorphological characters, including growth rates on CYA, MEA and CREA, reverse and soluble pigment colour on CYA, colony texture on MEA and acid production on CREA. Growth rates on especially CYA and MEA were taxonomically informative. For example, section Talaromyces species generally grow well on these media, species from section Trachyspermi grow poorly on CYA but well on MEA, while section Islandici species grow restrictedly on both CYA and MEA. Lack of and/or poor growth on CREA is characteristic of sections Trachyspermi and Subinﬂati. Acid produced on CREA is able to distinguish among closely related species. For example, T. allahadensis and T. radicus are 194 closely related, but T. allahabadensis produces acid in contrast to T. radicus, which lacks acid production. Species of medical importance includes T. amestolkiae, T. marneffei, T. helicus, T. piceus and T. stollii (Chiang et al. 1998, Horre et al. 2001, Tomlinson et al. 2011, Yilmaz et al. 2012), which are able to grow at 37 C. As a result, Table 9 sorts species based on growth rates on CYA at 37 C. Reverse colouration on CYA was taxonomically informative. For example, Talaromyces bacillisporus (especially at 30 and 37 C), T. derxii, T. euchlorocarpius, T. viridis and T. aerugineus all produce green CYA reverses. Colony texture on MEA was also useful for distinguishing species. For example, T. ruber produces velvety colonies compared to the loosely funiculose colonies of its closest relative T. amestolkiae, and T. rugulosus produces velvety colonies in contrast to its closest relative T. atricola, which has ﬂoccose colonies. Micromorphological characters are summarised in Table 4 and illustrated in Fig. 8. A diverse range of conidiophore branching patterns are observed in Talaromyces and range from having solitary phialides (Fig. 8A) to monoverticillate (Fig. 8B), biverticillate (Fig. 8C) and biverticillate conidiophores with additional subterminal branches (Fig. 8D, E). Phialide shape is a useful feature with most Talaromyces species having acerose phialides (Fig. 8F), while a small number of species have ﬂaskshaped phialides (Fig. 8G). Conidial sizes and ornamentations were taxonomically informative characters. Talaromyces bacillisporus and T. viridulus produces rod-shaped conidia (Fig. 8H), a small group of species produces roughened globose conidia (Fig. 8I), but most species produce ellipsoidal conidia that are ﬁnely roughened (Fig. 8J, K). Ascomata are produced in a number of Talaromyces species and these are included in Table 5. Generally, ascomata are yellow, but some species produce green (T. derxii, T. euchlorocarpius and T. viridis) or creamish white ascomata (T. assiutensis and T. trachyspermus). Size, shape and ornamentation of ascospores are also useful for distinguishing between species. Ascospores are generally spiny. Ascospores of T. stipitatus have single equatorial ridges, T. udagawae has numerous ornamented ridges on ascospores and T. helicus has smooth ascospores. In most species, ascospores are broadly ellipsoidal, but T. bacillisporus and T. rotundus have spiny, globose ascospores and T. tardifaciens produces smooth globose ascospores. A number of species typically produces synnemata and is included in Table 6. Synnemata can be either determinate or indeterminate. Determinate synnemata have a terminal, nonelongated conidiogenous zone, in contrast to indeterminate synnemata, which have elongated fertile zones, sometimes covering the whole conidioma (Seifert 1985). Mostly, synnemata are produced after more than one week of incubation. However, T. panamensis and T. duclauxii produce them within one week. The length of synnemata is an important character. In T. ramulosus and T. ﬂavovirens, synnemata are characteristically short (<1000 μm), whereas T. pseudostromaticus and T. calidicanius produce very tall synnemata (>6000 μm). Other characters distinguishing between synnemata producers, include growth rate on CYA and MEA at 25 and 30 C and acid production on CREA. Although red soluble pigment is not always consistently produced in strains from the same species, species such as T. albobiverticillius, T. atroroseus, T. marneffei, T. minioluteus and T. purpurogenus typically produce these pigments. Table 7 Table 3. Overview of macromorphological characters for the identiﬁcation of Talaromyces species. www.studiesinmycology.org Species name Ascomata Growth rate (mm) CYA reverse CYA soluble pigment MEA colony texture Acid production on CREA Light brown (7D4–7D5) at the centre fading into greyish yellow (4A3–4B3) Dark green (26F5) centre fading into greyish green (26B5–26B5) Greyish red to reddish brown (9C5–9F8) Orange to greyish orange (5A6–5B6) in the centre fading into light yellow to greyish yellow (4A5–4B5) Violet brown (11E8–11F8) Absent Floccose to funiculose Weak Absent Loosely funiculose to velvety Absent (no growth) Red (in some isolates absent) Velvety Absent Absent Velvety Present Very weak brownish red, in some strains absent Absent Floccose and overlaying funiculose Floccose Weak Absent (in some isolates very weak) Absent Floccose to strongly funiculose Weak Absent Floccose Absent Absent Floccose Absent CYA CYA (37 C) MEA 15–20 30–35 T. aerugineus Absent 26–27 35 49–50 T. albobiverticillius Absent 15–20 No growth 25–28 T. allahabadensis Absent 20–25 23–25 20–23 T. amestolkiae Absent 30–32 8–15 30–45 T. angelicus Absent 25–27 25–27 33–35 T. apiculatus Absent 38–41 25–35 40–42 T. assiutensis Present 15–21 25–30 20–23 T. atricola Absent 10 No growth 15 T. atroroseus Absent 28–40 23–25 30–35 T. aurantiacus Absent 30–32 19–21 38–40 T. austrocalifornicus Present 10–11 10–11 16–18 T. bacillisporus Present 10–15 33–37 15–20 T. bohemicus Absent Unknown Unknown T. boninensis Present 28 Unknown 40 (after 10 d) 30 T. brunneus Absent 19–20 No growth 17–19 T. calidicanius Absent 27–30 No growth 47–48 T. cecidicola Absent 33–34 2–3 37–38 T. chloroloma Absent 40–45 0–4 45–48 T. cinnabarinus Present Unknown Unknown Unknown T. cnidii Absent 25–30 20–28 38–40 T. coalescens Absent 32–34 2–4 43–45 Greyish red (7B3) centre fading into yellowish white (4A2) Greyish yellow (3B3) to yellowish white (3A2) Pale orange to brownish orange (5A3–5C5) Yellowish white to yellowish grey (4A2–4A3) Reddish brown (9E8) Red Velvety and ﬂoccose Absent Absent Floccose Absent Absent Floccose Absent (no growth) Absent Floccose Absent Unknown Floccose and loosely funiculose Unknown Absent Velvety and ﬂoccose Unknown Absent Velvety and ﬂoccose Present Absent Synnematous Present Absent Funiculose Absent Centre greyish orange (5B3), fading into dull greyish yellow (3B3) Unknown Absent Floccose and loosely funiculose Moderate Unknown Unknown Unknown Violet brown (11E8) fading into vivid red (11A8) Dark brown to violet brown (8F8–10F8) Red to yellow Velvety and ﬂoccose Absent Red Funiculose Very weak Pastel red (7A4) fading into orange white (6A2) Orange white to pale orange (5A2–5A3) Dull green (28F4) fading into greyish yellow to olive brown (4C5–4D5) (at 30 and 37 C reverse dark green (25F4)) Unknown Yellowish brown (5E5) centre fading into golden yellow (5B7) Yellowish brown (5E5) centre fading into golden yellow (5B7) Greyish yellow to olive brown (4C5–4D5) Dark brown to violet brown (8F8–10F8) 195 (continued on next page) TALAROMYCES 25–35 TAXONOMY OF Absent POLYPHASIC T. aculeatus Species name Ascomata Growth rate (mm) CYA soluble pigment MEA colony texture Acid production on CREA ET AL. CYA CYA reverse CYA (37 C) MEA T. columbinus Absent 13–14 42–43 20–21 Dark brown fading into beige (8F8) Light brownish orange Velvety and ﬂoccose Absent T. convolutus Present 10–12 6–8 13–15 Orange white to pale orange (5A2–5A3) Absent Floccose Absent T. dendriticus Absent 23–26 5–6 35–36 Absent Funiculose and synnematous Absent T. derxii Present 38–40 35–38 48–50 Dark brown (6F6), fading into light brown (6D6) and light yellow (2A5) Dark green (25F5) centre for both fading into light yellow (4A4) Blond to brownish orange (4C4–5C4) Absent Floccose Absent T. diversus Absent 7–10 5–8 25–35 T. duclauxii Absent 25–27 3–4 48–50 T. emodensis Present 8–10 4–6 8–10 T. erythromellis Absent 10–11 No growth 10–11 T. euchlorocarpius Present 15–18 No growth 38–40 T. ﬂavovirens Present 19–20 5–6 37–38 T. ﬂavus Present 9–10 19–20 31–32 T. funiculosus Absent (30–)38–45 38–50 30–45 T. galapagensis Present 15–17 25–26 25–28 T. hachijoensis Present 3 No growth 8–10 T. helicus Present 13–23 10–18 25–33 T. indigoticus Present 20–21 20–22 32–33 T. intermedius Present 15–16 No growth 48–50 T. islandicus Absent 20–27 8–17 21–26 T. liani Present 20–30 20–25 35–45 T. loliensis Absent 10–13 No growth 13–15 T. macrosporus Present 22–28 28–35 40–50 T. marneffei Absent 13–25 5–10 15–27 T. mimosinus Present 12–15 3–5 13–14 YILMAZ 196 Table 3. (Continued) Olive brown (4E8) fading into olive brown (4D7) and maize yellow (4A6) Brownish red to reddish brown (8C8–8D8) Dark red to brownish red (10C8–10D8) Reddish golden (6C7) centre fading into golden yellow (5B7) and deep yellow (4A8) Brown (7E6) centre fading into greyish yellow (2B6) Brownish orange (5C6) fading into light yellow (3A5) Light orange to greyish orange (5A4–5B4) Dark brown (6F6) centre fading into yellowish white (3A2) Brownish orange (5C6) Yellowish brown (5E4) centre fading into greyish orange (5B3) and yellowish white (4A2) Pale orange (5A3) centre fading into yellowish white (4A2) Greyish yellow (3B4) Reverse colouration brown (7E4) in the centre fading into light yellow to light orange (4A4–5A4) Light orange and light yellow (5A5–4A5) Centre deep yellow to deep orange (4A8–5A8) fading into light yellow to yellow (3A5–3A6) Yellowish brown (5E7–5F7) centre fading into brownish yellow to golden yellow (5C7–5B7) Reddish brown (9E8) fading into red (9A6) Light yellow (3A5) Absent Velvety Absent (no growth) Yellow Synnematous Weak Absent Floccose Absent Absent Loosely funiculose Absent Yellow to orange yellow Loosely funiculose and ﬂoccose Absent Yellow Velvety after 2 wk, covered with yellow mycelia Floccose Absent Absent Stringly funiculose Present Floccose Absent (no growth) Yellow Absent (in some isolates light red) Absent Unknown Velvety to ﬂoccose Unknown Absent Floccose Absent (no growth) Absent Floccose Absent Absent Floccose Absent (no growth) Absent Velvety and lossely funiculose Present Absent (in some isolates yellow) Velvety and ﬂoccose Absent (in some isolates very weak) Weak yellow Floccose and loosely funiculose Absent to very weak Absent (at 30 and 37 C brownish soluble pigment) Floccose Absent Red Loosely funiculose and ﬂoccose Absent Absent Floccose Absent (no growth) Table 3. (Continued) www.studiesinmycology.org Species name Ascomata Growth rate (mm) CYA reverse CYA soluble pigment MEA colony texture Acid production on CREA Brown (6E8) centre fading into brownish orange to light brown (6C8–6D8) Reddish brown (8E7) fading into brownish red (7C7–8C7) Greyish green (29B6–29C6) Weak brownish red Loosely funiculose and ﬂoccose Absent Absent Floccose Absent (no growth) Yellow Velvety and ﬂoccose Absent Absent Weak Present Unknown CYA CYA (37 C) MEA T. minioluteus Absent 17–18 No growth 21–22 T. muroii Present 15–16 19–20 30–32 T. oumae-annae Absent 16–18 10–11 29–30 T. palmae Absent 20–25 No growth 22–26 T. panamensis Absent 23–24 No growth 28–30 T. paucisporus Present 15–18 (after 14 d) No growth T. piceus Absent 20–27 30–35 19–20 (after 14 d) 25–27 T. pinophilus Absent 18–25 25–40 30–40 T. pittii Absent 34–36 No growth 42–44 T. primulinus Absent 5–6 No growth 20–25 T. proteolyticus Absent 20–22 No growth 20–21 T. pseudostromaticus Absent 25–34 No growth 38–43 T. ptychoconidium Absent 8–16 5–12 12–23 T. purpureus Present 2–4 No growth 15–16 Pastel yellow to greyish yellow (2A4–2B4) Brownish orange to yellowish brown (5C6–5D6) Violet brown (11E8) Absent Velvety and in the margins synnematous Synnematous Violet brown Velvety to tomentose Brown (6E6) Light brown Loosely funiculose to ﬂoccose Absent Greyish orange to orange (6B6–6B7) centre fading into pastel yellow (3A4) Reddish brown (8E7) at centre, fading into brownish red (8C6) Some isolates brownish orange to reddish brown (7C5–8C5) and in some isolates greenish grey to greyish green (1D2–1D3) Centre greyish red (9B5) fading into between light yellow (4A4) and light orange (5A4) Dark brown (7F8–8F8) Yellow, in some strains absent Loosely funiculose to ﬂoccose Moderate Red Yeast like slimy colonies, after 2 wk ﬂoccose and synnematous Floccose after 2 wk covered with yellow mycelia Absent Absent Absent Absent Floccose Moderate Absent Absent Absent (no growth) Absent (no growth) Centre light brown (5D4) fading into light yellow (3A5) Creamish beige Absent Velvety and loosely funiculose, after 2 wk synnematous Loosely funiculose Absent Floccose 20–25 16–25 30–45 Dark brown to violet brown (9F8–11F8) Red Velvety and ﬂoccose Absent 5–6 3 14–15 Pigmentation yellowish white (4A2) Absent Floccose Absent (no growth) T. radicus Absent 15–22 25–30 15–25 Yellowish brown (5E6–5F6) Absent Loosely funiculose to ﬂoccose Absent T. ramulosus Absent 32–40 5–8 45–48 Dark brown (6F4–6F7) Absent Funiculose and synnematous Absent T. rotundus Present 9–11 No growth 15–17 Absent Floccose Absent (no growth) T. ruber Absent 20–35 14–18 35–40 Greyish green (27E5) circle at centre fading into greenish grey (1B2 Brownish red (8E8–8F8) Velvety Absent T. rubicundus Absent 30–32 34–35 38–39 Very weak red, in some strains absent Absent Floccose Absent T. rugulosus Absent 15–17 No growth 17–20 Absent Velvety Absent to very weak 197 (continued on next page) TALAROMYCES Brownish orange (5C5) centre fading into orange white (5A2) Yellowish brown (5E4–5F4) fading into light yellow to greyish yellow (2A5–2B5 to 1A5–1B5) TAXONOMY OF Absent Absent POLYPHASIC T. purpurogenus T. rademirici Ascomata Growth rate (mm) CYA reverse CYA soluble pigment MEA colony texture Acid production on CREA CYA CYA (37 C) MEA T. ryukyuensis Present Unknown Unknown Unknown Unknown Unknown Unknown Unknown T. sayulitensis Absent 24–29 32–40 37–40 Absent Loosely funiculose to ﬂoccose Present T. scorteus Absent 8–16 No growth 10–15 Absent Velvety Absent T. siamensis Absent 20–22 15 32–33 Brown (6E6) in the centre, fading into brownish orange (6C7) and light yellow (4A5) Centre olive (2E4) centre fading into greyish yellow to olive (2C4–2D4) Blond to greyish yellow (4C4) Absent Velvety Absent T. solicola Absent 12–13 No growth 22–23 Absent Velvety and ﬂoccose Absent (no growth) T. stipitatus Present 32–38 28–32 45–48 T. stollii Absent 38–45 25–35 Yellow to orangish yellow Floccose Weak 45–50 Reddish brown to dark brown (9E6–9F6) Dark brown (6F6), fading into reddish golden brownish orange to light brown (6C7–6D7) Brownish red (8E8–8F8) Red, in some strains absent Floccose and lossely funiculose Present T. subinﬂatus Present 3–4 No growth 14–15 Yellowish white (4A2) Absent Floccose Absent (no growth) T. tardifaciens Present 9–10 No growth 13–15 Absent Floccose Absent (no growth) T. thailandensis Present 33–35 No growth 30–35 Absent Floccose Absent T. trachyspermus Present 13–24 32–40 17–25 Absent Floccose Absent (no growth) T. tratensis Present 10–12 No growth 15–20 Absent Present 10–20 7–17 18–24 Yellow and in some isolates absent Absent Loosely funiculose to ﬂoccose T. ucrainicus Floccose Absent (no growth) T. udagawae Present 6–8 No growth 10–11 Centre light orange (5A5–5A6) fading into greyish yellow (4C4) Yellowish red to red (8A6–9A6) centre with red (9B7) dots Light orange (5A5) fading into light yellow to yellow (3A5–3A6) Greyish yellow (4A5–4A6) to brownish orange (6C8) Olive to olive brown to brownish orange (1F6–4E5–5C5) Pale orange to light orange (5A3–5A4) Absent Floccose Absent (no growth) T. unicus Present 10.5–16 No growth 15–19.5 Pale red to reddish brown Floccose to funiculose Unknown T. varians Absent 22–24 27–28 30–32 Absent Floccose Absent T. verruculosus Absent 32–35 25–26 35–36 Absent Floccose and loosely funiculose Weak T. viridis Present 9 10–11 15 Absent Floccose Absent (no growth) T. viridulus Absent 12–15 6–7 37–38 Absent Floccose Absent T. wortmannii Present 18–28 0–7 15–25 Absent Velvety Absent to moderate T. yelensis Absent 20–22 25–26 15–16 Absent Floccose Absent Reddish white to violet brown (7A2–11E8) Dark green (27F6) centre fading into dull green (7D3) fading into pale orange to light orange (5A3–5A4) Greyish yellow to greyish orange (4C4–5C4) Greyish green (1C3) centre fading into yellowish white (4A2) Brownish red (9C7) centre fading into pale yellow (3A3) Reddish yellow (4A6) to greyish yellow (3B4–3C4) Yellowish white (2A2) to light yellow (3A5) to brown (5F6) ET AL. Species name YILMAZ 198 Table 3. (Continued) Table 4. Overview of micromorphological characters for the identiﬁcation of Talaromyces species. www.studiesinmycology.org Species name Conidiophore branching Conidial ornamentation Conidial shape Conidial size (μm) T. aculeatus Biverticillate, minor proportion with subterminal branches Rough to echinulate Globose 3–3.5 × 3–3.5 T. aerugineus Solitary phialides to monoverticillate Smooth Subglobose to ellipsoidal to fusiform 3–8.5 × 2.5–5 T. albobiverticillius Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Globose to subglobose 2–3.5(–4) × 1.5–2.5 T. allahabadensis Biverticillate Smooth Ellipsoidal to fusiform 2.5–4.5 × 1.7–2.5 T. amestolkiae Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Ellipsoidal 2–3 × 1.5–2.5 T. angelicus Mono- to biverticillate Finely rough Subglobose to ellipsoidal 2.5–3.5 × 2–3 T. apiculatus Biverticillate Rough to echinulate Globose 3–4(–5.5) × 3–4(–5) T. assiutensis Mono- to biverticillate Smooth Ovoidal to ellipsoidal 2–4 × 1.5–2.5 T. atricola Biverticillate having symmetrical subterminal branches Smooth Ellipsoidal to fusiform 2–5 × 2–5 T. atroroseus Biverticillate, minor proportion with subterminal branches Finely rough to rough Ellipsoidal 2–3.5 × 1.5–2.5 T. aurantiacus Biverticillate Smooth Cylindirical to ellipsoidal 3–5 × 1.5–2.5 T. austrocalifornicus Biverticillate Smooth Subglobose 1.5–3 × 1.5–2.5 T. bacillisporus Mono- to biverticillate Smooth Cylindrical, rod shaped to ellipsoidal 3–5(–6.5) × 1–2 T. bohemicus Monoverticillate, subterminal branching sometimes present Encrusted cell walls Fusiform 7–9 × 2.6–3 T. boninensis Biverticillate Smooth Ellipsoidal to fusiform 2–4 × 1.5–2.5 T. brunneus Biverticillate, minor proportion with subterminal branches Smooth Globose to subglobose 3–4(–7) × 2–4 T. calidicanius Biverticillate, minor proportion with subterminal branches Finely rough to rough with spiral striations Ellipsoidal to fusiform 2.5–4.5 × 2–3 T. cecidicola Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Ellipsoidal 2.5–4 × 1.5–3 T. chloroloma Biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal 2.5–4(–6) × 1.5–2.5 T. cinnabarinus Biverticillate Smooth Cylindirical 5.4–7.2 × 1.5–2 T. cnidii Biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal 2.5–4 × 2–2.5 Biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal to fusiform 2.5–3.5(–5.5) × 1.5–3 Biverticillate Smooth Ellipsoidal 2.5–3.5 × 3–4.5 T. convolutus Mono- to biverticillate Smooth Ellipsoidal (2–)3–4 × 1.5–2(–3) T. dendriticus Biverticillate Smooth Subglobose to ellipsoidal 2.5–3.5 × 2–2.5 T. derxii Mono- to biverticillate Smooth Cylindrical to ellipsoidal, sometimes curved and fusiform 4–8 × 1.5–3 T. diversus Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Subglobose to ellipsoidal 2–3(–5) × 2–3(–3.5) Smooth to ﬁnely rough Ellipsoidal 3–4 × 1.5–3.5(–4) Smooth Ovoidal to ellipsoidal 3–4 × 1.5–3 T. erythromellis Biverticillate having symmetrical subterminal branches Smooth Subglobose to ellipsoidal 2–3.5 × 1.5–2.5 T. euchlorocarpius Biverticillate Smooth Subglobose to ellipsoidal 2–4 × 2–3 T. ﬂavovirens Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Ellipsoidal 2.5–3.5(–5.5) × 2–2.5(–4) 199 (continued on next page) TALAROMYCES Biverticillate Monoverticillate, minor proportion with subterminal branches TAXONOMY OF T. duclauxii T. emodensis POLYPHASIC T. coalescens T. columbinus Conidiophore branching Conidial ornamentation Conidial shape Conidial size (μm) T. ﬂavus Monoverticillate Smooth Ellipsoidal 2–3 × 1.5–2.5 T. funiculosus Biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal 2–3(–5.5) × 1–2(–2.5) T. galapagensis Mono- to biverticillate Smooth Ovoidal to ellipsoidal 2.5–5 × 1.5–3 T. hachijoensis Absent Absent Absent Absent T. helicus Mono- to biverticillate Smooth Globose to subglobose 2.5–3.5(–4.5) × 2.2–3.5 T. indigoticus Mono- to biverticillate Smooth Ovoidal to ellipsoidal 2.4–4 × 2–3.2 T. intermedius Solitary phialides to monoverticillate Smooth Ellipsoidal 2.5–4.5 × 2.2–3.5 T. islandicus Biverticillate Smooth Ellipsoidal 2.5–6 × 2–4.5 T. liani Mono- to biverticillate Smooth Ellipsoidal 2.5–4(–4.5) × 2–3.5 T. loliensis Biverticillate Smooth, thick walled Subglobose to ellipsoidal 3–5 × 2.4–3.5 T. macrosporus Mono- to biverticillate Smooth Subglobose to ellipsoidal 2–4 × 2–3.5 T. marneffei Mono- to biverticillate Smooth Subglobose 2.5–4 × 2–3 T. mimosinus Mono- to biverticillate Smooth Globose to subglobose 2–3 × 2–2.5 T. minioluteus Biverticillate Smooth Ellipsoidal 2.5–4 × 1.5–2.5 T. muroii Mono- to biverticillate, minor proportion with subterminal branches Smooth Globose to subglobose 2–3 × 1.5–3 T. oumae-annae Biverticillate, minor proportion with subterminal branches Rough Ellipsoidal 3–3.5 × 2.5–3 T. palmae Biverticillate, minor proportion with subterminal branches Smooth Subglobose to ellipsoidal 3–4.5 × 2–3.5 T. panamensis Biverticillate having subterminal branches Smooth, thick walled Ellipsoidal to fusiform 3–5 × 2–3 T. paucisporus Absent Absent Absent Absent T. piceus Biverticillate Smooth Ellipsoidal 2–3.5 × 2–4 T. pinophilus Biverticillate, minor proportion with subterminal branches Smooth Globose to subglobose 2–3 × 2–3 T. pittii Mono- to biverticillate Smooth Ellipsoidal 2.5–4.5(–6.5) × 1.5–3.5 T. primulinus Biverticillate Smooth to ﬁnely rough Ellipsoidal to fusiform 2–4 × 1.5–3 T. proteolyticus Biverticillate, minor proportion with subterminal branches Smooth Globose to subglobose 2–3 × 1.5–2.5 T. pseudostromaticus Biverticillate, minor proportion with subterminal branches Smooth Subglobose to ellipsoidal 2.5–4 × 2–3 T. ptychoconidium Biverticillate Rough with spiral ridges Ellipsoidal 3–4.5(–5) × 2–3 T. purpureus Solitary phialides and monoverticillate Rough with spiral ridges Subglobose to ellipsoidal 3–4 × 2–3 T. purpurogenus Biverticillate Smooth Ellipsoidal 3–3.5 × 2–2.5 T. rademirici Mono- to biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal 2.5–4 × 1.5–2.5 T. radicus Biverticillate Finely rough with ridges Subglobose to ellipsoidal 2–3 × 2–2.5 T. ramulosus Biverticillate, minor proportion with subterminal branches Smooth Subglobose to ellipsoidal 2–3 × 1.5–2.5 T. rotundus Monoverticillate Smooth Ellipsoidal to fusiform 3–5(–6.5) × 1.5–2.5 T. ruber Biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal 2.5–3.5 × 1.5–2 ET AL. Species name YILMAZ 200 Table 4. (Continued) Table 4. (Continued) www.studiesinmycology.org Species name Conidiophore branching Conidial ornamentation Conidial shape Conidial size (μm) T. rubicundus Biverticillate, minor proportion with subterminal branches Smooth Subglobose to ellipsoidal 2–3 × 1.5–2.5 T. rugulosus Biverticillate having symmetrical subterminal branches Smooth to ﬁnely rough Ellipsoidal to fusiform 2.5–6 × 2.5–4 T. ryukyuensis Solitary phialides and monoverticillate Rough, thick walled Cylindrical to fusiform to ellipsoidal 3.5–12(–15) × 2.5–3.5 T. sayulitensis Biverticillate, minor proportion with subterminal branches Smooth Subglobose to ellipsoidal 2.5–3 × 2–2.5 T. scorteus Biverticillate having symmentrical subterminal branches Smooth Ellipsoidal 3–5.5 × 2–3 T. siamensis Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Ellipsoidal to fusiform 3–4 × 2–3 T. solicola Biverticillate Rough Globose to subglobose 2–3.5 × 2–2.5 T. stipitatus Mono- to biverticillate Smooth Ovoidal to ellipsoidal 2–7.5 × 2–4 T. stollii Biverticillate, minor proportion with subterminal branches Smooth to ﬁnely rough Ellipsoidal 2.5–4 × 2–2.5 T. subinﬂatus Biverticillate Smooth Ellipsoidal to fusiform 2.5–4 × 1.5–2 T. tardifaciens Solitary phialides and monoverticillate Smooth Cylindrical to ellipsoidal 3–6 × 1.5–2.5 T. thailandensis Biverticillate Smooth Broadly ellipsoidal to ovoidal 2–4 × 2–3 T. trachyspermus Mono- to biverticillate Smooth Ellipsoidal 2–3.5(–5) × 1.5–2.5 T. tratensis Biverticillate Smooth Broadly ellipsoidal to ovoidal 2–2.5 × 3–3.5 T. ucrainicus Mono- to biverticillate Smooth Broadly ellipsoidal to ovoidal 2–4(–5) × 1.5–2.5(–3) T. udagawae Biverticillate Smooth Subglobose to ellipsoidal 3–4 × 2–3 T. unicus Mono- to biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal to ovoidal 2.7–5 × 1.7–3.2 T. varians Mono- to biverticillate Smooth Ellipsoidal to cylindrical 2.5–4 × 1.5–2 T. verruculosus Biverticillate Rough to echinulate Globose 3–3.5 × 3–3.5 T. viridis Solitary phialides Smooth Fusiform to ellipsoidal 2.5–3.5(–4) × 1.5–2 T. viridulus Biverticillate Smooth Cylindrical to rod shaped 3.5–6 × 1–2 T. wortmannii Biverticillate, minor proportion with subterminal branches Smooth Ellipsoidal 2.5–6 × 1.5–3.5 T. yelensis Biverticillate, minor proportion with subterminal branches Rough Subglobose to ellipsoidal 2.5–3.5 × 2.5–3 POLYPHASIC TAXONOMY OF TALAROMYCES 201 YILMAZ 202 Table 5. Morphological characters distinguishing Talaromyces species producing ascoma. Ascoma colour Ascospores Ridges Ornamentation Shape Size (μm) Conidiophore branching Growth rate (mm) CYA MEA T. assiutensis Creamish white to pale yellow Absent Spiny Ellipsoidal 3.5–5 × 2–3 Mono- to biverticillate 15–21 20–23 T. austrocalifornicus Yellow Absent Finely spiny Broadly ellipsoidal 2.5–3.5 × 2–2.5 Biverticillate 10–11 16–18 T. bacillisporus Creamish white to pastel orange Absent Spiny Globose 3.5–5 × 3.5–5 Mono- to biverticillate 10–15 15–20 T. boninensis Grayish green Absent Spiny Broadly ellipsoidal (2.5–)3.5–4(4.5) × 2–3 Biverticillate 28 30 T. cinnabarinus Yellowish orange to orange red Present Broadly ellipsoidal, thick walled 4–5.5(–6) × 3–3.5 Biverticillate Unknown Unknown T. convolutus Sulphur yellow Absent Ornamented by irregular ridges Spiny Globose to subglobose 3–3.5 × 2.5–3(–3.5) Mono- to biverticillate 10–12 13–15 T. derxii Greyish green to dark bluish green Absent Spiny Ellipsoidal 3.5–5 × 2.5–3 Mono- to biverticillate 38–40 48–50 T. emodensis Creamish white to sulphur yellow Absent Spiny Globose to subglobose 2.5–4 × 2.5–3.5 8–10 8–10 T. euchlorocarpius Deep green Absent Spiny Ellipsoidal 3.5–5 × 2.5–3 Monoverticillate, subterminal branching sometimes present Biverticillate 15–18 38–40 T. ﬂavovirens Golden yellow with a reddish pigment Absent Spiny Ellipsoidal, thick walled (4–)4.5–6(–7) × 3–4 19–20 37–38 T. ﬂavus Deep yellow Absent Spiny Broadly ellipsoidal, thick walled 4–5.5 × 3–3.5 Biverticillate, minor proportion with subterminal branches Monoverticillate 9–10 31–32 T. galapagensis Creamish white to yellow Absent Spiny Broadly ellipsoidal 7–10 × 5.5–8 Mono- to biverticillate 15–17 25–28 T. hachijoensis Yellow Present Ellipsoidal 5.5–7 × 3.5–4.5 Absent 3 8–10 T. helicus Creamish white to yellow Absent Ellipsoidal 2.5–4 × 2–3 Mono- to biverticillate 13–23 25–33 T. indigoticus Yellow to orange Absent Ornamented by longitudinal, somewhat sinous ridges Smooth (some with minute spines) Spiny Ellipsoidal 3.5–5 × 2.5–3 Mono- to biverticillate 20–21 32–33 T. intermedius Creamish white to pastel pink Absent Spiny Broadly ellipsoidal, thick walled 4.5–7 × 3.5–5.5 15–16 48–50 T. liani Yellow to orange red Absent Spiny Broadly ellipsoidal 4–6 × 2.5–4 Solitary phialides to monoverticillate Mono- to biverticillate 20–30 35–45 T. macrosporus Yellow Absent Spiny 5–6.5 × 4.5–5.5 Mono- to biverticillate 22–28 40–50 T. mimosinus Pure yellow to sulphur yellow Present 7–8 × 6–7 Mono- to biverticillate 12–15 13–14 T. muroii Pastel yellow to yellow Absent Ornamented by conspicuous sinous ridges Spiny Subglobose to boradly ellipsoidal, thick walled Globose to subglobose Ellipsoidal, ﬁnely thick walled 3.5–6 × 3–4 15–16 30–32 T. paucisporus Yellow (sometimes red) Absent Spiny 14–18 × 12–16 15–18 (after 14 d) 19–20 (after 14 d) T. purpureus Yellow Absent Spiny Subglobose to boradly ellipsoidal, thick walled Ellipsoidal, thick walled Mono- to biverticillate, minor proportion with subterminal branches Absent Solitary phialides and monoverticillate Monoverticillate 2–4 15–16 9–11 15–17 Solitary phialides and monoverticillate Mono- to biverticillate Unknown Unknown 32–38 45–48 6.5–8 × 4.5–5.5 T. rotundus Yellow to orange Absent Spiny Broadly globose to ovoidal 4–5.5 × 4–5.5 T. ryukyuensis Deep reddish Absent Spiny Broadly ellipsoidal, thick walled 4–5 × 3–4 T. stipitatus Creamish white to yellow Present Smooth with single equatorial ridge Flattened ellipsoidal 3–5 × 2–3 ET AL. Species name Table 5. (Continued) www.studiesinmycology.org Species name Ascoma colour Ascospores Conidiophore branching Growth rate (mm) Ridges Ornamentation Shape Size (μm) CYA MEA T. subinﬂatus Sulphur yellow Absent Spiny Broadly ellipsoidal 3–4(–5.5) × 2–3(–5) Biverticillate 3–4 14–15 T. tardifaciens Creamish white to white yellow Absent Smooth Broadly ellipsoidal 3–3.5 × 2–3 9–10 13–15 T. thailandensis Yellow Absent Spiny Broadly ellipsoidal, thick walled 4–6 × 2.5–4.5 Solitary phialides and monoverticillate Biverticillate 33–35 30–35 T. trachyspermus Creamish white Absent Spiny Broadly ellipsoidal 3.5–5 × 2–3 Mono- to biverticillate 13–24 17–25 T. tratensis Yellow Absent Spiny Broadly globose to ovoidal 3.5–5.0 × 2.5–3.5 Biverticillate 10–12 15–20 T. ucrainicus Yellow Present Broadly ellipsoidal 3–5 × 2–3 Mono- to biverticillate 10–20 18–24 T. udagawae Yellow Present Ellipsoidal 3.5–6 × 2.5–3.5 Biverticillate 6–8 10–11 T. unicus Pastel yellow to yellow Present Ornamented with thin, somewhat jagged, irregular, mostly longitudinal ridges Ornamented with 3 to 5 regular, nearly parallel, transverse ridges, often spirally arranged Roughened to spiny, usually with a single ridge Ellipsoidal 3.3–6.7 × 2.8–4.2 10.5–16 15–19.5 T. viridis Dark green Present Ellipsoidal 3.7–4.5 × 2.5–3 9 15 T. wortmannii Yellow to orange Absent Finely ornamented with ridges Smooth to spiny Mono- to biverticillate, minor proportion with subterminal branches Solitary phialides Broadly ellipsoidal, thick walled 3.5–6.0 × 2.5–4 18–28 15–25 Biverticillate, minor proportion with subterminal branches POLYPHASIC TAXONOMY OF TALAROMYCES 203 YILMAZ ET AL. Table 6. Morphological characters distinguishing Talaromyces species producing synnemata. Species T. calidicanius Synnemata Shape Time of Length/ production height (μm) Determinate Prolonged Up to 6000 Acid on Growth rate (mm) CREA CYA 25 C CYA 30 C MEA 30 C MEA YES Moderate 27–30 33–35 30 47–48 40–41 T. cecidicola Determinate Prolonged Up to 1250 Absent 33–34 37–38 38–39 37–38 30–31 T. chloroloma Determinate Prolonged Up to 1200 40–45 50–55 55–60 45–48 35–40 T. coalescens Determinate Prolonged Up to 1200 Weak to moderate Very weak 32–34 40–41 45–46 43–45 34–35 T. dendriticus Determinate Prolonged Up to 5000 Absent 23–26 15–16 20 35–36 15–27 T. duclauxii Indeterminate After 7 d Up to 5000 Weak 25–27 34–35 40 48–50 43–44 T. ﬂavovirens Prolonged Up to 750 Absent 19–20 21–22 37–38 37–38 21–22 T. palmae Determinate, covered or masked by yellow mycelial covering Indeterminate Prolonged Up to 8000 Weak 20–25 20–22 25–27 22–26 25–26 T. panamensis Determinate After 7 d Up to 6800 Strong 23–24 20–21 27–28 28–30 30–32 T. pittii Determinate, cone shaped and often sterile Determinate, phototropic Determinate Prolonged Up to 1000 Absent 34–36 38–40 48–50 42–44 33–35 Prolonged Up to 8000 Absent 25–34 25–38 30–40 38–43 27–32 Prolonged Up to 500 Absent 32–40 39–50 48–52 45–48 40–43 T. pseudostromaticus T. ramulosus Table 7. Overview of morpholgical characters distinguishing soluble red pigment producers in the genus Talaromyces. Species name T. albobiverticillius Growth rate (mm) CYA CYA (37 C) MEA 15–20 No growth 25–28 MEA texture Acid CREA Velvety Absent T. amestolkiae 30–32 8–15 30–45 Floccose and overlaying funiculose Weak T. atroroseus 28–40 23–25 30–35 Velvety and ﬂoccose Absent T. cnidii 25–30 20–28 38–40 Velvety and ﬂoccose Absent T. coalescens 32–34 2–4 43–45 Funiculose Very weak T. marneffei 13–25 5–10 15–27 Loosely funiculose and ﬂoccose Absent T. minioluteus 17–18 No growth 21–22 Loosely funiculose and ﬂoccose Absent T. pittii 34–36 No growth 42–44 Yeast like slimy colonies, after 2 wk ﬂoccose and synnematous Absent T. purpurogenus 20–25 16–25 30–45 Velvety and ﬂoccose Absent T. ruber 20–35 14–18 35–40 Velvety Absent T. stollii 38–45 25–35 45–50 Floccose and lossely funiculose Present Table 8. Morphological characters distinguishing Talaromyces species which produce rough-walled and globose conidia. Species name Colony diameter (mm) CYA CYA 30 C CYA 37 C MEA Acid on CREA YES T. aculeatus 25–35 30–40 15–20 30–35 Moderate Dark green (30F3) to greyish yellow (4C5) Brownish orange (5C6) T. apiculatus 25–35 40–42 Weak Light brown (7D5) to brownish grey (7E2) Brownish orange (5C6) 5–8 25–35 Absent Light yellow (4A5–5A5) Brownish yellow (5C7–5C8) Colony reverse colour MEA 38–41 35–45 T. diversus 7–10 6–10 T. solicola 12–13 13–15 No growth 22–23 No growth Reddish brown to dark brown (9E6–9F6) Centre dark brown (7F7), fading into brown (7E7) T. verruculosus 32–35 37–38 25–26 35–36 Weak Greyish orange (5B5) Greyish orange (5C6) 204 POLYPHASIC TAXONOMY OF TALAROMYCES Table 9. Overview of growth rates on CYA at 37 C for Talaromyces species. Species name CYA 37 C (mm) Species name CYA 37 C (mm) T. chloroloma 0–4 T. varians 27–28 T. wortmannii 0–7 T. stipitatus 28–32 T. cecidicola 2–3 T. macrosporus 28–35 T. coalescens 2–4 T. piceus 30–35 T. rademirici 3 T. sayulitensis 32–40 T. duclauxii 3–4 T. trachyspermus 32–40 T. mimosinus 3–5 T. bacillisporus 33–37 T. emodensis 4–6 T. rubicundus 34–35 T. dendriticus 5–6 T. aerugineus 35 T. ﬂavovirens 5–6 T. derxii 35–38 T. diversus 5–8 T. funiculosus 38–50 T. ramulosus 5–8 T. columbinus 42–43 T. marneffei 5–10 T. albobiverticillius No growth T. ptychoconidium 5–12 T. atricola No growth T. viridulus 6–7 T. brunneus No growth T. convolutus 6–8 T. calidicanius No growth T. ucrainicus 7–17 T. erythromellis No growth T. amestolkiae 8–15 T. euchlorocarpius No growth T. islandicus 8–17 T. hachijoensis No growth T. austrocalifornicus 10–11 T. intermedius No growth T. oumae-annae 10–11 T. loliensis No growth T. viridis 10–11 T. minioluteus No growth T. helicus 10–18 T. palmae No growth T. ruber 14–18 T. panamensis No growth T. siamensis 15 T. paucisporus No growth T. aculeatus 15–20 T. pittii No growth T. purpurogenus 16–25 T. primulinus No growth T. ﬂavus 19–20 T. proteolyticus No growth T. muroii 19–20 T. pseudostromaticus No growth T. aurantiacus 19–21 T. purpureus No growth T. indigoticus 20–22 T. rotundus No growth T. liani 20–25 T. rugulosus No growth T. cnidii 20–28 T. scorteus No growth T. allahabadensis 23–25 T. solicola No growth T. atroroseus 23–25 T. subinﬂatus No growth T. galapagensis 25–26 T. tardifaciens No growth T. yelensis 25–26 T. thailandensis No growth T. verruculosus 25–26 T. tratensis No growth T. angelicus 25–27 T. udagawae No growth T. assiutensis 25–30 T. unicus No growth T. radicus 25–30 T. bohemicus Unknown T. apiculatus 25–35 T. cinnabarinus Unknown T. stollii 25–35 T. ryukyuensis Unknown T. pinophilus 25–40 T. boninensis Unknown (growth was reported, but no diam) www.studiesinmycology.org 205 YILMAZ ET AL. Fig 8. Micromorphological features of Talaromyces. A–E. Condiophore branching: A. Solitary phialides. B. Monoverticillate. C. Biverticillate. D–E. Biverticillate with extra subterminal branching. F. Acerose phialides G. Flask-shaped phialides. H–K. Shapes of conidia: H. Cylindrical and rod-shaped. G. Globose. H, I. Ellipsoidal. Scale bars: F, G = 10 μm; A = 10 μm, applies B–E and H–K. summarises the characters needed for distinguishing species that produce red soluble pigments and include growth rates on CYA and MEA at 25 and 37 C, colony texture on MEA and acid production on CREA. Talaromyces species generally produce acerose phialides and ellipsoidal to fusiform conidia. Some species in Talaromyces, however, produce rough-walled, globose conidia and these species are listed in Table 8. Species, which produce roughwalled globose conidia include T. aculeatus, T. apiculatus and T. verruculosus (classiﬁed in sect. Talaromyces), T. diversus and T. solicola (classiﬁed in sect. Trachyspermi). All these species can be distinguished by using characters like growth rate on CYA at 206 25, 30 and 37 C, acid production on CREA and reverse colour on YES and MEA (Table 5). Extrolite proﬁles Extrolite proﬁles are fundamental taxonomic criteria in ﬁlamentous fungi (Frisvad et al. 1990a, b). Talaromyces has unique and speciﬁc extrolites such as duclauxins, glauconic acids, mitorubrins, monascins, purpactins, rubratoxin, vermicellins, etc. (Samson et al. 2011). An overview of extrolites produced by Talaromyces is provided in Table 10. The extrolite proﬁles of each species are included POLYPHASIC TAXONOMY OF TALAROMYCES − − − − + + − − − − − − − − − − − − − − − − − − − − − − − − − − + − − − − − − − − − − T. allahabadensis − − − − − − − − − − − − + − − − + − − + − − − − −− − − − − T. apiculatus + + − − − − − − − − − − + + − − − − − − − − − − − − − − − T. assiutensis − − − − − − − − − − + − − − − − − − − − − − − − − − − − − T. atricola − − − − − − − − − − − − + − − − − − − + − − − − − − − − − Wortmannin − − Wortmannilactones ðF-H) − − Secalonic acid D & F Stipitatic acid Vermicellin Vermiculin Vermistatin & Funicones − − Rugulovasine − − Skyrin − − Rugulosin − − Rubratoxins Rubropunctatin and other monascus pigments − − Prugusenes=Ukulactones Duclauxin Epi-Austdiol Falconensis Glauconic=Glaucanic acid + T. albobiverticillius Penisimlissin Pentacecilides=Thailandolides Cyclochlorotine & Islanditoxin T. aculeatus Luteoskyrin Species name Alternariol Apiculides Asperphenamate Austin Bacillisporin Botryodiploidin Mitorubrins Penicillides=Purpactins=Vermixocins Table 10. An overview of extrolites produced by Talaromyces species (* Mycotoxins; + present; − absent). T. atroroseus − − − − − − − − − − + − + + − − − − + − − − − − − − − − − T. austrocalifornicus − − − − − − − − − − − − + + − − − − − − − − − − − − − − − T. bacillisporus − − − − + − − + − − − − − − − − − − − − − − − − − − − − − T. brunneus − − − − − − − − − − − − + + − − − - − + − − − − − − − − − T. calidicanius − − − − − − − + − − − − − − − −− − − − − − − − − − − − − − T. cecidicola − + − − − − − − − − − − − − − + − − − − − − − − − − − − T. coalescens − − − − − + − − − − − − − + − − − − − − − − − − − − − − − T. columbinus − − − − − − − − − − − − − − − − − − − + − − − − − − − − − − T. convolutus − − − − − − − − − − − − + + − − − - − − − − − − − − − − − T. dendriticus − − − − − − − − − − − − − + − − − + − − − − + − − + − − − T. diversus + − − + − - − − − − − − + − − − − − − − − − − − − −− − − − T. duclauxii − − − − − − − + − − − − − − − − − − − − − − − − − − − − − T. emodensis − − − − − − − + − − − − + − − − − − − − − − − − − − − − − T. erythromellis − − − − − − − − − − − − − − − − − − + − − − − − − − − − − T. ﬂavovirens − + − − − − − − − − − − + + − − − − − − − − + − − − − − − T. ﬂavus − + − − − − − − − − − − + + − − − − − − − − − − − + + − − T. funiculosus − − − − − − − − − − − − − − − − − − − − − − + − − − +(?) − − T. galapagensis − + − − − − − − − − − − − − − − − − − − − − − − − − − − − T. hachiojensis − − − − − − − − − − − − + − − − − − − − − − − − − − − − − T. helicus + − − − + − − − − − − − − − − − − − − − − − − − − − − − − T. islandicus − − − − − − + − − − − + + − − − − − − + − − − − − − − − − T. loliensis − − − − − − − − − − − − + − − − − − − − + − − − − − − − − T. macrosporus − − − − − − − + − − − − + − − − − − − − − − − − − − − − − T. marneffei − − − − − − − − − − − − + + − − − − + − − − − − − − − − − T. mimosinus − − − − − − − − − + − − − − − − − − − − − − − − − − − − − T. minioluteus − − − − − − − − − − − − + + − − − − − − − − + − − − − − − T. palmae − − − − − − − − − − − − + − − − − + − − − − − − − − − − − T. panamensis − − − − − − − − − − − − − + − − − − − − − − − − + − − − − T. piceus − − − − − − − − − − − − + − − − − − − + − − − − − − − − − T. pinophilus − − − + − − − − − − − − + + + − − − + − − − − − + + + − − T. primulinus − − − − − − − − − − − − + − − − − − − − − − − − − − − − − T. pseudostromaticus − − − − − − − − − − − − + − − − − − − − − − + − − + − − − T. purpurogenus − − − − − − − − − − + + + − − − + + − − + − − − + − − − − (continued on next page) www.studiesinmycology.org 207 YILMAZ ET AL. Species name Alternariol Apiculides Asperphenamate Austin Bacillisporin Botryodiploidin Cyclochlorotine & Islanditoxin Duclauxin Epi-Austdiol Falconensis Glauconic=Glaucanic acid Luteoskyrin Mitorubrins Penicillides=Purpactins=Vermixocins Penisimlissin Pentacecilides=Thailandolides Prugusenes=Ukulactones Rubratoxins Rubropunctatin and other monascus pigments Rugulosin Skyrin Rugulovasine Secalonic acid D & F Stipitatic acid Vermicellin Vermiculin Vermistatin & Funicones Wortmannilactones ðF-H) Wortmannin Table 10. (Continued). T. radicus − − − − − − − − − − − − + − − − + − − + − − − − − T. rotundus − − − − − − − − − − − − − − − − − − − + − − T. ruber − − − + − − − − − − − − − − − − − − − − − − − − − − − − − − − − − − − + − − − − T. rubicundus − − − − − − − − − − − − + − − − − + − − − − − − − − − − − T. rugulosus − − − − − − − − − − − − + − − − + − − + − − − − − − − − − T. scorteus − − − − − − − − − − − − + − − − − − − − − − − − − − − − − T. siamensis − − − − − − − − − − − − + + − − − − − − − − + − + − − − − T. stipitatus − − − − + + − + + − − − − + − − − − − − − − + + − − − − − T. tardifaciens − − − − − − − − − + − − + − − − − − − − + − − − − − − − − T. thailandensis − − + − − − − − − − − − − − + + − − − − − − − − − − − − − T. trachyspermus − − − − − − − − − − + − − − − − − − − − − − − − − − − − − T. tratensis − − − − − − − − − − − − + − − − − − − + − − − − + − − − + T. ucrainicus − − − − − − − − − − + − − − − − − − − − − − − − − − − − − T. udagawae − − − − − − − − − − − − + + − − − − − − − − − − − − − − − T. verruculosus + − − − − − − − − − − − + + − − − − − − − − − − + − + − − T. wortmannii − − − − − − − − − − − − + − − − − − − + − + − − − − − + − in the species descriptions (see Taxonomy). Mitorubrins, penicillides, purpactins and vermixocins are produced by all Talaromyces species in different ratios. Rugulosin and/or skyrin, which are known to be mycotoxins, are produced only by members of section Islandici. Some Talaromyces species produce other mycotoxins such as botryodiploidin (by T. coalescens and T. stipitatus), cyclochlorotine and islanditoxin (by T. islandicus), luteoskyrin (by T. islandicus and T. purpurogenus), rubratoxin (by T. purpurogenus), rugulovasine (by T. purpurogenus and T. wortmannii) and secalonic acid D & F (by T. dendriticus, T. ﬂavovirens, T. funiculosus, T. minioluteus, T. pseudostromaticus, T. siamensis and T. stipitatus). Red pigment producers such as, T. marneffei, T. albobiverticillius, T. purpurogenus, T. albobiverticillius in sections Talaromyces and Trachyspermi produce rubropunctatin and other Monascus pigments. Species producing globose rough-walled conidia also produce alternariol (eg. T. verruculosus, T. apiculatus, T. diversus). Glauconic acid and glaucanic acid are speciﬁc for section Trachyspermi. TAXONOMY (SPECIES LISTED IN ALPHABETICAL ORDER) Talaromyces aculeatus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 174. 2011. MycoBank MB560639. Fig. 9. ≡ Penicillium aculeatum Raper & Fennell, Mycologia 40: 535. 1948. 208 In: Talaromyces section Talaromyces Typus: IMI 040588, culture ex-type CBS 289.48 = CBS 136670 = ATCC 10409 = IMI 040588 = NRRL 2129 = NRRL A1474 = IBT 14259 = IBT 4185. ITS barcode: KF741995 (alternative markers: BenA = KF741929; CaM = KF741975; RPB2 = KM023271) Colony diam, 7 d (mm): CYA 25–35; CYA 30 C 30–40; CYA 37 C 15–20; MEA 30–35; MEA 30 C 40–42; DG18 15–17; CYAS 1–5; OA 28–33; CREA 15–20; YES 28–33. Colony characters: CYA 25 C, 7 d: Colonies moderately deep, radially and concentrically sulcate; margins low, narrow, entire; mycelia white, sometimes inconspicuously yellow; texture ﬂoccose; sporulation absent to sparse, conidia en masse greyish to dull green (27C4–27D4); soluble pigments absent; exudates minute red droplets; reverse light brown (7D4–7D5) at the centre, elsewhere pale to greyish yellow (4A3–4B3). MEA 25 C, 7 d: Colonies moderately deep, plane to sometimes radially sulcate, raised at centre; margins low to somewhat subsurface, wide (up to 5 mm), entire; mycelia white and inconspicuously yellow; texture ﬂoccose to funiculose; sporulation sparse to moderately dense, conidia en masse greyish green (27E5); soluble pigments absent, exudates minute red droplets; reverse brownish orange (5C6). YES POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 9. Morphological characters of Talaromyces aculeatus (CBS 289.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 209 YILMAZ ET AL. 25 C, 7 d: Colonies moderately deep, irregularly sulcate, raised at centre; margins low, narrow, entire; mycelia white, yellow, red; texture ﬂoccose to lightly funiculose; sporulation absent to sparse, conidia en masse greyish green (27D5–27E5); soluble pigments absent, exudates absent; reverse dark green (30F3), greyish yellow (4C5), pale yellow (4A3). DG18 25 C, 7 d: Colonies have a reddish reverse. CREA 25 C, 7 d: Weak acid produced. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 150–300 × 2.5–3.5 μm; branches 10–35 μm; metulae four to eight, divergent, 8.5–12 × 2.5–3.5 μm; phialides ﬂask-shaped, tapering into very thin neck, three to ﬁve phialides per metulae, 9–11 × 2.5–3.5 μm; conidia rough to echinulate, globose, 3–3.5 × 3–3.5 μm. Ascomata not observed. Extrolites: Altenusin, alternariol, alternariol momomethyl ether, cyclopeptin, MC-141, penicillide, purpactins, rubiginosin A, a rubropunctatin, secopenicillide C, vermicillin are produced by T. aculeatus (Fuska et al. 1979). Among the species treated here, T. aculeatus is the only species producing both cyclopeptin and MC-141. Strain NR 5165 = IFO 5689 = FAT 810 (not examined) produces the antibiotics penitricins (Okuda et al. 1984a, b, c). Distinguishing characters: Talaromyces aculeatus produces ﬂask-shaped phialides and rough-walled to echinulate, globose conidia (Fig. 9). These characters are similar to T. verruculosus and T. apiculatus. However, T. aculeatus grows slower than T. apiculatus on CYA (at 25 and 37 C) and MEA, and grows slower than T. verruculosus on CYA at 37 C. Talaromyces aerugineus (Samson) Yilmaz, Frisvad & Samson, comb. nov. MycoBank MB809553. Fig. 10. Basionym: Paecilomyces aerugineus Samson, Stud. Mycol. 6: 20. 1974. In: Talaromyces section Helici Typus: CBS H-7448, culture ex-type CBS 350.66 = BDUN 276 = IMI 105412. ITS barcode: AY753346 (alternative markers: BenA = KJ865736; CaM = KJ885285; RPB2 = JN121502) Colony diam, 7 d (mm): CYA 27–26; CYA 30 C 35–36; CYA 37 C 35; MEA 49–50; MEA 30 C 65–70; DG18 6–7; CYAS No growth; OA 45–50; CREA No growth; YES 29–30. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, sulcate, green appearance; margins low, plane, entire (1 mm); mycelia white; texture funiculose; sporulation absent; soluble pigments absent; exudates absent; reverse dark green (26F5) centre fading into greyish green (26B5–26B5). MEA 25 C, 7 d: Colonies low, sulcate, sterile white mycelia; margins low, plane, entire (1 mm); mycelia white; sporulation absent; texture loosely funiculose to velvety; soluble pigments absent; exudates absent; reverse greyish orange (5B5–5B6). YES 25 C, 7 d: Colonies raised at centre, slightly sulcate, green and orange appearance; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments 210 absent; exudates absent; reverse light yellow (4A4) with light orange (5A6) circle. DG18 25 C, 7 d: Colonies raised, plane; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse dull green (25D4–26D4); soluble pigments absent; exudates absent; reverse olive (1F3–1F4). OA 25 C, 7 d: Colonies low, plane, green (because of the reverse); margins low, plane, entire (3–5 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse greyish green to dull green (at 30 C greyish green (25E6)). CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores with solitary phialides or monoverticillate; stipes smooth walled, 20–75 × 2.5–5 μm; phialides acerose, one to three, 10–20 × 2.5–5 μm; conidia smooth, in various shapes subglobose to ellipsoidal to fusiform, 3–8.5 × 2.5–5 μm. Ascomata not observed. Distinguishing characters: Talaromyces aerugineus is characterised by fast growing colonies on MEA at 25 and 30 C. Conidiophores mainly have solitary phialides and sometimes monoverticillate conidiophores and produces large, smooth, globose to ellipsoidal conidia (3–8.5 × 2.5–5 μm) (Fig. 10). Talaromyces albobiverticillius (H.-M. Hsieh, Y.M. Ju & S.Y. Hsieh) Samson et al., Stud. Mycol. 71: 174. 2011. MycoBank MB560683. Fig. 11. ≡ Penicillium albobiverticillium H.-M. Hsieh, Y.M. Ju & S.Y. Hsieh, Fungal Sci. 25: 26. 2010. In: Talaromyces section Trachyspermi Typus: BCRC 34774, culture ex-type CBS 133440 = DTO 166E5 = YMJ 1292. ITS barcode: HQ605705 (alternative markers: BenA = KF114778; CaM = KJ885258; RPB2 = KM023310) Colony diam, 7 d (mm): CYA 15–20; CYA 30 C 13–17; CYA 37 C No growth; MEA 25–28; MEA 30 C 20–25; DG18 12–18; CYAS generally no growth some isolates up to 3; OA 20–25; CREA 2–5; YES 20–25. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, sulcate, in some isolates sunken at centre and crateriforme; margins low, plane, entire (<1 mm); mycelia white and in some isolates yellow; texture velvety to ﬂoccose; sporulation moderately dense to dense, conidia en masse only in the ex-type strain reddish or pinkish white (8A2), the other strains greenish grey (26C2) to dull green (26E3); soluble pigments in some isolates red, in some lacking; exudates in some isolates red droplets; reverse in some isolates reddish brown (9F8) and in some isolates greyish red (9C5). MEA 25 C, 7 d: Colonies slightly raised at centre, and in some isolates sunken in the centre, sulcate; margins low, plane, entire (1–2 mm); mycelia white and pastel yellow and also in some isolates yellow; texture velvety; sporulation moderately dense to dense, conidia en masse reddish or pinkish white (8A2) only in the the ex-type strain, and for the other isolates greyish green (25D5–26D5); soluble pigments absent; exudates in some isolates red droplets; reverse in some isolates reddish brown (9E8) and in some lack of red reverse and brownish orange (5C6). YES 25 C, 7 d: POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 10. Morphological characters of Talaromyces aerugineus (CBS 350.66T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B, C. Colony texture on MEA after 2 wk incubation. D–G. Conidiophores. H. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–H. www.studiesinmycology.org 211 YILMAZ ET AL. Fig. 11. Morphological characters of Talaromyces albobiverticillius (CBS 206.89). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on MEA after 1 wk incubation (CBS 113442T). C. Colony texture on MEA after 1 wk incubation (CBS 206.89). D–H Conidiophores. I. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–I. 212 POLYPHASIC Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white, pastel yellow and in some isolates yellow; texture velvety and ﬂoccose; sporulation moderately dense to dense, conidia en masse reddish or pinkish white (8A2) only in the ex-type strain, and for the other isolates greyish green (25D5–26D5); soluble pigments red (in some isolates absent); exudates in some isolates small red droplets; reverse in some isolates red (9B8), and in some isolates lacking of red reverse, greyish orange (5B5) centre, fading into brownish orange (5C3) and brownish grey (5C2). DG18 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white; texture velvety; sporulation sparse to moderately dense, conidia en masse reddish or pinkish white (8A2) only in the ex-type strain, and for the other isolates greenish grey to dull green (26D2–26D3); soluble pigments in some isolates red, generally absent; exudates in some isolates slimy clear droplets, in some isolates small red droplets; reverse in some isolates reddish brown (9E8), in some isolates reddish orange (7A8) centre fading into orange (6A8) and in some isolates dull yellow (3B4) centre fading into pale yellow (3A3). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia only in type white, pastel yellow and pastel red, in other strains white; texture velvety; sporulation moderately dense to dense, conidia en masse reddish or pinkish white (8A2) only in the ex-type strain, and for the other isolates dull green (26D4); soluble pigments in some isolates red, generally absent; exudates absent; reverse in some isolates dark red, in some lack of red reverse and pale green. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 200–400 × 2.5–4 μm; metulae three to eight, divergent, 8–13 × 1.5–4.5 μm; phialides acerose, three to eight per metulae, 8–13.5 × 2–3 μm; conidia smooth to ﬁnely rough, globose to subglobose and in some isolates fusiform, 2–3.5(–4) × 1.5–2.5 μm. Ascomata not observed. Extrolites: Talaromyces albobiverticillius produces mitorubrin, mitorubrinic acid, mitorubrinol, monascin, monascorubramin, penicillide, a purpactin, rubropunctatin, vermicillin (Frisvad et al. 2013). Distinguishing characters: The ex-type culture of T. albobiverticillius has pinkish white conidia, which is unique to this species. However, some strains do produce green conidia (Fig. 11). Talaromyces albobiverticillius produces intense red soluble pigment on CYA. It differs from other red pigment producing species by restricted growth on CYA at 25 C and no growth at 37 C. It also has stipes up to 400 μm long, as wide as 4 μm and divergent metulae. Talaromyces allahabadensis (B.S. Mehrotra & D. Kumar) Samson et al., Stud. Mycol. 71: 174. 2011. MycoBank MB560640. Fig. 12. ≡ Penicillium allahabadense B.S. Mehrotra & D. Kumar, Can. J. Bot. 40: 1399. 1962. = Penicillium zacinthae C. Ramírez & A.T. Martínez, Mycopathologia 74: 167. 1981. In: Talaromyces section Islandici Typus: University of Allahabad P-26, culture ex-type CBS 453.93 = ATCC 15067 = CBS 304.63 = NRRL 3397 = IBT 3926 = IBT 10824. www.studiesinmycology.org TAXONOMY OF TALAROMYCES ITS barcode: KF984873 (alternative markers: BenA = KF984614; CaM = KF984768; RPB2 = KF985006) Colony diam 7 d (mm): CYA 20–25; CYA 30 C 24–27; CYA 37 C 23–25; MEA 20–23; MEA 30 C 23–26; DG18 14–18; CYAS 14–18; OA 22–25; CREA 8–12; YES 22–23. Colony morphology: CYA 25 C, 7 d: Colonies 20–25 mm, slightly raised at centre, slightly sulcate; margins very narrow (1 mm), low, entire, plane; mycelium white and yellow; texture velvety and ﬂoccose; sporulation sparse to moderately dense; conidia en masse greyish green (27D5–27E5); exudates clear and yellow droplets; soluble pigment absent; reverse orange to greyish orange (5A6–5B6) in the centre fading into light yellow to greyish yellow (4A5–4B5). MEA 25 C, 7 d: Colonies 20–23 mm, slightly raised at centre, plane; margins narrow (1–2 mm), low, entire, plane; mycelium white and yellow; texture velvety; sporulation moderately dense to dense; conidia en masse dull green (26E4); exudates yellow droplets; soluble pigment absent; reverse dark brown (6F5) in the centre fading into light brown (6D5). YES 25 C, 7 d: Colonies 22–23 mm, slightly raised at centre, sulcate; margins narrow (1 mm), low, in some isolates entire and in some isolates not entire, plane; mycelium white and yellow; texture velvety; sporulation sparse to moderately dense; conidia en masse dull green (26D4–26E4); exudates absent (except DTO 67-F7, clear exudates); soluble pigment absent; reverse brownish orange (5C5–5C6) centre fading into greyish orange (5B4–5B6). DG18 25 C, 7 d: Colonies 14–18 mm, raised at centre, plane; margins narrow (1 mm), low, entire, plane; mycelium white (only CBS 137362 with yellow mycelia at centre); texture velvety and ﬂoccose; sporulation absent to dense (CBS 137362); conidia en masse greyish green to dull green (26D4–26D5); exudates clear and yellow droplets; soluble pigment absent; reverse greyish orange to orange (5A6–5B6) in the centre fading into light yellow to greyish yellow (4A5–4B5). OA 25 C, 7 d: Colonies 22–25 mm, in some isolates raised at centre and in some isolates plane plane; margins wide (2 mm), low, entire, plane; mycelium white and yellow; texture velvety and in some isolates loosely funiculose; sporulation moderately dense to dense; conidia en masse greyish green (30C6–30D6) and in some isolates dull green (28D4–28E4); exudates absent; soluble pigment absent; reverse bright orange yellow and in some isolates yellowish green. CREA, 25 C, 7 d: Colonies 8–12 mm, acid production. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 50–175 × 2.5–2.7 μm; metulae three to six, divergent, 8–11 × 2.4–3.2 μm; phialides acerose, three to six per metulae, 8.5–11 × 2–3 μm; conidia smooth, ellipsoidal to fusiform, 2.5–4.5 × 1.7–2.5 μm. Ascomata not observed. Extrolites: Talaromyces allahabadensis produces prugosenes/ ukulactones (Lang et al. 2007, Mori et al. 2011) and several azaphilones: mitorubrin, mitorubrinol, mitorubrinol acetate, mitorubrinic acid, 60 -hydroxy-30 -methoxy-mitorubrin, 40 -hydroxy30 -methoxy-(S)-mitorubrin and monomethyl-(S)-mitorubrin. Three isolates, IBT 12688, CBS 137397, CBS 441.89 also produced rugulosin A, while CBS 453.93 and CBS 178.81 did not. It seems that this species produces rugulosin A preferentially at 37 C, and the latter two species were only examined for extrolite production at 25 C. Talaromyces allahabadensis 213 YILMAZ ET AL. Fig. 12. Morphological characters of Talaromyces allahabadensis (CBS 453.93T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale Bar: B = 10 μm, applies to C–H. 214 POLYPHASIC shares prugosenes with T. rugulosus and the three latter mitorubrins listed above with T. radicus. Distinguishing characters: Talaromyces allahabadensis is characterised by colonies with bright yellow mycelia and velvety texture on most media (Fig. 12). The species grows well at 37 C. Talaromyces allahabadensis has ellipsoidal to fusiform conidia. Based on the multigene phylogeny, T. allahabadensis is closely related to T. radicus (Fig. 1). Talaromyces allahabadensis is distinguished from T. radicus by acid production on CREA, and denser sporulation on most media. Also, T. allahabadensis cannot grow at 40 C. Notes: Phylogenetically Penicillium zacinthae (CBS 178.81) is identical to T. allahabadensis (Fig. 7) and is considered a synonym. Talaromyces amestolkiae Yilmaz et al., Persoonia 29: 48. 2012. MycoBank MB801358. Fig. 13. In: Talaromyces section Talaromyces Typus: CBS H-21050, culture ex-type CBS 132696 = DTO 179-F5. ITS barcode: JX315660 (alternative markers: BenA = JX315623; CaM = KF741937; RPB2 = JX315698) Colony diam 7 d (mm): CYA 30–32; CYA 30 C 30–36; CYA 37 C 8–15; MEA 30–45; MEA 30 C 45; DG18 18–19; CYAS 3–8; OA 45–52; CREA 15–25; YES 25–35. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (2–3 mm); mycelia white, pastel yellow and pastel red; texture ﬂoccose in the centre and loosely funiculose; sporulation moderately dense to dense, conidia en masse greyish green (26E6–26E7); soluble pigments very weak brownish red, in some strains absent (at 37 C yellowish red); exudates absent; reverse violet brown (11E8–11F8). MEA 25 C, 7 d: Colonies slightly raised at centre, plane, in some colonies black sclerotia produced in longer incubation; margins low, plane, entire (3–5 mm); mycelia white and pastel red; texture ﬂoccose in centre with overlaying loosely funiculose; sporulation moderately dense to dense, conidia en masse greyish to dull green (25D4–26D4); soluble pigments absent; exudates absent; reverse dark brown (9F8) centre fading into reddish brown (9E8) and greyish yellow to greyish orange (3C5–5C5) at margins. YES 25 C, 7 d: Colonies slight raised at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white, pastel red and pastel orange; texture ﬂoccose and loosely funiculose; sporulation dense, conidia en masse greyish green (26E6–26E7); soluble pigments absent; exudates absent; reverse violet brown (11F8) to dark ruby (12F8). DG18 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose and loosely funiculose, aerial sterile hyphae at centre; sporulation dense, conidia en masse greyish green (26E6–26E7); soluble pigments absent; exudates red droplets; reverse dark brown (8F8) centre fading into brownish red (8C6) and margins greyish green (1C4). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (5–6 mm); mycelia white and in some isolates pastel yellow; texture velvety; sporulation dense, conidia en masse greyish www.studiesinmycology.org TAXONOMY OF TALAROMYCES green (25D4–25D6); soluble pigments absent; exudates small clear droplets; reverse red in centre fading into greyish green (in some isolates lack of red centre). CREA 25 C, 7 d: Weak acid production. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 90–180 × 2.5–3 μm; branches 15–50 μm; metulae three to six, divergent, 9–15 × 3–4 μm; phialides acerose, three to six per metulae, 9.5–12 × 2.5–3 μm; conidia smooth and sometimes ﬁnely rough, ellipsoidal, 2–3 × 1.5–2.5 μm. Ascomata not observed. Extrolites: Talaromyces amestolkiae produces berkelic acid, mitorubrin, mitorubrinic acid, mitorubrinol, Monascus red azaphilone pigments (not diffusing into the agar), pestalacin A, a purpactin and vermicellin (Yilmaz et al. 2012). Distinguishing characters: Fast growth on CYA and MEA, production of berkelic acid, red mycelia, loosely funiculose texture and sclerotia production after 2 wk of incubation make T. amestolkiae a distinct species. Red soluble pigment and dark to violet brown reverses are also important characters however most of the time this species was confused with other red pigment producers such as T. purpurogenus (Yilmaz et al. 2012). Talaromyces amestolkiae is distinguished from T. purpurogenus by producing acid on CREA. Phylogenetically it is closely related to T. ruber and T. stollii. Talaromyces ruber differs from T. amestolkiae by its velvety texture and acid production similar to that of T. purpurogenus. Talaromyces stollii is distinguished from T. amestolkiae by faster growth at 37 C. Talaromyces angelicus S.H. Yu, T.-J. An & H. Sang, J. Microbiol. 51: 707. 2013. MycoBank MB804807. Fig. 14. In: Talaromyces section Talaromyces Typus: KACC 46611, culture ex-type KACC 46611. ITS barcode: KF183638 (alternative markers: BenA = KF183640; CaM = KJ885259) Colony diam, 7 d (mm): CYA 25–27; CYA 30 C 30–32; CYA 37 C 25–27; MEA 33–35; MEA 30 C 40–41; DG18 13–15; CYAS 2–3; OA 30–32; CREA 12–13; YES 30–32. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate, white sterile appearance; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose (after 2 wk funiculose); sporulation absent (after 2 wk dense sporulation); soluble pigments absent; exudates absent (at 30 C clear droplets); reverse greyish red (7B3) centre fading into yellowish white (4A2), at 30 C reverse centre violet brown (1E6) fading into brownish red to brownish violet (11C6–11D6). MEA 25 C, 7 d: Colonies low, slightly sulcate, white and yellow sterile appearance; margins low, plane, entire (1–2 mm); mycelia white and yellow in the centre; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish orange (5B6). YES 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate, white sterile appearance; margins low, plane, entire (2–3 mm); mycelia white; sporulation absent; soluble pigments 215 YILMAZ ET AL. Fig. 13. Morphological characters of Talaromyces amestolkiae (CBS 132696T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 216 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 14. Morphological characters of Talaromyces angelicus (KACC 46611T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 217 YILMAZ ET AL. absent; exudates absent; reverse pale orange (5A3) centre fading into pastel yellow (3A4). DG18 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate, yellow sterile appearance; margins low, plane, entire (1–2 mm); mycelia yellow; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A5–5A6) centre fading into pastel yellow to light yellow (3A4–3A5). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia yellow; texture funiculose; sporulation moderately dense, conidia en masse greyish green (27C5–28C5); soluble pigments absent; exudates absent; reverse pale yellowish beige. CREA 25 C, 7 d: Acid production absent and in some isolates very weak acid production. Micromorphology: Conidiophores biverticillate also sometimes monoverticillate; stipes smooth walled, 15–120 × 2–3 μm; metulae three to six, divergent, 8–15 × 2.5–3.5 μm; phialides acerose, three to six per metulae, 8–14 × 2–3 μm; conidia ﬁnely roughed, subglobose to ellipsoidal, 2.5–3.5 × 2–3 μm. Ascomata not observed. Distinguishing characters: Talaromyces angelicus is characterised by ﬂoccose colonies after 1 wk, which become funiculose with time (Fig. 14). Phylogenetically it is closely related to T. pinophilus, T. muroii and T. liani (Fig. 2). It differs from T. muroii and T. liani by lacking ascomata. Morphologically, T. angelicus is similar to T. pinophilus and differs from the latter species by lack of acid production on CREA. Talaromyces apiculatus Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 174. 2011. MycoBank MB560641. Fig. 15. ≡ Penicillium aculeatum var. apiculatum Abe, S., 1956, J. Gen. Appl. Microbiol., Tokyo 2: 124. 1956 (nom. inval., Art. 36). In: Talaromyces section Talaromyces Typus: CBS H-20755, culture ex-typus: CBS 312.59 = ATCC 18315 = FRR 635 = IMI 068239 = ATCC 18315 = IFO 5728 = IBT 10894 = IBT 14261. ITS barcode: JN899375 (alternative markers: BenA = KF741916; CaM = KF741950; RPB2 = KM023287) Colony diam, 7 d (mm): CYA 38–41; CYA 30 C 35–45; CYA 37 C 25–35; MEA 40–42; MEA 30 C 52–57; DG18 19–22; CYAS 6–9; OA 36–40; CREA 20–22; YES 40–43. C, Colony characters: CYA 25 7 d: Colonies moderately deep, radially and concentrically sulcate; margins low, narrow, entire; mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse dull green (27D3–28D3); soluble pigments absent, exudates minute red droplets; reverse greyish yellow (3B3) to yellowish white (3A2) at margin. MEA 25 C, 7 d: Colonies moderately deep, plane, slightly raised at centre; margins low to somewhat subsurface, wide (up to 5 mm), entire; mycelia white and yellow; texture ﬂoccose to strongly funiculose; conidioge sporulation nesis sparse to moderately dense, conidia en masse greyish green (27D5–27E5); soluble pigments absent, exudates minute red droplets; reverse brownish orange (5C6). YES 25 C, 7 d: Colonies moderately deep, irregularly sulcate, raised at centre; margins low, narrow, entire; mycelia white, yellow; texture ﬂoccose; sporulation absent to sparse, conidia en masse greyish green (27D5–27E5); soluble pigments absent, 218 exudates absent; reverse light brown (7D5), sometimes areas of brownish grey (7E2), pale yellow (4A3). DG18 25 C, 7 d: Colonies have a bright orange reverse and produce abundant exudate. CREA 25 C, 7 d: Weak acid produced. Micromorphology: Conidiophores biverticillate; stipes smooth walled, (100–)150–350 × 2.5–3 μm; metulae four to eight, divergent, 7.5–11 × 2.5–3.5; phialides ﬂask-shaped, tapering into very thin neck, three to ﬁve per metulae, 9–11 × 2.5–3.5 μm; conidia echinulate, globose with a minor proportion apiculate, 3–4(–5.5) × 3–4(–5) μm. Ascomata not observed. Extrolites: Talaromyces apiculatus produces the macrocyclic polylactones (“apiculides”) NG-011 and NG-012 (Ito et al. 1992a, b), BK223-A (=NG-012), BK-223B, BK-223C (Breinholt et al. 1993), 15G256β, and 15G256α-2 (Gao et al. 2013). The latter ﬁve extrolites have been reported as antifungal (Breinholt et al. 1993, Schlingmann et al. 2002), while NG-011 and NG-012 are potentiators of nerve growth factors (Ito et al. 1992a, b). Furthermore, by implication, T. apiculatus produces bioxanthracene 2 = ES-242-2 (Toki et al. 1992, Gao et al. 2013), penicillide, prenpenicillide and prenxanthone (Gao et al. 2013). We detected altenusin (and alternariol & alternariol monomethyl ether in six strains, IBT 32315, IBT 13085, IMI 186297, IMI 191967, IMI 352119 & CBS 264.67), apiculides, bioxanthracene B, mitorubrins, purpactins, vermicillin. Decalpenic acid (Sakamoto et al. 2010) was tentatively identiﬁed in IMI 352119. Among the species treated here, T. apiculatus is the only species producing apiculides and bioxanthracene B. Distinguishing characters: Talaromyces apiculatus produces ﬂask-shaped phialides and rough-walled to echinulate, globose conidia (Fig. 15). These characters are also observed in T. aculeatus and T. verruculosus. However, T. apiculatus grows much faster than T. aculeatus on CYA (25 and 37 C) and on MEA. Talaromyces apiculatus also grows faster than T. verruculosus on CYA and MEA at 25 C. Talaromyces assiutensis Samson & Abdel-Fattah, Persoonia 9: 501. 1978. MycoBank MB324414. Fig. 16. ≡ Penicillium assiutense Samson & Abdel-Fattah, (simultaneously published). = Penicillium gossypii Pitt, The genus Penicillium: 500. 1980 ≡ Talaromyces gossypii Pitt, The genus Penicillium: 500. 1980. In: Talaromyces section Trachyspermi Typus: CBS 147.78, culture ex-type CBS 147.78. ITS barcode: JN899323 (alternative markers: BenA = KJ865720; CaM = KJ885260; RPB2 = KM023305) Colony diam, 7 d (mm): CYA 15–21; CYA 30 C 27–33; CYA 37 C 25–30; MEA 20–23; MEA 30 C 33–35; DG18 5–7; CYAS No growth; OA 18–21; CREA No growth to micro-colonies; YES 17–21. Colony characters: CYAA 25 C, 7 d: Colonies low to moderately deep, raised at centre, plane; margins low, narrow, entire (2–3 mm); mycelia white, inconspicuously yellow; texture ﬂoccose; sporulation absent to sparse, conidia en masse greenish grey (28B2); soluble pigments absent; exudates absent; reverse POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 15. Morphological characters of Talaromyces apiculatus (CBS 312.59T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 219 YILMAZ ET AL. Fig. 16. Morphological characters of Talaromyces assiutensis (CBS 116554) A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on OA after 2 wk incubation. C. Ascomata D. Asci and ascopsores. E–G. Conidiophores. H. Ascospores. I. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–I. 220 POLYPHASIC pale orange to brownish orange (5A3–5C5). MEA 25 C, 7 d: Colonies low to moderately deep, raised at centre, plane; margins low, narrow, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse greenish grey (28B2); soluble pigments absent; exudates absent; reverse brownish yellow to brown (5C8–5E7). YES 25 C, 7 d: Colonies low to moderately deep, raised at centre, plane; margins low, narrow, entire (2 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse greenish grey (28B2); soluble pigments absent; exudates absent; reverse greyish yellow to greyish orange (4C5–5B3). DG18 25 C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (2A2). OA 25 C, 7 d: Colonies low, plane, young ascomata visible; margins subsurface, wide, entire (3 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent, in some orange coloured inside media in colony periphery; exudates absent; reverse pale yellow to light orange (1A3–5A4). CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and also sometimes monoverticillate; stipes smooth walled, 15–100 × 1.5–3.5 μm; metulae two to ﬁve, divergent, 12–17 × 1.5–3 μm; phialides acerose, three to six per metulae, 9.5–16 × 2–3 μm; conidia smooth, ovoidal to ellipsoidal, 2–4 × 1.5–2.5 μm. Ascomata maturing after 1–2 wk of incubation on OA at 25 and 30 C, creamish white to pale yellow, globose to subglobose, 250–600 × 250–550 μm, asci 7–11 × 5.5–8.5 μm, ascospores ellipsoidal, spiny, 3.5–5 × 2–3 μm. Extrolites: Talaromyces assiutensis produces glauconic acid (Frisvad et al. 1990a). Distinguishing characters: Talaromyces assiutensis produces creamish white ascomata with spiny, ellipsoidal ascospores (Fig. 16). Talaromyces intermedius also produces creamish white ascomata, but differs from T. assiutensis by faster growth at 25 C and no growth at 37 C. Also, ascospores of T. intermedius are bigger than T. assiutensis. Phylogenetically, T. assiutensis is closely related to T. trachyspermus. Talaromyces assiutensis differs from T. trachyspermus by slower growth at 37 C. Notes: Phylogenetically, T. gossypii (CBS 645.80) is identical to T. assiutensis (Fig. 5) and is considered a synonym. Talaromyces atricola S.W. Peterson & Jurjevic, PLoS ONE 8: e78084-page 8. 2013. MycoBank MB804733. Fig. 17. TAXONOMY OF TALAROMYCES Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (<1 mm), low, entire, plane; mycelium white; sporulation absent to sparse; exudates absent; soluble pigment absent; reverse yellowish white to yellowish grey (4A2–4A3). MEA 25 C, 7 d: Colonies low, plane; margins narrow (1 mm), low, entire, plane; mycelium white; sporulation sparse to dense; texture ﬂoccose; conidia en masse greyish green to dull green (26C3–26D3); exudates absent; soluble pigment absent; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (<1 mm), low, entire, plane; mycelium white; sporulation absent; exudates absent; soluble pigment absent; reverse light orange to greyish orange (5A5–5B5). DG18 25 C, 7 d: Colonies low, plane; margins narrow (<1 mm), low, entire, plane; mycelium white; sporulation sparse to dense; texture ﬂoccose; conidia en masse greyish green to dull green (26C3–26D3); exudates absent; soluble pigment absent; reverse yellowish white to yellowish grey (4A2–4A3). OA 25 C, 7 d: Colonies low, plane; margins narrow (2 mm), low, entire, plane; mycelium white; sporulation sparse to dense; texture ﬂoccose; conidia en masse greyish green to dull green (26C3–26D3); exudates absent; soluble pigment absent; reverse beige. CREA, 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate often having symmetrical subterminal branches; stipes smooth walled, 20–200 × 2.0–3.0 μm; with extra branches 10–30 μm; metulae three to six, divergent, 6–13 × 1.5–3.0 μm; phialides acerose, three to six per metulae, 8–11 × 2–3 μm; conidia smooth, ellipsoidal, with one end connection to sometimes fusiform, 2–5 × 2–5 μm. Ascomata not observed. Extrolites: Talaromyces atricola produces questin, rugulosin A, skyrin, mitorubrin, mitorubrinol, mitorubrinol acetate, 60 -hydroxy40 -hydroxy-30 -methoxy-(S)-mitorubrin 30 -methoxy-mitorubrin, and monomethyl-(S)-mitorubrin. Distinguishing characters: Talaromyces atricola is characterised by restricted growth on most media and produces ﬂoccose colonies that sporulate poorly (Fig. 17). It is most similar to T. rugulosus. However, the ﬂoccose colonies and poor sporulation is in contrast to the velvety and well sporulating colonies of T. rugulosus. In addition, conidia of T. atricola are more ellipsoidal than those of T. rugulosus and colonies generally grow more restrictedly. Talaromyces atroroseus Yilmaz et al., PLoS ONE 8: e84102-page 8. 2013. MycoBank MB804901. Fig. 18. In: Talaromyces section Trachyspermi ≡ Penicillium rugulosum var. atricolum Thom, Penicillia: 474. 1930. In: Talaromyces section Islandici Typus: Unknown, culture ex-type CBS 255.31 = NRRL 1052 = FRR 1052 = Thom 4640.439 = ATCC 52257 = DTO 278F1 = IBT 4489. ITS barcode: KF984859 (alternative markers: BenA = KF984566; CaM = KF984719; RPB2 = KF984948) Colony diam, 7 d (mm): CYA 10; CYA 30 C 7–8; CYA 37 C No growth; MEA 15; MEA 30 C 5–6; DG18 15; CYAS 7; OA 15–17; CREA 8; YES 12. www.studiesinmycology.org Typus: CBS H-21790, culture ex-type CBS 133442 = IBT 32470 = DTO 178-A4. ITS barcode: KF114747 (alternative markers: BenA = KF114789; CaM = KJ775418; RPB2 = KM023288) Colony diam, 7 d (mm): CYA 28–40; CYA 30 C 33–40; CYA 37 C 23–25; MEA 30–35; MEA 30 C 38–40; DG18 20–25; CYAS Generally no growth, in some isolates colonies up to 10; OA 20–25; CREA 2–5; YES 33–35. Colony characters: CYA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white; texture velvety and 221 YILMAZ ET AL. Fig. 17. Morphological characters of Talaromyces atricola (CBS 255.31T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 222 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 18. Morphological characters of Talaromyces atroroseus (CBS 133442T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bars: B = 50 μm; C = 10 μm, applies to D–H. www.studiesinmycology.org 223 YILMAZ ET AL. Colony diam, 7 d (mm): CYA 30–32; CYA 30 C 43–45; CYA 37 C 19–21; MEA 38–40; MEA 30 C 50–52; DG18 4–5; CYAS No growth; OA 35–40; CREA 5–6; YES 10–12. some parts ﬂoccose; sporulation dense, conidia en masse dark green (27E5); soluble pigments red; exudates absent; reverse reddish brown (9E8). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white; texture ﬂoccose and in some isolates velvety; sporulation moderately dense to dense, conidia en masse greyish green (25C5) to dark green (27E5) (at 30 C dense and dark green); soluble pigments red (in some strains lacking at 25 C but present at 30 C); exudates red droplets (in some strains lacking); reverse reddish brown (9E8). YES 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white; texture velvety and ﬂoccose; sporulation dense, conidia en masse greyish green to dark green (25E5–25F5); soluble pigments red (in some isolates absent); exudates absent; reverse brownish red (10D8) centre fading into red (10B8). DG18 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1 mm); mycelia white; texture velvety and ﬂoccose; sporulation moderately dense, conidia en masse dull green (25D4–26D4); soluble pigments absent; exudates absent; reverse greyish green (1C3) centre fading into greenish grey (1B2) and pale yellowish white (2A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white; texture velvety; sporulation moderately dense to dense, conidia en masse dark green (26F8); soluble pigments absent; exudates absent; reverse in some isolates centre red, in some isolates centre lacking and pale green. CREA 25 C, 7 d: Acid production absent. Colony characters: CYA 25 C, 7 d: Colonies low, plane, sterile white aerial mycelia, pale red appearance because of reverse colouring; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose and funiculose; sporulation absent; soluble pigments absent; exudates absent; reverse pastel red (7A4) fading into orange white (6A2). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–4 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse greyish orange (5B5–5B6). YES 25 C, 7 d: Colonies deep, raised, slightly sulcate, light orange ﬂuffy appearance because of the reverse colour; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A5–6A5) with pastel red (8A5) circle close to margins. DG18 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse orange white (5A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (4–5 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse greyish green (27C4–27D4); soluble pigments absent; exudates absent; reverse pale orange. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 90–150 × 2.5–3 μm; branches 15–50 μm; metulae three to six, divergent, 8–15 × 3–4 μm; phialides acerose, three to six per metulae, 9.5–12.5 × 2.5–3 μm; conidia ﬁnely rough to rough, ellipsoidal, 2–3.5 × 1.5–2.5 μm. Ascomata not observed. Micromorphology: Conidiophores sporulation is best on OA after 2 wk incubation. Conidiophores biverticillate; stipes smooth walled, 50–100 × 2–3 μm; metulae two to ﬁve, divergent, 8–16 × 2.5–3.5 μm; phialides acerose, two to six per metulae, 10–17.5 × 2–3 μm; conidia smooth, cylindrical to ellipsoidal, 3–5 × 1.5–2.5 μm. Ascomata not observed. Extrolites: Talaromyces atroroseus produces glauconic acid, glaucanic acid, Monascus red azaphilone pigments, purpactins AC, purpuride, purpurogenone and ZG-1494α (Frisvad et al. 2013). Extrolites: Talaromyces aurantiacus produces duclauxin and mitorubrinic acid. Distinguishing characters: Talaromyces atroroseus is characterised by soluble red pigment and very dark green conidia on OA (Fig. 18). Talaromyces atroroseus produces glauconic acid, purpuride and ZG-1494α (Frisvad et al. 2013). Other species producing soluble red pigment include T. minioluteus, T. purpurogenus and T. albobiverticillius. Talaromyces atroroseus grows well on CYA at 37 C, in contrast to poor growth for T. albobiverticillius and T. minioluteus. Talaromyces atroroseus resembles T. purpurogenus, but differs from the latter by ﬁnely rough to rough, dull to dark green, thick walled, ellipsoidal conidia. Talaromyces aurantiacus (J.H. Mill., Giddens & A.A. Foster) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. MycoBank MB560642. Fig. 19. ≡ Penicillium aurantiacum J.H. Mill., Giddens & A.A. Foster, Mycologia 49: 797. 1957. Distinguishing characters: Talaromyces aurantiacus was considered synonymous with T. funiculosus by Pitt (1980), who considered it as nutritionally deﬁcient because of the poor growth on CYA. In our study we observed that T. aurantiacus grows restrictedly on YES and DG18. Micromorphologically, T. aurantiacus closely resembles T. funiculosus. However, acid production on CREA and strongly funiculose colonies on MEA and OA of T. funiculosus, distinguish T. funiculosus from T. auriantiacus. Talaromyces austrocalifornicus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 245. 1993. MycoBank MB361182. Fig. 20. ≡ Penicillium austrocalifornicum Yaguchi & Udagawa, (simultaneously published). In: Talaromyces section Trachyspermi Typus: CBM-PF 1117, culture ex-type CBS 644.95 = IBT 17522. In: Talaromyces section Talaromyces Typus: No. 1736 (A.A. Foster), culture ex-type CBS 314.59 = ATCC 13216 = IMI 099722 = NRRL 3398. ITS barcode: JN899380 (alternative markers: BenA = KF741917; CaM = KF741951) 224 ITS barcode: JN899357 (alternative markers: BenA = KJ865732; CaM = KJ885261) Colony diam, 7 d (mm): CYA 10–11; CYA 30 C 10–11; CYA 37 C 10–11; MEA 16–18; MEA 30 C 23–24; DG18 8; CYAS No growth; OA 25; CREA No growth; YES 18–20. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 19. Morphological characters of Talaromyces aurantiacus (CBS 314.59T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 225 YILMAZ ET AL. Fig. 20. Morphological characters of Talaromyces austrocalifornicus (CBS 645.95T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. 226 POLYPHASIC Colony characters: CYA 25 C, 7 d: Colonies raised, sunken at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse orange white to pale orange (5A2–5A3). MEA 25 C, 7 d: Colonies deep, plane, sterile white mycelia; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates orange droplets; reverse greyish orange (5B5–5B6). YES 25 C, 7 d: Colonies raised at centre, slightly sulcate, white appearance; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange to light orange (5A3–5A4). DG18 25 C, 7 d: Colonies raised at centre, plane; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates small clear droplets; reverse yellowish white (3A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–5 mm); mycelia white; sporulation absent; soluble pigments absent; exudates clear droplets; reverse brownish. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 30–100 × 2–3 μm; metulae three to eight, divergent, 8–17 × 2–3 μm; phialides acerose, three to eight per metulae, 7–12 × 1.5–2.5 μm; conidia smooth, subglobose, 1.5–3 × 1.5–2.5 μm. Ascomata not observed in our culture but ﬁde Yaguchi et al. (1993) maturing after 2–3 wk of incubation, yellow, globose to subglobose, 360–480 μm, asci 6–8 × 5–6 μm, ascospores broadly ellipsoidal, ﬁnely spiny, 2.5–3.5 × 2–2.5 μm. Extrolites: Talaromyces austrocalifornicus produces altenuene, mitorubrin, mitorubrinic acid and a purpactin. Distinguishing characters: Talaromyces austrocalifornicus is characterised by restricted growth on CYA and MEA, no growth on CREA, yellow ascomata that have small, broadly ellipsoidal, spiny ascopores and biverticillate conidiophores producing smooth subglobose conidia (Fig. 20). All these characters differentiate T. austrocalifornicus from other species that produce yellow ascomata wth ellipsoidal, spiny ascospores. The ex-type strain of T. austrocalifornicus was investigated during our study. Unfortunately, the ex-type strain was degenerated and did not produce ascomata. Talaromyces bacillisporus (Swift) C. R. Benj., [as ‘bacillosporus’], Mycologia 47: 682. 1955. MycoBank MB118745. Fig. 21. ≡ Penicillium bacillisporum Swift, Bull. Torrey Bot. Club 59: 221. 1932. TAXONOMY OF TALAROMYCES (30A5–30A6); margins low, plane, entire (1–2 mm); mycelia white and yellowish green; sporulation absent to sparse; soluble pigments absent; exudates absent; reverse centre dull green (28F4) fading into greyish yellow to olive brown (4C5–4D5) in some isolates lack of the centre colour; at 30 and 37 C reverse dark green (25F4). MEA 25 C, 7 d: Colonies raised at centre, slightly sunken at centre slightly sulcate; margins low, plane, entire (1–2 mm); mycelia white and yellow; texture ﬂoccose; sporulation sparse to moderately dense, conidia en masse dull green (25D4–25E4); soluble pigments absent; exudates absent; reverse brown (6E6) centre fading into caramel brown (6C6) and at 30 C reverse dark green (25F4). YES 25 C, 7 d: Colonies raised at centre, slightly sunken at centre, slightly sulcate; margins low, plane, entire (1–2 mm); mycelia white and yellow; texture ﬂoccose; sporulation sparse, conidia en masse greyish green (28C4); soluble pigments absent; exudates absent; reverse greyish yellow (4B5–4C5) centre fading into pale yellow (4A3). DG18 25 C, 7 d: Colonies raised at centre, pale; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pale yellow (4A3). OA 25 C, 7 d: Colonies low, plane, production of ascomata especially at 30 C; margins low, plane, entire (2 mm); mycelia white and pale pastel yellow; texture ﬂoccose; sporulation sparse, conidia en masse dull green (26D4); soluble pigments absent; exudates small clear droplets; reverse dark brown centre and at 30 C very dark green reverse. CREA 25 C, 7 d: Generally no growth colonies up to 3 mm acid production absent. Micromorphology: Conidiophores mostly monoverticillate and biverticillate with a minor proportion having subterminal branches; stipes rough-walled, 20–60 × 2–3.5 μm; branches 15–25 μm; metulae three to six, divergent, 9.5–13 × 3–3.5 μm; phialides acerose, two to ﬁve per metulae, 7–12 × 2–4 μm; conidia cylindrical, rod-shaped, occasionally ellipsoidal, 3–5(–6.5) × 1–2 μm (2.5–4 × 1.5–2.5 μm). Ascomata maturing after 1–2 wk of incubation on OA at 25 C and abundantly at 30 C, creamish white and longer incubation pastel orange, globose to subglobose, 80–150 μm, asci 8–12 × 9–14 μm, ascospores globose, spiny, 3.5–5 × 3.5–5 μm. Extrolites: Talaromyces bacillisporus produces pinselin and bacillisporin A-E (Yamazaki & Okuyama 1980, Dethoup et al. 2006) and talarotoxin (Ishii et al. 1995). Distinguishing characters: Talaromyces bacillisporus can easily be distinguished from other species by its globose, spiny ascospores, dark green reverse on CYA and MEA at 30 and 37 C, roughwalled stipes and cylindrical, rod-shaped conidia (Fig. 21). In: Talaromyces section Bacillispori Talaromyces bohemicus (Fassat. & Peckova) Yilmaz, Frisvad & Samson, comb. nov. MycoBank MB809554. Typus: CBS H-7813, culture ex-type CBS 296.48 = ATCC 10126 = IMI 040045 = NRRL 1025. Basionym: Sagenomella bohemica Fassat. & Peckova, Ceska Mykol. 44: 240. 1990. ITS barcode: KM066182 (alternative markers: BenA = AY753368; CaM = KJ885262; RPB2 = JF417425) In: Talaromyces section Helici Colony diam, 7 d (mm): CYA 10–15; CYA 30 C 25–30; CYA 37 C 33–37; MEA 15–20; MEA 30 C 35–38; DG18 3–7; CYAS No growth; OA 18–22; CREA Generally no growth in some strains colonies up to 4; YES 10–15. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly sulcate, colony appearance light green to yellowish green www.studiesinmycology.org Typus: Unknown, culture ex-type CBS 545.86 = CCF 2330 = IAM 14789. ITS barcode: JN899400 (alternative markers: BenA = KJ865719; CaM = KJ885286, RPB2 = JN121532) Colony diam, 10 d (mm): Fide Fassatiova & P^eckova (1990), MEA 40 mm. 227 YILMAZ ET AL. Fig. 21. Morphological characters of Talaromyces bacillisporus (CBS 298.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B Colony reverse on CYA after 1 wk incubation at 30 C. C. Colony texture and ascomata on OA after 2 wk incubation. D. Asci and ascopsores. E–G. Conidiophores. H. Ascospores. I. Conidia. Scale bars: C = 500 μm; D = 10 μm, applies to E–I. 228 POLYPHASIC Colony characters: Fide Fassatiova & P^eckova (1990) after 10 d of cultivation on MEA the colonies reach 4 cm diam., are very low and thin, mycelium is light brown, later cinnamon brown. The colonies grow in concentric zones. The centre of the colony is later tomentose, forming pink brown to red brown hyphae joining into narrow funicles. Three to four weeks later a heavy sporulation, grey green conidia; reverse at ﬁrst red brown and later diffusing into surrounding agar conspicuously colourating it. Micromorphology: Fide Fassatiova & P^eckova (1990), monoverticillate with extra branching up to 20 μm; phialides acerose, 13–25(–30) × 2.5–3 μm; conidia fusiform, encrusted cell wall, 7–9 × 2.6–3 μm. Notes: The ex-type strain of Talaromyces bohemicus was not in good condition and could not be studied. The description is based on Fassatiova & P^eckova (1990). Talaromyces boninensis (Yaguchi & Udagawa) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. MycoBank MB560643. ≡ Talaromyces helicus var. boninensis Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 33: 511. 1992. TAXONOMY OF TALAROMYCES it differs from T. helicus by the production of green ascomata and broadly ellipsoidal and spiny ascospores. Unfortunately the extype strain was unavailable and could not be investigated during our study. Based on Frisvad (unpublished data), T. boninensis (strain PF 1103 = IBT 17516): CYA 21–25 mm, no (poor) sporulation, cream yellow to light brown reverse (25 C, 7 d); MEA 25–28 mm, cream yellow reverse; YES 23–27 mm, yellow cream reverse, CREA weak growth (12 mm), weak acid production; CYA (at 37 C): 17–29 mm; OA (25 C) 32–33 mm, ascomata and T. boninensis (strain PF 1110 = IBT 17521): CYA 20 mm; MEA 31 mm; YES 23 mm; OA 32 mm, ascomata; CREA weak growth (13 mm), some acid production; CYA at 37 C 27 mm. Talaromyces brunneus (Udagawa) Samson, Yilmaz & Frisvad, Stud. Mycol. 70: 175. 2011. MycoBank MB560644. Fig. 22. ≡ Penicillium brunneum Udagawa, J. Agric. Sci. Tokyo Nogyo Daig. 5: 16. 1959. In: Talaromyces section Islandici Typus: NHL 6054, culture ex-type CBS 227.60 = ATCC 18229 = FRR 646 = IFO 6438 = IHEM 3907 = IMI 078259 = MUCL 31318 = IBT 4490. In: Talaromyces section Helici Typus: CBM PF-1103, culture ex-typus CBS 650.95 = IBT 17516. ITS barcode: JN899356 (alternative markers: BenA = KJ865721; CaM = KJ885263; RPB2 = KM023276) Colony diam, 7 d (mm): Fide Yaguchi et al. (1992), CYA 28; MEA 30; OA 32. Colony characters: Fide Yaguchi et al. (1992) colonies on CYA velvety, radially sulcate, consisting of a close basal felt in which ascomata slowly develop, reddish white to pastel green (7A2–29A4); sporulation limited; reverse light orange (5A4). At 37 C, mycelial growth somewhat rapid, but ascomata and conidia less formed. Colonies on MEA ﬂoccose, more or less plane, consisting of a thin basal felt, white to orange white (6A2), then becoming green to grayish green (28A6–28B6); ascomata and conidia not sufﬁciently produced to inﬂuence the colony appearance within 7 d; reverse uncoloured. Colonies on OA ﬂoccose to funiculose, consisting of a rather thin basal felt, at ﬁrst pale orange (5A3), then grayish green (28B6); ascomata abundantly produced, but conidia limited in number; reverse uncoloured to pale orange (5A3). Micromorphology: Fide Yaguchi et al. (1992) conidiophores biverticillate; stipe ﬁnely rough, 25–260 × 2.5–4 μm; metulae four to ten, divergent, 10–16(–20) × 2.5–3(–3.5); phialides two to six, acerose, 10–15 × 2–3.5 μm; conidia ellipsoidal to fusiform, sometimes globose, smooth, 2–4 × 1.5–2.5 μm. Ascomata ripening 14–21 d, grayish green, globose to subglobose, 280–550 × 240–480 μm; asci 8–10(–12) × 7–8 μm; ascospores broadly ellipsoidal, spiny, (2.5–)3.5–4(4.5) × 2–3 μm. Extrolites: Talaromyces boninensis produces talaroderxines. Notes: Talaromyces boninensis was described as a variety of T. helicus. Molecular results show T. boninensis as a unique species. Its macrmorphology is smilar to T. helicus (Yaguchi et al. 1992), but www.studiesinmycology.org ITS barcode: JN899365 (alternative markers: BenA = KJ865722; CaM = KJ885264; RPB2 = KM023272) Colony diam, 7 d (mm): CYA 19–20; CYA 30 C 18–20; CYA 37 C No growth; MEA 17–19; MEA 30 C 15–18; DG18 14–15; CYAS 6–10; OA 14–15; CREA 9–10; YES 24–25. Colony characters: CYA 25 C, 7 d: Colonies slightly raised, radially sulcate; margins low, plane, entire (1 mm); mycelia white and yellow; texture velvety and loosely funiculose and especially in the centre conidiophores born from sterile aerial hyphae; sporulation moderately dense, conidia en masse golden brown to yellowish brown (5D7–5E7); soluble pigments absent; exudates clear and yellow droplets; reverse yellowish brown (5E5) centre fading into golden yellow (5B7). MEA 25 C, 7 d: Colonies slightly raised, sulcate; margins low, plane, entire (1 mm); mycelia white and yellow; texture velvety and in the centre ﬂoccose; sporulation moderately dense, conidia en masse golden brown to yellowish brown (5D7–5E7); soluble pigments absent; exudates absent; reverse brown (6D7). YES 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white and yellow; sporulation absent; soluble pigments absent; exudates very small clear droplets; reverse greyish orange (5B5). DG18 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white, yellow; sporulation absent; soluble pigments absent; exudates absent; reverse orange (5A7) centre fading into light yellow (4A5). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1 mm) it does not have the transparent zone; mycelia white and yellow; texture velvety; sporulation dense, conidia en masse yellowish brown (5E5) centre fading into golden yellow (5B7); soluble pigments absent; exudates orange and clear droplets; reverse light orange. CREA 25 C, 7 d: Acid production. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 229 YILMAZ ET AL. Fig. 22. Morphological characters of Talaromyces brunneus (CBS 227.60T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 230 POLYPHASIC 60–130 × 2.5–4 μm; sometimes extra branches 15–25 μm; metulae three to six, 8–15(–18) × 2–4 μm; phialides acerose, three to ﬁve per metulae, 9–14 × 2–3.5 μm; conidia smooth, globose to subglobose 3–4(–7) × 2–4. Ascomata not observed. Extrolites: Talaromyces brunneus produces emodin, skyrin and (+)-rugulosin (Shibata & Ikekawa 1963, Shibata & Udagawa 1963, Sankawa et al. 1973, Seo et al. 1973). We detected rugulosin, skyrin, mitorubrin, mitorubrinol and mitorubrinol acetate and one tetracyclic compound in CBS 227.60, and in addition to several speciﬁc not yet structure elucidated extrolites. Distinguishing characters: Talaromyces brunneus is characterised by the producton of golden brown to yellowish brown conidia on CYA, MEA and OA (Fig. 22). The species is unable to grow at 37 C. These characters distinguish it from all other Talaromyces species. Talaromyces calidicanius (J.L. Chen) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. MycoBank MB560645. Fig. 23. ≡ Penicillium calidicanium J.L. Chen, Mycologia 94: 870. 2002. In: Talaromyces section Talaromyces Typus: CFC-7 (isotype TNM F12246), culture ex-type CBS 112002. ITS barcode: JN899319 (alternative markers: BenA = HQ156944; CaM = KF741934; RPB2 = KM023311) Colony diam, 7 d (mm): CYA 27–30; CYA 30 C 33–35; CYA 37 C No growth; MEA 47–48; MEA 30 C 30; DG18 15; CYAS No growth; OA 45; CREA 18–20; YES 40–41. C, 7 d: Colonies raised at centre, Colony characters: CYA 25 sulcate, young synnemata produced; margins low, plane, entire (<1 mm); mycelia white and pastel yellow; texture funiculose and ﬂoccose; sporulation moderately dense, conidia en masse greyish green (28D5–28E5); soluble pigments absent; exudates absent; reverse greyish yellow to olive brown (4C5–4D5). MEA 25 C, 7 d: Colonies raised at centre, sulcate, synnemata at centre sporulate, and young synnemata at margins; margins low, plane, entire (3 mm); mycelia white; texture synnematous, loosely funiculose to ﬂoccose; sporulation sparse to moderately dense, conidia en masse dark green (27F5) on the sporulated synnemata and on the rest of the colony greyish green (27D4); soluble pigments absent; exudates orange droplets; reverse golden brown to yellowish brown (5D7–5E7). YES 25 C, 7 d: Colonies deep, raised at centre, sulcate, young synnemata formation; margins low, plane, entire (2 mm); mycelia white and yellow; texture funiculose; sporulation moderately dense, conidia en masse greyish green (27E5–28E5); soluble pigments absent; exudates absent; reverse olive brown (4D8–4E8) centre fading into golden brown (4C6–4D6). DG18 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; texture velvety; sporulation dense, conidia en masse greyish green (26E5–27E5); soluble pigments absent; exudates small orange droplets; reverse golden brown (4C6) fading into pale yellow (2A3). OA 25 C, 7 d: Colonies low, www.studiesinmycology.org TAXONOMY OF TALAROMYCES plane, sporulated synnemata at centre and young synnemata at margins; margins low, plane, entire (4–5 mm); mycelia white; texture velvety; sporulation dense, conidia en masse dark green (28F5); soluble pigments absent; exudates small clear droplets; reverse pale brownish green. CREA 25 C, 7 d: Moderate acid production. Micromorphology: Synnemata 2000–6000 μm long. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 40–150(–250) × 2.8–4 μm; branches up to 20 μm; metulae two to six, divergent, 7–15 × 2.5–4 μm; phialides acerose, three to six per metulae, 8–13 × 2–3.5 μm; conidia ﬁnely rough to rough or spiral striate, ellipsoidal to fusiform, 2.5–4.5 × 2–3 μm. Ascomata not observed. Extrolites: Talaromyces calidicanius produces duclauxin and related compounds. Distinguishing characters: Talaromyces calidicanius produces synnemata after 1 wk of incubation (Fig. 23). It is characterised by fast growing colonies on MEA and grows well at 30 C. Determinate synnemata, up to 6 000 μm long, are produced after prolonged incubation. Talaromyces calidicanius produces conidia with spiral striations. Its morphology resembles T. panamensis, but T. panamensis grows slower at 30 C on MEA and produces more acid on CREA. Talaromyces cecidicola (Seifert, Hoekstra & Frisvad) Samson et al., Stud. Mycol. 71: 175. 2011. MycoBank MB560646. Fig. 24. ≡ Penicillium cecidicola Seifert, Hoekstra & Frisvad, Stud. Mycol. 50: 520. 2004. In: Talaromyces section Purpurei Typus: DAOM 233329, culture ex-type CBS 101419 = DAOM 233329. ITS barcode: AY787844 (alternative markers: BenA = FJ753295; CaM = KJ885287; RPB2 = KM023309) Colony diam, 7 d (mm): CYA 33–34; CYA 30 C 37–38; CYA 37 C 2–3; MEA 37–38; MEA 30 C 38–39; DG18 21–22; CYAS No growth; OA 31–33; CREA 6–8; YES 30–31. Colony characters: CYA 25 C, 7 d: Colonies low, raised at centre, plane; margins low, wide, entire (3–5 mm); mycelia white; texture ﬂoccose and strongly funiculose; sporulation moderately dense, conidia en masse dark green (26F7); soluble pigments absent; exudates absent; reverse very dark red at centre, dark brown to violet brown (8F8–10F8). MEA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (5 mm); mycelia white; texture strongly funiculose with some velvety areas; sporulation moderately dense to dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent; reverse dark brown (7F8–8F8). YES 25 C, 7 d: Colonies low, centrally raised surrounded by sunken in ridge; margins low, wide, entire (3 mm); mycelia white; texture funiculose, with some ﬂoccose areas; sporulation moderately dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent; reverse very dark red at 231 YILMAZ ET AL. Fig. 23. Morphological characters of Talaromyces calidicanius (CBS 112002T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 1 wk incubation. C–F. Conidiophores. G. Conidia. Scale bars: B = 1000 μm; C = 50 μm; D = 10 μm, applies to E–G. 232 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 24. Morphological characters of Talaromyces cecidicola (CBS 101419T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 1 wk incubation. C. Synnemata. D–F. Conidiophores. G. Conidia. Scale bars: B, C = 1000 μm; D = 10 μm, applies to E–G. www.studiesinmycology.org 233 YILMAZ ET AL. centre, dark brown to violet brown (8F8–10F8). DG18 25 C, 7 d: Colonies slightly raised in the centre, plane; margins low, narrow, entire (2–3 mm); mycelia white; texture funiculose; sporulation moderately dense, conidia en masse dark green (26F5); soluble pigments absent; exudates absent; reverse brown (7E8) at centre, olive (3D3) near margin. OA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (3–4 mm); mycelia white; texture strongly funiculose, with some velvety areas; sporulation moderately dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent; reverse grey to brownish grey (7B1–7C2). CREA 25 C, 7 d: Acid production absent. Micromorphology: Synnemata 250–1250 μm long. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 20–80(–200) × 2.5–4 μm; branches 8–25 μm; metulae three to eight, divergent, 9–15 × 2.5–3 μm; phialides acerose, three to six per metulae, 10–15 × 1.5–3 μm; conidia smooth to ﬁnely rough, ellipsoidal, 2.5–4 × 1.5–3 μm. Ascomata not observed. Distinguishing characters: Talaromyces cecidicola is characterised by colonies with dark red reverses and synnemata up to 1 250 μm long, which are produced after 1–2 wk of incubation (Fig. 24). It is most similar to T. coalescens. However, T. coalescens grows faster than T. cecidicola on MEA at 30 C. This species was isolated from insect galls (Seifert et al. 2004). Talaromyces chloroloma Visagie & K. Jacobs, Persoonia 28: 18. 2012. MycoBank MB564326. Fig. 25. In: Talaromyces section Purpurei Typus: PREM 60033, culture ex-type DAOM 241016 = CV 2802. ITS barcode: FJ160273 (alternative markers: BenA = GU385736; CaM = KJ885265; RPB2 = KM023304) Colony diam, 7 d (mm): CYA 40–45; CYA 30 C 50–55; CYA 37 C Generally no growth, sometimes colonies grow up to no 4; MEA 45–48; MEA 30 C 55–60; DG18 20–25; CYAS 20–23; OA 40–48; CREA 15–20; YES 35–40. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, lightly sulcate; margins low, plane, entire (2 mm); mycelia white; texture ﬂoccose, with some funicles present; sporulation moderately dense to dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates in some isolates absent, in some isolates small clear droplets; reverse centre greyish orange (5B3), fading into dull greyish yellow (3B3). MEA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (4–5 mm); mycelia white; texture ﬂoccose and loosely funiculose (especially in the centre); sporulation dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates very small clear droplets; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies low, radially sulcate; margins low, narrow, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent; reverse in some 234 isolates centre greyish red (9B6) to dull red (9B4) in some isolates lack of red centre, the rest pastel yellow (3A4). DG18 25 C, 7 d: Colonies low, plane; margins low, narrow, entire (2 mm); mycelia white; texture funiculose; sporulation absent to dense (no sporulation at centre, centre is slimy yeast like structure), conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent; reverse greyish green to dull green (27C4–27D4), in DTO 180F3 margins orange red (8A6). OA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (5–6 mm); mycelia white, especially in the centre synnemata like aerial hyphae growing; texture velvety and funiculose; sporulation moderately dense to dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates clear droplets in centre; reverse pale light green. (1B3). CREA 25 C, 7 d: Moderate acid production. Micromorphology: Synnemata produced on CYA where the colony surface scraped away after 2–3 wk, 700–1200 μm long. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 10–75 × 2.5–5 μm; branches 5–20 μm; metulae three to six, divergent, 7.5–10(–13) × 2–4 μm; phialides acerose, three to six per metulae, 7–11 × 1.5–3 μm; conidia smooth, ellipsoidal, 2.5–4(–6) × 1.5–2.5 μm. Ascomata not observed. Distinguishing characters: Talaromyces chloroloma grows rapidly on most media and produces synnemata after 2–3 wk, up to 1 250 μm long and similar to those of T. cecidicola and T. coalescens. However, T. chloroloma grows much faster on MEA at 30 C and produces acid on CREA, which is absent in its close relatives. Acid production also distinguishes T. choloroloma from T. ramulosus. Talaromyces cinnabarinus (S.C. Jong & E.E. Davis) Yilmaz, Samson & Frisvad. comb. nov. MycoBank MB809557. Basionym: Paecilomyces cinnabarinus S.C. Jong & E.E. Davis, Mycologia 67: 1144. 1975. = Aphanoascus cinnabarinus var. macrosporus Udagawa, Bull. Nat. Sci. Mus. Tokyo 16: 503. 1973. In: Talaromyces section Helici Typus: CBS H-6686, culture ex-type CBS 267.72 = ATCC 26215 = NHL 2673. ITS barcode: JN899376 (alternative markers: BenA = AY753377; CaM = KJ885256; RPB2 = JN121477) Colony characters: Fide Udagawa & Takada (1973) colonies on OA growing rapidly, plane, consisting of a thin mycelial felt in which scattered cleistothecia are embedded, surface appearing slightly ﬂocculent, light orange to reddish orange, conidial structures not produced; reverse pale reddish orange to dark brown. On malt agar spreading, orange to reddish orange, more or less ﬂoccose, ripening process of cleistothecia extremely delayed; colony reverse dark green to dark blue-green with surrounding agar similarly coloured. At 37 C grows slower than at 25 C, with production of fructiﬁcation much reduced and immatured. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 25. Morphological characters of Talaromyces chloroloma (DAOM 241016T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 2 wk incubation. C. Synnemata. D–G. Conidiophores. H. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–H. www.studiesinmycology.org 235 YILMAZ ET AL. Micromorphology: Fide Jong & Davis (1975) conidiophores short, smooth walled, up to 50 μm long; phialides two to four, 13.5–17.1 × 2.7–3.2 μm, smooth walled, consisting of a more or less cylindrical basal portion, tapering abruptly to a long thin neck 4.5–6.3 × 0.9–1.2 μm; conidia cylindrical with truncate ends, smooth walled, 5.4–7.2 × 1.5–2 μm. Fide Udagawa & Takada (1973) ascomata yellowish orange to orange red, subglobose to ovoidal, 350–600 μm; asci 9.5–11 μm; ascospores broadly ellipsoidal, thick walled, ornamented by irregular ridges, 4–5.5(–6) × 3–3.5 μm. Distinguishing characters: Fide Udagawa & Takada (1973) T. cinnabarinus is characterised by the production of its globose, red, non-ostiolate ascocarps, which are surrounded by loose wefts of encrusted hyphae and with a sclerotoid inner tissue, irregularly disposed globose asci, and ellipsoid, hyaline to reddish orange ascospores, ornamented with several narrow ridges. Notes: Aphanoascus cinnabarinus was described by Zukal (1890). The holotype and the ex-type strains were unavailable and Udagawa & Takada (1973) neotypiﬁed the species with CBS 267.72. However, this strain produced a Paecilomyces anamorph, which was described by Jong & Davis (1975) as Paecilomyces cinnabarinus, whereas Zukal's original description and illustrations clearly describes the structures of a Chrysosporium anamorph (Stolk & Samson 1983). Stolk & Samson (1983) placed Chromocleista cinnabaria (as A. cinnabarinus sensu Udagawa & Takada) in the Eurotiales. However, Ogawa & Sugiyama (2000), Houbraken & Samson (2011) and our results showed that Aphanoascus cinnabarinus belongs to Talaromyces and is classiﬁed here in Talaromyces section Helici. As such, we introduce the new combination T. cinnabarinus based on Paecilomyces cinnabarinus and consider A. cinnabarinus Zukal sensu Udagawa & Takada (J. Jpn. Bot. 48: 23. 1973) as a pseudonym. Aphanoascus cinnabarinus var. macrosporus is considered a synonym of T. cinnabarinus, based on the original description by Udagawa et al. (1973). The ex-type strain (NHL 2704) was, however, not examined in our study. Talaromyces cnidii S.H. Yu, T.-J. An & H. Sang, J. Microbiol. 51: 707. 2013. MycoBank MB804809. Fig. 26. In: Talaromyces section Talaromyces Typus: KACC 46617, culture ex-type KACC 46617. ITS barcode: KF183639 (alternative markers: BenA = KF183641; CaM = KJ885266; RPB2 = KM023299) Colony diam, 7 d (mm): CYA 25–30; CYA 30 C 30–37; CYA 37 C 20–28; MEA 38–40; MEA 30 C 45–50; DG18 10–16; CYAS 3–4; OA 30–35; CREA 9–15; YES 25–30. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1–2 mm); mycelia white and dull red; texture ﬂoccose; sporulation moderately dense to dense, conidia en masse greyish green (26C3–26C4) to dull green (26D3–26D4); soluble pigments red (DTO 270-B7 and DTO 303-E1) to yellow (DTO 270-A4 and DTO 270-A8) (at 30 C yellow to orange); exudates in some isolates absent, in some isolates small red droplets (DTO 303-E1 and 236 DTO 270-B7); reverse centre brownish violet (11D8) to violet brown (11E8) fading into vivid red (11A8) and orange red (8A8). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly concentrically sulcate; margins low, plane, entire (2–3 mm); mycelia white and in some isolates also yellow (DTO 303-E1); texture velvety and ﬂoccose especially at centre in some isolates just ﬂoccose; sporulation sparse (DTO 270-A8 and DTO 269-I2) to dense, conidia en masse dull green (26D4–26E4); soluble pigments absent; exudates in some isolates red (DTO 303-E1) and orange (DTO 270-B7) droplets in some isolates absent; reverse violet brown (11E8–11F8) centre fading into orange red (8A8) and light orange (5A5) (in some isolates lack of the centre pigmentation). YES 25 C, 7 d: Colonies raised at centre, in some isolates sunken at centre, slightly sulcate; margins low, plane, entire (2 mm); mycelia white (in some isolates red and yellow as well (DTO 270-B7); texture velvety and ﬂoccose; sporulation moderately dense to dense, conidia en masse dull green (26D4–26E4); soluble pigments in some isolates red (DTO 270-B7 and DTO 303-E1) and in some absent; exudates absent; reverse brownish red to violet red (10D8–10E8) centre fading into light yellow (4A4) and in some isolates centre reddish brown (8D6) to greyish red (8B6) and in some isolates brown (6E5). DG18 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white; texture velvety (DTO 303-E1) and in some isolates ﬂoccose; sporulation sparse and in some isolates dense (DTO 303-E1), conidia en masse dull green (26E4–27E4) (DTO 303-E1) and in some isolates greyish green (25D5–25E5); soluble pigments absent (except DTO 303-E1, red pigment); exudates absent (except DTO 303-E1 orange and red small droplets); reverse in some isolates (DTO 303-E1 and DTO 270-A4) violet brown (10E8) to reddish brown (10D8) centre fading into reddish orange (7A6) and in some isolates lack of centre colour and greyish red (8B5) (DTO 270-A8) and in some isolates pale orange (5A3). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture velvety and in the centre ﬂoccose; sporulation moderately dense to dense, conidia en masse dull green (27E4); soluble pigments absent; exudates absent; reverse brownish orange. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 60–230 × 2–3 μm; branches 15–40 μm; metulae three to eight, divergent, 10–15 × 2.5–4 μm; phialides acerose, three to six per metulae, 10–15 × 2–3.5 μm; conidia smooth, ellipsoidal, 2.5–4 × 2–2.5 μm. Ascomata not observed. Distinguishing characters: Talaromyces cnidii produces a red reverse on CYA and MEA (Fig. 26). It is phylogenetically closely related to T. siamensis. There are only minor morphological differences between these two species. Talaromyces cnidii grows slightly faster at 25 C and 37 C, but BenA sequences are necessary for a deﬁnitive identiﬁcation. Talaromyces coalescens (Quintan.) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. MycoBank MB560647. Fig. 27. ≡ Penicillium coalescens Quintan., Mycopathologia 84: 115. 1984. In: Talaromyces section Purpurei POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 26. Morphological characters of Talaromyces cnidii (KACC 46617T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 237 YILMAZ ET AL. Fig. 27. Morphological characters of Talaromyces coalescens (CBS 103.83T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on MEA after 2 wk incubation. C. Synnemata. D–G. Conidiophores. H. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–H. 238 POLYPHASIC TAXONOMY OF TALAROMYCES Typus: CBS 103.83, culture ex-type CBS 103.83. Typus: BPI 892668, culture ex-typus NRRL 58811. ITS barcode: JN899366 (alternative markers: BenA = JX091390; CaM = KJ885267; RPB2 = KM023277) ITS barcode: KJ865739 (alternative markers: BenA = KF196843; CaM = KJ885288; RPB2 = KM023270) Colony diam, 7 d (mm): CYA 32–34; CYA 30 C 40–41; CYA 37 C 2–4; MEA 43–45; MEA 30 C 45–46; DG18 19–20; CYAS No growth; OA 35–38; CREA 5–6; YES 34–35. Colony diam, 7 d (mm): CYA 13–14; CYA 30 C 30–31; CYA 37 C 42–43; MEA 20–21; MEA 30 C 32–33; DG18 9–11; CYAS 2–3; OA 22–23; CREA 7–8; YES 18–20. Colony characters: CYA 25 C, 7 d: Colonies low to moderately deep, raised at centre, plane; margins low, wide, entire (3 mm); mycelia white and yellow; texture ﬂoccose and strongly funiculose; sporulation moderately dense at centre, conidia en masse dark green (26F5); soluble pigments red; exudates absent; reverse dark brown to violet brown (8F8–10F8). MEA 25 C, 7 d: Colonies low, plane; margins low, wide, entire 5 mm); mycelia white to beige; texture strongly funiculose; sporulation moderately dense only at colony centre, conidia en masse dark green (26F5); soluble pigments red; exudates absent; reverse very dark red. YES 25 C, 7 d: Colonies low, plane; margins low, wide, entire (4 mm); mycelia white; texture funiculose, with some ﬂoccose areas, abundant aerial hyphae; sporulation moderately dense at colony centre, conidia en masse dark green (26F5); soluble pigments red; exudates absent; reverse very dark red. DG18 25 C, 7 d: Colonies low, plane; margins low, narrow, entire (2 mm); mycelia white; texture loosely funiculose; sporulation sparse, conidia en masse cannot be determined; soluble pigments red; exudates absent; reverse brown (7E7) at centre, fading into pale red (7A3). OA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (4 mm); mycelia white; texture strongly funiculose, with some velvety areas; sporulation moderately dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates clear droplets at centre; reverse brownish grey to greyish brown (5C2–5D3). CREA 25 C, 7 d: Very weak acid production. Colony characters: CYA, 25 C, 7 d: Colonies 11–12 mm, slightly raised in the centre, slightly sulcate; margins wide (2 mm), low, entire, plane; mycelium orange and white, colony appearance orange (6B7–6C7); sporulation absent; exudates absent; soluble pigment very pale light brownish orange around colonies; reverse dark brown fading into beige (8F8). MEA, 25 C, 7 d: Colonies 23–25 mm, slightly raised in the centre, sulcate; margins very narrow (<1 mm), low, entire, plane; mycelium white and yellow; texture velvety and ﬂoccose; sporulation dense, especially in the centre; conidia en masse greyish green (26E5–26E6); exudates absent; soluble pigment absent; reverse brown centre fading into brownish orange (8F8). YES, 25 C, 7 d: Colonies 18–20 mm, raised in the centre, concentrically sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white, pale yellow and orange; texture velvety and ﬂoccose; sporulation moderately dense; conidia en masse dull green (26E4); exudates absent; soluble pigment brownish; reverse dark brown (5F6). DG18, 25 C, 7 d: Colonies 14–15 mm, raised in the centre, slightly concentrically sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white and pale yellow in the centre; texture velvety; sporulation dense; conidia en masse greyish green to dark green (25E5–25F5); exudates absent; soluble pigment yellow around colonies; reverse olive green (2F6–3F6), light yellow margins (2A5). OA, 25 C, 7 d: Colonies 23–25 mm, low, plane; margins narrow (1–2 mm), low, entire, plane; mycelium white; texture velvety; sporulation dense; conidia en masse greyish green to dark green (25E5–25F5); exudates small yellow droplets; soluble pigment yellow around colonies; reverse yellowish brown fading into pastel yellow (centre 5D5 and fading into 2A4). CREA, 25 C, 7 d: Colonies 14–15 mm, acid production absent. Micromorphology: Synnemata 400–1200 μm long. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 30–140 × 2–5 μm; branches 10–45 μm; metulae three to six, divergent, 8–12(–16) × 2–3(–4) μm; phialides acerose, three to six per metulae, 10–15 × 1.5–2.5(–3.5) μm; conidia smooth, ellipsoidal with pointed end to fusiform, 2.5–3.5(–5.5) × 1.5–3 μm. Ascomata not observed. Extrolites: Talaromyces coalescens produces (−) botryodiploidin and related compounds (Cabedo et al. 2007). Botryodiploidin has been called a mycotoxin and also has antifungal, antibacterial and insecticidal effects (Cabedo et al. 2007). A purpactin was also detected and several nonadrides related to glauconic acid. Distinguishing characters: Talaromyces coalescens produces colonies with dark red reverses with abundant red soluble pigment (Fig. 27). Synnemata, up to 1 200 μm long, are produced after 2–3 wk incubation. It closely resembles T. cecidicola. However, T. coalescens grows faster than T. cecidicola on MEA at 30 C. Talaromyces columbinus S.W. Peterson & Jurjevic, PLoS ONE 8: e78084–page 6. 2013. MycoBank MB804732. Fig. 28. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 20–70 × 2–2.5 μm; terminating in vesicles up to 6 μm; metulae four to eight, 8–14 × 2–4 μm; phialides acerose, three to six per metulae, 7–12 × 1.5–2.2 μm; conidia smooth, ellipsoidal, 2.5–3.5 × 3–4.5 μm. Extrolites: Talaromyces columbinus (isolate CBS 137393) produces rugulosin A, skyrin and series of tetracyclic compounds. Distinguishing characters: Talaromyces columbinus is similar to T. piceus. Both species have biverticillate conidiophores with vesiculated stipes, grow fast at high temperatures and are able to grow at 40 C. However, T. columbinus produces a brown soluble pigment on YES that is absent in T. piceus. In addition, T. columbinus grows slower on CYA at 25 C. Talaromyces convolutus Udawaga, Mycotaxon 48: 141. 1993. MycoBank MB360474. Fig. 29. ≡ Penicillium convolutum Udagawa, (simultaneously published). In: Talaromyces section Islandici www.studiesinmycology.org In: Talaromyces section Trachyspermi 239 YILMAZ ET AL. Fig. 28. Morphological characters of Talaromyces columbinus (CBS 137393). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. 240 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 29. Morphological characters of Talaromyces convolutus (CBS 100537T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Ascomata from OA after 3 wk incubation. C. Initials. D–G. Conidiophores. H. Conidia. Scale bars: B = 500 μm; C = 10 μm; D = 10 μm, applies to E–H. www.studiesinmycology.org 241 YILMAZ ET AL. ITS barcode: JN899330 (alternative markers: BenA = KF114773; RPB2 = JN121414) Talaromyces dendriticus (Pitt) Samson et al., Stud. Mycol. 71: 175. 2011. MycoBank MB560648. Fig. 30. ≡ Penicillium dendriticum Pitt, The genus Penicillium: 413. 1980. Typus: CBM SUM-3018, culture ex-type CBS 100537 = IBT 14989. Colony diam, 7 d (mm): CYA 10–12; CYA 30 C 15–16; CYA 37 C 6–8; MEA 13–15; MEA 30 C 17–20; DG18 3–5; CYAS No growth; OA 15–18; CREA 2–3; YES 12–13. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate, white sterile appearance; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent (at 37 C clear droplets); reverse orange white to pale orange (5A2–5A3). MEA 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate, white sterile appearance; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates clear droplets (at 30 C bigger yellow droplets); reverse brownish orange (5B6). YES 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate, white sterile appearance; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange (5A3). DG18 25 C, 7 d: Colonies low, plane, slimy yeast like colony appearance; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse white (4A1). OA 25 C, 7 d: Colonies low, plane, white sterile appearance; margins low, plane, entire (2 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates clear droplets; reverse beige. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, 10–70 × 1.5–2.5 μm; metulae two to ﬁve, divergent, 8–12 × 1.5–3 μm; phialides acerose, three to ﬁve per metulae, 7.5–14 × 1.5–2.5 μm; conidia smooth, ovoidal to ellipsoidal, (2–)3–4 × 1.5–2(–3) μm. Ascomata not observed in our culture but ﬁde Udagawa (1993) maturing slowly after 4 wk of incubation, sulphur yellow, globose to subglobose, 120–360 μm, ascomatal initials composed of short clavate ascogonia 15–20 μm long and 4 μm wide, asci 8–11 × 7.5–9 μm, ascospores globose to subglobose, spiny, 3–3.5 × 2.5–3(–3.5) μm. Extrolites: Talaromyces convolutus produces talaroconvolution A-D and ZG-1494α (Suzuki et al. 2000) and (−)-mitorubrinal, (−)-mitorubrinic acid and anhydroﬂavomannin-9,10-quinone-6,60 O-methyl ether (Suzuki et al. 1999). We also found a purpactin in this species. ZG-1494α is also produced by T. atroroseus (Frisvad et al. 2013). Distinguishing characters: Talaromyces convolutus is characterised by its restricted growth, slowly maturing sulphur yellow ascoma, globose to subglobose, spiny ascospores and monoto biverticillate conidiophores. These characters distinguish T. convolutus from its close relatives in section Trachyspermi. The restricted growth of T. convolutus distinguishes it from other yellow ascoma producers such as T. ﬂavus, T. austrocalifornicus and T. tratensis. Unfortunately, the available strains of T. convolutus were degenerated and no longer produced ascomata. 242 In: Talaromyces section Purpurei Typus: IMI 216897, culture ex-type CBS 660.80 = IMI 216897. ITS barcode: JN899339 (alternative markers: BenA = JX091391; CaM = KF741965; RPB2 = KM023286) Colony diam, 7 d (mm): CYA 23–26; CYA 30 C 15–16; CYA 37 C 5–6; MEA 35–36; MEA 30 C 20; DG18 12–20; CYAS No growth; OA 30–37; CREA 6–8; YES 15–27. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly radially sulcate; margins low, plane, entire (2–3 mm); mycelia white and yellow; texture velvety and loosely funiculose; sporulation moderately dense in the centre, conidia en masse greyish green to dark green (27E4–27F4); soluble pigments absent; exudates absent; reverse centre dark brown (6F6), fading into light brown (6D6) and light yellow (2A5). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia yellow and white; texture loosely funiculose (colonies dominated with young yellow synnemata without sporulation); sporulation sparse to moderately dense (in the centre), conidia en masse greyish green to dark green (27E4–27F4); soluble pigments absent; exudates absent; reverse centre dark brown (8F6) fading into reddish brown (8D5–8E5) and brownish orange (5C5–5C6). YES 25 C, 7 d: Colonies slightly raised at centre, slightly radially sulcate; margins low, plane, entire (2–3 mm); mycelia white and yellow; texture velvety and loosely funiculose; sporulation moderately dense, conidia en masse greyish green to dark green (27E4–27F4); soluble pigments absent; exudates absent; reverse centre dark brown (6F6), fading into light brown (6D6) and light yellow (2A5). DG18 25 C, 7 d: Colonies slightly raised in the centre, slightly radially sulcate; margins low, plane, entire (3–4 mm); mycelia white and pale yellow; texture velvety; sporulation sparse to moderately dense in the centre, conidia en masse dark green (26F5–26F6); soluble pigments absent; exudates absent; reverse centre dark green (28F3), fading into pale yellowish white (2A2) to beige. OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire; mycelia white and yellow; texture velvety (colonies dominated with young yellow synnemata without sporulation); sporulation sparse to moderately dense in the centre, conidia en masse dark green (26F5–26F6); soluble pigments absent; exudates absent; reverse light brown centre, fading into light yellow. CREA 25 C, 7 d: Acid production absent. Micromorphology: Synnemata up to 5 000 μm long after 2 wk. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 20–200 × 2.5–4 μm; branches 12–20 μm; metulae four to six, divergent, 9.5–16 × 2–3.5 μm; phialides acerose, three to eight per metulae, 9.5–14 × 2–2.5 μm; conidia smooth, subglobose to ellipsoidal, 2.5–3.5 × 2–2.5 μm. Ascomata not observed. Extrolites: Talaromyces dendriticus produces mitorubrinic acid, secalonic acid D (Samson et al. 1989) in addition to a purpactin and nonadrides related to glauconic acid. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 30. Morphological characters of Talaromyces dendriticus (DTO 183-G3). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 2 wk incubation. C. Synnemata. D–G. Conidiophores. H. Conidia. Scale bars: B, C = 2000 μm; D = 10 μm, applies to E–H. www.studiesinmycology.org 243 YILMAZ ET AL. Distinguishing characters: Talaromyces dendriticus is recognised by colonies producing synnemata up 5 000 μm long after 1–2 wk of incubation. This species seems to be associated with Eucalyptus. Talaromyces dendriticus is distinguished from other synnema producers by its restricted growth on CYA at 25 and 30 C. Talaromyces derxii Takada & Udagawa, Mycotaxon 31: 418. 1988. MycoBank MB133755. Fig. 31. ≡ Penicillium derxii Takata & Udagawa, (simultaneously published). In: Talaromyces section Talaromyces Typus: NHL 2980, culture ex-type CBS 412.89 = NHL 2981. ITS barcode: JN899327 (alternative markers: BenA = JX494305; CaM = KF741959; RPB2 = KM023282) Colony diam, 7 d (mm): CYA 38–40 (12–13); CYA 30 C 43–45 (17–18); CYA 37 C 35–38 (15–16); MEA 48–50 (20–22); MEA 30 C 60–65 (30–32); DG18 14–15 (7–8); CYAS No growth; OA 10–20; CREA 2–6; YES 39–40 (15–16). Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, in some isolates (CBS 413.89) sunken at centre and crateriforme, slightly sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse; conidia en masse greyish green (25B3–26B3); soluble pigments absent; exudates absent; reverse dark green (25F5) centre for both fading into light yellow (4A4). MEA 25 C, 7 d: Colonies slightly raised at centre, in some isolates sunken at centre, low to deep, plane to sulcate; margins low, plane, entire (1–3 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse greyish green (25B3–26B3); soluble pigments absent; exudates absent; reverse in some isolates (CBS 412.89T) light brown to brown (6D6–6E6) centre fading into apricot (5B6) and in some isolates (CBS 413.89) greyish orange (5B5). YES 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (1–3 mm); mycelia white; texture ﬂoccose; sporulation absent to moderately dense, conidia en masse greyish green (25C5–25D5); soluble pigments absent; exudates absent; reverse in some isolates (CBS 412.89T) centre golden yellow (5B7) rest dark green (27F7) and in some isolates (CBS 413.89) centre greyish green (27D6) fading into light orange (5A4) and light yellow (4A4) with brownish orange (5C6) circle. DG18 25 C, 7 d: Colonies low to slightly raised at centre, plane to slightly sulcate, light orange appearance; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (6A5) to orange white (5A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture velvety and ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (27D5–27E5); soluble pigments absent; exudates absent; reverse dull green. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, (15–)30–60(–125) × 2.5–3.5 μm; metulae two to ﬁve, divergent, 10–20(–30) × 2–4 μm; phialides acerose, three to six per metulae, 10–20(–25) × 2–3.5 μm; conidia smooth, cylindrical to ellipsoidal and sometimes curved and fusiform, 4–8 × 1.5–3 μm. Ascomata not observed in our culture but ﬁde 244 Takada & Udagawa (1993) heterothallic. Superﬁcial, maturing within 1–2 wk at 37 C, greyish green to dark bluish green, globose to subglobose, asci 10–13 × 9–12 μm, ascospores ellipsoidal, spiny, 3.5–5 × 2.5–3 μm. Extrolites: Talaromyces derxii produces talaroderxine A & B (Suzuki et al. 1992), 4-[3-methyl-2-butenyl)oxy]benzoic acid, 4[(2,3-epoxy-3-methylbutyl)oxy]benzoic acid, (4E)-4-[(4-hydroxy3-methyl-2-butenyl)oxy]benzoic acid (Nozawa et al. 1989), and penicillide and dehydroisopenicillide (Suzuki et al. 1991). Distinguishing characters: Talaromyces derxii is a heterothallic species. It is characterised by a dark green reverse on CYA and cylindrical to ellipsoidal conidia (Fig. 31). These characters resemble T. bacillisporus. However, T. bacillisporus has globose, spiny ascospores whereas T. derxii produces green ascomata with spiny ellipsoidal ascospores. In addition, T. derxii has smooth walled stipes in contrast to the rough stipes of T. bacillisporus. Talaromyces diversus (Raper & Fennell) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011. MycoBank MB560649. Fig. 32. ≡ Penicillium diversum Raper & Fennell, Mycologia 40: 539. 1948. In: Talaromyces section Trachyspermi Typus: IMI 040579, culture ex-typus CBS 320.48 = ATCC 10437 = DSM 2212 = IMI 040579 = IMI 040579ii = NRRL 2121. ITS barcode: KJ865740 (alternative markers: BenA = KJ865723; CaM = KJ885268; RPB2 = KM023285) Colony diam, 7 d (mm): CYA 7–10; CYA 30 C 6–10; CYA 37 C 5–8; MEA 25–35; MEA 30 C 30–35; DG18 10–15; CYAS 2–5; OA 25–40; CREA No growth and sometimes up to 2; YES 8–10. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sunken at centre, slightly sulcate; margins low, plane, entire (<1 mm); mycelia white; texture velvety; sporulation moderately dense to dense, conidia en masse dull green (27D3–27D4); soluble pigments absent; exudates absent; reverse blond to brownish orange (4C4–5C4). MEA 25 C, 7 d: Colonies slightly raised, sunken at centre, sulcate; margins low, plane, entire (2 mm); mycelia white; texture velvety and in some isolates ﬂoccose (DTO 231-A1); sporulation dense, conidia en masse dull green (27D3–27D4); soluble pigments absent; exudates in some isolates small clear droplets; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies slightly raised at centre, sunken at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; texture velvety and in some isolates ﬂoccose; sporulation sparse to moderately dense, conidia en masse dull green (27D3–27D4); soluble pigments absent; exudates absent; reverse light yellow (4A5–5A5). DG18 25 C, 7 d: Colonies raised at centre, sunken in the centre, sulcate; margins low, plane, entire (1 mm); mycelia white; texture velvety; sporulation moderately dense to dense, conidia en masse dull green (27D3–27D4); soluble pigments absent; exudates in some isolates clear droplets at centre; reverse in some isolates centre orange white to pale orange (5A2–5A3), in some isolates centre greenish grey (1B2), in the margins greyish green to olive POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 31. Morphological characters of Talaromyces derxii (CBS 412.89T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. b. Colony texture of on OA after 2 wk incubation. C–F. Conidiophores. G. Conidia. Scale bars: B = 500 μm; C = 10 μm, applies to D–G. www.studiesinmycology.org 245 YILMAZ ET AL. Fig. 32. Morphological characters of Talaromyces diversus (CBS 320.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 246 POLYPHASIC (1C4–1D4–1E4). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–4 mm); mycelia white; texture velvety; sporulation sparse to moderately dense, conidia en masse dull green (27D3–27D4); soluble pigments absent; exudates absent; reverse very pale light greenish beige. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 200–300 × 2.5–4 μm; branches 10–35 μm; metulae three to eight, divergent, 7–14 × 2.5–3(–5) μm; phialides acerose, three to six per metulae, 8–12 × 2–3 μm; conidia smooth to verrucose, ellipsoidal to subglobose, 2–3(–5) × 2–3(–3.5) μm. Ascomata not observed. Extrolites: Talaromyces diversus have been reported to produce diversonol, α- and β-diversonol esters, isoaustin, austinol, alternariol monomethyl ether and lichexanthone (Turner 1978, Simpson et al. 1982, Holker et al. 1983). We have detected the alternariol monomethyl ether, diversonol, austins and mitorubrinic acid in this species. Distinguishing characters: Talaromyces diversus is distinguished from other species by weak, restricted growth on CYA, dull green (27D3–27D4) conidia, velvety texture on MEA and lack of growth on CREA (Fig. 32). Its restricted growth on CYA resembles T. rademirici, T. erythromellis and T. primulinus, however growth on CYA at 37 C and colony size on MEA at 25 C easily distinguish T. diversus from these species. Talaromyces duclauxii (Delacr.) Samson et al., Stud. Mycol. 71: 175. 2011. MycoBank MB560650. Fig. 33. ≡ Penicillium duclauxii Delacr., Bull. Soc. Mycol. France 7: 107. 1891. TAXONOMY OF TALAROMYCES reverse olive brown (4D6–4E6) centre fading into greyish yellow (4B6–4C6). DG18 25 C, 7 d: Colonies raised at centre, slightly deep, slightly sulcate; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates small clear and yellow droplets; reverse yellowish brown (5E6–5F6) and yellow (3A6). OA 25 C, 7 d: Colonies deep, raised, plane, sporulated synnemata produced; margins low, plane, entire (2–3 mm); mycelia white; texture synnemateus; sporulation only on synnemata densely, conidia en masse greyish green to dull green (26D3–26D4); soluble pigments absent; exudates absent; reverse brownish yellow. CREA 25 C, 7 d: Weak acid production. Micromorphology: Synnemata deep ﬂuffy structure, 2000–5000 μm long. Conidiophores biverticillate; stipes smooth walled, 15–50 × 3–4(–5.5) μm; metulae two to six, divergent, 8.5–15 × 2.5–5 μm; phialides acerose, three to eight per metulae, 9–15 × 2–3.5 μm; conidia smooth to ﬁnely rough, ellipsoidal, 3–4 × 1.5–3.5(–4) μm. Ascomata not observed. Extrolites: Talaromyces duclauxii produces duclauxin, xenoglaucin and crypptoclauxin and related compounds (Shibata et al. 1965, Ogihara et al. 1966). Distinguishing characters: Talaromyces duclauxii is characterised by its rapid growth, colonies that are deep and have ﬂuffy texture on MEA and synnemata up to 5 000 μm long (Fig. 33). Talaromyces palmae produces indeterminate synnemata like those of T. duclauxii. However, T. duclauxii grows faster on CYA at 30 C. Talaromyces emodensis Udagawa, Mycotaxon 48: 146. 1993. MycoBank MB360476. Fig. 34. ≡ Penicillium emodense Udagawa, (simultaneously published). In: Talaromyces section Talaromyces In: Talaromyces section Bacillispori Typus: IMI 24312, culture ex-type CBS 322.48 = ATCC 10439 = IMI 040044 = MUCL 28672 = MUCL 29094 = MUCL 29212 = NRRL 1030. Typus: CBM SUM-3025, culture ex-type CBS 100536 = IBT 14990. ITS barcode: JN899342 (alternative markers: BenA = JX091384; CaM = KF741955; RPB2 = JN121491) ITS barcode: JN899337 (alternative markers: BenA = KJ865724; CaM = KJ885269; RPB2 = JN121552) Colony diam, 7 d (mm): CYA 25–27; CYA 30 C 34–35; CYA 37 C 3–4; MEA 48–50; MEA 30 C 40; DG18 17–18; CYAS No growth; OA 48–50; CREA 18–20; YES 43–44. Colony diam, 7 d (mm): CYA 8–10; CYA 30 C 12–14; CYA 37 C 4–6; MEA 8–10; MEA 30 C 10–12; DG18 4; CYAS 2–3; OA 9–10; CREA 3–4; YES 9–10. Colony characters: CYA 25 C, 7 d: Colonies deep, raised at centre, slightly sulcate; margins low, plane, entire (2–3 mm); mycelia white and pastel yellow; texture ﬂuffy appearance, synnematous; sporulation absent to sparse, conidia en masse difﬁcult to determine; soluble pigments yellow; exudates absent; reverse olive brown (4E8) fading into olive brown (4D7) and maize yellow (4A6). MEA 25 C, 7 d: Colonies very deep, raised and sunken at centre, synnemata produced on the margins; margins low, plane, entire (2–3 mm); mycelia white; texture velvety; sporulation sparse, conidia en masse greyish green (25B5–25C5); soluble pigments absent; exudates absent; reverse brown (6E8) centre fading into brownish yellow (5C7). YES 25 C, 7 d: Colonies slightly raised at centre, concentrically sulcate; margins low, plane, entire (1–2 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; Colony characters: CYA 25 C, 7 d: Colonies raised at centre, sulcate, pastel yellow appearance; margins low, plane, entire (1 mm); mycelia white and pastel yellow; sporulation absent; soluble pigments absent; exudates absent; reverse brownish red to reddish brown (8C8–8D8). MEA 25 C, 7 d: Colonies raised, plane, white ﬂuffy appearance; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse reddish golden brown (6C7). YES 25 C, 7 d: Colonies raised at centre, sulcate, yellow and white ﬂuffy appearance; margins low, plane, entire (1 mm); mycelia white and yellow; sporulation absent; soluble pigments absent; exudates absent; reverse greyish orange to brownish orange (5B5–5C5). DG18 25 C, 7 d: Colonies low, plane, white appearance; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; www.studiesinmycology.org 247 YILMAZ ET AL. Fig. 33. Morphological characters of Talaromyces duclauxii (CBS 322.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 2 wk incubation. C–F. Conidiophores. G. Conidia. Scale bars: B = 1000 μm; C = 10 μm, applies to D–G. 248 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 34. Morphological characters of Talaromyces emodensis (CBS 100536T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D–E. Asci and ascopsores F. Ascospores. Scale bars: B = 1000 μm; C = 500 μm; D = 10 μm, applies to E–F. www.studiesinmycology.org 249 YILMAZ ET AL. reverse yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane, white appearance (at 30 C formation of pastel yellow ascomata); margins low, plane, entire (1–2 mm); mycelia white and yellow; sporulation absent; soluble pigments absent; exudates absent; reverse pastel yellow. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores not observed here, ﬁde Udagawa (1993), monoverticillate with a minor proportion having subterminal branches; stipes smooth walled, 18–60 × 2–2.5 μm; branches 15–25 μm; phialides acerose, one to three, 10–17.5 × 2–3 μm; conidia smooth, ovoidal to ellipsoidal, 3–4 × 1.5–3 μm. Ascomata maturing slowly within 4 wk on MEA at 25 C, 100–250 μm, sulphur yellow, pastel yellow and creamish white, globose to subglobose, asci 8–11 × 7–9.5 μm, ascospores globose to subglobose, spiny, 2.5–4 × 2.5–3.5 μm. Extrolites: Talaromyces emodensis produces duclauxin and mitorubrin. Distinguishing characters: Talaromyces emodensis grows restrictedly on all medium, produces small, globose to subglobose, spiny ascospores and conidiophores with solitary phialides to monoverticillate conidiophores (Fig. 34). Its restricted growth resembles T. tardifaciens, however, T. tardifaciens has smooth ascopsores. The typical characters of T. emodensis distinguish it from other species that produce globose to subglobose, spiny ascospores such as T. bacillisporus and T. rotundus. The asexual state was not observed in our study. Talaromyces erythromellis (A.D. Hocking) Samson et al., Stud. Mycol. 71: 175. 2011. MycoBank MB560652. Fig. 35. ≡ Penicillium erythromellis A.D. Hocking, The genus Penicillium: 459. 1980. pigments absent; exudates clear droplets; reverse orange white (5A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire, yeast like slimy texture (3–4 mm); mycelia white; texture velvety and loosely ﬂoccose; sporulation moderately dense, conidia en masse dull green (26D3–26D4); soluble pigments absent; exudates absent; reverse media colour, beige. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate having symmetrical subterminal branches; stipes smooth walled, 20–80 × 2–3 μm; branches 13–45 μm; metulae two to six, divergent, 9–20 × 1.5–3 μm; phialides acerose, three to six per metulae, 8.5–15 × 1.5–2.5 μm; conidia subglobose to ellipsoidal, 2–3.5 × 1.5–2.5 μm. Ascomata not observed. Extrolites: Talaromyces erythromellis produces rubropunctatin. Distinguishing characters: Talaromyces erythromellis grows restrictedly on all media studied. It produces red exudates on MEA and biverticillate conidiophores with additional subterminal branches (Fig. 35). Its restricted growth on CYA resembles T. rademirici, T. diversus and T. primulinus, however restricted growth on MEA and conidiophore branching pattern easily distinguish T. erythromellis from T. diversus and T. primulinus and growth on CREA distinguishes it from T. rademirici. Talaromyces euchlorocarpius Yaguchi, Somaya & Udagawa, Mycoscience 40: 133. 1999. MycoBank MB460481. Fig. 36. ≡ Penicillium euchlorocarpium Yaguchi, Someya & Udagawa, (simultaneously published). In: Talaromyces section Talaromyces In: Talaromyces section Trachyspermi Typus: PF 1203, culture ex-type PF 1203 = DTO 176-I3 = DTO 176-I4. Typus: IMI 216899, culture ex-typus: CBS 644.80 = FRR 1868 = IMI 216899. ITS barcode: AB176617 (alternative markers: BenA = KJ865733; CaM = KJ885271; RPB2 = KM023303) ITS barcode: JN899383 (alternative markers: BenA = HQ156945; CaM = KJ885270; RPB2 = KM023290) Colony diam, 7 d (mm): CYA 15–18; CYA 30 C 14–17; CYA 37 C No growth; MEA 38–40; MEA 30 C 25–32; DG18 12–13; CYAS No growth; OA 20–23; CREA 9–11; YES 20–25. Colony diam, 7 d (mm): CYA 10–11; CYA 30 C 13–14; CYA 37 C No growth; MEA 10–11; MEA 30 C 12–13; DG18 5–6; CYAS No growth; OA 15–16; CREA 4; YES 9. Colony characters: CYA 25 C, 7 d: Colonies raised at margins, sunken in the middle, pinkish colour; margins low, irregular (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates red small droplets; reverse dark red to brownish red (10C8–10D8). MEA 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; texture loosely funiculose; sporulation moderately dense, conidia en masse greyish green to dull green (26C3–26D3); soluble pigments absent; exudates small red droplets; reverse dark brown (7F7). YES 25 C, 7 d: Colonies raised, sulcate, pinkish red colour; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates small red droplets; reverse greyish orange (6B3–6B4). DG18 25 C, 7 d: Colonies slightly raised, plane; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble 250 Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, sulcate, in some isolates sunken at centre; margins low, plane, entire (1 mm); mycelia white and pastel yellow, especially in the centre; texture velvety; sporulation dense, conidia en masse greyish green to dull green (25C3–25D3); soluble pigments yellow to orange yellow; exudates absent; reverse reddish golden (6C7) centre fading into golden yellow (5B7) and deep yellow (4A8). MEA 25 C, 7 d: Colonies low, slightly sulcate; margins low, plane, entire (1–2 mm); mycelia white and yellow; texture loosely funiculose and ﬂoccose; sporulation moderately dense (at 30 C dense), conidia en masse greyish green (28B5–28C5); soluble pigments absent; exudates absent; reverse greyish orange (5B5–5B6) with red (9B7) dots. YES 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white and yellow; texture velvety and ﬂoccose; sporulation moderately dense to dense, conidia en masse greyish green (25D4–26D4); soluble pigments absent; exudates absent; reverse brownish yellow (5C7) centre fading POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 35. Morphological characters of Talaromyces erythromellis (CBS 644.80T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 251 YILMAZ ET AL. Fig. 36. Morphological characters of Talaromyces euchlorocarpius (PF 1203T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on Hay infusion agar after 1 wk incubation. C. Ascomata. D. Asci and ascospores. E. Initials. F–H. Conidiophores. I. Conidia. Scale bars: B = 500 μm; C = 10 μm; E = 10 μm; D = 10 μm, applies to F–I. 252 POLYPHASIC into light yellow (4A5). DG18 25 C, 7 d: Colonies low, plane, slimy yeast like sterile white mycelia. OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white; texture velvety; sporulation sparse, conidia en masse greyish green (25C4) and dull green (25E4); soluble pigments deep yellow; exudates absent; reverse deep yellow. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 100–150 × 1.5–2 μm; metulae three to six, divergent, 9–13 × 2–3.5 μm; phialides acerose, three to six per metulae, 8–14 × 2–3 μm; conidia smooth, subglobose to ellipsoidal, 2–4 × 2–3 μm. Ascomata maturing after 1–2 wk of incubation on hay infusion agar at 25 C and abundantly 30 C, deep green, globose to subglobose, 150–300 μm, asci 8–10 × 5–8 μm, ascospores ellipsoidal, spiny, 3.5–5 × 2.5–3 μm. TAXONOMY OF TALAROMYCES margins low, plane, entire (1 mm); mycelia white; texture velvety; sporulation dense, conidia en masse dull green (26D4–27D4); soluble pigments absent; exudates absent; reverse greyish red (9C5) centre fading into greyish yellow (4B3). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (4–5 mm); mycelia yellow; texture velvety and ﬂoccose, centre covered with yellow mycelia; sporulation moderately dense, conidia en masse greyish green to dark green (27E5–27F5); soluble pigments absent; exudates small yellow and clear droplets; reverse pale greyish red. CREA 25 C, 7 d: Acid production absent. Distinguishing characters: Talaromyces euchlorocarpius is characterised by yellow soluble pigment on CYA and OA, deep green ascomata and biverticillate conidiophores (Fig. 36). Green ascomata resembles T. derxii and T. viridis, however, lack of growth at 37 C distinguishes T. euchlorocarpius from both. In addition, T. viridis produces conidiophores with solitary phialides, whereas T. euchlorocarpius produces biverticillate conidiophores. Micromorphology: Synnemata up to 750 μm long, appearance to stipitate sporangia and covered with bright yellow layer. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 90–350 × 2–3 μm; branches 10–35 μm; metulae four to six, divergent, 9–15 × 2–3 μm; phialides acerose, three to eight per metulae, 8–12 × 2–3 μm; conidia smooth to ﬁnely rough, ellipsoidal, 2.5−3.5(–5.5) × 2–2.5(–4) μm. Ascomata production was reported by Visagie et al. (2012) from burned Quercus suber leaf litter. Ascomata golden yellow with a reddish pigment, only produced on Quercus suber leaf litter in nature, globose to subglobose, 150–380 μm, asci absent, ascospores ellipsoidal, thick walled, spiny, (4–)4.5–6(–7) × 3–4 μm. Talaromyces ﬂavovirens (Durieu & Mont.) Visagie, Llimona & Seifert, Mycotaxon 122: 404. 2012. MycoBank MB800438. Fig. 37. Extrolites: Talaromyces ﬂavovirens produces mitorubrin, mitorubrinol, mitorubrinic acid, a purpactin, secalonic acid D and vermicillin. ≡ Lasioderma ﬂavovirens Durieu & Mont., Ann. Sci. Nat., Bot. 4: 364. 1845. = Penicillium aureocephalum Munt.-Cvetk. et al., Fungal Divers. 7: 73. 2001. In: Talaromyces section Talaromyces Typus: Lectotype PC 0088796; epitype BCC 473 = BCN 473, culture ex-type CBS 102801 = IBT 27044. ITS barcode: JN899392 (alternative markers: BenA = JX091376; CaM = KF741933) Colony diam, 7 d (mm): CYA 19–20; CYA 30 C 21–22; CYA 37 C 5–6; MEA 37–38; MEA 30 C 37–38; DG18 12–13; CYAS No growth; OA 38–40; CREA 15; YES 21–22. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white; texture velvety; sporulation dense, conidia en masse greyish turquoise (24D4–25D4); soluble pigments yellow; exudates absent; reverse brown (7E6) centre fading into greyish yellow (2B6). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–4 mm); mycelia white; texture velvety after 2 wk of incubation covered with yellow mycelia layer; sporulation dense, conidia en masse dull green (26D3–27D3); soluble pigments absent; exudates absent; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (2 mm); mycelia white; texture velvety; sporulation dense, conidia en masse dull green (26D4–26E4); soluble pigments absent; exudates absent; reverse dark brown (6F6) centre fading into brownish yellow (5C7) and mustard yellow (3B6) and greyish green (1C5). DG18 25 C, 7 d: Colonies sunken at centre, low, slightly sulcate; www.studiesinmycology.org Distinguishing characters: Talaromyces ﬂavovirens produces yellow ascomata with spiny, ellipsoidal ascospores on Quercus suber and rarely on Q. ilex and Cistus salviifolius leaf litter and all cultures originate from Catalonia, Spain. No ascomata were observed in uncrossed or crossed cultures on general media. Colonies produce characteristic yellow mycelial layers that resemble T. marneffei and T. primulinus. However T. marneffei produces red soluble pigment on MEA and CYA, and it produces a yeast phase at 37 C. Talaromyces primulinus grows slower on all media studied. Notes: Visagie et al. (2012) introduced T. ﬂavovirens, describing it as the sexual state of P. aureocephalum and synonymising the latter. Talaromyces ﬂavus (Klöcker) Stolk & Samson, Stud. Mycol. 2: 10. 1972. MycoBank MB324416. Fig. 38. ≡ Gymnoascus ﬂavus Klöcker, Hedwigia 41: 80. 1902. = Penicillium vermiculatum P.A. Dang., Botaniste 10: 123. 1907 ≡ Talaromyces vermiculatus (P.A. Dang.) C.R. Benj., Mycologia 47: 684. 1955. = Arachniotus indicus Chattop. & C. Das Gupta, Trans. Brit. Mycol. Soc. 42: 72. 1959. = Arachniotus indicus Chattop. & C. Das Gupta var. major Chattop. & C. Das Gupta, Trans. Brit. Mycol. Soc. 42: 73. 1959. ≡ Penicillium dangeardii Pitt, The Genus Penicillium: 472. 1980. In: Talaromyces section Talaromyces Typus: CBS H-7820, culture ex-type CBS 310.38 = IMI 197477 = NRRL 2098. ITS barcode: JN899360 (alternative markers: BenA = JX494302; CaM = KF741949; RPB2 = J417426) 253 YILMAZ ET AL. Fig. 37. Morphological characters of Talaromyces ﬂavovirens (CBS 1102801T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on MEA after 3 wk incubation. C–F. Conidiophores. G. Conidia. Scale bars: B = 500 μm; C = 10 μm, applies to D–G. 254 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 38. Morphological characters of Talaromyces ﬂavus (CBS 310.38T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata D. Ascomata and asci. E. initials. F. Conidiophores. G. Asci and ascospores. H. Ascospores. Scale bars: B–D = 500 μm; E = 10 μm, applies to F–H. www.studiesinmycology.org 255 YILMAZ ET AL. Colony diam, 7 d (mm): CYA 9–10; CYA 30 C 17–18; CYA 37 C 19–20; MEA 31–32; MEA 30 C 40–45; DG18 13; CYAS No growth; OA 30–32; CREA 7–9; YES 24–26. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (2 mm); mycelia white and bright yellow; texture ﬂoccose; sporulation absent; soluble pigments yellow; exudates absent; reverse brownish orange (5C6) fading into light yellow (3A5). MEA 25 C, 7 d: Colonies low, plane, formation of yellow ascomata (at 30 C more abundant); margins low, plane, entire (2–3 mm); mycelia white, yellow and red; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse greyish orange (5B6–6B6). YES 25 C, 7 d: Colonies slightly raised at centre, sometimes sunken at centre, slightly sulcate; margins low, plane, entire (3–4 mm); mycelia white, yellow and red; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse orange (5A6) fading into reddish yellow (4A6). DG18 25 C, 7 d: Colonies raised at centre, plane; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose and funiculose; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange to greyish orange (5A3–5B3) centre fading into yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane, yellow ascomata formation; margins low, plane, entire (2 mm); mycelia white and yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish orange and fading into yellow. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores asexual state lacking, when present monoverticillate; stipes smooth walled, 15–20 × 1.9–20 μm; phialides acerose, one to three, 11–12 × 2–2.5 μm; conidia smooth, ellipsoidal, 2–3 × 1.5–2.5 μm. Ascomata maturing after 1–2 wk of incubation on OA and MEA at 25 C and abundantly 30 C, deep yellow, globose to subglobose, 150–400 μm, asci 9.5–13.5 × 8–11.5 μm, ascospores broadly ellipsoidal, thick walled, spiny, 4–5.5 × 3–3.5 μm. Extrolites: Talaromyces ﬂavus produces 3-hydroxymethyl-6,8dimethoxycoumarin, 2,5-dimethyl-7-hydroxychromone, (−)-trans2,3-epoxysuccinic acid, fosfonochlorin, 2-methylsorbic acid, 4,6dihydroxy-5-methylphthalide, funiculosic acid, hydroxyfuniculosic acid, vermistatin = ﬁjiensin and related compounds, altenusin, dehydroaltenusin, desmethyldehydroaltenusin, talaroﬂavon, deoxytalaroﬂavon, (−)-mitorubrin, (−)-mitorubrinol, (−)-mitorubrinic acid, (−)-diazaphilonic acid, vermixocin A (= penicillide) & vermixocin B (= purpac tin A), vermilutin, vermiculine, vermiculinic acid, vermiculic acid, 4-deoxyvermiculic acid, vermicillin, TAN-2177A & B (Proksa 2010). Talaromyces ﬂavus also produces unique polysaccharides such as talaron, which is fungicidal and a series of enzymes (Proksa 2010). The species has been used for biological pest control (Proksa 2010). Distinguishing characters: Talaromyces ﬂavus produces deep yellow ascomata and thick walled, spiny, broadly ellipsoidal ascospores (Fig. 38). It produces monoverticillate conidiophores that do not sporulate profusely. Colonies on CYA grow restrictedly (9–10 mm) after 7 d at 25 C. This distinguishes it from T. macrosporus, T. muroii and T. liani. Talaromyces tratensis, T. convolutus, T. austrocalifornicus also grow restrictedly on CYA, but bigger ascospores (4.2–5.2 × 3–3.6 μm) easily distinguish T. ﬂavus from the latter species. 256 Notes: Penicillium vermiculatum was described by Dangeard (1907) and transferred to Talaromyces by Benjamin (1955). Orr et al. (1963) considered Gymnoascus ﬂavus and T. vermiculatus as synonyms and this was followed by Stolk & Samson (1972) and Pitt (1980). Ghosh et al. (1961) re-evaluated the type strains of Arachniotus indicus and A. indicus var. major and both isolates proved to represent Talaromyces vermiculatus and therefore they synonymised it with T. ﬂavus. In our study we also concur with this. Talaromyces funiculosus (Thom) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560653. Fig. 39. ≡ Penicillium funiculosum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 69. 1910. In: Talaromyces section Talaromyces Typus: IMI 193019, culture ex-type CBS 272.86 = IMI 193019. ITS barcode: JN899377 (alternative markers: BenA = JX091383; CaM = KF741945; RPB2 = KM023293) Colony diam, 7 d (mm): CYA (30–)38–45; CYA 30 C (35–)45–55; CYA 37 C 38–50; MEA 30–45; MEA 30 C 40–55; DG18 13–20; CYAS 13–20, some strains no growth; OA 30–40; CREA (15–) 20–30; YES 35–45. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent to moderately dense, conidia en masse greyish green to dull green (26D3–26D4); soluble pigments only in DTO 60-E7 and DTO 50-F6 soluble light red pigment the rest absent; exudates clear droplets; reverse light orange to greyish orange (5A4–5B4) centre fading into greyish yellow (4B3), in some isolates, centre changes to greyish orange to brownish orange (5B5–5C5) and in red pigment producers brownish red (9C7) reverse. MEA 25 C, 7 d: Colonies moderately deep, plane; margins low, plane, entire (2 mm); mycelia white; texture funiculose; sporulation sparse to dense, conidia en masse greyish green to dull green (26D3–26D4); soluble pigments absent; exudates absent; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (2 mm); mycelia white; texture in some isolates sterile aerial mycelia grows like funiculose; sporulation absent; soluble pigments only in DTO 60-E7 light red; exudates in some isolates clear droplets; reverse greyish orange (5B5) centre fading into pastel yellow (3A4), and in red pigment producer (DTO 60-E7) greyish red (9B6) centre and in the margins red (9B7). DG18 25 C, 7 d: Colonies moderately deep, plane; margins low, plane, entire (1 mm); mycelia white; texture in some isolates funiculose, in some isolates no sporulation slimy yeast like colonies; sporulation in some isolates absent, in some isolates dense, conidia en masse greyish green to dull green (26D3–26D4); soluble pigments absent; exudates small clear droplets; reverse in some isolates centre greyish green (1C4–1D4) sometimes with light orange (5A5) circle; in some isolates light orange (5A5); in some isolates dark brown (6F6) centre fading into melon yellow (5A6). OA 25 C, 7 d: Colonies in some isolates low, in some isolates moderately deep, plane; margins low, plane, entire (2–3 mm); mycelia white; texture funiculose and in some isolates loosely funiculose to velvety; sporulation dense, conidia en masse greyish green to dull green POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 39. Morphological characters of Talaromyces funiculosus (CBS 883.70). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on MEA after 1 wk incubation. C–G. Conidiophores. H. Conidia. Scale bars: C = 50 μm; D = 10 μm, applies to E–H. www.studiesinmycology.org 257 YILMAZ ET AL. (26D3–26D4); soluble pigments absent; exudates in some isolates small clear droplets; reverse in some isolates greyish green, in some isolates brownish orange. CREA 25 C, 7 d: Strong acid production. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 15–100 × 2–3.5 μm; branches 8–18 μm; metulae three to six, divergent, 6.5–11 × 2–4 μm; phialides acerose, three to eight per metulae, 7.5–11 × 1.5–2.5 μm; conidia smooth, ellipsoidal, 2–3(–5.5) × 1–2(–2.5) μm. Ascomata not observed. Extrolites: Strain NRRL 1033 = CBS 169.91 produces secalonic acid D and nonadride metabolites, while NRRL 1035 = CBS 171.91 produces pestalacin A, while all strains examined produce a speciﬁc extrolite, that has not been structure elucidated. Distinguishing characters: Talaromyces funiculosus characteristically produces colonies that are strongly funiculose (Fig. 39). It shows fast growth on general media and at 37 C, and produces strong acid on CREA. All these characters distinguish T. funiculosus from other Talaromyces species. phialides acerose, three to ﬁve per metulae, 7.5–12(–15) × 2–3 μm; conidia smooth, ovoidal to ellipsoidal, 2.5–5 × 1.5–3 μm. Ascomata ﬁde Samson & Mahoney (1977), abundantly produced on OA at 30 C, within 2 wk, at ﬁrst white to cream after prolonged incubation yellow or reddish, globose to subglobose, 150–400 μm, asci 15–19 × 15–19 μm, ascospores broadly ellipsoidal, spiny, 7–10 × 5.5–8 μm. Extrolites: Apiculides have been found in T. galapagensis. Distinguishing characters: Talaromyces galapagensis produces distinct ascospores, which are broadly ellipsoidal, thick walled and ornamented with ridges. Its ascospores are much bigger (7–10 × 5.5–8 μm) than ascospores of T. stipitatus, T. unicus, T. ucrainicus and T. mimosinus, which have similar ridges as T. galapagensis. The ex-type strain of T. galapagensis was degenerated and did not produce ascomata. Talaromyces hachijoensis Yaguchi, Someya & Udagawa, Mycoscience 37: 157. 1996. MycoBank MB416016. In: Talaromyces section Proteolytcius Typus: PF 1174, culture ex-type PF 1174 = IFM 53624. Talaromyces galapagensis Samson & Mahoney, Trans. Brit. Mycol. Soc. 69: 158. 1977. MycoBank MB324417. Fig. 40. ≡ Penicillium galapagense Samson & Mahoney, (simultaneously published). In: Talaromyces section Talaromyces Typus: CBS H-7489, culture ex-type CBS 751.74 = IFO 31796. ITS barcode: JN899358 (alternative markers: BenA = JX091388; CaM = KF741966) Colony diam, 7 d (mm): CYA 15–17; CYA 30 C 23–27; CYA 37 C 25–26; MEA 25–28; MEA 30 C 33–35; DG18 4–5; CYAS No growth; OA 25–28; CREA No growth; YES 15–17. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse dark brown (6F6) centre fading into yellowish white (3A2). MEA 25 C, 7 d: Colonies low, sulcate; margins low, plane, entire (1–2 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse dark brown (6F6) centre fading into light yellow to light orange (4A4–5A5). YES 25 C, 7 d: Colonies raised, sulcate, pinkish red colour; margins low, plane, entire (1 mm); mycelia white and pale red; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse mustard brown (5E6) centre fading into brownish orange (5C3). DG18 25 C, 7 d: Colonies no sporulation colonies up to 4 mm. OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish orange. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, (40–)70–100(–200) × 1.6– 2.5 μm; metulae two to ﬁve, divergent, 8.5–12 × 2.5–3.5 μm; 258 ITS barcode: AB176620 Colony diam, 7 d (mm): Fide Yaguchi et al. (1996) CYA 3; CYA 37 C No growth; MEA 8–10; OA: 5–8. Colony characters: Fide Yaguchi et al. (1996) colonies on CYA growing very restrictedly, attaining a diameter of 3 mm in 7 d and 10–12 mm in 14 d at 25 C, velvety to somewhat funiculose, almost plane, conisisting of a compact basal felt, producing abundant ascomata on the felt within 21 d, light yellow (3A5), overgrown by pigmented aerial hyphae; margins entire, narrow; sporulation absent; exudates small, clear; reverse brownish orange (5C6). Colonies on MEA growing restrictedly, attaining a diameter of 8–10 mm in 7 d and 17–20 mm in 14 d at 25 C, velvety to ﬂoccose, centrally raised, conisisting of a thick basal felt; mycelia white to light yellow (2A5); ascomata very limited in number; sporulation absent; exudate abundant, clear to pale brown; margins entire; reverse greyish orange (5B4). Colonies on OA growing restrictedly, attaining diameter of 5–8 mm in 7 d and 15–17 mm in 14 d at 25 C, radially sulcate, more or less zonate, consisting of a thin basal felt with funiculose aerial hyphae, granular due to abundant production of ascomata within 21 d, pastel yellow (3A4); margins thin, broad, entire; sporulation absent; exudate abundant, clear, rather large; reverse greyish orange (5B4). Micromorphology: Fide Yaguchi et al. (1996) asexual state not observed. Ascomata discrete or often conﬂuent, yellow, maturing slowly within 21 d, globose to subglobose, 300–350 μm in diam; asci globose to ovoidal, 10.5–12.5 × 9–11 μm, evanescent. Ascospores ellipsoidal, 5.5–7 × 3.5–4.5 μm, ridges bearing 12–15 longitudinal, somewhat sinous ridges about 0.5 μm wide, usually covering at the two end. Notes: According to Yaguchi et al. (1996), the outstanding characters of T. hachijoensis are its extremely restricted growth on CYA, colony colouration on CYA and OA, production of yellow POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 40. Morphological characters of Talaromyces galapagensis (CBS 751.74T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on MEA after 2 wk incubation. C–G. Conidiophores. H. Conidia. Scale bars: B = 100 μm; C = 10 μm, applies to D–H. www.studiesinmycology.org 259 YILMAZ ET AL. ascomata with ellipsoidal, striate ornamentated ascospores and absence of conidiophores. Talaromyces helicus (Raper & Fennel) C.R. Benj., Mycologia 47: 684. 1955. MycoBank MB306715. Fig. 41. ≡ Penicillium helicum Raper & Fennell, Mycologia 40: 515. 1948. = Talaromyces helicus var. major Stolk & Samson, Stud. Mycol. 2: 19. 1972. = Talaromyces barcinensis Yaguchi & Udagawa Trans. Mycol. Soc. Japan 34: 15. 1993 ≡ Penicillium barcinense Yaguchi & Udagawa Trans. Mycol. Soc. Japan 34: 15. 1993. In: Talaromyces section Helici Typus: IMI 040593, culture ex-type CBS 335.48 = ATCC 10451 = DSM 3705 = IMI 040593 = NRRL 2106. ITS barcode: JN899359 (alternative markers: BenA = KJ865725; CaM = KJ885289; RPB2 = KM023273) Colony diam, 7 d (mm): CYA 13–23; CYA 30 C 18–28; CYA 37 C 10–18 (DTO 56-A8 no growth); MEA 25–33; MEA 30 C 35–40; DG18 5–12; CYAS No growth; OA 23–35; CREA No growth; YES 14–22. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1–2 mm); mycelia white and pale orange; texture ﬂoccose; sporulation absent to moderately dense, conidia en masse greyish green (25B3); soluble pigments absent; exudates absent; reverse in some isolates centre greyish red (9B6) and in some isolates yellowish brown (5E4) centre fading into greyish orange (5B3) and yellowish white (4A2). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (26B3); soluble pigments absent; exudates absent; reverse in some isolates brownish orange (6C6) and in some isolates yellowish brown (5D5) centre fading into brownish orange (5C5). YES 25 C, 7 d: Colonies sunken at centre, sulcate, sterile white appearance; margins low, plane, entire (1 mm); mycelia white; sporulation absent to sparse; soluble pigments absent; exudates absent; reverse shades of dull green (27E4) fading into between pale yellow (4A3) and pale orange (5A3). DG18 25 C, 7 d: Colonies low to slightly raised at centre, plane, white mycelia appearance; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane, start to produce white and in some isolates yellow ascomata at 30 C ascomata is more abundant; margins low, plane, entire (3–5 mm); mycelia white and bright yellow; texture ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (26B3); soluble pigments absent; exudates absent (at 30 C clear droplets); reverse beige and pale yellow (at 30 C bright orange). CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, 30–60(–80) × 2–2.5 μm; metulae two to ﬁve, divergent, 12–15 × 2–2.5 μm; phialides acerose, two to four per metulae, 8.5–12(–16) × 2.5–3 μm; conidia smooth, globose to subglobose, 2.5–3.5(–4.5) × 2.2–3.5 μm. Ascomata maturing within 1–2 wk on OA at 25 C and at 30 C abundantly, 100–300 μm, yellow, pastel yellow and creamish white, usually globose and 260 sometimes subglobose, asci 6–9 × 4.5–6 μm, ascospores ellipsoidal, usually smooth, in some with minute spines, 2.5–4 × 2–3 μm. Extrolites: Talaromyces helicus produces helicusins (Yoshida et al. 1995) and italinic acid. Distinguishing characters: Talaromyces helicus is distinguished by its creamish white to yellow ascomata that are covered with yellow mycelia and produces smooth, ellipsoidal ascospores and mono- to biverticillate conidiophores with a green stipe (Fig. 41). It grows relatively fast on CYA and MEA and does not grow on CREA. Its smooth, ellipsoidal ascospores distinguish T. helicus from the other creamish white to yellow ascoma producers. Its green stipes resemble T. varians, however, T. helicus produces ascomata and T. varians does not. Also, T. varians produces cylindrical to ellipsoidal conidia, whereas T. helicus produces globose to subglobose conidia. Notes: Talaromyces barcinensis (CBS 649.95) and T. helicus var. major (CBS 652.66) are phylogenetically identical to T. helicus (Fig. 1) and are considered synonyms. Talaromyces indigoticus Takada & Udagawa, Mycotaxon 46: 129. 1993. MycoBank MB359290. Fig. 42. ≡ Penicillium indigoticum Takada & Udagawa, (simultaneously published). In: Talaromyces section Talaromyces Holotypus: CBM SUM-3010, culture ex-type CBS 100534 = IBT 17590. ITS barcode: JN899331 (alternative markers: BenA = JX494308; CaM = KF741931) Colony diam, 7 d (mm): CYA 20–21; CYA 30 C 24–25; CYA 37 C 20–22; MEA 32–33; MEA 30 C 43–45; DG18 9–11; CYAS No growth; OA 15–17; CREA 13–15; YES 28–30. Colony characters: CYA 25 C, 7 d: Colonies low, plane, pinkish white appearance; margins low, plane, entire (1 mm); mycelia white and red pigmented (red crystal on mycelia); texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange (5A3) centre fading into yellowish white (4A2). MEA 25 C, 7 d: Colonies deep, plane, has a characteristic smell, ﬂuffy white appearance; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish red (10D8) centre fading into red (10B8). YES 25 C, 7 d: Colonies deep to low, plane, has a characteristic smell, ﬂuffy white and pale pink appearance; margins low, plane, entire (1 mm); mycelia white and pastel red; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange (5A3) centre fading into yellowish white (4A2). DG18 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation sparse, conidia en masse greyish green (25B4–25C4); soluble pigments absent; exudates absent; reverse pale yellow to light yellow (4A3–4A4). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 41. Morphological characters of Talaromyces helicus (CBS 335.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D. Asci and ascopsores. E. Initials. F–H. Conidiophores. I. Ascospores. J. Conidia. Scale bars: B = 500 μm; C = 250 μm; D = 10 μm, applies to E–J. www.studiesinmycology.org 261 YILMAZ ET AL. Fig. 42. Morphological characters of Talaromyces indogoticus (CBS 100534T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Initials. C–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. 262 POLYPHASIC reverse creamish white, beige. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, 10–75 × 2–3 μm; metulae two to four, divergent, 8–12 × 2–2.8 μm; phialides acerose, three to ﬁve per metulae, 8–14.5 × 2–3 μm; conidia smooth, ovoidal to ellipsoidal, 2.4–4 × 2–3.2 μm. Ascomata not observed in our culture however, ﬁde Takada & Udagawa (1993) maturing after 1–2 wk of incubation, yellow to orange, globose to subglobose, 350–550 μm, asci 9–12.5 × 7.5–10 μm, ascospores indigo blue, ellipsoidal, spiny, 3.5–5 × 2.5–3 μm. Distinguishing characters: Talaromyces indigoticus produces yellow to orange ascomata with indigo blue, spiny, ellipsoidal ascospores and mono- to biverticillate conidiophores with short stipes. It grows fast at 30 C. Blue ascospores have never been reported in other Talaromyces species. The T. indigoticus strain available for this study, has lost its ability to produce ascomata. Talaromyces intermedius (Apinis) Stolk & Samson, Stud. Mycol. 2: 21. 1972. MycoBank MB324418. Fig. 43. ≡ Penicillium intermedium Stolk & Samson, (simultaneously published). In: Talaromyces section Talaromyces Typus: CBS H-7828, culture ex-type CBS 152.65 = BDUN 267 = IFO 31752 = IMI 100874. ITS barcode: JN899332 (alternative markers: BenA = JX091387; CaM = KJ885290) Colony diam, 7 d (mm): CYA 15–16; CYA 30 C 7–8; CYA 37 C No growth; MEA 48–50; MEA 30 C 24–26; DG18 5–6; CYAS No growth; OA 34–35; CREA No growth; YES 28–30. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, plane, entire; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse greyish yellow (3B4). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (5 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse apricot yellow (5B6). YES 25 C, 7 d: Colonies sunken at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A5) fading into yellowish white (4A2). DG18 25 C, 7 d: Colonies low, plane; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (5–8 mm); mycelia white; texture ﬂoccose, young ascomata produced; sporulation absent; soluble pigments absent; exudates absent; reverse pastel pinkish white. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores lacking or sparse on all media, best development on hay-infusion agar. When present monoverticillate or with solitary phialides; stipes smooth walled, 4–30 × 1.5–2 μm; phialides ﬂask-shaped to acerose, one to three, 12–20 × 1.5–2 μm; conidia smooth, ellipsoidal, 2.5–4.5 × 2.2–3.5 μm. Ascomata maturing within 1–2 wk on OA www.studiesinmycology.org TAXONOMY OF TALAROMYCES at 25 C and at 30 C abundantly, 300–1000 μm, at ﬁrst creamish white and then becoming pastel pink, globose, asci 11–16 × 10–14 μm, ascospores broadly ellipsoidal, thick walled, spiny, 4.5–7 × 3.5–5.5 μm. Distinguishing characters: Talaromyces intermedius produces creamish white to pastel pink ascomata, grows restrictedly on CYA and DG18, has conidiophores with solitary phialides and does not grow on CREA (Fig. 43). It produces thick walled ascospores that are spiny and ellipsoidal. Its ascomata resemble T. trachyspermus and T. assiutensis, but T. intermedius differs from the latter species by conidiophores that have solitary phialides, as well as producing bigger ascospores (4.5–7 × 3.5–5.5 μm). Talaromyces islandicus (Sopp) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560654. Fig. 44. ≡ Penicillium islandicum Sopp, Skr. Vidensk.-Selsk. Christiania, Math.Naturvidensk. Kl. 11: 161. 1912. = Penicillium aurantioﬂammiferum C. Ramírez, A.T. Martínez & Berer, Mycopathologia 72: 28. 1980. In: Talaromyces section Islandici Typus: IMI 040042, culture ex-type CBS 338.48 = ATCC 10127 = IMI 040042 = MUCL 31324 = NRRL 1036 = IBT 14884 = IBT 4476. ITS barcode: KF984885 (alternative markers: BenA = KF984655; CaM = KF984780; RPB2 = KF985018) Colony diam, 7 d (mm): CYA 20–27; CYA 30 C 20–28; CYA 37 C 8–17; MEA 21–26; MEA 30 C 20–28; DG18 15–25; CYAS 10–20; OA 20–30; CREA 7–11; YES 22–30. Colony morphology: CYA 25 C, 7 d: Colonies raised at centre, crateriforme; margins narrow (1–2 mm), low, entire, plane; mycelium white, yellow and orange; texture velvety and in some isolates loosely funiculose; sporulation sparse to dense; conidia en masse greyish green to dull green (26C3–26D3) (and for DTO 2-C7 dark green (26F4)); exudates big clear droplets in the strains which are not sporulating well; soluble pigment absent; reverse brown (7E4) in the centre fading into light yellow to light orange (4A4–5A4), in some isolates lack of reverse colour (CBS 338.48T). MEA 25 C, 7 d: Colonies slightly raised at centre, crateriforme; margins narrow (1–2 mm), low, entire, plane; mycelium white and orange; texture velvety and in some isolates loosely funiculose, especially in the centre conidiophores born from aerial hyphae; sporulation sparse to dense; conidia en masse greyish green to dull green to dark green (26E4–26F4 to 26C3–26C4); exudates big clear and yellow droplets in the strains which are not sporulating well; soluble pigment absent; reverse brownish orange (6C7) and in some isolates brown (7E6) fading into brownish orange (7C7). YES 25 C, 7 d: Colonies raised at centre, crateriforme; margins narrow (1–2 mm), low, entire, plane; mycelium white, red and orange; texture velvety to ﬂoccose; sporulation absent to dense; conidia en masse greyish green to dull green to dark green (26E4–26F4 to 26C3–26C4); exudates big clear droplets in the strains which are not sporulating well; soluble pigment absent; reverse brownish orange (6C5–6C8). DG18 25 C, 7 d: Colonies very slightly raised at centre, slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white and orange; texture 263 YILMAZ ET AL. Fig. 43. Morphological characters of Talaromyces intermedius (CBS 152.65T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C–D. Ascomata. E. Conidiophores. F. Initials. G. Asci and ascospores. H. Ascospores. Scale bars: B, C = 500 μm; D = 100 μm; E = 10 μm, applies to F–H. 264 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 44. Morphological characters of Talaromyces islandicus (CBS 338.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. www.studiesinmycology.org 265 YILMAZ ET AL. velvety especially in the centre conidiophores born from aerial hyphae; sporulation sparse to dense; conidia en masse dark green (27F4); exudates absent; soluble pigment absent; reverse reddish brown (8E5) in the centre fading into golden yellow (5B7) to light yellow (4A5). OA 25 C, 7 d: Colonies low, plane; margins wide (2–3 mm), low, entire, plane; mycelium white and orange; texture velvety; sporulation dense; conidia en masse dull green to dark green (27F4 to 27D3–27D4); exudates big clear and yellow droplets in the strains which are not sporulating well; soluble pigment absent (except DTO 158-D6 and DTO 2-C7 weak red soluble pigment); reverse brownish orange (6C7) and in some isolates red and in some isolates reddish orange in the centre fading into green. CREA, 25 C, 7 d: Strong acid production. Talaromyces liani (Kamyschko) Yilmaz, Frisvad & Samson, comb. nov. MycoBank MB809555. Fig. 45. Micromorphology: Conidiophores biverticillate often additional branches occur; stipes smooth walled, 20–200 × 2.5–3.5 μm; with extra branches 13–30 μm; metulae three to six, divergent, 6–12 × 2.3–3.2 μm; phialides acerose, three to six per metulae, 7–10 × 1.5–3 μm; conidia smooth, ellipsoidal, 2.5–6 × 2–4.5 μm. Colony diam, 7 d (mm): CYA 20–30; CYA 30 C 25–37; CYA 37 C 20–25; MEA 35–45; MEA 30 C 50–55; DG18 10–17; CYAS No growth; OA 35–40; CREA 10–20; YES 35–40. Extrolites: 3-hydrophthalic acid (Gatenbeck 1957), islandicin = 1,4,5-trihydroxy-2-methylanthraquinone (Howard 1948, Howard & Raistrick 1949); chrysophanic acid (Howard & Raistrick 1950); skyrin and ﬂavoskyrin (Howard & Raistrick 1954a); iridoskyrin and (−)-rubroskyrin (Howard & Raistrick 1954b, Takeda et al. 1973), endocrocin (Gatenbeck 1959), emodin (Gatenbeck 1958, Sankawa et al. 1973, Kawai et al. 1984); (−)-luteoskyrin (Yamamoto et al. 1956, Ueno & Ishikawa 1969, Takeda et al. 1973, Ghosh et al. 1978a); (−)-rugulosin (Takeda et al. 1973); erythroskyrine (Howard & Raistrick 1954b, Shoji et al. 1965, Shibata et al. 1966, Kenkyusho 1983); 1,4,7,8-tetrahydroxy-2-anthraquinone, chrysophanol, catenarin and dimers of those and emodin and islandicin: dianhydrorugulosin, dicatenarin, roseoskyrin, auroskyrin, rhodoislandicin A & B, punicoskyrin, aurantioskyrin, oxyskyrin, skyrinol, (−)-deoxyrubroskyrin, (−)-deoxyluteoskyrin, (−)4a-oxyluteoskyrin (Ogihara et al. 1968, Takeda et al. 1973); islandic acid I & II (Fujimoto et al. 1982); pibasterol (Ghosh et al. 1978a); cyclochlorotine = islanditoxin = chlorine containing peptide = chloropeptide and simatoxin (Marumo & Sumiki 1955, Uraguchi et al. 1961, 1972, Ghosh et al. 1978b). Extrolites detected in this study (in CBS 338.48T; NRRL 1036; CBS 165.81; IBT 12697; IBT 15605; FRR 3445; FRR 3606; CBS 189.68 and CBS 117284): erythroskyrin, islandicin, luteoskyrin, mitorubrinic acid and other special mitorubrins, rugulosin & skyrin. UV spectra indicated that T. islandicus can also produce wortmannilactone E-H, in addition to some extrolites unique to this species. Distinguishing characters: Talaromyces islandicus is characterised by relatively fast growing colonies compared to other species in section Islandici. Talaromyces islandicus has bright orange mycelia and produces colonies with orange reverses on most media. Based on the ITS and BenA phylogenies, T. islandicus has T. loliensis as a close relative. Talaromyces islandicus differs from T. loliensis by the production of acid on CREA, its faster growth and wider conidiophores. Notes: Phylogenetically P. aurantioﬂammiferum (CBS 165.81) is identical to T. islandicus (Fig. 7) and is considered a synonym. 266 Basionym: Penicillium liani Kamyschko, Not. Syst. Crypt. Inst. bot. Acad. Sci. USSR 15: 86. 1962. = Talaromyces thermocitrinus Subrahm. & Gopalkr., Ind. Bot. Reporter 35: 35. 1984 [as ‘thermocitrinum’]. In: Talaromyces section Talaromyces Typus: Unknown, culture ex-type CBS 225.66 = ATCC 18325 = ATCC 18331 = IMI 098480 = NRRL 3380 = VKM F-301. ITS barcode: JN899395 (alternative markers: BenA = JX091380; CaM = KJ885257) Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly sulcate, white and pastel yellow appearance; margins low, plane, entire; mycelia white and pastel yellow; texture; sporulation absent to sparse; soluble pigments absent (except CBS 118885 produces yellow soluble pigment also at 30 and 37 C); exudates absent; reverse between light orange and light yellow (5A5–4A5), in some isolates (CBS 118885) pastel yellow (2D4) centre fading into greyish yellow to pale yellow (1B4 to 1A4). MEA 25 C, 7 d: Colonies low, plane, formation of yellow ascomata (at 30 C abundant yellow ascomata); margins low, plane, entire (2–3 mm); mycelia white and yellow; texture velvety to ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (26B4–26C4); soluble pigments absent; exudates absent; reverse brownish orange to greyish orange (5B6–5C6). YES 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white and pastel yellow; texture velvety to ﬂoccose; sporulation absent to dense (CBS 118885), conidia en masse dull green to greyish green (25D4–25D5); soluble pigments absent; exudates absent; reverse in some isolates centre between deep yellow and deep orange (4A8–5A8) fading into light yellow (4A5). DG18 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green to dull green (25D5–25E4); soluble pigments absent; exudates absent; reverse in some isolates dark green (28F5) centre fading into greyish green (28C5) and pastel green (28A4) and in some isolates (DTO 254-I1) light yellow (2A5). OA 25 C, 7 d: Colonies raised at centre, plane, formation of yellow to orange red ascomata (abundant at 30 C); margins low, plane, entire (2–3 mm); mycelia white and yellow; sporulation absent; soluble pigments absent; exudates absent; reverse pastel yellow. CREA 25 C, 7 d: Acid production absent (except CBS 118885 very weak). Micromorphology: Conidiophores monoverticillate and biverticillate; stipes smooth walled, 20–130 × 2.5–3.5 μm; branches 5–25 μm; metulae three to six, divergent, 9–20 × 2.5–3 μm; phialides acerose, three to six per metulae, 9–20 × 2–3.5 μm; conidia smooth, ellipsoidal, 2.5–4(–4.5) × 2–3.5 μm. Ascomata maturing after 1–2 wk of incubation on OA and MEA at 25 C, yellow to orange red, globose to subglobose, 150–550 × 150–545 μm, asci 9–13 × 7.5–11 μm, ascospores broadly ellipsoidal, spiny, 4–6 × 2.5–4 μm. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 45. Morphological characters of Talaromyces liani (CBS 118885). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D. Asci and ascopsores. E–G. Conidiophores. H. Ascospores. I. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–I. www.studiesinmycology.org 267 YILMAZ ET AL. Distinguishing characters: Talaromyces liani produces relatively fast growing colonies on MEA and CYA at 25, 30 and 37 C. It produces yellow to orange red ascomata and spiny ellipsoidal ascospores, similar to those of T. ﬂavus, T. convolutus, T. austrocalifornicus, T. ﬂavovirens, T. tratensis, T. macrosporus and T. muroii. Based on colony size at 25 C on CYA and MEA after 7 d, T. liani is more similar to T. muroii and T. macroporus. Talaromyces liani have ellipsoidal ascospores, whereas T. macrosporus has more subglobose to broadly ellipsoidal ascospores. Talaromyces muroii does not grow on CREA and produce yellow ascomata, whereas T. liani produces yellow to orange red ascomata. Notes: Samson et al. (2011) speculated that T. thermocitrinus might belong in the genus Eurotium. However, ITS and BenA sequences (unpublished data) from the ex-type strain are identical to T. liani strains and is considered to represent a synonym. Talaromyces loliensis (Pitt) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011. MycoBank MB560655. Fig. 46. ≡ Penicillium loliense Pitt, The genus Penicillium: 450. 1980. In: Talaromyces section Islandici Typus: IMI 216901, culture ex-type CBS 643.80 = ATCC 52252 = FRR 1798 = IMI 216901 = MUCL 31325 = IBT 4546. ITS barcode: KF984888 (alternative markers: BenA = KF984658; CaM = KF984783) Colony diam, 7 d (mm): CYA 10–13; CYA 30 C 11–12; CYA 37 C No growth; MEA 13–15; MEA 30 C 12–13; DG18 13–15; CYAS 7–8; OA 13–14; CREA 4–8; YES 13–15. Colony morphology: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins narrow (1 mm), low, entire, plane; mycelium white and yellow (CBS 172.91 with orange mycelia); sporulation absent; exudates small clear droplets; soluble pigment very weak yellow around colonies; reverse centre deep yellow to deep orange (4A8–5A8) fading into light yellow to yellow (3A5–3A6). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins narrow (1 mm), low, entire, plane; mycelium white and yellow; sporulation absent to sparse; texture ﬂoccose and loosely funiculose; conidia en masse greyish green to dark green (28D4–28F4); exudates small clear droplets; soluble pigment absent; reverse centre brownish orange to brown (6C7–6D8). YES 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins narrow (1 mm), low, entire, plane; mycelium white (in margins) and yellow (in centre); sporulation absent; exudates few very small clear droplets; soluble pigment absent; reverse centre deep yellow to deep orange (4A8–5A8) fading into light yellow to pastel yellow (3A4–3A5). DG18 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins narrow (1 mm), low, entire, plane; mycelium white and yellow; sporulation absent; exudates absent; soluble pigment very weak yellow around colonies; reverse centre deep yellow to deep orange (4A8–5A8) fading into light yellow to yellow (3A5–3A6). OA 25 C, 7 d: Colonies deep, plane; margins narrow (1 mm), low, entire, plane, slimy yeast like; mycelium white and yellow; sporulation moderately dense; texture velvety and ﬂoccose; conidia en masse greyish green to dark green (28D4–28F4); exudates small orange and clear droplets; soluble pigment absent; reverse orange yellow. CREA, 268 25 C, 7 d: In some isolates acid production absent and in some isolates very weak acid production. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 45–115 × 2.3–3.2 μm; metulae three to six, divergent, 6–12 × 2–3.6 μm; phialides acerose, three to six per metulae, 7–15 × 2–4 μm; conidia smooth, subglobose to ellipsoidal, some spores fusiform some spores just one ending has connections, 3–5 × 2.4–3.5 μm. Extrolites: Talaromyces loliensis produces mitorubrins and skyrin. No other known extrolites were detected, but two of the partly characterised extrolites were also found in other species in Talaromyces sect. Islandici. Distinguishing characters: Talaromyces loliensis is characterised by restricted growth on most media. Colonies are ﬂuffy and deep and have light yellow mycelia (Fig. 46). It resembles T. tratensis, however, T. tratensis produces yellow ascomata. Based on the ITS and BenA phylogenies, T. loliensis is closely related to T. islandicus. Talaromyces loliensis differs from T. islandicus by slower growth on most media. Talaromyces macrosporus (Stolk & Samson) Frisvad, Samson & Stolk, Antonie van Leeuwenhoek 57: 186. 1990. MycoBank MB126704. Fig. 47. ≡ Penicillium macrosporum Frisvad et al. nom. illegit. Art. 53 (non Penicillium macrosporum Berk. & Broome 1882, Ann. Mag. Nat. Hist. 9: 183. 1882) ≡ Talaromyces ﬂavus var. macrosporus Stolk & Samson Stud. Mycol. 2: 15. 1972. In: Talaromyces section Talaromyces Typus: CBS H-7822, culture ex-type CBS 317.63 = FRR 404 = IMI 197478. ITS barcode: JN899333 (alternative markers: BenA = JX091382; CaM = KF741952; RPB2 = KM023292) Colony diam, 7 d (mm): CYA 22–28; CYA 30 C 33–40; CYA 37 C 28–35; MEA 40–50; MEA 30 C 55–65; DG18 15–22; CYAS No growth; OA 40–45; CREA 15–20; YES 35–40. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, sulcate, pastel yellow and pinkish red appearance; margins low, plane, entire (1–2 mm); mycelia white, pastel yellow and pinkish red; sporulation absent to sparse; soluble pigments absent (at 30 and 37 C brownish soluble pigment); exudates absent; reverse yellowish brown (5E7–5F7) centre fading into brownish yellow to golden yellow (5C7–5B7). MEA 25 C, 7 d: Colonies low, plane, yellow ascomata formation (DTO 58-F4 & DTO 77-C5) (at 30 C, abundant ascomata except (DTO 108-A8)); margins low, plane, entire (2–3 mm); mycelia white, yellow and pinkish red; sporulation absent; soluble pigments absent; exudates absent (except DTO 58-F4, small red droplets at centre); reverse dark brown (8F8) centre fading into reddish brown (8E8–8D8) in the margins golden yellow (5B7). YES 25 C, 7 d: Colonies raised at centre, sulcate, light pinkish red and white appearance; margins low, plane, entire (3 mm); mycelia white and light pinkish red; sporulation absent; soluble pigments absent; exudates absent; reverse in some isolates centre dark brown (8F7) fading into orange POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 46. Morphological characters of Talaromyces loliensis (CBS 643.80T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 269 YILMAZ ET AL. Fig. 47. Morphological characters of Talaromyces macrosporus (CBS 317.63T). a. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D, E. Conidiophores. F. Conidia. G. Asci and ascospores. H. Ascospores. Scale bars: B = 1000 μm; C = 500 μm; D = 10 μm, applies to E–H. 270 POLYPHASIC (6B8), in some isolates centre brownish orange (5C4). DG18 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse dull green (25D4–25E4); soluble pigments absent; exudates absent; reverse in some isolates centre dark brown (6F5), fading into orange (6B7) to yellowish white (2A4). OA 25 C, 7 d: Colonies low, plane, yellow and white ascomata formation (at 30 C abundant ascomata except DTO 108-A8, yellow, orange in the centre and white in the margins); margins low, plane, entire (2–3 mm); mycelia white and yellow; sporulation absent; soluble pigments absent; exudates small clear and orange droplets; reverse pale brownish orange to orange and reddish orange. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, 15–50 × 2–3 μm; metulae three to ﬁve, divergent, 8–11 × 2–3(–4) μm; phialides acerose, three to six per metulae, (4–)8–12 × 2–3 μm; conidia smooth, subglobose to ellipsoidal, 2–4 × 2–3.5 μm. Ascomata maturing within 1–2 wk on OA at 25 C and 30 C abundantly, globose to subglobose, 100–400 × 100–350 μm, yellow, asci 13–15(–18) × 11.5–13.5 μm, ascospores subglobose to broadly ellipsoidal, thick walled, spiny, 5–6.5 × 4.5–5.5 μm. Extrolites: Talaromyces macrosporus produces duclauxin and mitorubrinic acid. Distinguishing characters: Talaromyces macrosporus produces yellow ascomata, thick walled, spiny, subglobose to broadly ellipsoidal ascospores and rapid growth on MEA (Fig. 47). Its yellow ascomata and rapid growth resemble T. muroii and T. liani. Talaromyces muroii and T. liani colonies sporulate better on MEA compared to T. macrosporus. Talaromyces muroii does not grow on CREA. Talaromyces liani produces yellow to orange red ascomata, whereas T. macrosporus produces only yellow ascomata. Additionally, T. macrosporus produces slightly bigger, subglobose to broadly ellipsoidal ascospores (5–6.5 × 4.5–5.5 μm) compared to T. liani (4–6 × 2.5–4 μm) and T. muroii (3.5–6 × 3–4 μm). The species is a well-known heat-resitant spoilage organism in various food products and beverages. Talaromyces marneffei (Segretain, Capponi & Sureau) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560656. Fig. 48. ≡ Penicillium marneffei Segretain, Capponi & Sureau, Bull. Soc. Mycol. France 75: 416. 1959 [1960]. In: Talaromyces section Talaromyces Typus: IMI 68794iii, culture ex-type CBS 388.87 = ATCC 18224 = CBS 334.59 = IMI 068794ii = IMI 068794iii. ITS barcode: JN899344 (alternative markers: BenA = JX091389; CaM = KF741958; RPB2 = KM023283) Colony diam, 7 d (mm): CYA 13–25; CYA 30 C 15–38; CYA 37 C 5–10; MEA 15–27; MEA 30 C 23–38; DG18 7–9; CYAS No growth; OA 20–25; CREA 8–15; YES 17–25. Colony characters: CYA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white and in some isolates pastel red; texture loosely funiculose and ﬂoccose (at 37 C yeast www.studiesinmycology.org TAXONOMY OF TALAROMYCES phase); sporulation moderately dense to dense, conidia en masse greyish green (25B4–26B4); soluble pigments red (in some isolates yellowish red in some isolates at 30 C lack of red soluble pigment); exudates absent; reverse reddish brown (9E8) fading into red (9A6). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (2 mm); mycelia white, yellow and pastel red; texture loosely funiculose and ﬂoccose, covered with yellow mycelia; sporulation moderately dense to dense, conidia en masse greyish green (29C6–30C6); soluble pigments red (also at 30 C); exudates absent; reverse dark brown (8F8). YES 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white; texture velvety and loosely funiculose; sporulation dense, conidia en masse greyish green (26B4–27B4); soluble pigments red; exudates absent; reverse reddish brown (8F8). DG18 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white and in some isolates yellow; texture velvety and loosely funiculose; sporulation dense, conidia en masse dull green (26D3–26E3); soluble pigments red (in some isolates weak); exudates absent; reverse reddish brown (8F8). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white and yellow; texture loosely funiculose and ﬂoccose, colonies covered with yellow mycelia; sporulation sparse to moderately dense, conidia en masse greyish green (29B4–29C4); soluble pigments red (in some isolates lack); exudates absent; reverse brownish red. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores generally biverticillate and sometimes monoverticillate; stipes smooth walled, 30–100 × 2.5–3 μm; metulae two to six, divergent, 7–11 × 2.5–3.5 μm; phialides acerose to ﬂask-shaped, four to seven per metulae, 6–8(–12) × 2.5–3.5 μm; conidia smooth, subglobose, 2.5–4 × 2–3 μm. Extrolites: Talaromyces marneffei produces Monascus red pigments including rubropunctatin, monascin, mitorubrin, mitorubrinol, mitorubrinic acid and a purpactin. Distinguishing characters: Talaromyces marneffei is the only dimorphic species of Talaromyces which grows as a yeast phase at 37 C (Fig. 48). It is a pathogen of immunocompromised patients, especially HIV positive individuals. It produces red soluble pigment on general media and conidiophores have ﬂask-shaped to acerose phialides. After 7 d, colonies on MEA are covered with sterile yellow mycelia, which resemble T. ﬂavovirens and T. primulinus. Talaromyces ﬂavovirens differs from T. marneffei by rapid growth on MEA and lacks the reddish appearance of T. marneffei on MEA. After 2 wk of incubation, T. primulinus also produces a yellow mycelial layer that covers the colony. However T. primulinus grows restrictedly on general media and cannot grow at 37 C. Talaromyces mimosinus A.D. Hocking, The genus Penicillium: 507. 1980. MycoBank MB116382. Fig. 49. ≡ Penicillium mimosinum A. D. Hocking, (simultaneously published). In: Talaromyces section Bacillispori Typus: IMI 223991, culture ex-type CBS 659.80 = FRR 1875 = IMI 223991. ITS barcode: JN899338 (alternative markers: BenA = KJ865726; CaM = KJ885272) 271 YILMAZ ET AL. Fig. 48. Morphological characters of Talaromyces marneffei (CBS 549.77). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Mycelia. C. Yeast phase. D–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 272 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 49. Morphological characters of Talaromyces mimosinus (CBS 659.80T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. www.studiesinmycology.org 273 YILMAZ ET AL. Colony diam, 7 d (mm): CYA 12–15; CYA 30 C 17–18; CYA 37 C 3–5; MEA 13–14; MEA 30 C 20–21; DG18 8–10; CYAS 4–5; OA 13–15; CREA No growth; YES 14–15. C, Colony characters: CYA 25 7 d: Colonies raised at centre, slightly sunken at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white and pale pastel yellow; sporulation absent; soluble pigments absent; exudates absent; reverse light yellow (3A5). MEA 25 C, 7 d: Colonies raised at centre, slightly sunken at centre, sulcate; margins low, plane, entire (1 mm); mycelia white and pale pastel yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A5–5A6). YES 25 C, 7 d: Colonies raised at centre, slightly sunken at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white and pale pastel yellow; sporulation absent; soluble pigments absent; exudates absent; reverse maize yellow (4A6). DG18 25 C, 7 d: Colonies low, plane, slimy yeast like; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pale pastel yellow. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores lacking on all media. After 3 wk incubation best development on OA. Conidiophores biverticillate and monoverticillate with a minor proportion having subterminal branches; stipes smooth walled, 15–50(–90) × 1.8–2.2 μm; branches 8–22 μm; metulae two to ﬁve, divergent, 7–10(–13) × 1.8–2.5 μm; phialides acerose, two to ﬁve per metulae, 7–9(–12) × 1.8–2.5 μm; conidia smooth, globose to subglobose, 2–3 × 2–2.5 μm. Ascomata not observed in our culture but ﬁde Pitt (1980) maturing within 2–3 wk on MEA, pure yellow and sulphur yellow, 2.5 mm, asci 13–16 × 13–16 μm, Ascospores globose to subglobose, ornamented on all surfaces by conspicuous sinous ridges about 0.5 μm high, 7–8 × 6–7 μm. Extrolites: Talaromyces mimosinus produces some mitrorubrinlike extrolites. Distinguishing characters: Talaromyces mimosinus produces big (2.5 mm) pure yellow to sulphur yellow ascomata; very big (7–8 × 6–7 μm), globose to subglobose ascospores, ornamented with ridges; mono- to biverticillate conidiophores with globose to subglobose conidia (Fig. 49). These characters distinguish it from all other species. Conidiophores were only observed on OA after 3 wk of incubation. Talaromyces minioluteus (Dierckx) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560657. Fig. 50. ≡ Penicillium minioluteum Dierckx, Ann. Soc. Sci. Bruxelles 25: 87. 1901. = Penicillium gaditanum C. Ramírez & A.T. Martínez, Mycopathologia 74: 165. 1981. = Penicillium purpurogenum var. rubrisclerotium Thom, Mycologia 7: 142. 1915. = Penicillium samsonii Quintan., Mycopathologia 91: 69. 1985. In: Talaromyces section Trachyspermi Typus: CBS 642.68, culture ex-type CBS 642.68 = IMI 089377 = MUCL 28666. 274 ITS barcode: JN899346 (alternative markers: BenA = KF114799; CaM = KJ885273; RPB2 = JF417443) Colony diam, 7 d (mm): CYA 17–18; CYA 30 C 10; CYA 37 C No growth; MEA 21–22; MEA 30 C 9–10; DG18 14–15; CYAS No growth; OA 22–23; CREA 2–3; YES 18. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly concentrically sulcate; margins low, plane, entire (1 mm); mycelia white; sporulation absent to sparse; soluble pigments weak brownish red; exudates absent; reverse brown (6E8) centre fading into brownish orange to light brown (6C8–6D8). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly concentrically sulcate; margins low, plane, entire (1 mm); mycelia yellow and white; texture loosely funiculose and ﬂoccose, conidiophores born from sterile aerial hyphae; sporulation dense, conidia en masse greyish green to dull green (27E4–27E6); soluble pigments absent; exudates absent; reverse brownish orange (7C8). YES 25 C, 7 d: Colonies raised at centre, sulcate, crateriform; margins low, plane, entire (1 mm); mycelia white; texture velvety; sporulation dense, conidia en masse greyish green to dull green (26D4–26D5); soluble pigments absent; exudates absent; reverse orange (6B8). DG18 25 C, 7 d: Colonies slightly raised at centre, very slightly concentrically sulcate; margins low, plane, entire (1 mm); mycelia white; sporulation absent to sparse; soluble pigments absent; exudates clear droplets; reverse brownish orange (7C8) centre fading into orange (7B8). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture slimy hyphae; sporulation absent; soluble pigments absent; exudates absent; reverse light pale orange. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 80–300 × 2–3 μm; metulae three to ﬁve, divergent, 10–20 × 2–3.5 μm; phialides acerose, three to ﬁve per metulae, 10–17 × 2–3 μm; conidia smooth, ellipsoidal, 2.5–4 × 1.5–2.5 μm. Ascomata not observed. Extrolites: Talaromyces minioluteus produces mitorubrin, mitorubrinic acid, a purpactin, secaslonic acid D and vermicellin. It has also been reported to produce minioluteic acid and spiculisporic acid (Birkinshaw & Raistrick 1934), 22-epoxyberkeleydione, miniolutelide A & B (Iida et al. 2008), HY558-1 (Haeyoung et al. 2004), NG-061 (Ito et al. 1999) and BE-31405 (Okada et al. 1998). Some of these metabolites may be produced by Penicillium minioluteum sensu Pitt (which can be Talaromyces amestolkiae or T. ruber). It is also an efﬁcient producer of dextranase (Roca et al. 1996). Distinguishing characters: Talaromyces minioluteus grows moderately on general media, produces weak acid on CREA and cannot grow at 37 C. Both Penicillium samsonii and P. purpurogenum var. rubisclerotiorum resemble T. minioluteus and ITS and BenA phylogenies also show that latter two species are synonyms of T. minioluteus. Notes: van Reenen-Hoekstra et al. (1990) considered P. gaditanum (CBS 169.81) synonymous with T. minioluteus based on morphological and chemical data, which we follow here. Our ITS and BenA data show that P. samsonii (CBS 137.84) and P. purpurogenum var. rubisclerotium (CBS 270.35) are POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 50. Morphological characters of Talaromyces minioluteus (CBS 642.68T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. www.studiesinmycology.org 275 YILMAZ ET AL. phylogenetically identical to T. minioluteus (Fig. 5) and are considered synonyms. Talaromyces muroii Yaguchi, Someya & Udagawa, Mycoscience 35: 252. 1994. MycoBank MB362930. Fig. 51. = Penicillium victoriae Szilvinyi, Arch. Hydrobiol. 14, (suppl. 3): 538. 1936. Distinguishing characters: Talaromyces muroii produces relatively fast growth on CYA at 30 and 37 C, MEA at 25 and 30 C, and does not grow on CREA. It produces yellow ascomata and spiny, ellipsoidal ascospores (Fig. 51). Its yellow ascomata and rapid growth resemble T. macrosporus and T. liani. Talaromyces muroii produces ellipsoidal ascospores, whereas T. macrosporus has more subglobose to broadly ellipsoidal ascospores. Talaromyces liani grows on CREA and produces yellow to orange red ascomata. In: Talaromyces section Talaromyces Typus: CBM PF-1153, culture ex-type CBS 756.96 = PF 1153. Notes: Penicillium victoriae (CBS 274.26) is phylogenetically identical to T. muroii and is considered a synonym. ITS barcode: JN899351 (alternative markers: BenA = KJ865727; CaM = KJ885274) Talaromyces oumae-annae Visagie et al., Stud. Mycol. 78: 130. 2014. MycoBank MB809187. Colony diam, 7 d (mm): CYA 15–16; CYA 30 C 20–22; CYA 37 C 19–20; MEA 30–32; MEA 30 C 37–40; DG18 10–13; CYAS 2–3; OA 27–30; CREA No growth; YES 25–26. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (1 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (25C3–25C4); soluble pigments absent; exudates absent; reverse reddish brown (8E7) fading into brownish red (7C7–8C7) and in some isolates also greyish green (1C3). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (3–5 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (25C3–25C4); soluble pigments absent; exudates sticky clear droplets; reverse brownish orange (5C6) to light brown (5D6) and in some isolates reddish brown (9E7) dots. YES 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white, pastel yellow and pastel red; texture velvety and ﬂoccose; sporulation sparse to dense, conidia en masse dull green (26D4–26E4); soluble pigments absent; exudates clear and yellow droplets; reverse olive (2F3–2F4) reverse with shades (dots) of reddish brown (9E8). DG18 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (1–2 mm); mycelia white; texture velvety; sporulation dense, conidia en masse dull green (26E3–27E3); soluble pigments absent; exudates small clear droplets; reverse greyish green (26E5–27E5). OA 25 C, 7 d: Colonies slightly raised at centre, plane, yellow ascomata formation (at 30 C more abundat 1 wk later); margins low, plane, entire (2–3 mm); mycelia white; texture velvety and ﬂoccose; sporulation dense, conidia en masse greyish green (26D4–26E4); soluble pigments absent; exudates small clear droplets; reverse red. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate with a minor proportion having subterminal branches; stipes smooth walled, 20–50(–100) × 2.5–3 μm; branches 5–20 μm; metulae three to six, divergent, 10–16 × 2–3 μm; phialides acerose, three to six per metulae, 11.5–15(–20) × 2–3 μm; conidia smooth, globose to subglobose, 2–3 × 1.5–3 μm. Ascomata maturing after 1–2 wk of incubation on OA at 25 C and abundantly 30 C, yellow and pastel yellow, globose to subglobose, 140–290 × 160–330 μm, asci 9.5–13 × 7.5–9.5 μm, ascospores ellipsoidal, ﬁnely thick walled, spiny, 3.5–6 × 3–4 μm. 276 In: Talaromyces section Talaromyces Typus: CBS H-21797, culture ex-type CBS 138208 = DTO 269E8. ITS barcode: KJ775720 (alternative markers: BenA = KJ775213; CaM = KJ775425) Colony diam, 7 d (mm): CYA 16–18; CYA 30 C 16–17; CYA 37 C 10–11; MEA 29–30; MEA 30 C 35–37; DG18 14–17; CYAS No growth; OA 30–35; CREA 5–6; YES 20–23. Colony characters: Fide Visagie et al. (2014) CYA 25 C, 7 d: Colonies low, slightly raised at centre, plane; margins low, narrow, entire; mycelia white; texture ﬂoccose and velvety; sporulation moderately dense to dense, conidia en masse dull green (25D4–25E4); soluble pigments yellow; exudates absent; reverse greyish green (29B6–29C6). MEA 25 C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white and pastel yellow; texture velvety, centrally ﬂoccose with sterile aerial mycelia; sporulation dense, conidia en masse greyish green (27D5–27E5); soluble pigments absent; exudates absent; reverse light brown (7D6) in the centre fading into brownish orange (6C6). YES 25 C, 7 d: Colonies low, raised at centre, lightly sulcate; margins low, narrow, entire; mycelia white to yellow; texture velutinous to ﬂoccose; sporulation dense, conidia en masse dull green (25D4–25E4); soluble pigments yellow; exudates absent; reverse centre light yellow to greyish yellow (2A5–2B5), at margins greyish green (27E5). DG18 25 C, 7 d: Colonies moderately deep, lightly sulcate; margins low, narrow, entire; mycelia white to yellow; texture ﬂoccose; sporulation dense, conidia en masse greyish green (26E5–27E5); soluble pigments absent; exudates absent; reverse light orange (6A4–6A5). OA 25 C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white; texture velvety; sporulation dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent. CREA 25 C, 7 d: Acid production absent. Micromorphology: Fide Visagie et al. (2014) conidiophores biverticillate, subterminal branches sometimes present; stipes smooth walled, 85–240 × 2.5–3.5 μm; branches up to 30 μm long; metulae appressed, 8–11(–12.5) × 2.5–3.5 μm; phialides acerose, 9–11.5 × 2–3 μm; conidia rough, ellipsoidal, 3–3.5 × 2.5–3 μm. Notes: Fide Visagie et al. (2014) Talaromyces oumae-annae grows restrictedly on CYA and DG18, but grows moderately on POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 51. Morphological characters of Talaromyces muroii (CBS 756.96T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D. Asci and ascopsores. E–G. Conidiophores. H. Ascospores. I. Conidia. Scale bars: B = 500 μm; C = 250 μm; D = 10 μm, applies to E–I. www.studiesinmycology.org 277 YILMAZ ET AL. other media. It produces biverticillate conidiophores with some subterminal branches, smooth-walled stipes and has roughened, ellipsoidal conidia. It is phylogenetically resolved as a close relative to T. verruculosus and T. viridulus (Fig. 2). However, T. oumae-annae produces ellipsoidal conidia compared to spheroid conidia of T. verruculosus, which also grows faster on CYA at all temperatures (CYA 32–35; CYA 30 C 37–38; CYA 37 C 25–26). Talaromyces viridulus, originally described as Geosmithia viridis (Pitt & Hocking 1985), produces rod-shaped conidia, making identiﬁcation of the species rather easy. For an illustration of the species, readers are referred to Visagie et al. (2014). Distinguishing characters: Talaromyces palmae characteristically produces indeterminate synnemata, especially at colony margins. Conidiophores typically have short stipes (up to 85 μm) and colonies grow restricted on DG18 and do not grow at 37 C. Talaromyces duclauxii also produces indeterminate synnemata, but T. palmae grows slower on CYA at 30 C. Talaromyces palmae (Samson, Stolk & Frisvad) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560658. Fig. 52. Talaromyces panamensis (Samson, Stolk & Frisvad) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560659. Fig. 53. ≡ Penicillium palmae Samson, Stolk & Frisvad, Stud. Mycol. 31: 135. 1989. ≡ Penicillium panamense Samson, Stolk & Frisvad, Stud. Mycol. 31: 136. 1989. conidia smooth, subglobose to ellipsoidal, 3–4.5 × 2–3.5 μm. Ascomata not observed. Extrolites: Talaromyces palmae produces mitorubrin, mitorubrinol and mitorubrinol acetate (Samson et al. 1989). In: Talaromyces section Subinﬂati In: Talaromyces section Talaromyces Typus: CBS 442.88, culture ex-typue CBS 442.88 = IMI 343640. Typus: CBS 128.89, culture ex-type CBS 128.89 = IMI 297546. ITS barcode: JN899396 (alternative markers: BenA = HQ156947; CaM = KJ885291; RPB2 = KM023300) ITS barcode: JN899362 (alternative markers: BenA = HQ156948; CaM = KF741936; RPB2 = KM023284) Colony diam, 7 d (mm): CYA 20–25; CYA 30 C 20–22; CYA 37 C No growth; MEA 22–26; MEA 30 C 25–27 mm; DG18 4–5; CYAS No growth; OA 25–27; CREA 7–11; YES 25–26. Colony diam, 7 d (mm): CYA 23–24; CYA 30 C 20–21; CYA 37 C No growth; MEA 28–30; MEA 30 C 27–28; DG18 10–11; CYAS 2–3; OA 33–35; CREA 22–25; YES 30–32. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, crateriforme; margins low, plane, entire (1 mm); mycelia white and yellow; texture velvety; sporulation dense, conidia en masse dull green (28D4–29D4); soluble pigments absent; exudates absent; reverse pastel yellow to greyish yellow (2A4–2B4). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1 mm); mycelia white and yellow; texture velvety in centre aerial hyphae, production of synnemata started; sporulation dense, conidia en masse dull green (28D4–29D4); soluble pigments absent; exudates absent; reverse brownish orange (5B5–5B6). YES 25 C, 7 d: Colonies raised at centre, sunken at centre and sometimes crateriforme, sulcate; margins low, plane, entire (1–2 mm); mycelia white and yellow; texture velvety, early synnemata formation; sporulation dense, conidia en masse dull green (28D4–29D4); soluble pigments absent; exudates absent; reverse light yellow to yellow (3A5–3A6). DG18 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1 mm); mycelia white and light pastel yellow; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (3A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white and yellow; texture velvety and in the margins early synnemata production; sporulation moderately dense, conidia en masse dull green (28D4–29D4); soluble pigments absent; exudates absent; reverse pastel yellow. CREA 25 C, 7 d: Weak acid production. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white; sporulation absent to sparse, conidia en masse difﬁcult to determine; soluble pigments absent; exudates small clear droplets; reverse brownish orange to yellowish brown (5C6–5D6). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate, synnemata at centre sporulate; margins low, plane, entire (1 mm); mycelia white, yellow and orange; texture synnematous; sporulation sporulation only on synnemata densely, conidia en masse dark green (28F5); soluble pigments absent; exudates orange droplets; reverse yellowish brown (5E7) fading into golden brown to brownish yellow (5C7–5D7). YES 25 C, 7 d: Colonies slightly raised at centre, concentrically sulcate; margins low, plane, entire (1–2 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse brown (6E7) with shades of golden brown (5D7) fading into brownish yellow (5C6) with dots of brownish red (9C8). DG18 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (<1 mm); mycelia white and yellow; texture velvety; sporulation dense, conidia en masse greyish green (26D5–26E5); soluble pigments absent; exudates absent; reverse light brown (7D6) centre fading into brownish orange (6C7) and greyish orange (5B6). OA 25 C, 7 d: Colonies low, plane, sporulated synnemata at centre and young synnemata at margins; margins low, plane, entire (2–3 mm); mycelia white, yellow, orange; texture synnematous; sporulation only on synnemata densely, conidia en masse dark green (28F5); soluble pigments absent; exudates small clear droplets; reverse red. CREA 25 C, 7 d: Strong acid production. Micromorphology: Synnemata formed at the edge of the colony on MEA up to 8 000 μm long after 1–2 wk. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 25–85 × 3–4 μm; branches 10–15 μm; metulae three to ﬁve, divergent, 10–17 × 2.5–4 μm; phialides acerose, three to six per metulae, 13–20 × 2–3.5 μm; 278 Micromorphology: Synnemata formed after 7 d on MEA 1 800–5 000 μm long and on OA 800–6800 μm long. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 52. Morphological characters of Talaromyces palmae (CBS 442.88T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–E. Conidiophores. F. Conidia. Scale bar: B = 10 μm, applies to C–F. www.studiesinmycology.org 279 YILMAZ ET AL. Fig. 53. Morphological characters of Talaromyces panamensis (CBS 128.89T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 1 wk incubation. C–E. Conidiophores. F. Conidia. Scale bar: C = 10 μm, applies to D–F. 280 POLYPHASIC Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 40–85 × 3–4 μm; branches 15–25 μm; metulae three to six, divergent, 9.5–14 × 2.5–3.5(–4) μm; phialides acerose, three to six per metulae, 9.5–13.5 × 2–3 μm; conidia smooth, thick walled, ellipsoidal to fusiform, 3–5 × 2–3 μm. Ascomata not observed. Extrolites: Talaromyces panamensis produces mitorubrinic acid, mitorubrin and vermicillin (Samson et al. 1989). Distinguishing characters: The most striking characteristic of T. panamensis is the production of determinate synnemata with yellow stalks up to 5 000 μm (MEA) and 6 800 μm (OA) in length after 7 d of growth (Fig. 53). It produces acid on CREA. Morphologically, T. panamensis resembles T. calidicanius. However, T. calidicanius grows faster at 30 C on MEA and produces less acid on CREA. Talaromyces paucisporus (Yaguchi, Someya & Udagawa) Samson & Houbraken, Stud. Mycol. 71: 176. 2011. MycoBank MB560684. ≡ Erythrogymnotheca paucispora Yaguchi, Someya & Udagawa, Mycoscience 35: 219. 1994. Typus: PF 1150, culture ex-type PF 1150 = IFM 53616. Notes: Erythrogymnotheca paucispora was described by Yaguchi et al. (1994) introducing the new genus Erythrogymnotheca. It produces asci that have one to two and sometimes four ascospores, whereas Talaromyces generally produces asci with eight ascospores. However, molecular studies show that Erythrogymnotheca paucispora belongs in Talaromyces (Samson et al. 2011). Unfortunately, the ex-type strain was not available for this study. Macro- and micromophological characters are thus based on the original description (Yaguchi et al. 1994). According to Frisvad (unpublished data): Colony diameters (Strain IBT 17569): CYA 2–3 mm, cream reverse; MEA 8–10 mm orange red reverse and orange red exudate; YES 7–9 mm, cream yellow reverse; CYA at 37 C: 0–1 mm; OA 8–11 mm; OA at 30 C 32 mm, yellow mycelium and reddish reverse; OA at 37 C 16 mm; CREA weak growth (13 mm), weak acid under colony. Talaromyces piceus (Raper & Fennell) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560661. Fig. 54. ≡ Penicillium piceum Raper & Fennell, Mycologia 40: 533. 1948. = Penicillium ilerdanum C. Ramírez & A.T. Martínez, Mycopathologia 72: 32. 1980. In: Talaromyces section Islandici Typus: IMI 040038, culture ex-type CBS 361.48 = ATCC 10519 = IMI 040038 = NRRL 1051 = QM 7609 = IBT 4460. ITS barcode: AB176603 Colony diam, 14 d (mm): CYA 15–18; CYA 37 C No growth; MEA 19–20; OA 15–18. Colony characters: Fide Yaguchi et al. (1994) colonies on CYA growing restrictedly, attaining a diam of 15–18 mm in 14 d at 25 C, ﬂoccose, centrally convolute and raised, radially sulcate, more or less zonate, consisting of a compact basal felt, at ﬁrst white, then becoming greyish yellow to pastel pink (1B4–11A3), producing abundant ascomata on the felt, with a thin irregular margin; exudate very abundant red; odour indistinct; reverse and agar violet brown (11E8). Colonies on MEA agar growing restrictedly, attaning a diam of 19–20 mm in 14 d at 25 C, velvety to tomentose, almost plane, consisting of a thin basal felt, dull yellow to pastel pink (3B4–11A3), producing abundant ascomata on the felt, with a narrow entire margin; exudate lacking; reverse and agar ruby (11E8). Colonies on OA growing restrictedly, attaining a diam of 15–18 mm in 14 d at 25 C, velvety, plane, consisting of a thin basal felt, becoming granular due to development of abundant ascomata, pale yellow (4A3); exudates small, pink; reverse and agar greyish green. Micromorphology: Fide Yaguchi et al. (1994) asexual state lacking. Ascomata yellow to sometimes red, globose to subglobose, 200–500 μm in diam, maturing within 21 d; asci 1–2(–4) spored, globose to subglobose, 22–25(–26) × 18–22(–25) μm, ascospores subglobose to broadly ellipsoidal, thick walled (up to 2 μm), spiny (spines up to 1.5 μm), 14–18 × 12–16 μm. www.studiesinmycology.org TALAROMYCES ITS barcode: KF984792 (alternative markers: BenA = KF984668; CaM = KF984680; RPB2 = KF984899) In: Talaromyces section Talaromyces Extrolites: Talaromyces paucisporus dictyphenone and purpactins. TAXONOMY OF produces mono- Colony diam, 7 d (mm): CYA 20–27; CYA 30 C 28–33; CYA 37 C 30–35; MEA 25–27; MEA 30 C 30–33; DG18 15–20; CYAS 7–15; OA 20–27; CREA 11–20; YES 15–20. Colony morphology: CYA, 25 C, 7 d: Colonies raised in the centre, crateriforme; margins narrow (1–2 mm), low, entire, plane; mycelium white and pale yellow, in some strains also pale orange, colony has golden colour appearance; texture ﬂoccose; sporulation absent to sparse; conidia en masse greyish green (28B4–28C4); exudates clear and pale orange droplets in the centre; soluble pigment light brown; reverse brown (6E6). MEA, 25 C, 7 d: Colonies slightly raised in the centre, slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white and pale yellow, in some strains also orange; texture loosely funiculose and in some isolates ﬂoccose; sporulation sparse to dense; conidia en masse greyish green (28B4–28C4); exudates clear droplets in the centre; soluble pigment absent; reverse brown (6D7). YES, 25 C, 7 d: Colonies low, sulcate; margins narrow (1 mm), low, entire, plane; mycelium white and yellow, colonies with butter yellow to maize yellow (4B5–4B6) appearance; texture funiculose to ﬂoccose; sporulation absent to sparse; conidia en masse grayish green to dull green; exudates absent; soluble pigment absent; reverse orange to brownish orange in centre, fading into yellowish brown (5D5). DG18, 25 C, 7 d: Colonies low, slightly sulcate; margins narrow (1 mm), low, entire, plane; mycelium white and yellow; texture loosely funiculose to ﬂoccose; sporulation absent to moderately dense; conidia en masse grayish green to dull green (28B4–28C4–28D4); exudates absent; soluble pigment absent; reverse yellowish brown in centre (5D5), fading into light yellow (2A5). OA, 25 C, 7 d: Colonies low to slightly raised, plane; 281 YILMAZ ET AL. Fig. 54. Morphological characters of Talaromyces piceus (CBS 137377). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 282 POLYPHASIC margins narrow (1–2 mm), low, entire, plane; mycelium white and yellow; texture velvety to loosely funiculose; sporulation sparse to dense; conidia en masse grayish green to dull green (29D3–29D4); exudates big clear droplets; soluble pigment absent, in some isolates around colonies light yellow to orange; reverse in the centre greyish yellow (2C3) fading into pale yellow in the margins (2A3). CREA, 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 10–65 × 2.5–3.5 μm; terminating in vesicles up to 5.5 μm; metulae four to eight, 8–14 × 2–4 μm; phialides acerose, three to six per metulae, 7–10 × 2.5–3.5 μm; conidia smooth, ellipsoidal, ranging in size from 2–3.5 × 2–4 μm. Extrolites: Talaromyces piceus produces emodin, mitorubrin, mitorubrinol, rugulosin A and other rugulosins, and skyrin. It also produces diverse other anthraquinones and extrolites with chromophores suggesting tetracyclic compounds, citreoisocoumarin and restricticin. Distinguishing characters: Talaromyces piceus is characterised by biverticillate conidiophores having vesiculated stipes (Fig. 54). The species is able to grow at high temperatures, growing faster on CYA at 37 C than 25 C. The species can also grow at 40 C. These characters are very similar to its close relative T. columbinus (Fig. 7). Compared to T. columbinus, T. piceus grows faster on CYA and YES at 25 C, but slower on CYA at 37 C. Notes: Penicillium ilerdanum is phylogenetically identical to T. piceus and is considered a synonym. Talaromyces pinophilus (Hedgc.) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560662. Fig. 55. ≡ Penicillium pinophilum Hedgc. apud Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 37. 1910. = Penicillium korosum J.N. Rai, Wadhwani & J.P. Tewari, Antonie van Leeuwenhoek 35: 430. 1969. = Acremonium cellulolyticus nom. nud., Japanese patent, 1317660. 1985; US Patent 4562150. 1985. = Talaromyces cellulolyticus T. Fujii et al., FEMS Microbiology Letters 351: 38. 2013. In: Talaromyces section Talaromyces Typus: IMI 114933, culture ex-type CBS 631.66 = ATCC 36839 = CECT 2809 = DSM 1944 = IAM 7013 = IMI 114933. ITS barcode: JN899382 (alternative markers: BenA = JX091381; CaM = KF741964; RPB2 = KM023291) Colony diam, 7 d (mm): CYA 18–25; CYA 30 C 30–40; CYA 37 C 25–40; MEA 30–40; MEA 30 C 45–55; DG18 10–20; CYAS 3–7; OA 30–40; CREA 15–22; YES 25–40. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white, yellow and in some strains red; texture loosely funiculose and ﬂoccose especially in the centre; sporulation sparse to moderately dense, conidia en masse greyish green to dull green (26C3–26D4); soluble pigments only in DTO 60-C5 weak yellow soluble pigments at 37 C except DTO 60-F4 they all produce yellow pigments; exudates small clear and red droplets; reverse greyish orange to orange (6B6–6B7) centre fading into pastel www.studiesinmycology.org TAXONOMY OF TALAROMYCES yellow (3A4). MEA 25 C, 7 d: Colonies slightly raised, slightly sulcate; margins low, plane, entire (4–5 mm); mycelia white, yellow and red; texture loosely funiculose to ﬂoccose especially in the centre; sporulation sparse to dense, conidia en masse greyish green to dull green (27D4–29D4); soluble pigments absent; exudates in some isolates small red droplets; reverse brownish orange (5C6). YES 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (3–4 mm); mycelia white, yellow and in some isolates red; texture ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (28B5–28C4); soluble pigments absent; exudates in some isolates small clear droplets; reverse in some isolates centre dark brown to reddish golden brownish orange (6F6–6C7) fading into yellow (3A7) and in some isolates yellowish orange (4A7). DG18 25 C, 7 d: Colonies raised at centre, plane; margins low, plane, entire (2–3 mm); mycelia white only in DTO 60-C7 yellow; texture loosely funiculose to ﬂoccose, especially in the centre conidiophores born from sterile aerial hyphae; sporulation moderately dense, conidia en masse dull green (26E3–26E4); soluble pigments absent; exudates absent; reverse olive (3F3) centre fading into pastel yellow to light yellow (3A4–3A5) and greyish yellow to greyish green (1B3–1C4). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–5 mm); mycelia white and yellow; texture funiculose and ﬂoccose, especially in the centre aerial mycelia; sporulation dense, conidia en masse greyish green (28D5–28E5); soluble pigments absent; exudates in some isolates small clear droplets; reverse 60F4 bright orange, the rest pale brownish orange to pale light brownish green. CREA 25 C, 7 d: Acid production moderate (no acid production DTO 60-F4). Micromorphology: Conidiophores biverticillate; stipes smooth walled, 30–200 × 2–3 μm; metulae three to eight, divergent, 10–11 × 2.5–3 μm; phialides acerose, three to eight per metulae, 8.5–12 × 2–3 μm; conidia smooth, globose to subglobose, 2–3 × 2–3 μm. Ascomata not observed. Extrolites: Talaromyces pinophilus has been reported to produce secopenicillide C, penicillide, MC-141, pestalacin A and stromemycin (Nonaka et al. 2011), dinapinone A1 & A2 (Uchida et al. 2012), monoapinone A-E (Kawamoto et al. 2011), 3-O-methylfunicone and 3-O-methyl-5,6-epoxyfunicone (de Stefano et al. 1999a, b). We have found mitorubrin, mitorubrinic acid, vermicillin, a purpactin, penicillide, pestalacin A and Monascus red pigments in T. pinophilus. Distinguishing characters: Talaromyces pinophilus grows relatively fast on general media at 25, 30 and 37 C, produces weak acid on CREA. Colonies have white, yellow and red mycelia and ﬂoccose to loosely funiculose texture (Fig. 55). These characters resemble T. angelicus, T. aculeatus, T. apiculatus, T. verruculosus, T. amestolkiae and T. stollii. Talaromyces aculeatus, T. apiculatus and T. verruculosus produce ﬂask-shaped phialides and rough-walled, globose conidia, which differ from T. pinophilus conidia. Talaromyces angelicus differs from T. pinophilus by lack of acid production on CREA. Talaromyces amestolkiae colonies have red reverses on CYA and MEA, black sclerotia after 2 wk of incubation and grows more restrictedly at 37 C compared to T. pinophilus. Talaromyces pinophilus micro- and macromorphologically closely resembles T. stollii. However T. stollii grows slightly faster than T. pinophilus on CYA. Sequence data is, however, needed for reliable identiﬁcation of the species, 283 YILMAZ ET AL. Fig. 55. Morphological characters of Talaromyces pinophilus (DTO 183-I6). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bars: B = 50 μm; C = 10 μm, applies to D–H. 284 POLYPHASIC TAXONOMY OF TALAROMYCES since Visagie et al. (2014) introduced the phylogenetically distinct, but morphologically almost identical, T. sayulitensis. often sterile. Conidiophores have short stipes up to 60 μm long (Fig. 56). These characters distinguish it from other species. Notes: Strain Y-94, an important cellulase producer for biotechnology, was given the name Acremonium cellulolyticus in a patent application (Yamanobe et al. 1985a, b), but the species was never formally described. Fujii et al. (2013) subsequently described and introduced the name T. cellulolyticus for strain Y94. However, sequence data (ITS: A474749; RPB1: AB856422; BenA: AB773823) show that the species is a synonym of T. pinophilus. Penicillium korosum (CBS 762.68) was also found to be identical to T. pinophilus (Fig. 2) and is considered a synonym. Talaromyces primulinus (Pitt) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011. MycoBank MB560664. Fig. 57. Talaromyces pittii (Quintan.) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560663. Fig. 56. ≡ Penicillium pittii Quintan., Mycopathologia 91: 69. 1985. In: Talaromyces section Purpurei Typus: CBS 139.84, culture ex-type CBS 139.84 = IMI 327871. ITS barcode: JN899325 (alternative markers: BenA = KJ865728; CaM = KJ885275; RPB2 = KM023297) Colony diam, 7 d (mm): CYA 34–36; CYA 30 C 38–40; CYA 37 C No growth; MEA 42–44; MEA 30 C 48–50; DG18 21–23; CYAS No growth; OA 35–36; CREA 4–5; YES 33–35. Colony characters: CYA 25 C, 7 d: Colonies low, raised at centre, plane, yeast-like appearance at centre, this character is not observed at 30 C; margins low, narrow, entire (1 mm); mycelia white; texture velvety, after 2 wk of incubation ﬂoccose and synnematous, indeterminate synnemata produced centrally; sporulation moderately dense, conidia en masse greyish green (25C4); soluble pigments red; exudates clear to light red; reverse reddish brown (8E7) at centre, fading into brownish red (8C6). MEA 25 C, 7 d: Colonies brownish red to reddish brown (8D6–8E8) yeast-like slimy colonies; sporulation absent; soluble pigments red; reverse brownish red to reddish brown (8C8–8D8). YES 25 C, 7 d: Colonies yellowish grey (3B2–4B2) yeast-like slimy colonies; sporulation absent; soluble pigments absent; reverse yellowish grey (3B2–4B2). DG18 25 C, 7 d: Colonies yellowish grey (3B2) yeast-like slimy colonies; sporulation absent; soluble pigments absent; reverse pale (3A2). OA 25 C, 7 d: Colonies low, plane, having a reddish colour underneath sporulating areas; margins low, wide, entire (4 mm); mycelia white; texture velvety; sporulation moderately dense, conidia en masse greyish green (25C4); soluble pigments absent; exudates absent; reverse reddish brown (9E7). CREA 25 C, 7 d: Acid production absent. Micromorphology: Synnemata formed after 2–3 wk on MEA up to 1 000 μm long. Conidiophore biverticillate and monoverticillate; stipes smooth walled, 10–60 × 1.8–2.5 μm; metulae two to ﬁve, divergent, 9–15 × 1.8–3 μm; phialides acerose, three to ﬁve per metulae, 11–15 × 2–3 μm; conidia smooth, ellipsoidal, 2.5–4.5(–6.5) × 1.5–3.5 μm. Ascomata not observed. Distinguishing characters: Talaromyces pittii grows fast on general media and colonies have a slimy texture and produce a typical red reverse. Synnemata are produced after 2–3 wk of incubation and have a greyish colour, are narrowly conical and www.studiesinmycology.org ≡ Penicillium diversum var. aureum Raper & Fennell, Mycologia 40: 541. 1948 (nom. inval., Art. 39, no latin diagnosis). ≡ Penicillium primulinum Pitt, The genus Penicillium: 455. 1980. In: Talaromyces section Talaromyces Typus: IMI 040031, culture ex-type CBS 321.48 = ATCC 10438 = CBS 439.88 = FRR 1074 = IMI 040031 = MUCL 31321 = MUCL 31330 = NRRL 1074. ITS barcode: JN899317 (alternative markers: BenA = JX494305; CaM = KF741954; RPB2 = KM023294) Colony diam, 7 d (mm): CYA 5–6; CYA 30 C 5–6; CYA 37 C No growth; MEA 20–25; MEA 30 C 13–16; DG18 6–8; CYAS No growth; OA 22–25; CREA 2–3; YES 8–10. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose and velvety; sporulation sparse to dense, conidia en masse greyish green (27E5–28E5) (on DTO 270-F3); soluble pigments absent; exudates absent; reverse in some isolates brownish orange to reddish brown (7C5–8C5) (DTO 250-F2) and in some isolates greenish grey to greyish green (1D2–1D3) (DTO 250-F3). MEA 25 C, 7 d: Colonies slightly raised at centre, sunken at centre, sulcate, degraded, white and dull red appearance; margins low, plane, entire (1–2 mm); mycelia white and dull red; texture ﬂoccose and after 2 wk of incubation colonies covered with yellow mycelia; sporulation absent; soluble pigments absent; exudates absent; reverse violet brown (10E7) centre fading into greyish orange (5B5). YES 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose (DTO 250F2) and velvety (DTO 250-F3); sporulation moderately dense to dense, conidia en masse dull green (26D3) (DTO 250-F2) and greyish green (25E5–26E5) (DTO 250-F3); soluble pigments absent; exudates absent; reverse olive (2F5) (DTO 250-F3) and light orange to greyish orange (5A5–5B5) (DTO 250-F2). DG18 25 C, 7 d: Colonies raised at centre, plane; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation absent to sparse, conidia en masse greyish green (28B5) (DTO 250-F3); soluble pigments absent; exudates absent; reverse greyish green (1C5) centre fading into greyish yellow (1B5) (DTO 250-F3) and in some isolates yellowish white (4A2) (DTO 250F2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white and in some isolates pastel yellow as well (DTO 250-F3); texture velvety; sporulation moderately dense, conidia en masse greyish green (25C4 for DTO 250-F2 and for DTO 250-F3 26E5–27E5); soluble pigments absent; exudates absent; reverse dull reddish brown (DTO 250-F2) and olive green (DTO 250-F3). CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 70–200 × 1.5–3 μm, slightly vesiculate; branches 15 μm; metulae three to eight, divergent, 6–12 × 2–3.5; phialides 285 YILMAZ ET AL. Fig. 56. Morphological characters of Talaromyces pittii (CBS 139.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 3 wk incubation. C–G. Conidiophores. H. Conidia. Scale bars: B = 1000 μm; C = 10 μm, applies to D–H. 286 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 57. Morphological characters of Talaromyces primulinus (CBS 321.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 287 YILMAZ ET AL. acerose, three to eight per metulae, 7.5–12 × 1.5–2.5 μm; conidia smooth to ﬁnely roughed, ellipsoidal to fusiform, 2–4 × 1.5–3 μm. Extrolites: According to the literature T. primulinus can produce sclerone, isoclserone, juglone and herqueinone (Fujimoto et al. 1986). We detected herqueinone, mitorubrin, mitorubrinic acid in NRRL 1704 = IBT 3805. Distinguishing characters: Talaromyces primulinus grows restrictedly on all media and does not grow at 37 C. After 2 wk of incubation, yellow mycelia cover the colony. This is also observed in T. ﬂavovirens and T. marneffei. However, T. marneffei grows at 37 C as a yeast form and T. ﬂavovirens grows faster on all media. The restricted growth of T. primulinus on CYA resembles T. rademirici, T. diversus and T. erythromellis. Talaromyces primulinus differs from T. erythromellis by its biverticillate conidiophores. Talaromyces diversus grows faster on MEA, while T. rademirici grows slower on MEA. Also, T. primulinus has slightly vesiculated stipes. Both T. primulinus strains were nutritionally deﬁcient (Pitt 1980). Talaromyces proteolyticus (Kamyschko) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011. MycoBank MB560665. Fig. 58. ≡ Penicillium proteolyticum Kamyschko, Nov. Sist. niz. Rast. Sist. niz. Rast. 14: 227. 1961. In: Talaromyces section Bacillispori mycelia white; texture velvety; sporulation moderately dense to dense, conidia en masse dull green (26D3–27D3); soluble pigments absent; exudates small clear droplets; reverse pale and light green). CREA 25 C, 7 d: Acid production moderate. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 150–250 × 2–3 μm; branches 13–26 μm; metulae three to eight, divergent, 9–15 × 2–3 μm; phialides acerose, three to eight per metulae, 8–11 × 1.8–2.6 μm; conidia smooth, globose to subglobose, 2–3 × 1.5–2.5 μm. Ascomata not observed. Extrolites: We detected monorden and compounds with the same chromophore as emodin, herqueinone and neosartorin in T. proteolyticus NRRL 3378 = IBT 4481 = IBT 14325. Chemically T. proteolyticus is very different from T. verruculosus. Distinguishing characters: Talaromyces proteolyticus produces deep, yellow colonies and grows relatively fast on general media, except on DG18 and on CYA at 37 C. It produces smooth, globose to subglobose conidia (Fig. 58). Pitt (1980) considered T. proteolyticus synonymous with T. verruculosus. However, T. verruculosus produces globose to subglobose, rough-walled conidia, ﬂask-shaped phialides and grows at 37 C. Talaromyces pseudostromaticus (Hodges, G.M. Warner & Rogerson) Samson et al., Stud. Mycol. 71: 176. 2011. MycoBank MB560666. Fig. 59. ≡ Penicillium pseudostromaticum Hodges, G.M. Warner & Rogerson, Mycologia 62: 1106. 1970. Typus: CBS 303.67, culture ex-type CBS 303.67 = ATCC 18326 = NRRL 3378. In: Talaromyces section Purpurei ITS barcode: JN899387 (alternative markers: BenA = KJ865729; CaM = KJ885276; RPB2 = KM023301) Typus: Warner 18 (NY), culture ex-type CBS 470.70 = ATCC 18919 = FRR 2039. Colony diam, 7 d (mm): CYA 20–22; CYA 30 C 19–20; CYA 37 C No growth; MEA 20–21; MEA 30 C 20; DG18 10; CYAS 9–10; OA 20–22; CREA 15; YES 20–21. ITS barcode: JN899371 (alternative markers: BenA = HQ156950; CaM = KJ885277; RPB2 = KM023298) C, Colony characters: CYA 25 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white, pastel yellow and pastel orange; texture ﬂoccose; sporulation sparse, conidia en masse pastel green (25B4–26B4); soluble pigments absent; exudates small clear droplets; reverse centre greyish red (9B5) fading into between light yellow (4A4) and light orange (5A4). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation sparse, conidia en masse pastel green (25B4–26B4); soluble pigments absent; exudates absent; reverse greyish orange to brownish orange (5B6–5C6). YES 25 C, 7 d: Colonies raised at centre, slightly sulcate, ﬂuffy appearance; margins low, plane, entire (1 mm); mycelia white and pale pastel yellow; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A5) with pastel red (8A5) circle close to margins. DG18 25 C, 7 d: Colonies raised at centre, pale, ﬂuffy white appearance with sterile aerial hyphae; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (3A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); 288 Colony diam, 7 d (mm): CYA 25–34; CYA 30 C 25–38; CYA 37 C No growth; MEA 38–43; MEA 30 C 30–40; DG18 19–21; CYAS No growth; OA 35–37; CREA 4–6; YES 27–32. Colony characters: CYA 25 C, 7 d: Colonies low, slightly raised at centre, plane, red colour underneath sporulating areas; margins low, narrow, entire (2–3 mm); mycelia white, some yellow present; texture velvety; sporulation moderately dense to dense, conidia en masse dull green (26E4); soluble pigments absent; exudates absent; reverse dark brown (7F8–8F8). MEA 25 C, 7 d: Colonies low, slightly raised at centre, plane, red colour underneath sporulating areas; margins low, narrow, entire (2 mm); mycelia white, some yellow present; texture velvety, funicles present, after 2 wk of incubation synnemata produced; sporulation moderately dense, conidia en masse greyish green (25E5–25E6); soluble pigments absent; exudates absent; reverse dark brown (7F8–8F8). YES 25 C, 7 d: Colonies low, slightly raised at centre, red colour underneath sporulating areas; margins low, narrow, entire (2–3 mm); mycelia white, yellow and pink; texture velvety, some loose funicles present; sporulation moderately dense, conidia en masse greyish green (25E5); soluble pigments red; exudates absent; reverse very dark red at centre, dark brown to violet brown (8F8–10F8) elsewhere. DG18 25 C, 7 d: POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 58. Morphological characters of Talaromyces proteolyticus (CBS 303.67T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 289 YILMAZ ET AL. Fig. 59. Morphological characters of Talaromyces pseudostromaticus (CBS 470.70T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 2 wk incubation. C. Synnemata. D–F. Conidiophores. G. Conidia. Scale bars: B, C = 2000 μm; D = 10 μm, applies to E–G. 290 POLYPHASIC Colonies low, slightly raised at centre, plane, having a reddish colour in non sporulating areas; margins low, wide, entire (4–5 mm); mycelia white; texture velvety; sporulation sparse, conidia en masse greyish green (25D5); soluble pigments inconspicuously yellow; exudates pink; reverse reddish brown (8D7) at centre, fading into greyish red (8B6). OA 25 C, 7 d: Colonies low, plane, having a reddish colour underneath sporulating areas; margins low, wide, entire (4 mm); mycelia white; texture velvety; sporulation moderately dense, conidia en masse dull green (26E4); soluble pigments absent; exudates absent; reverse greyish red (8C5). CREA 25 C, 7 d: Acid production absent. Micromorphology: Synnemata phototropic and formed after 2–3 wk on MEA up to 8 000 μm long. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 30–180 × 2.5–4.5 μm; branches 16–50 μm; metulae two to six, divergent, 7.5–17 × 2.5–4.5 μm; phialides acerose, three to six per metulae, 10–16 × 2–3.5 μm; conidia smooth, subglobose to ellipsoidal, 2.5–4 × 2–3 μm. Ascomata not observed. TAXONOMY OF TALAROMYCES and the rest brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies low, slightly radially sulcate; margins low, narrow, entire (1 mm); mycelia white; texture loosely funiculose and especially in the centre conidiophores born from sterile aerial hyphae; sporulation sparse to moderately dense, conidia en masse dull green (26E4); soluble pigments absent; exudates small, clear droplets; reverse centre olive (2F5) fading into pastel yellow (2A4). DG18 25 C, 7 d: Colonies slightly raised in the centre, plane; margins low, narrow, entire; mycelia white; sporulation absent to sparse (difﬁcult to determine); soluble pigments absent; exudates clear and yellow droplets; reverse light yellow (4A5). OA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (3–6 mm); mycelia yellow and white; texture velvety and loosely ﬂoccose especially in the centre conidiophores born from aerial mycelia; sporulation moderately dense to dense, conidia en masse dull green to dark (26E6–26F6); soluble pigments absent; exudates small clear droplets; reverse light pale brown. CREA 25 C, 7 d: No growth. Extrolites: Talaromyces pseudostromaticus produces mitorubrinic acid and secalonic acid D (Samson et al. 1989). It also produces a Monascus red pigment (azaphilone). Micromorphology: Conidiophores biverticillate; stipes smooth walled, green pigmented, 40–95 × 2.5–3.5 μm; metulae three to six, divergent, 10–12.5 × 2–2.5 μm; phialides acerose, three to six per metulae, 10–12 × 2–2.5 μm; conidia rough with spiral ridges, ellipsoidal to apiculate, 3–4.5(–5) × 2–3 μm. Ascomata not observed. Distinguishing characters: Talaromyces pseudostromaticus grows relatively fast on general media, but does not grow on CYA at 37 C. It produces reddish brown to dark brown reverses on general media (Fig. 59). After 2–3 wk of incubation, it produces determinate, phototropic synnemata up to 8 000 μm long. These characters easily distinguish T. pseudostromaticus from other synnemata producers. Distinguishing characters: Talaromyces ptychoconidium grows restrictedly on CYA and MEA. It produces slimy, clear exudates and synnema like structures after 2 wk of incubation (Fig. 60). It characteristically produces spirally roughened conidia and thus resembles T. purpureus. However, T. purpureus grows restrictedly, has vesiculated stipes and produces red soluble pigments on MEA and OA. Talaromyces ptychoconidium Visagie & K. Jacobs, Persoonia 28: 18. 2012. MycoBank MB564327. Fig. 60. Talaromyces purpureus (E. Müll. & Pacha-Aue) Stolk & Samson, Stud. Mycol. 2: 57. 1972. MycoBank MB324420. Figs 61, 62. In: Talaromyces section Purpurei Typus: PREM 60041, culture ex-type DAOM 241017 = CV 2808 = DTO 180-E7. ITS barcode: FJ160266 (alternative markers: BenA = GU385733; CaM = JX140701; RPB2 = KM023278) Colony diam, 7 d (mm): CYA 8–16; CYA 30 C 15–20; CYA 37 C 5–12; MEA 12–23; MEA 30 C 30–35; DG18 6–12; CYAS No growth; OA 18–30; CREA No growth; YES 8–16. Colony characters: CYA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white; texture loosely funiculose, abundant aerial hyphae produced at colony centre; sporulation sparse, conidia en masse greyish green (30D4–30E4); soluble pigments absent; exudates very small clear droplets; reverse centre light brown (5D4) fading into light yellow (3A5). MEA 25 C, 7 d: Colonies low, plane; margins low, narrow, entire (1–2 mm); mycelia white; texture loosely funiculose and especially in the centre conidiophores born from sterile aerial hyphae; sporulation sparse to moderately dense, conidia en masse dull green (26E4); soluble pigments absent; exudates slimy, clear and yellow droplets; reverse centre raw umber (5F8) www.studiesinmycology.org ≡ Arachniotus purpureus E. Müll. & Pacha-Aue, Nova Hedwigia 15: 552. 1968. ≡ Penicillium purpureum Stolk & Samson, Stud. Mycol. 2: 57. 1972. In: Talaromyces section Purpurei Typus: CBS H-7832, culture ex-type CBS 475.71 = ATCC 24069 = ATCC 52513 = FRR 1731 = IMI 181546. ITS barcode: JN899328 (alternative markers: BenA = GU385739; CaM = KJ885292; RPB2 = JN121522) Colony diam, 7 d (mm): CYA 2–4; CYA 30 C No growth (sometimes up to 3); CYA 37 C No growth; MEA 15–16; MEA 30 C 17–18; DG18 1–2; CYAS No growth; OA 18–20; CREA No growth; YES 7–8. Colony characters: CYA 25 C, 7 d: Colonies germinating but sterile. MEA 25 C, 7 d: Colonies low, plane; margins low, narrow, entire (2 mm); mycelia white in the margins, red in the centre; colony is red appearance; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse reddish brown to dark brown (8F8–9F8). YES 25 C, 7 d: Colonies germinating but sterile. DG18 25 C, 7 d: Colonies germinating but sterile. OA 25 C, 7 d: Colonies low, plane; 291 YILMAZ ET AL. Fig. 60. Morphological characters of Talaromyces ptychoconidium (DAOM 241017T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B, C. Colony texture on MEA after 2 wk incubation. D–G. Conidiophores. H. Conidia. Scale bars: B, C = 500 μm; D = 100 μm; E = 10 μm, applies to F–H. 292 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 61. Morphological characters of Talaromyces purpureus (CBS 475.71T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on MEA after 2 wk incubation. C. Colony texture on OA after 2 wk incubation. D. Initials. E–J. Conidiophores. K. Conidia. Scale bars: B = 500 μm; C = 100 μm; D = 10 μm, applies to E–K. www.studiesinmycology.org 293 YILMAZ ET AL. Fig. 62. Micromorphological characters of Talaromyces purpureus (CBS 475.71T) from Stolk & Samson (1972). A. Conidiogenous structures. B. Conidia. C. Initials surrounded by thin hyphae. D. Initials. E. Asci produced in chains. F. Ascospores. G. Loosely branched hyphae covering ascoma. Scale bar 10 μm. margins low, wide, entire (1–2 mm); mycelia red; sporulation absent; soluble pigments absent; exudates absent; reverse dark cherry red. CREA 25 C, 7 d: No growth. roughened conidia (Figs 61, 62). Talaromyces ptychoconidium also produces spirally roughened conidia, but produces biverticillate conidiophores. Micromorphology: Conidiophores with solitary phialides or monoverticillate; stipes smooth walled, sometimes vesiculate up to 4 μm, 7–22 × 2–2.5 μm; phialides ﬂask-shaped and acerose, one to four, 4.5–8(–16) × 2–3 μm; conidia rough with spiral ridges, thick walled, subglobose to broadly ellipsoidal, 3–4 × 2–3 μm. Ascomata not observed in our culture however ﬁde Stolk & Samson (1972) maturing after 3 wk of incubation on MEA at 30 C, yellow, globose to subglobose 20–60 μm, asci 9–12 μm, ascospores ellipsoidal, thick walled, spiny over their entire surface, 6.5–8 × 4.5–5.5 μm. Talaromyces purpurogenus (Stoll) Samson et al., Stud. Mycol. 71: 177. 2011. MycoBank MB560667. Fig. 63. ≡ Penicillium purpurogenum Stoll, Beitr. Morph. Biol. Char. Penicill.: 32. 1904. = Penicillium sanguineum Sopp, Skr. Vidensk.-Selsk. Christiana, Math.Naturvidensk. Kl. 11: 175. 1912. = Penicillium crateriforme J.C. Gilman & E.V. Abbott, Iowa State Coll. J. Sc. 1: 293. 1927. = Penicillium vanilliae Bouriquet, Bull. Acad. Malgache 24: 68. 1941 (nom. inval., Art. 36). In: Talaromyces section Talaromyces Distinguishing characters: Talaromyces purpureus grows very restrictedly on general media and produces red soluble pigments on MEA and OA. It produces yellow ascomata with big, thick walled, spiny, ellipsoidal (6.5–8 × 4.5–5.5 μm) ascospores. Its conidiophores often have solitary phialides, but are mostly monoverticillate and have vesiculated stipes and spirally 294 Typus: IMI 091926, culture ex-type CBS 286.36 = IMI 091926. ITS barcode: JN899372 (alternative markers: BenA = JX315639; CaM = KF741947; RPB2 = JX315709) POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 63. Morphological characters of Talaromyces purpurogenus (CBS 132707). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 295 YILMAZ ET AL. Colony diam, 7 d (mm): CYA 20–25; CYA 30 C 18–27; CYA 37 C 16–25; MEA 30–45; MEA 30 C 50–52; DG18 10–15; CYAS 5; OA 28–35; CREA 7–15; YES 25–40. Colony characters: CYA 25 C, 7 d: Colonies moderately deep, radially sulcate; margins low, plane, entire (1 mm); mycelia white and pastel red; texture ﬂoccose; sporulation sparse to moderately dense, conidia en masse dull green (26D3–26D4); soluble pigments bright red (also at 30 C); exudates absent; reverse dark brown to violet brown (9F8–11F8). MEA 25 C, 7 d: Colonies raised at point of inoculation, plane; margins low, plane, entire (3–4 mm); mycelia white and light orange; texture velvety and ﬂoccose, especially in the centre; sporulation moderately dense to dense, in some isolates absent, conidia en masse dull green (26E4–26E5); soluble pigments absent; exudates absent; reverse brownish yellow to brownish orange (5C7–6C7). YES 25 C, 7 d: Colonies low, sulcate; margins low, plane, entire (1–2 mm); mycelia white and light orange; texture ﬂoccose; sporulation moderately dense to dense, conidia en masse dull green to greyish green (26E4–26E5); soluble pigments absent; exudates absent; reverse centre brown (6D7) fading into light yellow (4A5), in some isolates dark red to dark brown (8F4). DG18 25 C, 7 d: Colonies raised at the point of inoculation, slightly sulcate, fruity odour; margins low, plane, entire (1 mm); mycelia white and bright orange; texture velvety and in some isolates ﬂoccose; sporulation sparse to dense, conidia en masse dark green (27F5); soluble pigments absent; exudates absent; reverse light to brownish orange (5A4–5C4). OA 25 C, 7 d: Colonies low, plane, fruity odour; margins low, plane, entire (2–3 mm); mycelia white and light orange; texture velvety and in some isolates ﬂoccose; sporulation sparse to dense, conidia en masse dull green to dark green (26E4–27F5); soluble pigments absent; exudates absent; reverse dull red and in some isolates colour lacking. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 70–200 × 1.5–3 μm; metulae three to ﬁve, divergent, 12–15 × 2.5–4; phialides acerose, three to six per metulae, 12–14 × 3 μm; conidia smooth, 3–3.5 × 2–2.5 μm. Ascomata not observed. Extrolites: Talaromyces purpurogenus produces rugulovasine A & B, rubratoxin A & B, spiculisporic acid, mitorubrin, mitorubrinic acid, mitorubrinol, purpactin A-C, Monascus red azaphilone pigments, including N-glutarylrubropunctamine and luteoskyrin (Yilmaz et al. 2012). Distinguishing characters: Talaromyces purpurogenus colonies produce red soluble pigments on CYA at 25 and 30 C and have orange mycelia on DG18 (Fig. 63). Talaromyces purpurogenus resembles T. atroroseus, but T. atroroseus has ﬁnely rough to rough, dull to dark green, thick walled, ellipsoidal conidia and produces darker conidia especially on OA. Notes: Yilmaz et al. (2012) synonymised Penicillium crateriforme, P. sanguineum and P. vanilliae with T. purpurogenus and this is followed here. Talaromyces rademirici (Quintan.) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. MycoBank MB560668. Fig. 64. ≡ Penicillium rademirici Quintan., Mycopathologia 91: 69. 1985. In: Talaromyces section Purpurei Typus: CBS 140.84, culture ex-type CBS 140.84 = CECT 2771 = IMI 282406 = IMI 327870. ITS barcode: JN899386 (alternative markers: BenA = KJ865734; RPB2 = KM023302) Colony diam, 7 d (mm): CYA 5–6; CYA 30 C 5–7; CYA 37 C 3; MEA 14–15; MEA 30 C 14–16; DG18 4–5; CYAS No growth; OA 9–10; CREA No growth; YES 5–6. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2). MEA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate, white appearance; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish orange (5C6) centre fading into greyish orange (5B6). YES 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pale yellow (4A3). DG18 25 C, 7 d: Colonies low, plane, slimy yeast like colony appearance; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse white (4A1). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white; texture velvety; sporulation moderately dense, conidia en masse greyish green (27C5–27D5); soluble pigments absent; exudates absent; reverse dull beige. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate and monoverticillate with a minor proportion having subterminal branches; stipes smooth walled, 25–95 × 1.5–2.5 μm; branches 10–15 μm; metulae two to ﬁve, divergent, 7–11 × 2–2.5 μm; phialides acerose, two to six per metulae, 7.5–11.5 × 1.5–3 μm; conidia smooth, ellipsoidal, 2.5–4 × 1.5–2.5 μm. Ascomata not observed. Distinguishing characters: Talaromyces rademirici grows restrictedly on all media and does not grow on CREA (Fig. 64). Its restricted growth on CYA resembles T. erythromellis, T. diversus and T. primulinus. Talaromyces diversus grows much faster on MEA. Talaromyces erythromellis produces unusual biverticillate conidiophores with subterminal branches, whereas T. rademirici produces mono- to biverticillate conidiophores. Talaromyces primulinus grows slightly faster and has slightly vesiculated stipes. The ex-type strain of T. rademirici is nutritionally deﬁcient. Talaromyces radicus (A.D. Hocking & Whitelaw) Samson et al., Stud. Mycol. 71: 177. 2011. MycoBank MB560669. Fig. 65. ≡ Penicillium radicum A.D. Hocking & Whitelaw, Mycol. Res. 102: 802. 1998. In: Talaromyces section Islandici 296 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 64. Morphological characters of Talaromyces rademirici (CBS 140.84T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 297 YILMAZ ET AL. Fig. 65. Morphological characters of Talaromyces radicus (CBS 100489T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–G. Conidiophores. H. Conidia. Scale bar: B = 10 μm, applies to C–H. 298 POLYPHASIC Typus: DAR 72374, culture ex-type CBS 100489 = FRR 4718 = IBT 14379. ITS barcode: KF984878 (alternative markers: BenA = KF984599; CaM = KF984773; RPB2 = KF985013) Colony diam, 7 d (mm): CYA 15–22; CYA 30 C 22–27; CYA 37 C 25–30; MEA 15–25; MEA 30 C 24–27; DG18 10–15; CYAS 14–20; OA 15–22; CREA 4–8; YES 22–25. Colony morphology: CYA 25 C, 7 d: Colonies low, slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white and yellow; texture velvety to loosely funiculose; sporulation absent to sparse; conidia en masse greyish green (30C7–30E7); exudates clear and yellow droplets; soluble pigment absent; reverse centre yellowish brown (5E6–5F6) in the centre fading into light brown (5D6) and pale yellow (4A3). MEA 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate; margins narrow (1 mm), low, entire, plane; mycelium white and yellow; texture loosely funiculose to ﬂoccose; sporulation absent to sparse; conidia en masse greyish green (29C5–29E5) exudates in some colonies clear small droplets; soluble pigment absent; reverse yellowish brown (5F5) centre. YES 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (1 mm), low, entire, plane; mycelium white, yellow and pale orange; texture ﬂoccose; sporulation absent to sparse; conidia en masse dull green (26D4–26E4); exudates absent; soluble pigment absent; reverse light brown to olive brown (4D6–6D6) centre fading into reddish yellow to orange (4A6–6A6). DG18 25 C, 7 d: Colonies raised at centre, plane; margins narrow (1 mm), low, entire, plane; mycelium white; texture ﬂoccose; sporulation absent to sparse; conidia en masse greyish green (29E5); exudates clear and yellow droplets; soluble pigment absent; reverse golden yellow (5B7) fading into olive brown (4D6). OA 25 C, 7 d: Colonies raised at centre, plane; margins narrow (1–2 mm), low, entire, plane, in some isolates yeast like slimy margins; mycelium white and yellow; texture loosely funiculose; sporulation moderately dense to dense; conidia en masse greyish green (30C6–30D6) and in some isolates dull green (28D4–28E4); exudates clear small droplets in the centre; soluble pigment absent; reverse brownish orange centre fading into yellow. CREA, 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches up to 15 μm; stipes smooth walled, 80–270 × 2–2.5 μm; metulae three to six, divergent, 9–12 × 2–3 μm; phialides acerose, three to six per metulae, 8–11 × 2–2.5 μm; conidia ﬁnely rough-walled in ridges, globose to ellipsoidal, 2–3 × 2–2.5 μm. Extrolites: Talaromyces radicus produces xanthoradone A, B & C (Yamazaki et al. 2009a, b, 2010a), which has only found in this species, and we also detected emodin, skyrin and rugulosin A and a series of azaphilones, including 60 -hydroxy-30 -methoxymitorubrin, 40 -hydroxy-30 -methoxy-(S)-mitorubrin and monomethyl-(S)-mitorubrin in agreement with Yamazaki et al. (2010b). Furthermore CBS 137382 produced traces of one of the ukulactones. We detected two anti-MRSA antibiotics rugulosin A and C (Yamazaki et al. 2010c) and the latter authors also isolated rugulosin B. 60 -Hydroxy-30 -methoxy-mitorubrin is a potentiator of miconazol activity (Yamazaki et al. 2010a). www.studiesinmycology.org TAXONOMY OF TALAROMYCES Distinguishing characters: Talaromyces radicus is characterised by yellow ﬂuffy colonies on CYA and MEA (Fig. 65). Colonies are deep and consist of bright yellow mycelia that form a ﬂuffy texture, with the production of funicles on MEA (Hocking et al. 1998). It also has the ability to grow at 37 and 40 C. Phylogenetically it is closely related to T. allahabadensis. However, T. radicus does not produce acid on CREA and weakly sporulates on general media, as well as growing faster at 37 C, compared to T. allahabadensis. Talaromyces ramulosus (Visagie & K. Jacobs) Samson et al., Stud. Mycol. 71: 177. 2011. MycoBank MB560670. Fig. 66. ≡ Penicillium ramulosum Visagie & K. Jacobs, Mycologia 101: 890. 2009. ITS barcode: EU795706 (alternative markers: BenA = FJ753290; CaM = JX140711; RPB2 = KM023281) Typus: PREM 59947, culture ex-type DAOM 241660 = CV 2837 = DTO 184-B8. In: Talaromyces section Purpurei Colony diam, 7 d (mm): CYA 32–40; CYA 30 C 39–50; CYA 37 C 5–8; MEA 45–48; MEA 30 C 48–52; DG18 23–25; CYAS 15–18; OA 40–45; CREA 10–14; YES 40–43. Colony characters: CYA 25 C, 7 d: Colonies low, slightly raised at centre, radially sulcate; margins low, plane, entire (3–5 mm); mycelia white and in some isolates orange to red; texture funiculose and ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (25D5–25E5); soluble pigments absent; exudates absent; reverse centre dark brown (6F4–6F7), in some isolates fading into greyish orange (6B6). MEA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (5–6 mm); mycelia white; texture funiculose (especially in the centre sterile aerial hyphae), formation of synnemata after 2 wk of incubation on scraped agar; sporulation dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates very small clear droplets; reverse centre dark brown to reddish brown (7F7–8F7), in some isolates fading into orange red (8B7). YES 25 C, 7 d: Colonies slightly raised at centre, radially sulcate; margins low, narrow, entire (3 mm); mycelia white and in some isolates orange red centre; texture ﬂoccose and funiculose; sporulation sparse to moderately dense, conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates absent; reverse centre dark brown (6F8–7F8), fading into greyish yellow to light blonde (4B3–4C3), in some isolates golden yellow (5B7). DG18 25 C, 7 d: Colonies slightly raised in the centre, plane; margins low, narrow, entire (3–5 mm); mycelia white; texture funiculose; sporulation moderately dense to dense (no sporulation at centre, centre is slimy yeast like structure), conidia en masse greyish green to dark green (26E5–26F5); soluble pigments absent; exudates clear droplets at centre; reverse greyish green to olive (1C3–1E4), in some isolates (DTO 181-F6) greyish orange (5B6). OA 25 C, 7 d: Colonies low, plane; margins low, wide, entire (5–6 mm); mycelia white, especially in the centre synnemata like aerial hyphae growing, and there is an exudate in the centre makes a structure like synnemata; texture velvety and funiculose; sporulation dense, conidia en masse greyish green to dark green (26E5–26F5); 299 YILMAZ ET AL. Fig. 66. Morphological characters of Talaromyces ramulosus (DAOM 241660T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 3 wk incubation. C. Synnemata. D–G. Conidiophores. H. Conidia. Scale bars: B, C = 1000 μm; D = 10 μm, applies to E–H. 300 POLYPHASIC soluble pigments absent; exudates clear droplets in centre growing to air; reverse pale light green, DTO 183-A7 light red centre. CREA 25 C, 7 d: Acid production absent. Micromorphology: Synnemata formed after 2 wk on MEA up to 50–250 μm long. Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 10–60 × 2.5–3.5 μm; branches 10–20 μm; metulae three to six, divergent, 8–11 × 2–3 μm; phialides acerose, three to six per metulae, 8.5–11 × 2–2.5 μm; conidia smooth, subglobose to ellipsoidal, 2–3 × 1.5–2.5 μm. Ascomata not observed. Distinguishing characters: This species produces fast growing colonies on most media, with synnemata up to 500 μm long, produced after 2 wk incubation on MEA, similar to T. cecidicola and T. chloroloma. However, the latter two species produce longer synnemata. Talaromyces ramulosus and T. chloroloma grows faster on MEA than T. cecidicola, while T. chloroloma produces acid on CREA, which is absent in both T. ramulosus and T. choloroloma. Macromorphologically T. ramulosus resembles T. pinophilus, T. amestolkiae and T. stollii. However shorter stipes (up to 60 μm) and extra branches on the conidiophores can easily differentiate T. ramulosus from these species. Talaromyces rotundus (Raper & Fennell) C.R. Benj., Mycologia 47: 683. 1955. MycoBank MB306719. Fig. 67. ≡ Penicillium rotundum Raper & Fennell, Mycologia 40: 518. 1948 ≡ Penicillium sphaerum Pitt, Gen. Penicil.: 494. 1980. In: Talaromyces section Islandici Typus: IMI 040589, culture ex-typus CBS 369.48 = IMI 040589 = NRRL 2107 = FRR 2107 = ATCC 10493 = IBT 4829. ITS barcode: JN899353 (alternative markers: BenA = KJ865730; CaM = KJ885278; RPB2 = KM023275) Colony diam, 7 d (mm): CYA 9–11; CYA 30 C 12–15; CYA 37 C No growth; MEA 15–17; MEA 30 C 18–19; DG18 14–15; CYAS 2–3; OA 20; CREA No growth; YES 9–10. Colony characters: CYA 25 C, 7 d: Colonies slightly raised, sulcate; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse greyish green (27E5) circle at centre fading into greenish grey (1B2). MEA 25 C, 7 d: Colonies slightly raised, slightly sulcate; margins low, plane, entire (<1 mm); mycelia white and yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies slightly raised at centre, sunked at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse light yellow (4A4). DG18 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; texture yeast like slimy hyphae; sporulation absent; soluble pigments absent; exudate absent; reverse yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white and orange yellow; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange centre fading into pale light yellowish beige. CREA 25 C, 7 d: No growth. www.studiesinmycology.org TAXONOMY OF TALAROMYCES Micromorphology: Conidiophores irregular monoverticillate and biverticillate; stipes smooth walled, 20–80 × 1–2 μm; metulae one to three, divergent, 6–12 × 1–2 μm; phialides acerose, two to ﬁve, 5–17 × 1–2 μm; conidia smooth, ellipsoidal to fusiform, 3–5(–6.5) × 1.5–2.5 μm. Ascomata yellow to orange, globose to subglobose, ripening within 2–3 wk on OA at 25 C, 100–300 μm, covering consisting of a few layers of well-developed networks of white and light yellow hyphae; asci broadly globose to ovoidal 11–17 × 9.5–13 μm; ascospores globose, spinose, 4–5.5 × 4–5.5 μm. Extrolites: Talaromyces rotundus produces emodin, rugulosin and an extrolite with the same UV spectrum as talaroxanthone (Koolen et al. 2013). Distinguishing characters: Talaromyces rotundus is characterised by restricted growth on general media. Colonies lack sporulation after 7 d of incubation at 25 C and form irregular monoverticillate and biverticillate conidiophores (Fig. 67). Based on an ITS and BenA phylogeny, T. rotundus is consistently resolved in a separate clade, closely related to T. tratensis and T. wortmannii (Fig. 7). Talaromyces rotundus differs from the rest of the ascoma producers in this clade by the production of globose spinose ascospores. It resembles T. bacillisporus because of the ascospore shape and ornamentation, but T. bacillisporus produces cylindrical and rod-shaped conidia and has a dark green colony reverses on CYA at 30 and 37 C. Talaromyces ruber (Stoll) Yilmaz et al., Persoonia 29: 48. 2012. MycoBank MB801360. Fig. 68. ≡ Penicillium rubrum Stoll, Beitr. Morph. Biol. Char. Penicill.: 35. 1904. In: Talaromyces section Talaromyces Typus: CBS H-21052, culture ex-type CBS 132704 = DTO 193H6 = IBT 10703 = CBS 113137. ITS barcode: JX315662 (alternative markers: BenA = JX315629; CaM = KF741938; RPB2 = JX315700) Colony diam, 7 d (mm): CYA 20–35; CYA 30 C 25–35; CYA 37 C 14–18; MEA 35–40; MEA 30 C 45–48; DG18 14–18; CYAS 4–5; OA 40–45; CREA 8–15; YES 20–35. Colony characters: CYA 25 C, 7 d: Colonies sunken at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white, pastel yellow and pastel red; texture velvety; sporulation moderately dense to dense, conidia en masse olive green to greyish green (26D4–27D4); soluble pigments weak red (also at 30 C), in some isolates lack of pigments; exudates absent; reverse brownish red (8E8–8F8). MEA 25 C, 7 d: Colonies raised at point of inoculation, slightly sunken at centre; margins low, plane, entire (3–5 mm); mycelia white and yellow (especially at centre); texture velvety; sporulation dense, conidia en masse greyish green (26D4–26E4); soluble pigments absent; exudates absent; reverse brownish red to dark brown centre, in some isolates lack of centre pigmentation, greyish yellow to greyish orange (4B4–5B4). YES 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white and yellow; texture velvety and ﬂoccose in some isolates; sporulation sparse to dense, conidia en masse greyish green (27C5–27E6–27E7); soluble pigments absent; exudates absent; 301 YILMAZ ET AL. Fig. 67. Morphological characters of Talaromyces rotundus (CBS 369.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Texture an ascomata on OA after 2 wk incubation. C. Asci and ascospores. D. Ascospores. E–H. Conidiophores. I. Conidia. Scale bar: C = 10 μm, applies to D–I. 302 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 68. Morphological characters of Talaromyces ruber (CBS 132704T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 303 YILMAZ ET AL. reverse greyish brown to brown (5F3–5F8) centre fading into orange to light brown (5C4–5D4). DG18 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (27D4); soluble pigments absent; exudates small clear droplets; reverse greyish green (30D6–30E6) centre fading into greenish white (30A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (4–5 mm); mycelia white and light orange, in some isolates light yellow; texture velvety and in some isolates ﬂoccose; sporulation moderately dense to dense, conidia en masse dull green (27D4); soluble pigments generally absent in some isolates orange soluble pigments around colony; exudates absent; reverse reddish orange. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 100–250 × 2.5–3 μm, branches 14–15 μm; metulae three to ﬁve, 7–12 × 2–3 μm; phialides acerose, three to six per metulae, 9–12 × 2–2.5 μm; conidia smooth, ellipsoidal, 2.5–3.5 × 1.5–2 μm. Extrolites: Talaromyces ruber produces austin, berkelic acid, mitorubrin, mitorubrinic acid, Monascus red azaphilone pigments (not diffusing into agar), pestalacin A, a purpactin and vermicillin (Yilmaz et al. 2012). Distinguishing characters: Talaromyces ruber grows rapidly on general media, produces red colony reverses and velvety texture on MEA (Fig. 68). These characters distinguish T. ruber from other species. Talaromyces rubicundus (J.H. Mill., Giddens & A.A. Foster) Samson et al., Stud. Mycol. 71: 177. 2011. MycoBank MB560671. Fig. 69. ≡ Penicillium rubicundum J.H. Mill., Giddens & A.A. Foster, Mycologia 49: 797. 1957. In: Talaromyces section Talaromyces Typus: No. 2531 (A.A. Foster), culture ex-type CBS 342.59 = ATCC 13217 = IMI 099723 = NRRL 3400. ITS barcode: JN899384 (alternative markers: BenA = JX494309; CaM = KF741956; RPB2 = KM023296) Colony diam, 7 d (mm): CYA 30–32; CYA 30 C 38–40; CYA 37 C 34–35; MEA 38–39; MEA 30 C 50–52; DG18 15; CYAS 5–6; OA 40–45; CREA 17–18; YES 30–35. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, concentrically sulcate; margins low, plane, entire (2 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish orange (5C5) centre fading into orange white (5A2). MEA 25 C, 7 d: Colonies deep, plane; margins low, plane, entire (2 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates clear droplets; reverse coffee brown (5F7) centre fading into brownish orange (6C6). YES 25 C, 7 d: Colonies low, slightly sulcate; margins low, plane, entire (2–3 mm); mycelia white, pastel red and pastel yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates 304 absent; reverse light yellow (4A4) centre fading into light yellow (4A2). DG18 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose and funiculose; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish brown (5E8). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish beige. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 10–85 × 2–3 μm; branches 12–15 μm; metulae three to six, divergent, 8–17 × 2–3 μm; phialides acerose, three to six per metulae, 7.5–15.5 × 1.5–2.5 μm; conidia smooth, subglobose to ellipsoidal, 2–3 × 1.5–2.5 μm. Ascomata not observed. Extrolites: Talaromyces rubicundus produces a compound with islandicin chromophore, mitorubrin, mitorubrinic acid and rubratoxins. Distinguishing characters: Talaromyces rubicundus grows fast on general media and colonies are deep and have ﬂuffy texture on MEA (Fig. 69). It has relatively short stipes up to 85 μm long. Pitt (1980) considered T. rubicundus, T. varians and T. aurantiacus as synonyms of T. funiculosus. However, T. funiculosus has a strongly funiculose texture and strong acid production on CREA, whereas T. rubicundus does not produce acid. Talaromyces varians has pigmented stipes and T. aurantiacus grows restrictedly on DG18 and YES, and produces cylindrical to ellipsoidal conidia. Talaromyces rugulosus (Thom) Samson et al., Stud. Mycol. 71: 177. 2011. MycoBank MB560672. Fig. 70. = Penicillium elongatum Bainier, Bull. Trimest. Soc. Mycol. Fr. 23: 17. 1907 (non Dierckx 1901). ≡ Penicillium rugulosum Thom, U.S.D.A. Bur. Animal Industr. Bull. 118: 60. 1910. = Penicillium chrysitis Biourge, Cellule 33: 252. 1923. = Penicillium tardum Thom, Penicillia: 485. 1930. = Talaromyces echinosporus (Nehira) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 175. 2011 ≡ Penicillium echinosporum Nehira, J. Ferment. Technol., Osaka 11: 861. 1933. In: Talaromyces section Islandici Typus: IMI 040041, culture ex-type CBS 371.48 = ATCC 10128 = IMI 040041 = MUCL 31201 = NRRL 1045 = IBT 4485. ITS barcode: KF984834 (alternative markers: BenA = KF984575; CaM = KF984702; RPB2 = KF984925) Colony diam, 7 d (mm): CYA 15–17; CYA 30 C (6–)10–15; CYA 37 C No growth; MEA 17–20; MEA 30 C (3–)13–15; DG18 13–15; CYAS 8–12; OA 15–16; CREA 8–10; YES 15–20. Colony characters: CYA 25 C, 7 d: Colonies in some isolates slightly raised at centre and some isolates low, plane to slightly concentrically sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium generally only white, in some isolates in the centre very light yellow (DTO 225-I6, DTO 180-A4, DTO 254-A2, DTO 179-I3, DTO 278-E9) and in some pale light orange (14A2); POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 69. Morphological characters of Talaromyces rubicundus (CBS 342.59T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 305 YILMAZ ET AL. Fig. 70. Morphological characters of Talaromyces rugulosus (DTO 180-A4). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 306 POLYPHASIC texture velvety; sporulation dense; conidia en masse dull green to dark green (26E4–26F4); exudates absent; soluble pigment absent; reverse centre yellowish brown (5E4–5F4) fading into light yellow to greyish yellow (2A5–2B5 to 1A5–1B5). MEA 25 C, 7 d: Colonies low, plane; margins narrow (1 mm), low, entire, plane; mycelium in margins white, in centre aerial yellow; texture velvety; sporulation dense; conidia en masse dull green to dark green (26E4–26F4); exudates absent; soluble pigment absent; reverse centre yellowish brown (5E6–6E6). YES 25 C, 7 d: Colonies, slightly raised at centre, some isolates crateriforme and some isolates slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium generally only white, in some isolates in the centre very light yellow (CBS 137360, CBS 101423 and DTO 269-G4); texture velvety; sporulation dense; conidia en masse dull green to dark green (26E4–26F4); exudates absent; soluble pigment absent; reverse centre orange (6B7–6B8) fading into pastel yellow to light yellow (1A4–2A5) and greyish green (1C5–1D5), in some isolates lack of centre colour. DG18 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white; texture velvety; sporulation dense; conidia en masse dull green to dark green (26E4–26F4); exudates absent; soluble pigment absent; reverse centre yellowish brown (5E4–5F4) fading into light yellow to greyish yellow (2A5–2B5 to 1A5–1B5). OA 25 C, 7 d: Colonies low, plane; margins narrow (1 mm), low, entire, plane; mycelium white and yellow; texture velvety to loosely funiculose, especially at centre; sporulation dense; conidia en masse dull green to dark green (26E4–26F4); exudates clear, yellow and orange droplets; soluble pigment absent; reverse yellowish green and in some isolates brownish orange centre. CREA, 25 C, 7 d: Colonies in some isolates acid production absent and in some isolates very weak acid production. Micromorphology: Conidiophores biverticillate having symmetrical subterminal branches; stipes smooth walled, 40–110 × 2–3 μm; with extra branches 10–25 μm; metulae three to six, divergent, 7–15 × 2–3.5 μm; phialides acerose to ﬂask-shaped, three to six per metulae, 7–14 × 2–3.5 μm; conidia smooth to ﬁnely rough, ellipsoidal to fusiform, 2.5–6 × 2.5–4 μm. Extrolites: Talaromyces rugulosus produces emodin, endocrocin and skyrin (Tatsuno et al. 1975); (+)-rugulosin = rugulosin A (Howard & Raistrick 1954a, b, Takeda et al. 1973), rugulin (Sedmera et al. 1978), OF4949-I, -II, III, IV, -D and -F (Sano et al. 1986a, b) and prugosin (prugosene = ukulactones) A1, A2, A3, B1, B2, C1, C2 (Lang et al. 2007, Mori et al. 2011). We detected berkelic acid, two mitorubrins, rugulosin, skyrin, one of the OF4949 compounds, and three prugosins (= ukulactones). Distinguishing characters: Talaromyces rugulosus grows slow and produces compact and velvety colonies with dark green conidia (Fig. 70). Its conidiophores have acerose to ﬂask-shaped phialides and additional symmetrical subterminal branches. Based on ITS and BenA phylogenies, T. rugulosus is resolved as a close relative of T. atricola (Fig. 7). In addition, Penicillium chrysitis, P. tardum and T. echinosporus are synonomous with T. rugulosus (Fig. 7). Talaromyces rugulosus differs from its close relatives by the production of dark green, compact and velvety colonies. Notes: Samson et al. (2011), accepted T. echinosporus (CBS 344.51) as a valid species. However our ITS and BenA www.studiesinmycology.org TAXONOMY OF TALAROMYCES phylogenies show, that T. echinosporus (CBS 344.51), P. tardum (NRRL 1073), P. chrysitis (CBS 137366), and P. elongatum (NRRL 1073) are identical with T. rugulosus (Fig. 7) and are considered synonyms. A detailed study of this species complex is being prepared in a separate publication. Talaromyces ryukyuensis (S. Ueda & Udagawa) Arx, Persoonia 13: 282. 1987. MycoBank MB132096. ≡ Sagenoma ryukyuense S. Ueda & Udagawa, Mycotaxon 20: 499. 1984. In: Talaromyces section Helici Typus: Unknown, culture ex-type NHL 2917 = DTO 176-I6. ITS barcode: AB176628 Colony characters: Fide Ueda & Udagawa (1984) colonies on OA spreading rapidly at 37 C, attaining a diameter of 8.5 cm within 10 d, consisting of thin layer of numerous ascomata coral to dark vinaceous (Rayner 1970) in colour; conidia limited; exudate lacking; odour faintly mouldy; reverse amber to coral (Rayner 1970). Colonies on malt agar spreading, 7.8 cm in 14 d at 37 C, thin with reduced formation of ascomata, showing a marked tendency to develop sectors; conidia abundantly produced, saffron to yellow-green or malachite green (Rayner 1970). Colonies on czapek agar growing restrictedly, very thin, vegetative mycelium submerged, pale luteous (Rayner 1970); ascomata not produced. Micromorphology: Fide Ueda & Udagawa (1984) conidiophores with solitary phialides or monoverticillate, stipe smooth, 12.5–50 × 3–4 μm; phialides four to six, acerose, 10–17.5(–20) × 2.5–5 μm; conidia formed in long chains, thick walled, usulally rough sometimes smooth, cylindrical to fusiform, ellipsoidal or ovoidal, 3.5–12(–15) × 2.5–3.5 μm. Ascomata ripening within 2 to 3 wk, deep reddish, globose to subglobose, 80–350 μm in diam; asci 8.5–10.5 μm in diam; ascospores yellowish orange, reddish in mass, broadly ellipsoidal, 4–5 × 3–4 μm, thick walled, usually spiny with spines up to 0.5 μm long. Notes: Talaromyces ryukyuensis is a thermotolerant species (Ueda & Udagawa 1984) with an optimum growth temperature of 30–37 C, while at 25 and 40 C a reduction in sporulation occur (Ueda & Udagawa 1984). It was originally described as a Sagenoma species, but based on molecular results, it is classiﬁed in Talaromyces. Unfortunately, the ex-type strain was not available for our study, and data were taken from Ueda & Udagawa (1984). Talaromyces sayulitensis Visagie et al., Stud. Mycol. 78: 132. 2014. MycoBank MB809188. In: Talaromyces section Talaromyces Typus: CBS H-21798, culture ex-type CBS 138204 = DTO 245-H1. ITS barcode: KJ775713 (alternative markers: BenA = KJ775206; CaM = KJ775422) 307 YILMAZ ET AL. Colony diam, 7 d (mm): Fide Visagie et al. (2014) CYA 24–29; CYA 30 C 35–43; CYA 37 C 32–40; MEA 37–40; MEA 30 C 24–27; DG18 18–22; CYAS 5–8; OA 40–42; CREA 15–18; YES 37–40. Colony characters: Fide Visagie et al. (2014) CYA 25 C, 7 d: Colonies low, raised at centre, slightly sulcate; margins low, narrow, entire; mycelia white to yellow to red; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent to clear in some isolates; reverse brown (6E6) in the centre, fading into brownish orange (6C7) and light yellow (4A5). MEA 25 C, 7 d: Colonies low, slightly raised at centre, plane; margins low, narrow, entire; mycelia white, pastel yellow and pastel red; texture loosely funiculose to ﬂoccose; sporulation sparse, conidia en masse greyish green (27D5–E5); soluble pigments absent; exudates absent; reverse brownish orange (6C6–6C7). YES 25 C, 7 d: Colonies low, raised at centre, sulcate; margins low, narrow, entire; mycelia white to yellow; texture loosely funiculose to ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (27D5–27E5); soluble pigments absent; exudates absent; reverse brownish orange (6C6–6C7). DG18 25 C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white to yellow; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (26D5–26E5); soluble pigments absent; exudates absent; reverse light yellow (3A5–4A5). OA 25 C, 7 d: Colonies low, slightly raised at centre, plane; margins low, wide, entire; mycelia white to yellow; texture loosely funiculose and ﬂoccose, especially in the centre sterile aerial hyphae; sporulation dense, conidia en masse greyish green (27C5–27D5); soluble pigments absent; exudates absent. CREA 25 C, 7 d: Acid strongly produced. Micromorphology: Fide Visagie et al. (2014) conidiophores biverticillate, subterminal branches sometimes present; stipes smooth walled, (40–)85–300 × 2–3.5 μm; branches up to 40 μm long; metulae divergent, 8–11.5(–14) × 2.5–3 μm; phialides acerose, 8–11 × 2.5–3 μm; conidia smooth, subglobose to broadly ellipsoidal, 2.5–3 × 2–2.5. Notes: Talaromyces sayulitensis is a close relative of T. pinophilus and T. liani. Talaromyces liani lacks the acid production observed in T. sayulitensis, produces larger conidia (2.5–4 μm) and typically produces a sexual state. Talaromyces pinophilus also produces acid on CREA. Other colony characters are very similar between T. pinophilus and T. sayulitensis with only minor differences observed in colony growth rates. However, this is not considered to be sufﬁcient for morphological identiﬁcation. As a result, sequence data is needed for making correct identiﬁcations. For an illustration of the species, readers are referred to Visagie et al. (2014). Talaromyces scorteus (Nakazawa, Takeda & Suematsu) S.W. Peterson & Jurjevic, PLoS ONE e78084-page 8. 2013. MycoBank MB492647. Fig. 71. ≡ Penicillium scorteum Nakazawa, Takeda & Suematsu, J. Agric. Chem. Soc., Japan 10: 103. 1934. = Talaromyces phialosporus (Udagawa) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 176. 2011 ≡ Penicillium phialosporum Udagawa, J. Agric. Sci. Tokyo Nogyo Daig. 5: 11. 1959. In: Talaromyces section Islandici 308 Typus: Unknown, culture ex-typus: CBS 340.34 = NRRL 1129 = FRR 1129. ITS barcode: KF984892 (alternative markers: BenA = KF984565; CaM = KF984684; RPB2 = KF984916) Colony diam, 7 d (mm): CYA 8–16; CYA 30 C 8–9; CYA 37 C No growth; MEA 10–15; MEA 30 C 15–16; DG18 8–17; CYAS Generally no growth, in some isolates colonies up to 8; OA 8–16; CREA 4–10; YES 7–16. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (12 mm), low, in some isolates entire and in some isolates not entire, plane; mycelium white and yellow; sporulation sparse to dense; texture velvety and in some isolates ﬂoccose in the centre; conidia en masse dark green (26F4–26F6); exudates absent; soluble pigment absent (except CBS 500.75 yellow soluble pigments around colonies); reverse centre olive (2E4) centre fading into greyish yellow to olive (2C4–2D4) (CBS 500.75 reverse reddish yellow (4A6)). MEA 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (1 mm), low, in some isolates entire and in some isolates not entire, plane; mycelium white and yellow; sporulation sparse to dense; texture velvety and in some isolates ﬂoccose in the centre; conidia en masse dark green (26F4–26F6); exudates absent; soluble pigment absent; reverse centre yellowish brown (5F6) fading into brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (1–2 mm), low, in some isolates entire and in some isolates not entire, plane; mycelium white and yellow; sporulation sparse to dense; texture velvety and in some isolates ﬂoccose in the centre; conidia en masse dark green (26F4–26F6); exudates absent; soluble pigment absent; reverse centre olive (3E5–3F5). DG18 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins narrow (1 mm), low, in some isolates entire and in some isolates not entire, plane; mycelium white and yellow; sporulation sparse to dense; texture velvety and in some isolates ﬂoccose in the centre; conidia en masse dark green (26F4–26F6); exudates absent; soluble pigment absent; reverse centre olive (2E4) centre fading into greyish yellow to olive (2C4–2D4) (except DTO 270-A6 and CBS 340.34 reverse orange (6A7) and light yellow (4A4–4A5), respectively). OA 25 C, 7 d: Colonies low, plane; margins narrow (1 mm), low, entire, plane; mycelium white and yellow; sporulation sparse to dense; texture velvety to ﬂoccose; conidia en masse greyish green (29D4–29E4); exudates absent (except DTO 270A6 clear droplets); soluble pigment absent; reverse dark green to yellowish green. CREA, 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate having symmetrical subterminal branches; stipes smooth walled, 40–110 × 2.5–3 μm; with extra branches 10–25 μm; metulae three to six, divergent, 7–12 × 1.5–3.0 μm; phialides ﬂaskshaped to acerose, three to six per metulae, 6–13 × 2–3.5 μm; conidia smooth, ellipsoidal, 3–5.5 × 2–3 μm. Extrolites: Talaromyces scorteus produced mitorubrin and compounds with a chromophore like rubiginosin. Distinguishing characters: Talaromyces scorteus is characterised by restricted growth and deep, compact and densely sporulating colonies (Fig. 71). It cannot grow at 37 C. Talaromyces scorteus is most similar to T. rugulosus, both having biverticillate POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 71. Morphological characters of Talaromyces scorteus (CBS 233.60). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 309 YILMAZ ET AL. conidiophores, commonly with subterminally branches, and ﬂask-shaped to acerose phialides. However, T. scorteus has shorter stipes, smaller colony sizes on most media and darker conidia. Notes: We are following Peterson & Jurjevic (2013) who considered that T. phialosporus (CBS 233.60) is a synonym of T. scorteus (Fig. 7). Talaromyces siamensis (Manoch & C. Ramírez) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. MycoBank MB560674. Fig. 72. ≡ Penicillium siamense Manoch & C. Ramírez, Mycopathologia 101: 32. 1988. Extrolites: Talaromyces siamensis produces mitorubrin, mitorubrinic acid, a purpactin, secalonic acid D and vermicillin. Distinguishing characters: The ex-type strain of T. siamensis is deteriorated and no longer produces the reported characteristic purple red exudates and reverse (Manoch & Ramírez 1988). Talaromyces siamensis is closely related to T. cnidii and only minor morphological differences between these two species exist. Talaromyces cnidii grows slightly faster at 25 and 37 C compare to T. siamensis, but for reliable identiﬁcation, BenA should be sequenced. Talaromyces solicola Visagie & K. Jacobs, Persoonia 28: 20. 2012. MycoBank MB564328. Fig. 73. In: Talaromyces section Talaromyces In: Talaromyces section Trachyspermi Typus: CBS 475.88, cultures ex-type CBS 475.88 = IMI 323204. Typus: PREM 60037, cultures ex-type DAOM 241015 = CV 2800 = CBS 133445 = DTO 180-D4. ITS barcode: JN899385 (alternative markers: BenA = JX091379; CaM = KF741960; RPB2 = KM023279) Colony diam, 7 d (mm): CYA 20–22; CYA 30 C 30–32; CYA 37 C 15; MEA 32–33; MEA 30 C 40–42; DG18 10–12; CYAS No growth; OA 30–32; CREA 7–8; YES 27–28. Colony characters: CYA 25 C, 7 d: Colonies sunken at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation sparse, conidia en masse greyish turquoise (25D4–25D5); soluble pigments absent; exudates absent; reverse blond to greyish yellow (4C4) and at 30 C centre reddish brown (9F7) fading into light yellow (4A5). MEA 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (2 mm); mycelia white; texture velvety; sporulation sparse, conidia en masse dull green (26D4–26E4); soluble pigments absent; exudates clear droplets at centre; reverse light brown (5D6) centre with an orange (5A7) circle and fading into greyish orange (5B5). YES 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (2 mm); mycelia white and yellow; texture ﬂoccose; sporulation sparse, conidia en masse greyish green (25E5–26E5); soluble pigments absent; exudates absent; reverse yellowish brown (5E6) centre fading into greyish orange (5B6) to reddish yellow (4A6). DG18 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1–2 mm); mycelia white; texture velvety; sporulation sparse, conidia en masse greyish green (25E5–26E5); soluble pigments absent; exudates clear droplets; reverse yellowish white (1A2) centre with greyish green (1C3) circle. OA 25 C, 7 d: Colonies slightly raised at centre, plane, in centre yeast like colony appearance; margins low, plane, entire (3 mm); mycelia white; texture velvety; sporulation moderately dense, conidia en masse greyish green (26E5); soluble pigments absent; exudates absent; reverse brownish orange. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches; stipes smooth walled, 150–450 × 2.5–3.5 μm; branches 10–25 μm; metulae three to six, divergent, 11–16 × 2.5–4 μm; phialides acerose, three to eight per metulae, 10–13 × 2–3 μm; conidia smooth to ﬁnely rough, ellipsoidal to fusiform, 3–4 × 2–3 μm. Ascomata not observed. 310 ITS barcode: FJ160264 (alternative markers: BenA = GU385731; CaM = KJ885279; RPB2 = KM023295) Colony diam, 7 d (mm): CYA 12–13; CYA 30 C 13–15; CYA 37 C No growth; MEA 22–23; MEA 30 C 24–26; DG18 11–12; CYAS No growth; OA 20; CREA No growth; YES 12–15. Colony characters: CYA 25 C, 7 d: Colonies slightly raised, crateriforme; margins low, plane, entire (1 mm); mycelia white and red; texture ﬂoccose; sporulation sparse, conidia en masse greyish green (25C5); soluble pigments absent; exudates clear droplets; reverse reddish brown to dark brown (9E6–9F6). MEA 25 C, 7 d: Colonies slightly raised, crateriforme; margins low, plane entire (1 mm); mycelia white and very pale light yellow; texture velvety and ﬂoccose in centre, with aerial sterile hyphae; sporulation dense, conidia en masse dark green (25F6–25F7); soluble pigments absent; exudates light red droplets; reverse centre dark brown (7F7), fading into brown (7E7). YES 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (1 mm); mycelia white and red; texture velvety; sporulation sparse, conidia en masse difﬁcult to determine but greyish green (25C5); soluble pigments just around colonies red soluble pigments; exudates red droplets; reverse reddish brown to dark brown (9E6–9F6). DG18 25 C, 7 d: Colonies slightly raised at centre, very slightly sulcate; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates big and small clear droplets; reverse orange white (5A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–4 mm); mycelia white; texture velvety; sporulation moderately dense to dense, conidia en masse dark green (27F4–27F5); soluble pigments absent; exudates very small clear droplets; reverse beige. CREA 25 C, 7 d: No growth (Fig. 73). Micromorphology: Conidiophores biverticillate; stipes smooth walled, 90–230 × 2.5–3.5 μm; metulae three to six, divergent, 8.5–11 × 2.5–3.5 μm; phialides acerose, three to eight per metulae, 9–11 × 2–2.5 μm; conidia rough, globose to subglobose, 2–3.5 × 2–2.5 μm. Ascomata not observed. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 72. Morphological characters of Talaromyces siamensis (CBS 475.88T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 311 YILMAZ ET AL. Fig. 73. Morphological characters of Talaromyces solicola (CBS 133445T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 312 POLYPHASIC Distinguishing characters: Talaromyces solicola is characterised by its slow growth on CYA, YES and DG18, and no growth on CREA and CYA at 37 C. Colonies typically produce red reverses on CYA and MEA, conidiophores are biverticillate and produce rough-walled globose to subglobose conidia (Fig. 73). Micromorphological characters resemble T. verruculosus, T. aculeatus and T. apiculatus. However, its slow growth, lack of growth at 37 C and colony reverse colour distinguish T. solicola from the latter species. Talaromyces stipitatus (Thom) C.R. Benj., Mycologia 47: 684. 1955. MycoBank MB306722. Fig. 74. TAXONOMY OF TALAROMYCES 12–15 × 2–3 μm; conidia ellipsoidal to ovoidal, 2–7.5 × 2–4 μm. Ascomata maturing after 1–2 wk of incubation on OA at 25 C and abundantly 30 C, creamish white, pastel yellow and yellow, globose to subglobose, 180–370 × 150–400 μm, asci 6–8.5 × 5–7 μm, ascospores ﬂattened ellipsoidal with single equatorial ridge, smooth, 3–5 × 2–3 μm. Extrolites: Talaromyces stipitatus produces botryodiploidin (Fuska et al. 1988), 7-epiaustdiol, 8-methoxy-epiaustdiol, duclauxin (Kuhr et al. 1973), compounds related to glauconic acid or rubratoxins, secalonic acid A, skyrin, stipitalide, stipitatic acid, stipitatonic acid and triacetic acid lactone. ≡ Penicillium stipitatum Thom, Mycologia 27: 138. 1935. In: Talaromyces section Talaromyces Typus: CBS H-7835, culture ex-type CBS 375.48 = ATCC 10500 = NRRL 1006 = IMI 39805. ITS barcode: JN899348 (alternative markers: BenA = KM111288; CaM = KF741957; RPB2 = KM023280) Colony diam, 7 d (mm): CYA 32–38; CYA 30 C 36–41; CYA 37 C 28–32; MEA 45–48; MEA 30 C 65–70; DG18 10–15; CYAS No growth; OA 30–35; CREA 8–13; YES 40–45. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate, bright yellow appearance; margins low, plane, entire (2–3 mm); mycelia white and bright yellow; sporulation absent; soluble pigments yellow to orange yellow (also at 30 and 37 C); exudates absent; reverse centre in some isolates brown (6E6) and in some isolates dark brown (6F6), fading into reddish golden brownish orange to light brown (6C7–6D7). MEA 25 C, 7 d: Colonies slightly raised at centre, plane, light yellow appearance, start to produce light yellow ascomata (at 30 C more abundant); margins low, plane, entire (2–3 mm); mycelia light yellow and white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brown (7E6) centre fading into brownish orange (7C6). YES 25 C, 7 d: Colonies sunken at centre, sulcate, bright yellow appearance; margins low, plane, entire (3–4 mm); mycelia bright yellow and white; sporulation absent; soluble pigments yellow to orange yellow; exudates absent; reverse brown (7D7–7E7) centre fading into sunﬂower yellow (4A7). DG18 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white and pastel yellow; texture ﬂoccose; sporulation sparse to moderately dense in the centre, conidia en masse greyish green (30D5); soluble pigments absent; exudates absent; reverse centre light yellow to yellow (2A5–2A6), in some isolates with olive brown (4F4) dot and fading into pale yellow (2A3). OA 25 C, 7 d: Colonies low, plane, formation of yellowish white ascomata (at 30 C more abundant); margins low, plane, entire (2 mm); mycelia yellowish white and white; sporulation absent; soluble pigments absent (except DTO 108-A2 produces yellow soluble pigments); exudates absent; reverse brownish orange to brownish yellow. CREA 25 C, 7 d: Acid production weak. Micromorphology: Conidiophores asexual state lacking, ﬁde Stolk & Samson (1972), biverticillate and monoverticillate with a minor proportion having subterminal branches; stipes smooth walled, 10–100 × 2–2.5 μm; metulae two to ﬁve, divergent, 11–16 × 2.2–3 μm; phialides acerose, two to six per metulae, www.studiesinmycology.org Distinguishing characters: Talaromyces stipitatus is distinguished by its smooth, ﬂattened, ellipsoidal ascospores with single equatorial ridge (Fig. 74). It grows fast on general media, except on DG18. The asexual state was not observed during our study. Talaromyces stollii Yilmaz et al., Persoonia 29: 52. 2012. MycoBank MB801359. Fig. 75. In: Talaromyces section Talaromyces Typus: CBS H-21053, culture ex-type CBS 408.93. ITS barcode: JX315674 (alternative markers: BenA = JX315633; CaM = JX315646; RPB2 = JX315712) Colony diam, 7 d (mm) CYA 38–45; CYA 30 C 40–50; CYA 37 C 25–35; MEA 45–50; MEA 30 C 53–55; DG18 18–22; CYAS 4–5; OA 45–50; CREA 20–35; YES 35–45. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (2–3 mm); mycelia white and pastel red; texture ﬂoccose; sporulation moderately dense, conidia en masse dull green (26E4–27E4); soluble pigments red and in some isolates absent; exudates absent; reverse brownish red (8E8–8F8) in the centre fading into greyish red (7B3). MEA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–5 mm); mycelia white, pastel yellow and pastel red; texture ﬂoccose and loosely funiculose; sporulation moderately dense, conidia en masse greyish to dull green (27C4–27D4); soluble pigments absent; exudates absent; reverse brownish orange to brownish yellow (5C6–6C7). YES 25 C, 7 d: Colonies raised at centre, sulcate; margins low, plane, entire (3–4 mm); mycelia white; texture ﬂoccose; sporulation sparse, conidia en masse greyish green to dull green (27C4–27D4); soluble pigments absent; exudates absent; reverse melon yellow (5A6). DG18 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2–3 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A4–5A5). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (5–6 mm); mycelia white and pale yellow; texture ﬂoccose and loosely funiculose, in the centre sterile aerial mycelia; sporulation moderately dense, conidia en masse greyish green to dull green (27C4–27D4); soluble pigments absent; exudates absent; reverse in some isolates reddish centre and green elsewhere, in some strain yellowish beige. CREA 25 C, 7 d: Acid production. Micromorphology: Conidiophores biverticillate with a minor proportion having subterminal branches, smooth walled, arising 313 YILMAZ ET AL. Fig. 74. Morphological characters of Talaromyces stipitatus (CBS 236.60). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D, E. Asci and ascospores. F. Ascospores. Scale bars: B = 1000 μm; C = 500 μm; D = 10 μm, applies to E, F. 314 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 75. Morphological characters of Talaromyces stollii (CBS 408.93T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 315 YILMAZ ET AL. from aerial hyphae and overgrowing the ascomata, stipes 90–250 × 3–4.5 μm; metulae three to ﬁve, divergent 11–15 × 2–3.5 μm; phialides acerose, three to six per metulae, 13–17 × 2.0–2.5 μm; conidia, smooth to ﬁnely rough, ellipsoidal, 2.5–4 × 2–2.5 μm. 7.5–11 × 2–3 μm; conidia smooth, ellipsoidal to fusiform, 2.5–4 × 1.5–2 μm. Ascomata not observed in our culture however, ﬁde Yaguchi et al. (1993) maturing after 2–3 wk of incubation, sulphur yellow, globose to subglobose, 300–480 μm, asci 7–9.5 × 6.5–8 μm, ascospores broadly ellipsoidal, spiny, 3–4(–5.5) × 2–3(–5) μm. Extrolites: Talaromyces stollii produces austins (Yilmaz et al. 2012). Distinguishing characters: Talaromyces stollii is characterised by fast growth on general media and colonies with ﬂoccose or loosely funiculose textures on MEA, with white, yellow and red mycelia (Fig. 75). These characters resemble T. angelicus, T. aculeatus, T. apiculatus, T. verruculosus, T. amestolkiae and T. pinophilus. Talaromyces aculeatus, T. apiculatus and T. verruculosus produce rough-walled, globose conidia and ﬂask-shaped phialides, which distinguish them from Talaromyces stollii. Talaromyces angelicus differs from T. stollii by the lack of acid production on CREA. Talaromyces amestolkiae produces red colony reverses on CYA and MEA, black sclerotioa after 2 wk of incubation and grows more restrictedly on CYA at 37 C, when compared to T. stollii. Talaromyces stollii micro- and macromorphologically most closely resembles T. pinophilus and T sayulitensis. However, T. stollii grows slightly faster than both these species on CYA. Sequence data is, however, recommended for reliable identiﬁcation. Talaromyces subinﬂatus Yaguchi & Udagawa, Trans. Mycol. Soc. Japan 34: 249. 1993. MycoBank MB361184. Fig. 76. ≡ Penicillium subinﬂatum Yaguchi & Udagawa, (simultaneously published). In: Talaromyces section Subinﬂati Typus: CBM PF-1113, culture ex-type CBS 652.95 = IBT 17520. ITS barcode: JN899397 (alternative markers: BenA = KJ865737; CaM = KJ885280; RPB2 = KM023308) Colony diam, 7 d (mm): CYA 3–4; CYA 30 C 3–4; CYA 37 C No growth; MEA 14–15; MEA 30 C 10–11; DG18 No growth; CYAS No growth; OA 13–15; CREA No growth; YES 3–4. Colony characters: CYA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2). MEA 25 C, 7 d: Colonies slightly raised, plane, ﬂuffy; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse centre is dark brown (6F5) fading into brownish orange (5B5–5B6). YES 25 C, 7 d: Colonies low, plane; margins low, plane, entire (<1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pastel yellow (3A4). DG18 25 C, 7 d: Colonies No growth. OA 25 C, 7 d: Colonies raised, plane, ﬂuffy; margins low, plane, entire (<1 mm) it does not have the transparent zone; mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse pale pastel yellow. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate; stipes smooth walled, 150–350 × 2–2.5 μm; metulae two to ﬁve, divergent, 7.5–10.5 × 2.5–3 μm; phialides acerose, two to ﬁve per metulae, 316 Distinguishing characters: Talaromyces subinﬂatus grows very restrictedly on common media, MEA being the exception on which it reaches 15 mm, and does not grow on DG18, CREA or CYA at 37 C (Fig. 76). These growth rates distinguish T. subinﬂatus from all other Talaromyces species. It produces large sulphur yellow ascomata and rather small, broadly ellipsoidal, spiny ascospores. When Yaguchi et al. (1993) described T. subinﬂatus they mentioned that it produced a Merimbla-like conidiophore, with swollen apices of stipes and strongly divergent metulae. The ex-type strain of T. subinﬂatus is unfortunately deteriorated and we did not observe the Merimbla-like conidiophores or ascomata. Talaromyces tardifaciens Udagawa, Mycotaxon 48: 150. 1993. MycoBank MB360478. Fig. 77. ≡ Penicillium tardifaciens Udagawa, (simultaneously published). In: Talaromyces section Islandici Typus: CBM SUM 3017, culture ex-type CBS 250.94 = DTO 247D6 = IBT 14986. ITS barcode: JN899361 (alternative markers: BenA = KC202954; CaM = KF984682; RPB2 = KF984908) Colony diam, 7 d (mm): CYA 9–10; CYA 30 C 7–8; CYA 37 C No growth; MEA 13–15; MEA 30 C 10–13; DG18 7–8; CYAS No growth; OA 13–15; CREA No growth; YES 9–10. Colony morphology: CYA, 25 C, 7 d: Colonies 9–10 mm, slightly raised in the centre, plane; margins very narrow (<1 mm), low, not entire, plane; mycelium white; sporulation absent; exudates very small clear droplets; soluble pigment absent; reverse centre light orange (5A5–5A6) fading into greyish yellow (4C4). MEA, 25 C, 7 d: Colonies 13–15 mm, slightly raised in the centre, plane; margins very narrow (1 mm), low, not entire, plane; mycelium white; texture ﬂoccose; sporulation absent; exudates absent; soluble pigment absent; reverse centre golden yellow to brownish yello (5B7–5C7) colour. YES, 25 C, 7 d: Colonies 9–10 mm, slightly raised in the centre, plane; margins very narrow (<1 mm), low, not entire, plane; mycelium white and light yellow; sporulation absent; exudates very small bright orange droplets; soluble pigment absent; reverse deep orange to orange (5A8–5B8). DG18, 25 C, 7 d: Colonies 7–8 mm, slightly raised in the centre, plane; margins very narrow (1 mm), low, not entire, plane; mycelium white and yellow; sporulation absent; exudates absent; soluble pigment absent; reverse centre orange (5A6) fading into butter yellow to maize yello (4A5–4A6). OA, 25 C, 7 d: Colonies 13–15 mm, slightly raised in the centre, plane; margins narrow (1–2 mm), low, not entire, plane; mycelium white; after 3 wk incubation white ascomata production; sporulation absent; exudates very small clear droplets; soluble pigment absent; reverse centre light orange fading into light yellow and has a characteristic mouldy soil smell. CREA, 25 C, 7 d: No growth. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 76. Morphological characters of Talaromyces subinﬂatus (CBS 652.95T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 317 YILMAZ ET AL. Fig. 77. Morphological characters of Talaromyces tardifaciens (CBS 250.94T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Asci. C. Ascospores. D. Initials. E–H. Conidiophores. I. Conidia. Scale bar: B = 10 μm, applies to C–I. 318 POLYPHASIC TAXONOMY OF TALAROMYCES Micromorphology: Conidiophores with solitary phialides or monoverticillate, smooth walled, arising from aerial hyphae and overgrowing the ascomata, stipes 15–40 × 2–2.5 μm; metulae absent; phialides one to three, acerose, 12.5–20 × 2.0–2.5 μm; conidia, smooth walled, cylindrical to ellipsoidal, 3–6 × 1.5–2.5 μm. Ascomata creamish white to white yellow, globose to subglobose, ripening within 3 wk or more, 200–400 μm. Covering consisting of a few layers of welldeveloped networks of white and light yellow hyphae. Asci broadly globose to ovoidal, 6–8 × 6–7 μm. Ascospores, broadly ovoidal, smooth walled, 3–3.5 × 2–3 μm. orange (6A6) with a deep orange (6A8) circle in the margins. DG18 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates clear droplets; reverse light orange (5A4) centre with a pastel red (8A4) circle in the margins. OA 25 C, 7 d: Colonies low, plane, formation of yellow and orange ascomata (30 C more abundant); margins low, plane, entire (2–3 mm); mycelia dominantly yellow and also white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse orange yellow. CREA 25 C, 7 d: Acid production absent (Fig. 78). Extrolites: Talaromyces tardifaciens produces skyrin, 60 -hydroxy30 -methoxy-mitorubrin related extrolites, and some extrolites that are unique for this species (structures not yet elucidated, but with characteristic UV spectra). Micromorphology: Conidiophores biverticillate; stipes smooth walled, 200–400 × 2.5–3.5 μm; metulae three to six, divergent 9.5–17 × 2.5– 3.5 μm; phialides acerose, three to six per metulae, 9–14 × 2–3.5 μm; conidia smooth, broadly ellipsoidal, sometimes ovoidal, 2–4 × 2–3 μm. Ascomata maturing after 1–2 wk of incubation on MEA and OA at 25 C and abundantly 30 C, yellow, globose to subglobose, 160–520 μm, asci 10–13.5 × 8.5–11 μm, ascospores broadly ellipsoidal, thick walled, spiny, 4–6 × 2.5–4.5 μm. Distinguishing characters: Talaromyces tardifaciens is characterised by restricted growth on general media, with colonies that lack sporulation after 7 d of incubation at 25 C. Based on the ITS and BenA phylogenies, T. tardifaciens is closely related to T. tratensis and T. wortmannii. Talaromyces tardifaciens and T. tratensis and some strains of T. wortmannii produce the sexual state; however, the latter two species differ from T. tardifaciens by fast maturing ascomata (1–2 wk), whereas T. tardifaciens ascomata maturation take 21 d (Udagawa 1993). Also, T. tardifaciens produces monoverticillate conidiophores and ascospores (2.5–3 × 2–3 μm), compared to the biverticillate conidiophores and bigger ascospores of T. tratensis (3–5 × 2.5–3.5 μm) and T. wortmannii (3.5–6 × 2.5–4 μm). Extrolites: Talaromyces thailandensis produces asperphenamate, thailandolide A & B, 3-methyl-6-hydroxy-8-methoxy-3,4dihydroisocoumarin, penisimplicissin, vermistatin and hydroxydihydrovermistatin (Dethoup et al. 2007). We found mitorubrin, mitorubrinol, mitorubrinic acid, a purpactin, vermicillin, and a compound with an emodin chromophore. In: Talaromyces section Talaromyces Distinguishing characters: Talaromyces thailandensis grows fast on general media and produces yellow ascomata with spiny, thick walled, broadly ellipsoidal ascospores (Fig. 78). Its characters resemble those of T. muroii, T. macrosporus and T. liani. However, T. thailandensis is distinguished from these species by the absence of growth on CYA at 37 C. Typus: CBS H-21075, culture ex-type CBS 133147 = KUFC 3399. Talaromyces trachyspermus (Shear) Stolk & Samson, Stud. Mycol. 2: 32. 1972. MycoBank MB324421. Fig. 79. Talaromyces thailandensis Manoch, Dethoup & Yilmaz, Mycoscience 54: 339. 2013. MycoBank MB801737. Fig. 78. ITS barcode: JX898041 (alternative markers: BenA = JX494294; CaM = KF741940; RPB2 = KM023307) Colony diam, 7 d (mm): CYA 33–35; CYA 30 C 38–40; CYA 37 C No growth; MEA 30–35; MEA 30 C 35–36; DG18 16–20; CYAS No growth; OA 38–40; CREA 10–12; YES 35. ≡ Arachniotus trachyspermus Shear, Science 16: 138. 1902. = Penicillium spiculisporum Lehman, Mycologia 12: 268. 1920 ≡ Talaromyces spiculisporus (Lehman) C.R. Benj., Mycologia 47: 683. 1955. = Talaromyces trachyspermus var. macrocarpus J.E. Wright & Loewenb., Bol. Soc. Argent. Bot. 15: 100. 1973. ≡ Penicillium lehmani Pitt, The genus Penicillium: 497. 1980. In: Talaromyces section Trachyspermi Colony characters: CYA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (1–2 mm); mycelia white and yellow; texture ﬂoccose; sporulation sparse, conidia en masse greyish turquoise to greyish green (24E5–25E5); soluble pigments absent; exudates absent (at 30 C clear droplets); reverse yellowish red to red (8A6–9A6) centre with red (9B7) dots. MEA 25 C, 7 d: Colonies low, plane, entire; margins low, plane, entire (1 mm); mycelia dominantly yellow and also white; texture ﬂoccose; sporulation sparse to moderately dense, conidia en masse greyish green (25D5); soluble pigments absent; exudates absent; reverse greyish orange (5B6–6B6). YES 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (1–2 mm); mycelia white, yellow and pink; texture ﬂoccose; sporulation sparse, conidia en masse greyish green (25D5); soluble pigments absent; exudates absent; reverse www.studiesinmycology.org Typus: IMI 040043, culture ex-type CBS 373.48 = ATCC 10497 = IMI 040043 = NRRL 1028. ITS barcode: JN899354 (alternative markers: BenA = KF114803; CaM = KJ885281; RPB2 = JF417432) Colony diam, 7 d (mm): CYA 13–24; CYA 30 C 30–40; CYA 37 C 32–40; MEA 17–25; MEA 30 C 32–40; DG18 Generally no growth some colonies up to 8; CYAS No growth; OA 20–24; CREA Generally no growth some colonies up to 4; YES 15–23. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate; margins low, plane, entire (2–3 mm); mycelia white and yellow; texture ﬂoccose; sporulation 319 YILMAZ ET AL. Fig. 78. Morphological characters of Talaromyces thailandensis (CBS 133147T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata D. Asci and ascospores. E–G. Conidiophores. H. Conidia. I. Ascospores. Scale bars: B, C = 1000 μm; D = 10 μm, applies to E–I. 320 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 79. Morphological characters of Talaromyces trachyspermus (CBS 373.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata D. Asci and ascospores. E–G. Conidiophores. H. Ascospores. I. Conidia. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–I. www.studiesinmycology.org 321 YILMAZ ET AL. absent to sparse, conidia en masse greyish green to dull green (25C4–25D3); soluble pigments absent; exudates absent; reverse in some isolates centre yellowish brown (5D5–5E5) fading into pale orange to greyish orange (5A3–5B3), in some isolates centre light orange (5A5) fading into light yellow to yellow (3A5–3A6). MEA 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse centre brownish orange to light brown (6C6–6D6–6E6), fading into brownish orange (5C5–5C6). YES 25 C, 7 d: Colonies raised at centre, slightly sulcate; margins low, plane, entire (5 mm); mycelia white; texture ﬂoccose; sporulation sparse, conidia en masse greyish green to dull green (25C4–25D3); soluble pigments absent; exudates absent; reverse light orange (5A5–5A6) centre fading into yellowish white (4A2). DG18 25 C, 7 d: Colonies slightly raised in the centre, very slightly radially sulcate; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–4 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse beige only in DTO 228-D9 yellow. CREA 25 C, 7 d: No growth (only DTO 56-A4 up to 4 mm). Micromorphology: Conidiophores generally monoverticillate; stipes smooth walled, 15–45 × 1–2 μm; metulae two to three, divergent, 9–12 × 2–2.5 μm; phialides acerose, three to ﬁve per metulae, 10–15 × 1.3–2.5 μm; conidia smooth, ellipsoidal, 2–3.5(–5) × 1.5–2.5 μm. Ascomata maturing after 1–2 wk of incubation on OA at 25 C and abundantly 30 C, creamish white, globose to subglobose, 190–650 × 170–650 μm, asci 7.5–11.5 × 6.5–8 μm, ascospores broadly ellipsoidal, spiny, 3.5–5 × 2–3 μm. Extrolites: Talaromyces trachyspermus produces trachyspic acid (Shiozawa et al. 1995), spiculisporic acid (Clutterbuck et al. 1931) and glauconic acid. Distinguishing characters: Talaromyces trachyspermus produces creamish white ascomata with spiny, ellipsoidal ascospores (Fig. 79) and closely resembles T. intermedius and T. assiutensis. Talaromyces intermedius differs from T. trachyspermus by faster growth at 25 and 30 C and no growth at 37 C. Also, ascospores of T. intermedius are larger than T. trachyspermus. Phylogenetically and morpohologically, T. assiutensis is very similar to T. trachyspermus, with them having identical ITS sequences. However, T. trachyspermus grows slightly faster than T. assiutensis on CYA at 37 C and both species have unique BenA sequences. Notes: Talaromyces spiculisporus and T. trachyspermus var. macrocarpus were synonymised with T. trachyspermus by Stolk & Samson (1972) and Pitt (1980), and we concur with this. Talaromyces tratensis Manoch, Dethoup & Yilmaz, Mycoscience 54: 337. 2013. MycoBank MB801738. Fig. 80. In: Talaromyces section Islandici Typus: CBS H-21074, culture ex-type CBS 133146 = KUFC 3383. 322 ITS barcode: KF984891 (alternative markers: BenA = KF984559; CaM = KF984690; RPB2 = KF984911) Colony diam, 7 d (mm): CYA 10–12; CYA 30 C 13–20; CYA 37 C No growth; MEA 15–20; MEA 30 C 16–20; DG18 12–17; CYAS No growth; OA 15–20; CREA 4–8; YES 12–18. Colony characters: CYA, 25 C, 7 d: Colonies 10–12 mm, slightly raised in the centre, slightly plane; margins very narrow (<1 mm), low, entire, plane; mycelium white and very pale light orange yellow; sporulation sparse; texture loosely funiculose to ﬂoccose; conidia en masse greyish green to dull green (26C3–26D3 and in some isolates 29B5); exudates clear droplets; soluble pigment in some isolates very pale yellow, in some isolates absent; reverse greyish yellow (4A5–4A6) and in some isolates brownish orange (6C8) centre fading into beige. MEA, 25 C, 7 d: Colonies 15–20 mm, raised in the centre, plane; margins narrow (1 mm), low, entire, plane; mycelium white and predominately orange yellow (3A6); formation of yellow ascomata; sporulation absent to sparse; exudates very small clear droplets; soluble pigment absent; reverse brownish yellow (5C7–5C8) and in some isolates brown centre (6E4). YES, 25 C, 7 d: Colonies 12–18 mm, slightly raised in the centre, slightly sulcate; margins very narrow (<1 mm), low, not entire, plane; mycelium white and yellow; sporulation dense; texture ﬂoccose; conidia en masse greyish green (25C4–26C3); exudates in some isolates clear droplets (CBS 133146T); soluble pigment absent; reverse greyish orange (5B5–5B6). DG18, 25 C, 7 d: Colonies 12–17 mm, slightly raised in the centre, slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white and yellow; sporulation dense; texture velvety to loosely funiculose; conidia en masse greyish green to dull green (26C3–26D3 and in some isolates 29B5); exudates absent; soluble pigment absent; reverse in centre greyish yellow (2C5) and in some isolates yellow (3A7) fading into greyish yellow (2B5). OA, 25 C, 7 d: Colonies 15–20 mm, low, slightly sulcate; margins narrow (1–2 mm), low, entire, plane; mycelium white and predominately yellow; formation of yellow to orange ascomata; sporulation sparse, difﬁcult to determine; texture loosely funiculose, conidiophores raised at centre; exudates small yellow and orange droplets; soluble pigment absent; reverse pale to bright brownish orange. CREA, 25 C, 7 d: Colonies 4–8 mm, generally acid production absent, in some isolates very weak acid production (CBS 137401). Micromorphology: Conidiophores biverticillate, smooth walled stipes, arising from aerial hyphae and overgrowing the ascomata, stipes 120–200 × 2.5–3.5 μm; metulae three to six, divergent, 8.5–12.0 × 1.7–3.0 μm; phialides three to seven, acerose, 10–13.5 × 2.0–3.0 μm; conidia, smooth walled, ellipsoidal, 2.0–2.5 × 3.0–3.5 μm. Ascomata bright yellow, globose, ripening within 1–2 wk, covering consisting of a few layers of well-developed networks of yellow hyphae; asci broadly globose to subglobose, 7.5–11 × 8–11 μm; ascospores ovoidal to broadly ellipsoidal, thick walled, ﬁnely rough, 3.5–5.0 × 2.5–3.5 μm. Extrolites: Talaromyces tratensis produces mitorubrin, mitorubrinol, mitorubrinic acid, rugulosin, skyrin, vermicillin, wortmannin and a tetracyclic compound. POLYPHASIC TAXONOMY OF TALAROMYCES Fig 80. Morphological characters of Talaromyces tratensis (CBS 113146T) a. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Texture and ascomata on OA after 2 wk incubation. C. Asci and ascospores. D. Ascospores. E–H. Conidiophores. I. Conidia. Scale bar: C = 10 μm, applies to D–I. www.studiesinmycology.org 323 YILMAZ ET AL. Distinguishing characters: Talaromyces tratensis is characterised by restricted growth on general media and cannot grow at 37 C. Colonies are deep and consist of light yellow mycelia that form a ﬂuffy texture (Fig. 80). It produces yellow ascomata with spiny, ellipsoidal ascospores. The characters most closely resemble T. ﬂavus, T. austrocalifornicus and T. convolutus. On MEA, T. ﬂavus grows slighty faster than T. tratensis. Talaromyces tratensis ascopore size (3.5–5.0 × 2.5–3.5), yellow deep ﬂuffy colony texture on MEA and lack of growth at 37 C, distinguishes it from T. convolutus and T. austrocalifornicus. Talaromyces ucrainicus (Panas.) Udagawa, Trans. Mycol. Soc. Japan 7: 94. 1966. MycoBank MB449587. Fig. 81. = Penicillium ucrainicum Panas., Mycologia 56: 59. 1964. = Penicillium ohiense Pitt, The genus Penicillium: 502. 1980 ≡ Talaromyces ohiensis Pitt, The genus Penicillium: 502. 1980. = Talaromyces panasenkoi Pitt, The genus Penicillium: 482. 1980 ≡ Penicillium panasenkoi Pitt, The genus Penicillium: 482. 1980. In: Talaromyces section Trachyspermi Typus: Unknown, culture ex-type CBS 162.67 = ATCC 22344 = FRR 3462 = NHL 6086. ITS barcode: JN899394 (alternative markers: BenA = KF114771; CaM = KJ885282; RPB2 = KM023289) C Colony diam, 7 d (mm): CYA 10–20; CYA 30 20–26; CYA 37 C 7–17; MEA 18–24; MEA 30 C 25–33; DG18 5–7; CYAS No growth; OA 15–20; CREA No growth; YES 13–19. C, Colony characters: CYA 25 7 d: Colonies low to moderately deep, slightly raised at centre, plane; margins low, narrow, entire (1 mm); mycelia white, inconspicuously yellow; texture ﬂoccose; sporulation absent to sparse, conidia en masse greenish grey (28B2); soluble pigments absent; exudates absent; reverse olive to olive brown to brownish orange (1F6–4E5–5C5). MEA 25 C, 7 d: Colonies low to moderately deep, raised at centre, plane; margins low, narrow, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brown (6E6) at centre, greyish orange (5B5) elsewhere. YES 25 C, 7 d: Colonies low to moderately deep, raised at centre, plane; margins low, narrow, entire (2 mm); mycelia white to inconspicuously yellow; texture ﬂoccose; sporulation absent to sparse, conidia en masse turquoise grey (24B2); soluble pigments absent; exudates absent; reverse olive to brownish orange (3F6–5C4). DG18 25 C, 7 d: Colonies low, plane; margins low, narrow, entire; mycelia white; texture ﬂoccose; sporulation sparse, conidia en masse greenish grey (26C2); soluble pigments absent; exudates absent; reverse greyish green (1C4–29E5). OA 25 C, 7 d: Colonies low, plane, young ascomata visible; margins subsurface, wide, entire (6 mm); mycelia white and yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse pale yellow (1A3). CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, 14–80 × 1.5–2.6 μm; metulae three to six, divergent, 8–13 × 2–2.6 μm; phialides acerose, three to ﬁve per metulae, 8–13 × 1.5–3 μm; conidia smooth, broadly ellipsoidal to ovoidal, 2–4(–5) × 1.5–2.5(–3) μm. 324 Ascomata maturing after 1–2 wk of incubation on OA at 25 C and abundantly 30 C, yellow, globose to subglobose, 430–490 × 400–410 μm, asci 6.5–9.5 × 5–7.5 μm, ascospores broadly ellipsoidal, ornamented with thin, somewhat jagged, irregular, for the greater part longitudinal ridges, 3–5 × 2–3 μm. Extrolites: Talaromyces ucrainicus produces spiculisporic acid (Fujimoto et al. 1988). Spiculisporic acid was claimed to be a mycotoxin by Fujimoto et al. (1988), as it was lethal to mice, but the toxicity of the compound was based on interperitoneal injection, and can there not be regarded as a real mycotoxin. In fact spiculisporic acid can be produced in a yield of 110 g/l of fermentation medium and can be used as a biosurfectant (Ishigami et al. 2000). We also found alternariol in CBS 118436 and glauconic acid in CBS 583.72C and NRRL 2103. Distinguishing characters: Talaromyces ucrainicus grows restrictedly on general media and cannot grow on CREA. It produces yellow ascomata and ascospores, which are ornamented with thin and irregular ridges (Fig. 81). These characters easily distinguish T. ucrainicus from other species, which produce ascospores with ridges. Notes: Phylogenetically T. ohiensis (CBS 127.64) is identical to T. ucrainicus (Fig. 5) and is considered a synonym. Talaromyces udagawae Stolk & Samson, Stud. Mycol. 2: 36. 1972. MycoBank MB324424. Fig. 82. ≡ Penicillium udagawae Stolk & Samson, (simultaneously published). In: Talaromyces section Trachyspermi Typus: CBS H-7841, culture ex-type CBS 579.72 = FRR 1727 = IMI 197482. ITS barcode: JN899350 (alternative markers: BenA = KF114796) Colony diam, 7 d (mm): CYA 6–8; CYA 30 C 7–8; CYA 37 C No growth; MEA 10–11; MEA 30 C 15–17; DG18 No growth; CYAS No growth; OA 12; CREA No growth; YES 8–9. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (<1 mm); mycelia white and very pale light yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse pale orange to light orange (5A3–5A4). MEA 25 C, 7 d: Colonies raised at centre, sulcate, yellow appearance after 3 wk of incubation yellow ascomata produced; margins low, plane, entire (1 mm); mycelia white and yellow; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies slightly raised at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white and light orange (5A5); texture light orange appearance; sporulation absent; soluble pigments absent; exudates absent; reverse light orange (5A4) fading into pale orange (5A3). DG18 25 C, 7 d: No growth. OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (2 mm); mycelia white and yellow; texture yellow appearance after 2 wk of incubation yellow ascomata is produced; sporulation absent; soluble pigments absent; exudates absent; reverse light yellow. CREA 25 C, 7 d: No growth. POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 81. Morphological characters of Talaromyces ucrainicus (CBS 583.72C). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on OA after 2 wk incubation. C. Ascomata. D. Asci and ascospores. E–G. Conidiophores. H. Conidia. I. Ascospores. Scale bars: B, C = 500 μm; D = 10 μm, applies to E–I. www.studiesinmycology.org 325 YILMAZ ET AL. Fig. 82. Morphological characters of Talaromyces udagawae (CBS 579.72T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture on OA after 1 wk incubation. C. Colony texture and ascomata on OA after 3 wk incunation. D. Initials. E. Asci and ascospores. F. Ascospores. Scale bars: B, C = 500 μm; D = 10 μm, applies to E, F. 326 POLYPHASIC Micromorphology: Conidiophores asexual state lacking, ﬁde Stolk & Samson (1972), abundantly developed on hay-infusion agar at the colony margin. Conidiophores biverticillate; stipes smooth walled, 50–200 × 2.5–3.5 μm; metulae two to ﬁve, divergent, 7.5–10 × 2–2.4 μm; phialides acerose, three to six per metulae, 12–15 × 2.2–2.5 μm; conidia smooth, subglobose to ellipsoidal, 3–4 × 2–3 μm. Ascomata maturing after 2–3 wk of incubation on OA at 25 C and abundantly 30 C, yellow, globose to subglobose, 200–400 μm, asci 8–13.5 × 6–9.5 μm, ascospores ellipsoidal, ornamented with 3 to 5 regularly transverse, nearly parallel ridges, 3.5–6 × 2.5–3.5 μm. Extrolites: Talaromyces udagawae strain CBS 579.72 produces mitorubrin, mitorubrinol acetate, mitorubrinic acid and a purpactin. TAXONOMY OF TALAROMYCES Micromorphology: Fide Tzean et al. (1992) conidiophores on MEA, monoverticillate to biverticillate with a minor proportion having subterminal branches; stipes ﬁnely rough to rough, 35–180 × 2.6–4.2 μm; metulae ﬁnely rough to rough, two to ﬁve, divergent, 8.3–27.4 × 2.1–3.3 μm; phialides acerose, two to seven; smooth to ﬁnely rough, 6.7–22 × 2.1–3 μm; conidia mostly ellipsoidal to ovoidal, sometimes subglobose, smooth, 2.7–5 × 1.7–3.2 μm. Ascomata soft, abundant on MEA, usually discrete, globose, subglobose, or ellipsoidal, pastel yellow or yellow (3A4–3A6), 280–450 × 270–410 μm, maturing mostly in 8 wk; asci 8.5–10.8 × 7.3–9.2 μm; ascospores ellipsoidal, 3.3–6.7 × 2.8–4.2 μm, very rough to spiny; usually with a single ridge up to 1.3 μm wide. Extrolites: Talaromyces unicus produces an extrolite with a chromophore similar to talaroderxine. Distinguishing characters: Talaromyces udagawae is distinguished from the other Talaromyces species by its ascospores, which are ornamented with three to ﬁve regularly transverse, nearly parallel ridges (Fig. 82). It grows restrictedly on general media and does not grow on CREA, DG18 or CYA at 37 C. Conidiophores were not observed during our study. Pitt (1980) synonymised T. udagawae with T. luteus. However, Samson et al. (2011) showed that they are distinct species with T. luteus that does not belong in Talaromyces but is a close relative of Thermomyces. Further studies are needed on this species. Distinguishing characters: Talaromyces unicus grows restrictedly on CYA and does not grow at 37 C. Its most striking feature is its rough-walled ascospores that have a single ridge. Ascomata mature in 4 wk. Its single equatorial ridge resembles T. stipitatus, but ascospores of T. unicus are rough, whereas T. stipitatus ascospores are smooth. The ex-type strain of T. unicus was badly degraded and did not produce its characteristic features as originally described and as a result macro- and micromorphological characters were taken from Tzean et al. (1992). Talaromyces unicus Tzean, J.L. Chen & S.H. Shiu, Mycologia 84: 739. 1992. MycoBank MB360172. Talaromyces varians (G. Sm.) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. MycoBank MB560677. Fig. 83. ≡ Penicillium unicum Tzean, J.L. Chen & S.H. Shiu, (simultaneously published). ≡ Penicillium varians G. Sm., Trans. Brit. Mycol. Soc. 18: 89. 1933. In: Talaromyces section Helici In: Talaromyces section Bacillispori Typus: PPH 16, holotype Nat. Taiwan Univ., culture ex-type CBS 100535 = CCRC 32703 = IBT 18385. ITS barcode: JN899336 (alternative markers: BenA = KJ865735; CaM = KJ885283) Colony diam, 7 d (mm): Fide Tzean et al. (1992) CYA 10.5–16; CYA 37 C No growth; MEA 15–19.5; OA: 23–28. Colony characters: Fide Tzean et al. (1992) colonies on CYA restrictedly, dense, raised, ﬂoccose, white to reddish grey (10–11B2); mycelium white and reddish grey (10–11B2); sporulation not observed on CYA; soluble pigment pale greyish, pale red to reddish brown (8C–D5, 9D6); exudate absent; reverse reddish white, reddish orange, greyish red to reddish brown or violet brown or violet brown (7A2–6, 7B4–6, 8B–D4–6, 10–11E7–8). Colonies on MEA dense, ﬂoccose to funiculose; margins lower ﬁbrillous; mycelium whte to yellowish white (3–4A2); ascomata discrete or occasionally conﬂuent; sporulation moderate, white; soluble pigment pale greyish red; exudate, clear to pale yellow; reverse pastel (light) yellow, yellow (3A4–6, 5A–B6–7) or light orange becoming dark orange (6A4–8, 6B7–8) to reddish orange (7A–B6) in age. Colonies on OA thin and ﬂat, centre somewhat ﬂoccose, submargin to margin velvety, white; mycelium white; sporulation conspicuous; reverse white to yellowish white; exudate and soluble pigment absent; ascomata produced after 4 wk of incubation. www.studiesinmycology.org Typus: IMI 040586, culture ex-type CBS 386.48 = ATCC 10509 = IMI 040586 = NRRL 2096. ITS barcode: JN899368 (alternative markers: BenA = KJ865731; CaM = KJ885284; RPB2 = KM023274) Colony diam, 7 d (mm): CYA 22–24; CYA 30 C 26–30; CYA 37 C 27–28; MEA 30–32; MEA 30 C 40–43; DG18 8–12; CYAS 2–4; OA 25–28; CREA 9–10; YES 27–28. Colony characters: CYA 25 C, 7 d: Colonies slightly raised at centre, slightly sulcate; margins low, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (25B3); soluble pigments absent; exudates absent; reverse dark green (27F6) centre fading into dull green (7D3) fading into pale orange to light orange (5A3–5A4). MEA 25 C, 7 d: Colonies slightly raised at centre, plane; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (25C4); soluble pigments absent; exudates absent; reverse dark green (26F6) centre fading into between light yellow (4A5) and light orange (5A5). YES 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate; margins low, plane, entire (2 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse pastel green (25A4); soluble pigments absent; exudates absent; reverse shades of dull green (27E4) fading into between pale yellow (4A3) and pale orange (5A3). DG18 25 C, 7 d: Colonies raised at centre, slightly concentrically sulcate; margins low, plane, 327 YILMAZ ET AL. Fig. 83. Morphological characters of Talaromyces varians (CBS 386.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 328 POLYPHASIC entire (<1 mm); mycelia white; sporulation absent to sparse; soluble pigments absent; exudates absent; reverse yellowish white (4A2) with shades of olive brown (4F4). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3–4 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green to dull green (26C3–26D3); soluble pigments absent; exudates absent; reverse dark green. CREA 25 C, 7 d: Acid production absent. Micromorphology: Conidiophores biverticillate and monoverticillate; stipes smooth walled, pigmented, 35–65 × 1.5–3 μm; metulae two to four, divergent, 9–12 × 1.5–2.5 μm; phialides acerose, three to ﬁve per metulae, 10–16.5 × 1.5–2.5 μm; conidia smooth, ellipsoidal to cylindrical, 2.5–4 × 1.5–2 μm. Ascomata not observed. Distinguishing characters: Talaromyces varians characteristically produces a strongly pigmented, smooth walled stipe (Fig. 83). It grows relatively fast at 30 and 37 C. Pitt (1980) considered T. varians synonymous with T. funiculosus, but its colony texture is not funiculose and its pigmented stipe can easily distinguish T. varians from T. funiculosus. Talaromyces verruculosus (Peyronel) Samson et al., Stud. Mycol. 71: 177. 2011. MycoBank MB560678. Fig. 84. ≡ Penicillium verruculosum Peyronel, Germi Atmosf. Fung. Micel.: 22. 1913. TAXONOMY OF TALAROMYCES 8.5–10.5 × 2.5–3.5 μm; conidia echinulate, globose, 3–3.5 × 3–3.5 μm. Extrolites: Altenuene, altenusin, alternariol, berkelic acid, hypomiltin, MC-141, mitorubrin, mitorubrinic acid, penicillide, pestalacin A (only detected in IMI 193912), purpactins, rubiginosin A, vermicillin. Distinguishing characters: Talaromyces verruculosus produces ﬂask-shaped phialides and rough to echinulate, globose conidia (Fig. 84). These characters are also observed in T. aculeatus and T. apiculatus. Talaromyces verruculosus grows slower than T. apiculatus on CYA and MEA at 25 C. Compared to T. aculeatus, T. verruculosus grows faster on CYA at 37 C. Talaromyces viridis (Stolk & G.F. Orr) Arx, Persoonia 13: 2821. 1987. MycoBank MB132097. Fig. 85. ≡ Sagenoma viride Stolk & G.F. Orr, Mycologia 66: 677. 1974. In: Talaromyces section Talaromyces Typus: CBS H-7732 (isotype), CBS H-7733 (isotype), CBS H7734 (isotype), culture ex-type CBS 114.72 = ATCC 22467 = NRRL 5575. ITS barcode: AF285782 (alternative markers: BenA = JX494310; CaM = KF741935; RPB2 = JN121430) In: Talaromyces section Talaromyces Typus: IMI 040039, culture ex-type CBS 388.48 = CBS 136671 = ATCC 10513 = DSM 2263 = IMI 040039 = NRRL 1050 = FRR 1050 = IBT 10891 = IBT 32644. Colony diam, 7 d (mm): CYA 9; CYA 30 C 10; CYA 37 C 10–11; MEA 15; MEA 30 C 16–17; DG18 6–7; CYAS 3; OA 14–15; CREA No growth; YES 9. Colony characters: CYA 25 C, 7 d: Colonies moderately deep, radially sulcate; margins low, narrow, entire; mycelia white and yellow; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (26D5–27D5); soluble pigments absent, exudates absent; reverse greyish yellow to greyish orange (4C4–5C4) at centre, pale elsewhere. MEA 25 C, 7 d: Colonies moderately deep, plane; margins low to somewhat subsurface, narrow, entire; mycelia white and yellow; texture ﬂoccose, with some loosely funiculose areas; sporulation moderately dense, conidia en masse greyish green (26D5–27D5); soluble pigments absent, exudates absent; reverse greyish orange (5C6). YES 25 C, 7 d: Colonies moderately deep, irregularly sulcate, raised at centre; margins low, narrow, entire; mycelia white, yellow and pink; texture ﬂoccose; sporulation absent; soluble pigments absent, exudates absent; reverse greyish orange (5B5) at centre, pale elsewhere. DG18 25 C, 7 d: Colonies consist of white mycelium and produce clear exudate. CREA 25 C, 7 d: Weak acid produced. Colony characters: CYA 25 C, 7 d: Colonies low, slightly sulcate; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse greyish green (1C3) centre fading into yellowish white (4A2). MEA 25 C, 7 d: Colonies slightly raised at centre, sulcate, with a soily smell; margins low, plane, entire (1 mm); mycelia white and yellow; texture ﬂoccose; sporulation sparse, conidia en masse greyish green to dull green (26C4–26D4); soluble pigments absent; exudates absent; reverse brown (6E7) centre fading into reddish golden (6C7). YES 25 C, 7 d: Colonies raised at centre, sunken at centre, sulcate; margins low, plane, entire (<1 mm); mycelia white; texture ﬂoccose; sporulation dense, conidia en masse greenish grey (26B2–26C2); soluble pigments absent; exudates absent; reverse greyish green (1D3 fading into 1C3). DG18 25 C, 7 d: Colonies sunken at centre, sulcate, with a greyish green appearance; margins raised, plane, entire (1 mm); mycelia white; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates clear droplets; reverse greyish green (1C4–1D4). OA 25 C, 7 d: Colonies low, plane, formation of yellow ascomata (at 30 C more abundant); margins low, plane, entire (3–4 mm); mycelia white and yellow; texture velvety; sporulation moderately dense, conidia en masse greyish green (27C5) (at 30 C greyish green to dark green (27E5–27F5)); soluble pigments absent; exudates absent; reverse greyish yellow. CREA 25 C, 7 d: No growth. Micromorphology: Conidiophores biverticillate; stipes smooth walled, (50–)150–300(–400) × 2.5–3 μm; metulae four to eight, divergent, 8.5–12.5 × 2.5–3.5 μm; phialides ﬂask-shaped, tapering into very thin neck, three to ﬁve per metulae, Micromorphology: Conidiophores with solitary phialides; phialides usually solitary, 10–25 × 1.5–2.5 μm; conidia smooth, fusiform to ellipsoidal, 2.5–3.5(–4) × 1.5–2 μm. Ascomata maturing after 1–2 wk of incubation on MEA and OA at 25 C ITS barcode: KF741994 (alternative markers: BenA = KF741928; CaM = KF741974; RPB2 = KM023306) Colony diam, 7 d (mm): CYA 32–35; CYA 30 C 37–38; CYA 37 C 25–26; MEA 35–36; DG18 15–16; CYAS 11–12; OA 38–40; CREA 24–26; YES 36–38. www.studiesinmycology.org 329 YILMAZ ET AL. Fig. 84. Morphological characters of Talaromyces verruculosus (CBS 388.48T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. 330 POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 85. Morphological characters of Talaromyces viridis (CBS 114.72T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and synnemata on MEA after 2 wk incubation. C. Ascomata. D–E. Asci and ascospores. F. Peridial hyphae. G–I. Conidiophores. J. Conidia. K. Ascospores. Scale bars: B = 500 μm; C = 50 μm; D = 10 μm, applies to E–K. www.studiesinmycology.org 331 YILMAZ ET AL. and abundantly 30 C, dark green, globose to subglobose, 200–400 μm. Peridial hyphae brownish green, thick walled, richly branched 8–35 × 1–1.5 μm, asci 7.5–8 × 6–8.5 μm, ascospores ellipsoidal, ﬁnely ridges, 3.5–4.5 × 2.5–3 μm. Extrolites: Talaromyces viridis (IBT 31150 = CBS 114.72) produces a purpactin and several extrolites only found in Talaromyces. Distinguishing characters: Talaromyces viridis grows restrictedly on general media and does not grow on CREA. It produces dark green ascomata, which are covered with rough-walled, brownish green peridial hyphae (Fig. 85). Ascospores of T. viridis are ﬁnely striate and ellipsoidal. It produces reduced conidiophores consisting of solitary phialides. These features can easily distinguish T. viridis from other Talaromyces species. Talaromyces viridulus Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011. MycoBank MB560679. Fig. 86. ≡ Geosmithia viridis Pitt & A.D. Hocking, Mycologia 77: 822. 1985. = Penicillium viride (Pitt & A.D. Hocking) Frisvad, Samson & 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, Persoonia 13: 2821. 1987). ITS barcode: JN899314 (alternative markers: BenA = JX091385; CaM = KF741943; RPB2 = JF417422) Micromorphology: Conidiophores biverticillate; stipes smooth walled, 25–80 × 2.5–3 μm; metulae three to six, divergent, 9–15 × 2–3 μm; phialides acerose, two to six per metula, 8–12 × 1.5–3 μm; conidia smooth, cylindrical, rod-shaped, 3.5–6 × 1–2 μm. Ascomata not observed. Distinguishing characters: Talaromyces viridulus grows restrictedly on general media, except on MEA, OA and YES. It produces deep colonies with a white, pastel yellow and pastel red appearance. Its most striking character is its smooth, cylindrical, rod-shaped conidia (Fig. 86). Similar conidia are observed in T. bacillisporus, but T. bacillisporus produces dark green colony reverses on CYA at 30 C, and globose, spiny ascospores. Talaromyces wortmannii (Klöcker) C.R. Benj. Mycologia 47: 683. 1955. MycoBank MB344294. Fig. 87. ≡ Penicillium wortmannii Klöcker, Compt. Rend. Lab. Carlsberg, Physiol. 6: 100. 1903. ≡ Penicillium kloeckeri Pitt, The genus Penicillium: 491. 1980. = Talaromyces sublevisporus (Yaguchi & Udagawa) Samson, Yilmaz & Frisvad, Stud. Mycol. 71: 177. 2011 ≡ Talaromyces wortmannii var. sublevisporus Yaguchi & Udagawa, Mycoscience 35: 63. 1994. = Talaromyces variabilis (Sopp) Samson et al., Stud. Mycol. 71: 177. 2011 ≡ Penicillium variabile Sopp, Skr. Vidensk.-Selsk. Christiania, Math.-Naturvidensk. Kl. 11: 169. 1912. = Penicillium concavorugulosum S. Abe, J. Gen. Appl. Microbiol., 2: 127. 1956 (nom. inval., Art. 36). In: Talaromyces section Islandici Typus: FRR 1863, culture ex-type CBS 252.87 = FRR 1863 = IMI 288716. Typus: IMI 040047, culture ex-type CBS 391.48 = ATCC 10517 = IMI 040047 = NRRL 1017 = IBT 4838. In: Talaromyces section Talaromyces Colony diam, 7 d (mm): CYA 12–15; CYA 30 C 16–18; CYA 37 C 6–7; MEA 37–38; MEA 30 C 39–41; DG18 16–17; CYAS No growth; OA 35–38; CREA 2–4; YES 23–25. Colony characters: CYA 25 C, 7 d: Colonies raised at centre, slightly sulcate, white sterile appearance; margins low, plane, entire (1 mm); mycelia white; sporulation absent; soluble pigments absent; exudates absent; reverse brownish red (9C7) centre fading into pale yellow (3A3). MEA 25 C, 7 d: Colonies deep, plane, sterile white, pastel yellow and pastel red appearance; margins low, plane, entire (2–3 mm); mycelia white, pastel yellow and pastel red; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse brownish orange to light brown (5C6–5D6). YES 25 C, 7 d: Colonies slightly raised, deep, plane, sterile white and red appearance; margins low, plane, entire (3–4 mm); mycelia white, pastel red and dull red; sporulation absent; soluble pigments absent; exudates absent; reverse red (9B8) fading into orange red (8B8) and pale red (8A3). DG18 25 C, 7 d: Colonies low, slightly sulcate; margins low, plane, entire (1–2 mm); mycelia white; texture ﬂoccose; sporulation moderately dense, conidia en masse greyish green (25C4); soluble pigments absent; exudates absent; reverse reddish brown (9E8) centre fading into pastel red (9A5). OA 25 C, 7 d: Colonies low, plane; margins low, plane, entire (3 –4 mm); mycelia white; texture velvety and in the centre ﬂoccose; sporulation moderately dense to dense, conidia en masse dull green (27D4); soluble pigments absent; exudates absent; reverse beige. CREA 25 C, 7 d: Acid production absent. 332 ITS barcode: KF984829 (alternative markers: BenA = KF984648; CaM = KF984756; RPB2 = KF984977) Colony diam, 7 d (mm): CYA 18–28; CYA 30 C 18–26; CYA 37 C Generally no growth, in some isolates colonies up to 7; MEA 15–25; MEA 30 C 17–27; DG18 13–18; CYAS 10–12; OA 20–25; CREA 5–15; YES 20–30. Colony characters: CYA 25 C, 7 d: Colonies 18–28 mm, slightly raised at centre, sulcate; margins narrow (up to 2 mm), low, entire, plane; mycelium generally only white, in some isolates yellow dominant (CBS 100258; CBS 293.53, CBS 319.63, CBS 391.48T, CBS 137376); texture velvety; sporulation absent (CBS 391.48T, CBS 319.63, CBS 293.53), moderately dense to dense; conidia en masse greyish green to dull green (25D4–25E4); exudates absent; soluble pigment absent; reverse in some isolates centre brown (6E6–6E7) fading into in some isolates reddish yellow (4A6), in some isolates greyish orange to orange (6B6–6B7) (NRRL 2125; CBS 293.53), in some isolates centre yellowish brown (5F5) fading into in some isolates olive (3D5–3E5) and in some isolates greyish yellow (3B4–3C4) (CBS 100258; DTO 67-G1; DTO 189-D9; CBS 137384; DTO 127-I4) in some isolates yellow (3A6) with dark blonde (5D4) dots in centre (CBS 137376, CBS 319.63). MEA 25 C, 7 d: Colonies 15–25 mm, in some isolates low, in some isolates slightly raised at centre, sulcate; margins very narrow (up to 1 mm), low, entire, plane; mycelium generally only white, in some isolates yellow dominant (CBS 100258; CBS 319.63, CBS 391.48T, CBS 137376); texture velvety; sporulation absent (CBS 391.48T, CBS 293.53), sparse (CBS 137376) to dense; conidia en masse POLYPHASIC TAXONOMY OF TALAROMYCES Fig. 86. Morphological characters of Talaromyces viridulus (CBS 252.87T). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B–F. Conidiophores. G. Conidia. Scale bar: B = 10 μm, applies to C–G. www.studiesinmycology.org 333 YILMAZ ET AL. Fig. 87. Morphological characters of Talaromyces wortmannii (CBS 319.63). A. Colonies from left to right (top row) CYA, MEA, DG18 and OA; (bottom row) CYA reverse, MEA reverse, YES and CREA. B. Colony texture and ascomata on MEA after 2 wk incubation. C. Asci and ascopsores. D–G. Conidiophores. H. Conidia. Scale bar: C = 10 μm, applies to D–H. 334 POLYPHASIC greyish green to dull green (25D4–25E4); in some isolates yellow ascomata production with the longer incubation (up to 9 d) (CBS 319.63, CBS 137376); exudates absent (except CBS 319.63, CBS 100258, yellow droplets); soluble pigment absent; reverse in some isolates centre brown (6E5) fading into brownish orange to brown (6C8–6D6), in some isolates lack of reverse brownish yellow (5C7–5C8). YES 25 C, 7 d: Colonies 20–30 mm, raised at centre, some isolates crateriforme and some isolates sulcate; margins narrow (up to 2 mm), low, entire, plane; mycelium generally only white, in some isolates yellow dominant (CBS 319.63, CBS 391.48T, CBS 137376); texture velvety and crusty; sporulation absent (CBS 293.53, CBS 391.48T) to dense; conidia en masse greyish green to dull green (25D4–25E4); exudates absent; soluble pigment absent; reverse in some isolates centre light yellow to greyish orange (4A5–5B6) fading into greyish yellow to olive (2C4–2D4) (DTO 127-I4, DTO 189-D9, DTO 67-G1, CBS 137384, NRRL 2125), in some isolates olive (2F4) in the centre fading into greyish yellow to olive (2C4–2D4) (CBS 100258, CBS 137394), in some isolates centre orange to golden yellow (5A6–5B6) fading into yellow (2A6). DG18 25 C, 7 d: Colonies 13–18 mm, raised at centre, sulcate; margins very narrow (up to 1 mm), low, entire, plane; mycelium generally only white, in some isolates yellow dominant (CBS 293.53; CBS 319.63, CBS 137376); texture velvety; sporulation absent (CBS 391.48T, CBS 293.53) to dense; conidia en masse greyish green to dull green (25D4–25E4); exudates absent; soluble pigment absent (except CBS 100258, CBS 137376, CBS 319.63 yellow); reverse in some isolates centre orange (5B8) fading into greyish green (26B3–26C3) (NRRL 2125, CBS 137384, DTO 127-I4), in some isolates centre pale yellow (1A3) centre fading into greyish yellow (1B3) (DTO 189-D9, DTO 67G1), in some isolates yellowish white to yellow to vivid yellow (2A2–2A6–2A8) (CBS 293.53, CBS 319.63, CBS 137376, CBS 100258), in some isolates olive (3E4) centre fading into dull yellow (3B3) (CBS 137394) and in some isolates pale yellow (3A3) (CBS 391.48T). OA 25 C, 7 d: Colonies 20–25 mm, low, plane; margins wide (2–4 mm), low, entire, plane; mycelium generally only white and pale yellow, in some isolates bright yellow dominant (CBS 319.63, CBS 391.48T, CBS 137376); texture velvety in some isolates centre ﬂoccose; sporulation sparse (CBS 137376, CBS 391.48T, DTO 108-A4, CBS 293.53) to dense; conidia en masse greyish green to dull green (25D4–25E4); in some isolates yellow ascomata production with the longer incubation (up to 9 d) (CBS 319.63, CBS 137376; CBS 391.48T; CBS 293.53); exudates in some isolates absent and in some isolates yellow and clear droplets; soluble pigment absent; reverse in some isolates orange brown centre fading into orange (NRRL 2125, CBS 391.48T, CBS 137376, CBS 319.63), in some very pale orange pink (5A3, CBS 293.53) and rest pale green to beige. CREA, 25 C, 7 d: Colonies 5–15 mm, in some isolates acid production absent and in some isolates moderate production (NRRL 2125, CBS 137384, CBS 319.63). Micromorphology: Conidiophores biverticillate, sometimes with additional branches, smooth walled stipes, arising from aerial hyphae and overgrowing the ascomata, stipes 100–400 × 2.5–4 μm, extra branches 15–30 μm; metulae three to six, divergent, 7–15 × 12.5–4.0 μm; phialides three to six, acerose to almost ﬂask-shaped, 7–15 × 2–3.5 μm; conidia, smooth walled, ellipsoidal, 2.5–6 × 1.5–3.5 μm. Ascomata bright yellow, to orange globose, ripening within 1 to 2 wk, covering consisting of a few layers of well-developed networks of yellow www.studiesinmycology.org TAXONOMY OF TALAROMYCES hyphae; asci broadly globose to subglobose, 8–14 × 7–11 μm; ascospores, broadly ellipsoidal, 3.5–6.0 × 2.5–4 μm, thick walled, spiny, to smooth (DTO 176-I6). Extrolites: Talaromyces wortmannii is reported to produce wortmannilactones E-H (Dong et al. 2009), closely related to the prugosins/prugosenes/ukulactones produced by T. allahabadense and T. rugulosus. Furthermore it produces rugulovasine A and B (Abe et al. 1969, Yamatodani et al. 1970, Zhelifonova et al. 2006, 2010, Antipova et al. 2008), chanoclavine-I (Abe et al. 1969); wortmannilactones A-D (Dong et al. 2006); mitorubrinol acetate (Suzuki et al. 1999); β-caryophyllene (Yamagiwa et al. 2011); Talaromin A and B (Bara et al. 2013); atrovirin B1 and B2 (Gill & Morgan 2004); We detected rugulosin, skyrin, rugulovasine A, wortmannilactones E, F, G, H, mitorubrin, mitorubrinol, mitorubrinol acetate and other mitorubrins, and a tetracyclic compound. Wortmannin, virone and wortmannolone have been reported to be produced by T. wortmannii (Brian et al. 1957, Blight & Grove 1986), but we did not detect these compounds in any T. wortmannii sensu stricto isolates. Distinguishing characters: Talaromyces wortmannii is characterised by velvety colonies that sporulate densely. Many strains also produce ascomata with ascospores (Fig. 87). Acid production and growth at 37 C varies among strains of T. wortmannii. Talaromyces wortmannii was previously believed to be closely related to T. rugulosus. However, T. wortmannii grows faster on most media and some strains produce yellow to orange ascomata (CBS 319.63, CBS 137376, CBS 293.53). Conidiophores are also often much wider. Ascomata and ascospores of T. wortmannii closely resemble those of T. tratensis. However, T. tratensis produces dense ﬂoccose colonies and conidiophores are more appressed than T. wortmannii. Notes: Samson et al. (2011) accepted T. variabilis (CBS 385.48) and T. sublevisporus (CBS 137376) as valid species. However, based on ITS, BenA, CaM and RPB2 phylogenies, P. concavorugulosum, T. sublevisporus (CBS 137376) and T. variabilis (CBS 385.48) form a coherent clade with T. wortmannii (Yilmaz et al., unpubl.). Talaromyces yelensis Visagie et al., Stud. Mycol. 78: 134. 2014. MycoBank MB809189. In: Talaromyces section Islandici Typus: CBS H-21799, culture ex-type CBS 138209 = DTO 268E5. ITS barcode: KJ775717 (alternative markers: BenA = KJ775210) Colony diam, 7 d (mm): Fide Visagie et al. (2014) CYA 20–22; CYA 30 C 25–26; CYA 37 C 14–16; MEA 15–16; MEA 30 C 21–22; DG18 16–17; CYAS 13–14; OA 18–20; CREA 9–10; YES 20–21. Colony characters: Fide Visagie et al. (2014) CYA 25 C, 7 d: Colonies moderately deep; margins low, narrow, entire; mycelia white to yellowish to orange; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates clear and sticky; reverse yellowish white (2A2) to light yellow (3A5) to brown (5F6). MEA 335 YILMAZ ET AL. 25 C, 7 d: Colonies very deep, plane; margins deep, narrow, entire; mycelia white to yellow to orange; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates yellow; reverse brownish yellow to yellowish brown to brown (5C8–E8). YES 25 C, 7 d: Colonies very deep, plane; margins low, narrow, entire; mycelia white to yellow to orange; texture ﬂoccose; sporulation absent; soluble pigments absent; exudates absent; reverse yellowish white (4A2) to greyish orange (5B5). DG18 25 C, 7 d: Colonies deep, plane; margins low, narrow, entire; mycelia white to yellow; texture ﬂoccose; sporulation sparse, conidia en masse greyish green (26C3); soluble pigments absent; exudates yellow and sticky; reverse yellowish white to yellow (3A2–3A6). OA 25 C, 7 d: Colonies moderately deep, plane; margins low, narrow, entire; mycelia white to yellow; texture ﬂoccose; sporulation moderately dense, conidia en masse dark green (26F6); soluble pigments absent; exudates clear and sticky. CREA 25 C, 7 d: Acid production absent. Micromorphology: Fide Visagie et al. (2014) Conidiophores biverticillate, subterminal branches sometimes present; stipes smooth walled, 60–190 × 2.5–3.5 μm; branches up to 30 μm long; metulae appressed, 8–11 × 2.5–3.5 μm; phialides ﬂaskshaped, ending in a ﬁne apical pore, 8–10 × 2.5–3 μm; conidia rough, subglobose to broadly ellipsoidal, 2.5–3.5 × 2.5–3 μm. Distinguishing characters: Talaromyces yelensis produces very dense, deep and yellow colonies on general media. Colonies closely resemble those produced by T. tratensis. However, T. tratensis typically produces ascomata with roughened ascospores and conidiophores that produce ellipsoidal smooth walled conidia, in contrast to T. yelensis that lacks ascomata and produces subglobose to broadly ellipsoid conidia that have rough walls. For an illustration of the species, readers are referred to Visagie et al. (2014). DOUBTFUL SPECIES LIST Penicillium guizhouanum H.Z. Kong, Mycosystema 19: 1. 2000. [MB464324]. — Herb.: HMAS 74529. Ex-type: AS 3.5215. Note: Sequence data from Visagie et al. (2014) show that this species is a synonym of T. funiculosus, which we follow here. Penicillium krugeri C. Ramirez, Mycopathologia 110: 23. 1990. [MB126186]. — Herb.: TISTR3377. Ex-type: No culture data available. Note: Culture was described from South Africa, but no type material is available. Penicillium mirabile Beliakova & Milko, Mikol. Fitopatol. 6: 145. 1972. [MB319286]. — Herb.: VKM F-1328. Ex-type: CBS 624.72 = CCRC 31665 = FRR 1959 = IMI 167383 = LCP 72.2193 = MUCL 31206 = VKM F1328. Note: The ex-type culture for this species (obtained from CBS and NRRL) is contaminated and additional work is needed to determine its exact taxonomic position. Penicillium oblatum Pitt & A. D. Hocking, Mycologia 77: 819. 1985. [MB104603]. — Herb.: FRR 2234. Ex-type: CBS 258.87 = FRR 2234. Note: Samson et al. (2011) published sequence data showing a close relationship with T. dendriticus. In our opinion CBS 258.87 is contaminated and not representing the ex-type culture. Penicillium pachmariensis A.K. Gupta & S. Chauhan, Indian For. 122: 1182. 1996. [MB459628]. — Herb.: IMI 278556. Ex-type: IMI 278556. Note: According to the IMI, the species is synonymous with Talaromyces purpurogenus. However, we were not able to examine this strain. Penicillium pascuum (Pitt & A. D. Hocking) Frisvad et al., Persoonia 14: 229. 1990 ≡ Paecilomyces pascuus Pitt & A. D. Hocking, Mycologia 77: 822. 1985. [MB492634]. — Herb.: Unknown. Ex-type: CBS 253.87 = FRR 1925. Note: Samson et al. (2011) published sequence data showing a close relationship with T. dendriticus. In our opinion CBS 253.87 is contaminated and not representing the ex-type culture. 336 Penicillium resedanum McLennan & Ducker, Aust. J. Bot. 2: 360. 1954. [MB302422]. — Herb.: IMI 062877. Ex-type: CBS 181.71 = ATCC 22356 = FRR 578 = IMI 062877 = NRRL 578. Note: ITS sequence data (AF033398) show that this species belongs in Talaromyces, and is not related to any section we accept here. Unfortunately CBS 181.71 is not viable and the species will be re-examined in a future study. Talaromyces lagunensis Udagawa, Uchiy. & Kamiya, Mycoscience 35: 403. 1994 ≡ Penicillium lagunense Udagawa, Uchiy. & Kamiya, Mycoscience 35: 403. 1994. [MB414250]. — Herb.: CBM BF-49341. Ex-type: Unknown. Note: We have not examined authentic material for this species and did not treat species in this study. Talaromyces malagensis (Thüm.) Stalpers & Samson, Stud. Mycol. 24: 69. 1984. [MB107328]. — Herb.: Unknown. Ex-type: Unknown. Note: No type culture information is available and we can not determine its taxonomic position. Talaromyces retardatus Udagawa, Kamiya & Kaori Osada, Trans. Mycol. Soc. Japan 34: 9. 1993 ≡ Penicillium retardatum Udagawa, Kamiya & Kaori Osada, Trans. Mycol. Soc. Japan 34: 9. 1993. [MB360062]. — Herb.: CBM BF-24811. Ex-type: Unknown. Note: We have not examined authentic material for this species and did not treat it in this study. Talaromyces versatilis P.F. Cannon, Bridge & Buddie nom. inval., Art. 38.1(a). Note: Talaromyces versatilis was never published. However, Delmas et al. (2014) and Lafond et al. (2014) have incorrectly used this name, based on IMI 378536, in their publication as a “basionym” of Penicillium funiculosum (≡ T. funiculosus). We did not examine this strain, but based on the BenA sequence for IMI 378536 (KC992272), it is distinct from all accepted Talaromyces species and is closely related to T. angelicus. Further studies are needed for this taxon. Paecilomyces tenuis Y.F. Yan & Z.Q. Liang, Mycotaxon 102: 54. 2007. [MB510919]. — Herb.: GZUIFR-C43-1. Ex-type: Not available. Note: This species has an ITS sequence (EU004812) identical to T. assiutensis and T. trachyspermus. No strain is available for examination and thus its correct classiﬁcation cannot be determined. EXCLUDED SPECIES LIST Penicillium emersonii Stolk, Antonie van Leeuwenhoek 31: 262. 1965 ≡ Talaromyces emersonii Stolk, Antonie van Leeuwenhoek 31: 262. 1965 ≡ Rasamsonia emersonii (Stolk) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 417. 2011. [MB335725]. — Herb.: CBS H-7817. Ex-type: CBS 393.64 = ATCC 16479 = CECT 2607 = IFO 31232 = IMI 116815 = IMI 116815ii. Penicillium lignorum Stolk, Antonie van Leeuwenhoek 35: 264. 1969. [MB335743]. — Herb.: CBS H-7505. Ex-type: CBS 709.68 = ATCC 22051 = FRR 804 = IMI 151899 = UPSC 3184. Note: A preliminary phylogenetic analysis indicates that this species does not belong to Talaromyces and might represent a new genus (Samson et al. 2011). Penicillium sabulosum Pitt & A.D. Hocking in Mycologia 77: 818. 1985 ≡ Talaromyces sabulosus (Pitt & A.D. Hocking) Samson, Yilmaz & Frisvad, Stud. Mycol. 70: 177. 2011. [MB104604]. — Herb.: FRR 2743. Extype: CBS 261.87 = FRR 2743. Note: This species was incorrectly transferred to Talaromyces by Samson et al. (2011) and belongs in Penicillium section Exilicaulis. Penicillium striatum Raper & Fennell, Mycologia 40: 521. 1948 ≡ Talaromyces striatus (Raper & Fennell) C. R. Benj., Mycologia 47: 682. 1955 ≡ Hamigera striata (Raper & Fennell) Stolk & Samson, Persoonia 6: 347. 1971. [MB289109]. — Herb.: IMI 039741. Ex-type: CBS 377.48 = ATCC 10501 = IFO 6106 = IMI 039741 = NRRL 717 = QM 1857 = VKM F-2044. Talaromyces avellaneus (Thom & Turesson) C.R. Benj. Mycologia 47: 682. 1955 ≡ Penicillium avellaneum Thom & Turesson, Mycologia 7: 284. 1915 ≡ Hamigera avellanea (Thom & Turesson) Stolk & Samson, Stud. Mycol. 6: 345. 1971. [MB306712]. — Herb.: IMI 040230. Ex-type: CBS 295.48 = ATCC 10414 = CECT 2265 = DSM 2208 = IMI 040230 = NRRL 1938. Talaromyces brevicompactus Kong, Mycosystema 18: 9. 1999 ≡ Merimbla brevicompacta Kong, Mycosystema 18: 9. 1999. [MB460109]. — Herb.: HMAS 62770. Ex-type: CBS 102661 = AS 3.4676. Note: Molecular data show that this species belongs to the Hamigera clade and not Talaromyces (Houbraken & Samson 2011). Talaromyces byssochlamydoides Stolk & Samson, Stud. Mycol. 2: 45. 1972 ≡ Paecilomyces byssochlamydoides Stolk & Samson, Stud. Mycol. 2: 45. 1972 ≡ Rasamsonia byssochlamydoides (Stolk & Samson) Houbraken POLYPHASIC & Frisvad, Antonie van Leeuwenhoek 101: 415. 2011. [MB324415]. — Herb.: CBS H-7816. Ex-type: CBS 413.71 = IMI 178524 = JCM 12813 = NRRL 3658. Talaromyces cejpii Milko in Novosti Sist. Nizs: Rast.: 208. 1964 ≡ Dichotomomyces cejpii (Milko) D.B. Scott in Trans. Brit. Mycol. Soc. 55: 313. 1970 ≡ Aspergillus cejpii (Milko) Samson et al., Stud. Mycol. 78: 141. MycoBank MB809582.— Herb.: CBS H-7011. Ex-type: CBS 157.66. Note: Houbraken & Samson (2011) showed that this species is phylogenetically in Aspergillus and is transferred in Samson et al. (2014). Talaromyces eburneus Yaguchi et al., Mycoscience 35: 249. 1994 ≡ Geosmithia eburnea Yaguchi et al. ≡ Rasamsonia eburnea (Yaguchi et al.) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 416. 2011. [MB362928]. — Herb.: CBM PF-1151. Ex-type: CBS 100538 = IBT 17519. Talaromyces emersonii Stolk, Antonie van Leeuwenhoek 31: 262. 1965 ≡ Penicillium emersonii Stolk, Antonie van Leeuwenhoek 31: 262. 1965 ≡ Rasamsonia emersonii (Stolk) Houbraken & Frisvad, Antonie van Leeuwenhoek 101: 417. 2011. [MB339920]. — Herb.: CBS H-7817. Ex-type: CBS 393.64 = ATCC 16479 = CECT 2607 = IFO 31232 = IMI 116815 = IMI 116815ii. Talaromyces leycettanus H.C. Evans & Stolk, Trans. Brit. Mycol. Soc. 56: 45. 1971 ≡ Penicillium leycettanus H.C. Evans & Stolk ≡ Paecilomyces leycettanus (H.C. Evans & Stolk) Stolk et al., Persoonia 6: 342. 1971. [MB324419]. — Herb.: CBS 398.68. Ex-type: CBS 398.68 = ATCC 22469 = IMI 178525 = JCM 12814. Note: Houbraken & Samson (2011) showed that this species is phylogenetically unrelated to Talaromyces and belongs in the Hamigera clade. Talaromyces luteus (Zukal) C.R. Benj., Mycologia 47: 681. 1955 ≡ Penicillium luteum Zukal, Sitzungsber Kaiserl. Akad. Wiss. Math-Naturwiss. C1., Abt. 1, 98: 561. 1890. [MB306716]. — Herb.: IMI 89305. Ex-type: CBS 348.51 = CECT 2950 = IFO 31753 = IMI 089305 = LSHB BB228. Note: Although the morphological characters of this species resemble species of Talaromyces, Houbraken et al. (2014) showed that this species is basal to Thermomyces and probably represents a distinct genus. Talaromyces ocotl Bills & Heredia, Mycologia 90: 533. 1998 ≡ Sagenomella ocotl (Bills & Heredia) Samson et al., Stud. Mycol. 71: 179. 2011. [MB467796]. — Herb.: GB 6125. Ex-type: CBS 102855. 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. [MB361677]. — Herb.: CBM SUM3030. Ex-type: CBS 101075 = ATCC 90900 = FRR 5219 = JCM 12815. Talaromyces striatus (Raper & Fennell) C.R. Benj., Mycologia 47: 682. 1955 ≡ Penicillium striatum Raper & Fennell, Mycologia 40: 521. 1948 ≡ Byssochlamys striata (Raper & Fennell) Arx, Mycotaxon 26: 120. 1986 ≡ Hamigera striata (Raper & Fennell) Stolk & Samson, Persoonia 6: 347. 1971. [MB306723]. — Herb.: IMI 039741. Ex-type: CBS 377.48 = ATCC 10501 = IFO 6106 = IMI 039741 = NRRL 717 = QM 1857 = VKM F-2044. Talaromyces thermophilus Stolk, Antonie van Leeuwenhoek 31: 268. 1965 ≡ Thermomyces dupontii (Griffon & Maublanc) Houbraken & Samson, Adv. Appl. Microbiol. 86: 218. 2014. [MB805186]. — Herb.: CBS 236.58. Extype: CBS 236.58 = ATCC 10518 = ATCC 16461 = ATCC 52514 = FRR 2155 = IFO 31798 = IMI 048593 = NRRL 2155 = QM 1851 = VKM F-2043. 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