TAXONOMIC DESCRIPTION Polburee et al., Int J Syst Evol Microbiol 2017;67:2377–2382 DOI 10.1099/ijsem.0.001961

ist of ublications for 17 hanok AAAACA todaengensis f.a., sp. nov., a yeast isolated from a 1. Phitsuwan, P., Permsriburasuk, C., Baramee, S., Teeravivattanakit, T., and peat swamp forest in Thailand Ratanakhanokchai, K. (2017). Structural analysis of alkaline pretreated rice straw for 1,5 1 2 3 1,4, ethanol production. International Journal of Polymer Science 17: Article ID Pirapan Polburee, Noppon Lertwattanasakul, Pitayakon Limtong, Marizeth Groenewald and Savitree Limtong * 4876969, 9 pages. https:doi.org10.115520174876969. Abstract

2. Teeravivattanakit, T., Baramee, B., Phitsuwan, P., Sornyotha, S., Waeonukul, R., T Pason, P., Tachaapaikoon, C., Poomputsa, K., Kosugi, A., Sakka, K., and Strain DMKU-PS11(1) was isolated from peat in a swamp forest in Thailand. DNA sequence analysis showed that it belonged to a novel species that was most closely related to Nakazawaea laoshanensis. However, it differed from the type strain of Ratanakhanokchai, K. (2017). Chemical pretreatment-independent saccharifications N. laoshanensis (NRRL Y-63634T) by 2.3 % nucleotide substitutions in the D1/D2 region of the large subunit (LSU) rRNA gene, of xylan and cellulose of rice straw by bacterial weak lignin-binding xylanolytic and 1.0 % nucleotide substitutions in the small subunit (SSU) rRNA gene and 8.0 % nucleotide substitutions in the internal cellulolytic enzymes. Applied and Environmental Microbiology : no.22, transcribed spacer (ITS) region. The phylogenetic analyses based on the combined sequences of the SSU and the D1/D2 region e01522-17. and that of the SSU sequences alone confirmed the placement of the novel species in the Nakazawaea clade and its close affinity with N. laoshanensis. Hence, the species Nakazawaea todaengensis f.a., sp. nov. is proposed. The type strain is DMKU- T T T 3. Sonklin, C., aohakunjit, N., Kerdchoechuen, ., and Ratanakhanokchai, K. (2018). PS11(1) (=CBS 14555 =TBRC 6559 ). The MycoBank number for Nakazawaea todaengensis f.a., sp. nov. is MB 819513 olatile flavour compounds, sensory characteristics and antioxidant activities of mungbean meal protein hydrolysed by bromelain. Journal of Food Science and Technology : 265-277. DI 10.1007s13197-017-2935-7. Nakazawaea is an ascomycetous yeast in the order N. wyomingensis [4]. One of these new combinations, N. . It was first proposed by Yamada et al. ernobii, which was suggested to be conspecific with N. hol- [1] to transfer Pichia holstii to a new genus, Nakazawaea, stii due to similar sequences of the D1/D2 region [5] showed 4. Apiwatanapiwat, W., aithanomsat, P., Thanapase, W., Ratanakhanokchai, K., and with N. holstii as the type species. N. holstii could be sepa- notable differences in the EF-1a, RPB1 and RPB2 sequences Kosugi, A. (2018). ylan supplement improves 1,3-propanediol fermentation by rated from the other hat-shaped, ascospore-forming and [4]. However, Candida ernobii was listed as a synonym of Clostridium butyricum. Journal of Bioscience and Bioengineering 1: 662-668. nitrate-assimilating species of the genus Pichia based on the N. holstii in the latest version of The Yeasts, a Taxonomic partial sequences of the D1/D2 region of the large subunit Study [6]. Since then, one additional anamorphic species, N. (LSU) and the small subunit (SSU) rRNA genes. The pro- siamensis, which was isolated from the external surface of a 5. Aikawa, S., Baramee, S., Sermsathanaswadi, J., Thianheng, P., Tachaapaikoon, C., posal of this genus was later supported by phylogenetic sugarcane leaf in Thailand, was proposed by Kaewwichian Shikata, A., Waeonukul, R., Pason, P., Ratanakhanokchai, K., and Kosugi, A. (2018). analysis of the sequences of multiple protein-coding genes and Limtong [7]. Characterization and high-quality draft genome sequence of Herbivorax saccincola including actin (ACT1), the RNA polymerase largest sub- A tropical peat swamp forest is a unique ecosystem that is unit (RPB1) and second largest subunit (RPB2), the second A7, an anaerobic, alkaliphilic, thermophilic, cellulolytic, and xylanolytic bacterium. always flooded. Recent research showed the significance of Systematic and Applied Microbiology 41: 261-269. subunit of the mitochondrial cytochrome oxidase (COX2) and the D1/D2 region of the LSU rRNA gene [2]. According this ecosystem as a global carbon store for soil organic mat- to the new International Code of Nomenclature for algae, ter, but its value for maintaining biodiversity remains poorly fungi, and plants, it is possible to assign the teleomorphic understood [8]. Only a few yeasts isolated from peat lands and related anamorphic species to the same genus [3]. have been reported, of which most were obtained from peat – Therefore, 10 anamorphic species of the genus Candida in lands of Canada and Russia [9 12]. Recently, an investiga- the Nakazawaea clade were transferred to the genus Naka- tion on yeast diversity in peat of a swamp forest in the zawaea based on the phylogenetic analysis of the sequences southern part of Thailand was carried out by using an of the D1/D2 region of the LSU rRNA, the SSU rRNA, enrichment isolation technique [13]. In this investigation translation elongation factor-1a (EF-1a), and the RPB1 five species in the phylum , namely Cyberlind- and RPB2 gene regions. They were Nakazawaea anatomiae, nera subsufficiens, Debaryomyces fabryi, Meyerozyma guil- N. ernobii, N. ishiwadae, N. laoshanensis, N. molendini-olei, liermondii, Saturnispora diversa and Schwanniomyces N. peltata, N. pomicola, N. populi, N. wickerhamii and polymorphus var. africanus, and five species in the phylum

Author affiliations: 1Department of Microbiology, Faculty of Science, Kasetsart University, Bangkok, Thailand; 2Land Development Department, Ministry of Agriculture and Cooperatives, Bangkok, Thailand; 3Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; 4Royal Society of Thailand, Bangkok, Thailand; 5Rattanakosin College for Sustainable Energy and Environment (RCSEE), Rajamangala University of Technology Rattanakosin, Nakhon Pathom, Thailand. *Correspondence: Savitree Limtong, [email protected] Keywords: Nakazawaea todaengensis sp. nov.; ascomycetous yeast; new species; peat; swamp forest; Thailand. Abbreviations: LSU, large subunit; SSU, small subunit. The GenBank/EMBL/DDBJ accession numbers of the SSU, the ITS region and the D1/D2 region of the LSU rRNA gene sequences of strain DMKU- PS11(1)T are LC171727, LC171728 and LC171729, respectively. The accession number for the SSU rRNA gene sequence of Nakazawaea siamensis is LC171726. The Mycobank number for Nakazawaea todaengensis f.a., sp. nov. is MB 819513.

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Basidiomycota, namely Piskurozyma taiwanensis, Cutaneo- PS11(1)T, which was isolated from a peat sample collected trichosporon mucoides, Papiliotrema flavescens, Papiliotrema from the rhizosphere of Calophyllum teysmannii var. inophyl-   laurentii and Rhodotorula mucilaginosa were reported. loide at the sampling site (6 7¢ 52.3† N 101 58¢ 8.4† E) in the Su-ngai Padi district on 14 August 2015, was found to repre- In the current study, the yeast diversity in peat from a swamp sent a novel species of the genus Nakazawaea. forest in the Princess Sirindhorn Wildlife Sanctuary (Pru To Daeng Wildlife Sanctuary), that includes areas of four districts Genomic DNA extraction of strain DMKU-PS11(1)T was intheNarathiwatprovinceinthesouthernpartofThailand performed using the method described by Limtong et al. viz. Mueang, Tak Bai, Su-ngai Kolok and Su-ngai Padi, was [14]. The SSU rRNA gene, the ITS region and the D1/D2 investigated. During this investigation, a novel species of the region of the LSU rRNA gene were amplified using the pri- genus Nakazawaea was found. Yeasts were isolated from 12 mers SSU1f, SSU4r, SSU3f and SSU2r [15], ITS1 and ITS4 peat samples by using a dilution plate method. Peat (2 g) was [16], and NL1 and NL4 [5], respectively. The PCR products added to 50 ml 0.85 % saline solution in a 250 ml Erlenmeyer were purified by using the HiYield Gel/PCR DNA Frag- flask and mixed well by shaking on a rotary shaker at ments Extraction kit (RBC Bioscience) according to the  150 r.p.m. at room temperature (30±2 C) for 1 h. The solution manufacturer’s instructions. The purified products were (0.1 ml) was then spread on yeast malt extract (YM) agar sequenced by Macrogen (Seoul, Korea) using the PCR (0.3 % yeast extract, 0.3 % malt extract, 0.5 % peptone, 1 % glu- amplification primers. The sequences were edited, assem- cose and 2 % agar) supplemented with 0.025 % sodium propi- bled, concatenated and aligned with the MEGA software ver-  onate and 0.02 % chloramphenicol and incubated at 25 C sion 7 [17]. In the present study, the SSU rRNA gene until yeast colonies appeared. Yeast colonies of different mor- sequence of N. siamensis [7] was also determined (Gen- phologies were picked and purified by cross streaking on YM Bank/EMBL/DDBJ accession number LC171726) and used agar. Purified yeast cultures were preserved in YM broth sup- in the phylogenetic analyses together with sequences  plemented with 10 % (v/v) glycerol at À80 C. Fifty yeast obtained from GenBank (Fig. 1) for the additional species. stains were obtained. Among these strains, strain DMKU- The phylogenetic tree was constructed using the neighbour-

100 Nakazawaea populi CBS 7351T (EU011726/EU011646) Nakazawaea anatomiae CBS 5547T (EU011725/EU011645) 98 Nakazawaea wyomingensis CBS 8703T (KM065919/AF153673) 60 Nakazawaea wickerhamii CBS 2928T (EU011727/EU011647) 0.0050 Nakazawaea ernobii CBS 1737T (EU011728/EU011648) 100 Nakazawaea holstii *CBS 4140T (JQ698919/JQ689055) 92 Nakazawaea molendini-olei CBS 12508T (HE574679/JN688665) 61 Nakazawaea ishiwadae CBS 6022T (EU011729/EU011650) T 76 Nakazawaea pomicola CBS 4242 (KM065915/AF245400) Nakazawaea peltata CBS 5576T (EU011730/EU011651) 100 Nakazawaea siamensis CBS 12569T (LC171726/AB772177) 100 Nakazawaea todaengensis DMKU-PS11(1)T (LC171727/LC171729) 100 Nakazawaea laoshanensis CBS 11389T (KM065912/KM065901) Pachysolen tannophilus CBS 4044T (JQ698920/JQ689056) T 97 100 Peterozyma toletana CBS 2504 (JQ698921/JQ689057) Peterozyma xylosa CBS 2286T (EU011732/EU011653) 99 Candida natalensis CBS 2935T (KM065913/KM065902) 100 Candida quercitrusa CBS 4412T (KM065916/KM065904) 98 Kurtzmaniella cleridarum CBS 8793T (JQ698907/Q689038) 98 Debaryomyces hansenii CBS 767T (JQ698910/JQ689041) T 61 99 Candida temnochilae CBS 9938 (KM065918/KM065906) Candida heliconiae CBS 10000T (KM065911/KM065900) Cephaloascus fragrans CBS 121.29T (JQ698916/JQ689052) Trigonopsis variabilis CBS 1040T (JQ698933/U45827)

Fig. 1. Phylogenetic tree based on the combined sequences of the SSU rRNA and the D1/D2 region of the LSU rRNA genes showing the position of DMKU-PS11(1)T with respect to strains of closely related species. The phylogenetic tree was constructed from evolu- tionary distance data with maximum composite likelihood correction, using the neighbour-joining method with MEGA version 7.0. Num- bers at nodes indicate percentages of bootstrap sampling, derived from 1000 datasets. Trigonopsis variabilis NRRL Y-1579T was used

as the outgroup. The teleomorphic species, N. holstii, is marked with an asterisk. Bar, 0.005 Knuc distance.

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joining method [18], and the evolutionary distances were recommendation of Lachance [25]. The name Nakazawaea computed using the maximum composite likelihood todaengensis f.a., sp. nov. (MB 819513) is proposed. method [19]. Confidence levels of the clades were estimated from bootstrap analysis with 1000 replicates [20]. DESCRIPTION OF NAKAZAWAEA Strain DMKU-PS11(1)T was characterized morphologically, TODAENGENSIS POLBUREE, M. GROENEW. biochemically and physiologically by standard methods AND S. LIMTONG F.A., SP. NOV. (MB 819513) [21]. Formation of pseudohyphae and true hyphae was Nakazawaea todaengensis (to.daeng.en¢sis. N.L. fem. adj. investigated by cultivation on corn meal agar (2 % corn  todaengensis referring to Pru To Daeng Wildlife Sanctuary meal infusion and 2 % agar) in slide culture at 25 C for up where the type strain was isolated). to 14 days. Ascospore formation was investigated on 5 %  malt extract agar (5 % malt extract and 2 % agar), Gorod- Growth in YM broth: After 5 days at 25 C, cells are globose kowa agar (0.1 % glucose, 0.5 % sodium chloride, 1 % pep- to subglobose (2–3Â3–4 μm), and occur singly or in pairs tone and 2 % agar), Fowell’s acetate agar (0.5 % sodium (Fig. 2). Budding is multilateral. Pseudohyphae and true  acetate trihydrate and 2 % agar) and corn meal agar at 25 C hyphae are not formed in slide culture on corn meal agar   and 15 C for up to 6 weeks. Assimilation of carbon and after 14 days at 25 C. Ascospores are not produced on 5 % nitrogen compounds was tested in liquid medium, and malt extract agar, Fowell’s acetate agar, Gorodkowa agar or  starved inoculums were used in the nitrogen assimilation corn meal agar at 15 and 25 C in up to 6 weeks. The streak tests [21]. Growth at various temperatures was determined culture on YM agar is white with smooth margins. Glucose by cultivating the strain in YM broth. Ubiquinones were is fermented but D-galactose, lactose, maltose, melibiose, extracted from cells that were cultivated in a 500 ml Erlen- raffinose, sucrose, trehalose and xylose are not fermented. meyer flask containing 250 ml yeast extract peptone glucose D-Glucose, D-galactose, L-sorbose, N-acetylglucosamine, D- (YPG) broth (1 % yeast extract, 2 % peptone and 2 % glu- ribose (slowly), D-xylose, L-arabinose (weakly), D-arabinose  cose) on a rotary shaker at 28 C for 72 h and purified (weakly), L-rhamnose, sucrose, maltose, trehalose, methyl according to the methods described by Yamada and Kondo a-D-glucoside (slowly), cellobiose, salicin, melezitose, inulin [22] and Kuraishi et al. [23]. Isoprenologues were identified (slowly), soluble starch, glycerol, ribitol (slowly), D-glucitol by HPLC as described previously [14]. (slowly), D-mannitol, D-glucono-1,5-lactone, 2-ketogluco- nate, succinate (slowly), citrate (slowly), ethanol (slowly) and xylitol (weakly) are assimilated but melibiose, lactose, NOVEL SPECIES DELINEATION AND raffinose, erythritol, galactitol, myo-inositol, 5-ketogluco- IDENTIFICATION nate, D-gluconate, D-glucuronate, D-galacturonate, DL-lac- tate and methanol are not. Ammonium sulfate, potassium Analysis of the sequences of the D1/D2 region of the LSU nitrate, sodium nitrite ethylamine hydrochloride, L-lysine, rRNA revealed that strain DMKU-PS11(1)T was most cadaverine and creatine are assimilated. Growth in vitamin- closely related to Nakazawaea (Candida) laoshanensis T free medium is not present. Growth on medium containing NRRL Y-63634 (KM065901) but differed by 2.3 % nucleo- 50 % glucose or 10 % sodium chloride/5 % glucose is absent. tide substitutions (13 nucleotide substitutions and 1 gap out Growth with 0.01 % cycloheximide and 0.1 % cycloheximide of 562 nt). Strain DMKU-PS11(1)T differed from NRRL Y- 63634T by 8.0 % nucleotide substitutions (41 nucleotide sub- stitutions out of 572 nt) in the ITS region (FJ613523) and 1.0 % nucleotide substitutions (17 nucleotide substitutions and 1 gap out of 1720 nt) in the SSU region (KM065912). The phylogenetic tree reconstructed from the combined sequences of the SSU rRNA and the D1/D2 region of the LSU rRNA genes (Fig. 1) was in agreement with the tree reconstructed from the sequences of the SSU rRNA gene alone (data not shown). The trees confirmed the separation of strain DMKU-PS11(1)T from the other species of the genus Nakazawaea and its close relatedness to N. laosha- nensis, which is one of the basal members of the genus Nakazawaea (Fig. 1). On the basis of a genetic species con- cept, we concluded that strain DMKU-PS11(1)T represents a novel species. As the proposed ‘one one name’ nomenclature rule for fungi was included in the Interna- tional Code of Nomenclature for algae, fungi, and plants, [24] the novel species is assigned to the genus Nakazawaea Fig. 2. Budding cells of Nakazawaea todaengensis f.a., sp. nov. DMKU- although the formation of ascospores was not observed. The PS11(1)T in YM broth after 5 days at 25 C. Bar, 5 mm. designation forma asexualis (f.a.) was included following the

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Table 1. Phenotypic characteristics that differentiae N. todaengensis from the other species of the genus Nakazawaea Species: 1, N. todaengensis; 2, N. laoshanensis; 3, N. anatomiae; 4, N. holstii; 5, N. ishiwadae; 6, N. molendini-olei; 7, N. peltata; 8, N. pomicola; 9, N. populi; 10, N. siamensis; 11, N. wickerhamii; 12, N. wyomingensis. Data for species 2 are from Wang et al. [28], for species 3, 5, 7–9 and 11–12 are from Lachance et al. [29], for species 4 are from Kurtzman [6], for species 6 are from Cadez et al. [30] and for species 10 are from Kaewwichian and Limtong [7]. +, Positive; À, negative; d; delayed positive; S, slow; W, weakly positive; V, variable; ND, no data available.

Characteristic 1 2 3 4 5 6 7 8 9 10 11 12

Fermentation Galactose À + À W/SS/ÀÀ S ÀÀ À ÀÀ Maltose ÀÀÀ À + ÀÀÀÀ À ÀÀ Sucrose ÀÀÀ ÀS/ÀÀÀÀÀ À ÀÀ Trehalose À ND À + SSÀ V ÀÀ V À Assimilation of carbon compounds D-Galactose + + À + V À + S À S + À L-Sorbose + ÀÀ + S À + V À S ÀÀ N-Acetylglucosamine + V À S + S +++ + ++ D-Ribose S + À ++S + W ++VV D-Xylose + + À ++S +++ + S + L-Arabinose WVÀ + S ++++ WWÀ D-Arabinose WVÀ + V À + S + SV+ Sucrose + + À ++À +++ + ÀÀ Maltose + + À ++À +++ + ÀÀ Methyl a-D-glucoside S d À + V À + SV + ÀÀ Melibiose À d À À ÀÀÀÀÀ À ÀÀ Melezitose + S À + V À +++ + ÀÀ Inulin S ÀÀ À ÀÀÀÀÀ + ÀÀ Soluble starch + + À ++À ++À + ÀÀ Glycerol + + À + +++++ + ++ Erythritol ÀÀÀ ++À VWÀ + ÀÀ Ribitol S + À + +++++ + +V D-Glucitol S + À + +++++ + ++ D-Mannitol + + À + +++++ + ++ Galactitol À V À VVÀ V ÀÀ W ÀÀ D-Glucono-1,5-lactone + ND S ND + S +++++À 2-Ketogluconate + ND À VSÀÀÀÀ W VV D-Gluconate À ND W V + ÀÀ++ SVÀ D-Glucuronate À ND À ND ÀÀÀÀÀ + ÀÀ D-Galacturonate À ND ND ND ND À + À ND À ND ND DL-Lactate À + ÀÀ V + ÀÀÀÀÀV Succinate SVÀ + +++++ + ++ Citrate S + À + +++++ + +À Ethanol S ++ + +++++ + S + Xylitol W ND À ND V ++++ + ÀÀ Assimilation of nitrogen compounds Potassium nitrate + ÀÀ + ++À ++ À ++ Sodium nitrite + À S ND ++À ++ À + S Creatine + ND ND ND ND À ND ND ND ND ND ND Additional growth tests and other characteristics  Growth at 30 C+ND À ND ++ND W ++++  Growth at 35 C S ND À ND +++À ND + ND ND  Growth at 37 C S + À V À W + ÀÀ + ÀÀ  Growth at 40 C ÀÀÀ À ÀND ND ÀÀ + ÀÀ 50 % Glucose medium À ND À ND + ÀÀ S À + ÀÀ 60 % Glucose medium À ND À ND ND ND ÀÀÀ + ÀÀ 10 % NaCl/5 % glucose À ND À +/W ++ND + À + ÀÀ Acid formation + ND W ND À ND ÀÀÀND ÀÀ

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Table 1. cont.

Characteristic 1 2 3 4 5 6 7 8 9 10 11 12

Pseudohyphae ÀÀW + ÀÀÀÀÀ À À+ Septate hyphae ÀÀÀ + ÀÀÀÀÀ À À+ Ascospores ÀÀÀ + ÀÀÀÀÀ À ÀÀ Major ubiquinone Q-7 ND Q-8 Q-8 Q-8 ND Q-7 ND Q-8 Q-7 Q-7 ND

 is present. Growth at 15, 30 and 37 C is present. Formation species of the genus Nakazawaea in peat swamp forests has of acid is positive. Starch-like compounds are not produced. never been reported before to our knowledge. Diazonium blue B colour and urease reactions are negative. The major ubiquinone is Q-7. T Funding information The type strain, DMKU-PS11(1) , was isolated from peat in This work was supported by the Office of The Royal Development Pro- the rhizosphere of Calophyllum teysmannii var. inophylloide jects Board, Thailand, The Pikunthong Royal Development Study Cen- in Princess Sirindhorn Wildlife Sanctuary (Pru To Daeng tre, Land Development Department, Thailand. Wildlife Sanctuary), Narathiwat province, Thailand. This Acknowledgements strain has been deposited as the holotype in the CBS collec- The authors would like to thank Dr Pisoot Vijarnsorn, Ms Saiyud Petsik tion of the Westerdijk Fungal Biodiversity Institute, Utrecht, and Mr Anurak Buakleeklay, Land Development Department,Thailand, the Netherlands as CBS 14555T. The isotype has been for their comments and peat collection. deposited in the Thailand Bioresource Research Centre Conflicts of interest (TBRC), National Centre for Genetic Engineering and Bio- The authors declare that there are no conflicts of interest. technology (BIOTEC), Pathumthani, Thailand, as TBRC T 6559 . The Mycobank registration number is MB 819513. References 1. Yamada Y, Maeda K, Mikata K. The phylogenetic relationships of N. todaengensis can be distinguished from its closest phylo- the hat-shaped ascospore-forming, nitrate-assimilating Pichia genetic relative, N. laoshanensis, not only on the basis of the species, formerly classified in the genus Hansenula Sydow et Sydow, based on the partial sequences of 18S and 26S ribosomal gene sequences of the SSU rRNA, the D1/D2 region of the RNAs (Saccharomycetaceae): the proposals of three new genera, LSU rRNA and the ITS region but also by some phenotypic Ogataea, Kuraishia, and Nakazawaea. Biosci Biotechnol Biochem characteristics (Table 1). N. todaengensis ferments only D- 1994;58:1245–1257. glucose where N. laoshanensis ferments both D-glucose and 2. Tsui CK, Daniel HM, Robert V, Meyer W. Re-examining the phylog- eny of clinically relevant Candida species and allied genera based D-galactose. The novel species assimilates inulin (slowly), L- on multigene analyses. FEMS Yeast Res 2008;8:651–659. sorbose, potassium nitrate and sodium nitrite while 3. McNeill J, Barrie FR, Buck WR, Demoulin V, Greuter W et al. N. laoshanensis does not. The phenotypic differences International Code of Nomenclature for Algae, Fungi, and Plants between the novel species and the other species of the genus (Melbourne Code). Regnum Veg, 154. Koenigstein, Germany: Gant- Nakazawaea are shown in Table 1. ner Verlag;2012. 4. Kurtzman CP, Robnett CJ. Description of Kuraishia piskuri f.a., sp. Species of the genus Nakazawaea have been isolated from a nov., a new methanol assimilating yeast and transfer of phyloge- wide variety of substrates. However, some members of this netically related Candida species to the genera Kuraishia and Nakazawaea as new combinations. FEMS Yeast Res 2014;14:1028– genus seem to be primarily associated with plants. Many 1036. strains of N. holstii, the type species of this genus, were iso- 5. Kurtzman CP, Robnett CJ. Identification and phylogeny of ascomy- lated from bark beetles infesting pine, spruce and fir trees cetous yeasts from analysis of nuclear large subunit (26S) ribo- whereas the type strain of N. ishiwadae was obtained from a somal DNA partial sequences. Antonie van Leeuwenhoek 1998;73: – deep core of stratigraphic drillings in Japan [6]. N. populi 331 371. 6. Kurtzman CP. Nakazawaea Yamada, Maeda and Mikata (1994). In: was at first described as Candida populi from 24 strains Kurtzman CP, Fell JW and Boekhout T (editors). The Yeasts, A recovered from sap fluxes of trembling aspen (Populus trem- Taxonomic Study, 5th ed. Amsterdam: Elsevier; 2011. pp. 637–639. uloides), black cottonwood (Populus trichocarpa) and birch 7. Kaewwichian R, Limtong S. Nakazawaea siamensis f.a., sp. nov., a (Betula sp.) in the Pacific Northwest region of North Amer- yeast species isolated from phylloplane. Int J Syst Evol Microbiol 2014;64:266–270. ica [26]. N. (Candida) wickerhamii was described previously 8. Posa MRC, Wijedasa LS, Corlett RT. Biodiversity and conservation as Torulopsis wickerhamii based on strains obtained from of tropical peat swamp forests. Bioscience 2011;61:49–57. olive husks used for silage making [27]. The type strain of 9. Thormann MN, Rice AV, Beilman DW. Yeasts in peatlands: a N. siamensis was isolated from the external surface of review of richness and roles in peat decomposition. Wetlands a sugarcane leaf collected in Thailand [7]. The closest rela- 2007;27:761–773. tive of N. todaengensis, N. laoshanensis, was isolated from 10. Kachalkin AV, Glushakova AM, Iurkov AM, Chernov NIu. [Charac- terization of yeast groupings in the phyllosphere of Sphagnum decayed wood in China [28]. In this study, N. todaengensis mosses]. Mikrobiologiia 2008;77:474–481. was isolated from peat in the rhizosphere of a plant in a 11. Kachalkin AV, Yurkov AM. Yeast communities in Sphagnum phyl- peat swamp forest in Thailand. The occurrence of strains of losphere along the temperature-moisture ecocline in the boreal

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