Proposal of one new order, two new families, nine new genera and seventy-eight new basidiomycetous yeast species isolated from China

doi:10.21203/rs.3.rs-2300767/v1

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Proposal of one new order, two new families, nine new genera and seventy-eight new basidiomycetous yeast species isolated from China

https://doi.org/10.21203/rs.3.rs-2300767/v1

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More than 2,000 yeast strains isolated from 1200 samples mostly collected from Tibet and Yunnan provinces were identified as 466 species according to the internal transcribed spacer including the 5.8S rDNA (ITS) and the D1/D2 domains of the large subunit rDNA (LSU) sequence analyses. Among them, 78 new basidiomycetous yeast species were proposed based on the multi-locus phylogenetic analyses including the D1/D2 domains, the ITS, the small subunit rDNA (SSU), the largest subunit of RNA polymerase II (RPB1), the second largest subunit of RNA polymerase II (RPB2) and translation elongation factor 1-α (TEF1), as well as the phenotypic comparisons. Fifty-three new species belong to 17 genera in the Agaricomycotina, 19 new species occur in 12 genera in the Pucciniomycotina and six new species distribute in four genera in the Ustilaginomycotina, and eleven new species were classified in nine newly established genera. Two new families and one new order were also proposed based on two of these new genera. The genus Guomyces and the species Guomyces nicotianae were validated.

90 new taxa

Basidiomycetous yeasts

species diversity

molecular phylogeny

The number of yeast species has dramatically increased, i.e., 166, 362, 499, 725 and 1301 species are accepted in the five editions of The Yeasts, a Taxonomic Study, respectively, and now 1958 species are listed in The Yeasts Trust database (http://theyeasts.org/). The rapid growth of new species number in the last 20 years was mainly due to the progressive taxonomic survey in Asia (Boekhout et al. 2022). The yeast diversity from various substrates in China has been investigated over the past decades, and nearly 800 yeast species including more than 280 new species have been reported (Boekhout et al. 2022, Li et al. 2022). Among the described new species, nearly 200 are classified in three subphyla of Basidiomycota, namely Agaricomycotina, Pucciniomycotina and Ustilaginomycotina (Boekhout et al. 2022, Li et al. 2020, 2022). Our recent description of more than 120 yeast species covering the 20 years of yeast collection in our group (Li et al. 2020, 2022) increased the species diversity of basidiomycetous yeasts by 20% (Boekhout et al. 2022). However, the described species from China or worldwide presents only the tip of the iceberg of yeast species diversity in nature. Fell (2012) and (Lachance 2006) estimated that only 1% of yeast species have been described, which indicated that the estimated total number of yeast species might reach 200,000. Therefore, a major effort is needed to document unknown yeast species that might exist in nature (Boekhout et al. 2022).

Our studies have shown that Tibet and Yunnan provinces are the hotpots of yeast species diversity in China (Zhao et al. 2003, Wang et al. 2007, Wang et al. 2012, Bing et al. 2014, Wang and Wang 2015, Han et al. 2016, Li et al. 2020). Ninety-two new species have been discovered from the above two regions (51 species from Tibet and 41 species from Yunnan province), accounting for 32.9% (92/280) of all new species discovered in China. However, the approximate number of species in these two regions is still unknown. In order to give some hint about the yeast species diversity in these two regions, we carried out surveys twice collecting 1200 samples including leaves, flowers, barks, soil and rotten wood from Tibet and Yunnan provinces during the last three years. More than 2,000 strains were isolated from these samples, and 466 species have been identified by the internal transcribed spacer including the 5.8S rDNA (ITS) and the D1/D2 domains of the large subunit rDNA (LSU) sequence analyses. Besides those two regions, 14 strains isolated from Hainan, Hubei and Qinghai provinces were included in this study. Among them, 95 strains were classified into 78 groups representing the undescribed basidiomycetous yeast species in nature. The multi-locus phylogenetic analyses based on the small subunit rDNA (SSU), the D1/D2, the ITS, the largest subunit of RNA polymerase II (RPB1), the second largest subunit of RNA polymerase II (RPB2) and translation elongation factor 1-α (TEF1) were used to clarify the taxonomic positions of those new species, and nine new genera, two new families and one new order were proposed based on these taxa.

Strains and phenotypic characterization

A total of 95 strains collected in this study were listed in Table 1. Three isolation methods were used for yeast collection. Strains were isolated from soil and rotten wood by an enrichment method as described by Li et al. (2020). Isolation of strains from the surface of plant leaves and flowers was performed by the ballistoconidia-fall method as described by Nakase and Takashima (1993) and the washing and diluting method described in detail below. The leaves and flowers were cut into pieces with sterile scissors, put into a 10 ml sterile centrifuge tube, added sterile water containing 0.05% Tween 80. After shaking for about 10 min with a shaker, diluted washing samples to 1*10^-2, then 200 μL of each dilution was spread out on potato dextrose agar containing 200ug/ml chloramphenicol (PDA, 20% potato infusion, 2.0% glucose, 2.0% agar, Difco) plates at 17 °C for 5–7 days. The strains were characterized morphologically, biochemically, and physiologically according to standard methods (Kurtzman et al. 2011) and the results were recorded after 7, 14 and 21 days of incubation. The potential sexual cycles of all new species were assessed on YM, PDA, V8 (10% V8 juice, 2% agar) and corn meal agar (CM, 5% infusion corn meal, 1.5% agar, Difco). The ballistoconidium-forming activity of all new species was observed by the inverted-plate method (do Carmo-Sousa and Phaff, 1962) using CM agar at 17 °C.

DNA extraction and ribosomal DNA sequencing

The method for nuclear DNA extraction was described previously by Wang and Bai (2008). The sequences of three rDNA loci including the ITS (including 5.8S rDNA) region and D1/D2 domains of the LSU rDNA gene and the small subunit (SSU) rDNA were amplified and sequenced using the methods described previously (Wang and Bai 2004, Wang et al. 2003). The three nuclear protein-coding genes, RPB1, RPB2 and TEF1, were determined according to Wang et al. (2014). GenBank accession numbers for all sequences newly determined in this study were given in Table 1.

Sequences of six genes, ITS, D1/D2, SSU, RPB1, RPB2 and TEF1, were aligned with the MAFFT program (Standley 2013) using the G-INS-i algorithm, respectively. The minor gaps in all alignments were trimmed manually. Then two combined six-gene and ITS+D1/D2 datasets were concatenated with the aligned single genes. The best nucleotide substitution models were determined with jModeltest version 3.04 (Posada and Crandall 1998) using the Akaike information criterion (AIC). The maximum likelihood (ML) phylogeny of six-gene, ITS+D1/D2 and single D1/D2 datasets were inferred using RAxML-HPC 7.2.8 (Stamatakis 2006), and the bootstrap values were calculated with 500 replicates. BI analysis was conducted using MrBayes 3.1.2 (Ronquist et al. 2012) with 5 000 000 to 10 000 000 generations using the parameter settings described previously (Wang et al. 2015). A bootstrap percentage (BP) of ≥70% or a Bayesian posterior probability (PP) of ≥0.9 was considered as significantly supported in all constructed trees in this study. The alignments and trees were deposited in TreeBASE (www.treebase.org, No. 29910).

New taxon delineation and phylogenetic placement

Sequence analyses of the D1/D2 domains and the ITS region have been widely used for yeast species delineation (Kurtzman and Robnett 1998, Fell et al. 2000, Scorzetti et al. 2002, Boekhout et al. 2011, 2021, Kurtzman 2011, Vu et al. 2016, Li et al. 2020). A sequence similarity cut-off (threshold), less than 1 % in either the ITS or D1/D2 regions among conspecific strains, for yeast species identification has been recommended by Kurtzman and Robnett (1998), Fell et al. (2000), Scorzetti et al. (2002), Kurtzman and Fell (2006), Kurtzman (2014, 2015) and Kurtzman et al. (2015). Based on the barcode data of ca. 9 000 yeasts generated at the Westerdijk Fungal Biodiversity Institute, Vu et al. (2016) recommended a similarity threshold to predict species boundaries, it was 98.41% and 99.51% for ITS and D1/D2, respectively. The threshold has been updated by Boekhout et al. (2021) with taxonomic improvement, 98.4% and 99% for ascomycetous yeasts and basidiomycetous yeasts using ITS sequence, respectively, while 99.6% for both yeast groups using D1/D2 barcode. However, it is not reliable to recognize yeast species using a single barcode (gene) and the same threshold, because different lineages may have various rates of nucleotide substitution in different barcodes or genes (Fell et al. 2000, Scorzetti et al. 2002, Boekhout et al. 2021). In our previous study, a case-by-case sequences similarity analysis of nucleotide variation in the ITS region and D1/D2 domains among most yeast genera in Agaricomycotina and Pucciniomycotina were determined (Li et al. 2020), which give more helpful than the general threshold to delimit yeast species. The optimal thresholds predicted for genus discrimination for all yeasts using D1/D2 and ITS sequences were also recommended by Vu et al. (2016) and Boekhout et al. (2021), 96.31% for ITS and 97.11% for D1/D2 in the former study, 93.7% for ITS and 98.9% for D1/D2 in the latter study. The thresholds for basidiomycetous yeast genus identification were 95.2% for ITS and 98.9% for D1/D2, for ascomycetous yeasts were 91.2% for ITS and 94.0% for D1/D2 (Boekhout et al. 2021).

Boekhout et al. (2021) suggested that sequence similarity cut-offs should not be used as a sole criterion for the delineation of yeast species and genera. Consequently, a combined approach including a case-by-case sequences similarity analysis of both D1/D2 and ITS used in Li et al. (2020), multi-locus phylogenetic analysis and physiological properties comparison was used to recognize species and discriminate genus in this study.

Ninety-five yeast strains mostly isolated from Tibet and Yunnan provinces were grouped into 78 undescribed species using the species identification approach suggested by Fell et al. (2000), Scorzetti et al. (2002), Kurtzman and Fell (2006), Kurtzman (2014, 2015) and Kurtzman et al. (2015) as well as the taxonomic thresholds of yeast species recommended by Vu et al. (2016), Li et al. (2020) and Boekhout et al. (2021). Fifty-three new species distribute in 17 genera in the Agaricomycotina, 19 new species occur in 12 genera in the Pucciniomycotina and six new species locate in four genera in the Ustilaginomycotina (Figs 1–21, Table 1). Eleven new species have high sequence diversity both in D1/D2 and ITS regions and the BLASTn results using both regions showed that those new taxa had low sequence similarity with various known genera, which indicated that they did belong to any known genera. Nine new genera should be proposed to accommodate those species.

In order to illustrate the phylogenetic positions of these undescribed species, the multi-loci phylogenetic trees were constructed from two datasets, the combined ITS (including 5.8S) and D1/D2 dataset and the six-gene dataset combined ITS, D1/D2, SSU, RPB1, RPB2 and TEF1 genes. A total of 95 SSU, 95 ITS, 95 D1/D2, 93 RPB1, 94 RPB2 and 92 TEF1 new sequences were generated (Table 1). The six-gene phylogenetic trees (Figs 1–3) were used to determine the phylogenetic placement for each new species. The trees (Figs 4–21) constructed from the ITS+D1/D2 or single D1/D2 datasets were used to calculate the similarity between the new species and their closely related described species.

Part of describing new species in this study includes one strain from our isolations. In order to find potentially conspecific strains for those single-strain species, we searched GenBank and The Yeasts Trust database to find identical or similar sequence strains or uncultured clones using their ITS and D1/D2 sequences as queries. 101 sequences including 74 strains and 27 uncultured fungus clones found from different origins were added to the new species delimitation below (Table 1).

New taxa identification in the Tremellomycetes (Agaricomycotina)

Derxomyces (Bulleribasidiaceae, Tremellales)

Six strains isolated from Motuo county in Tibet formed four groups and were located in the Derxomyces (Figs 1A and 4). Strains MT-2A-35 and MT-3B-3 have an identical sequence in the D1/D2 domains and two nucleotides (nt) difference in the ITS region, which indicated that they are conspecific. This group was closely related to D. cylindricus and D. xingshanicus (Figs 1A and 4), and differed from them by zero to one nt substitutions in the D1/D2 domains, but by 21–28 nt (~3–4%) mismatches (including substitutions and deletions) in the ITS region. Strain MT-5C-1 was closely related to D. ovatus and D. taiwanicus (Figs 1A and 4), and differed from them by nine to ten nt (~1.8%) and 52–54 nt (~8.5%) in the D1/D2 domains and ITS region, respectively. Strains MT-1B-10 and MT-4A-7 formed a separate branch in the six-gene and ITS+D1/D2 trees (Figs 1A and 4), and clustered with three strains isolated from the leaf in Vietnam, Derxomyces sp. VY-55, VY-142 and VY-65 (Fig. 4). They differed from one another by zero to one nt in the D1/D2 domain and two to six nt in the ITS region, which indicated that they belong to the same species. MT-3B-4 clustered with strains Derxomyces sp. 2851, 2357, 1612, 3032, 2213 and 2533, and has zero to one nt D1/D2 sequence differences and eight to ten nt (~1.2–1.5%) ITS sequence differences, which indicated that they may be different species.

Hannaella (Bulleribasidiaceae, Tremellales)

Four strains isolated from Yunnan province, YN14M1, YN25-1, YN28-3 and YN33-4, formed three groups in the genus Hannaella (Figs 1A and 4). Strain YN28-3 has the same D1/D2 sequence and four nt ITS sequence differences with Hannaella sp. DSM 101917 isolated from beans in Germany (Prior et al. 2017), which indicated that they are conspecific. Those two strains were related to H. taiwanensis (Fig. 4), and differed from it by 12 nt (~2%) and 20 nt (~4%) in the D1/D2 domains and ITS region, respectively. Strains YN25-1 and YN33-4 have identical ITS and D1/D2 sequences with Hannaella sp. strains NYNU 224166 and NYNU 224240. They were closely related to H. pagnoccae and H. dianchiensis (Fig. 4), and differed from them by more than 4% sequence diversity in both D1/D2 and ITS regions. Strain YN14M1 has an identical or similar sequence with Hannaella sp. NYUN 211096 and NYNU 22867 in the D1/D2 and ITS regions, and closely related to H. surugaensis (Figs 1A and 4). Those three Chinese strains differed from H. surugaensis by 20 nt (~3.6%) and 46 nt (~8.4%) in the D1/D2 domains and ITS region, respectively.

Note: ‘Cryptococcus’ sp. MB146852 (KF830205) isolated from hummingbird saliva in Costa Rica (Belisle et al. 2014.) and an uncultured fungus clone ClerRtSt1199 (JX242210) isolated from rotting stems from Clermontia arborescens in the USA (Ort et al. 2012) have the same sequences with YN14M1, NYNU 22867 and NYUN 211096 in the D1/D2 domain, which indicated that they may be conspecific. An uncultured ‘Cryptococcus’ clone 10D10C29 (HG937180) isolated from Zea mays field bulk soil in Germany (Moll et al. 2016) has the identical ITS sequence with YN28-3 and DSM 101917, which indicated that the species represented by YN28-3 was also distributed in Germany. ‘Cryptococcus’ sp. SM3S12 (EF612202) and SM13L10 (FJ873462) isolated from soil in Taiwan, China and Fungal sp. YT-2014-ARM5 (AB976614) isolated from pyroloid root tip in Japan have the same ITS sequence with YN25-1, YN33-4, NYNU 224166 and NYNU 224240, which indicated that they are conspecific. ‘Cryptococcus’ sp. MB1425 (KF830199) isolated from hummingbird saliva in Costa Rica and ‘Cryptococcus’ sp. SM3S05 (EF460579) isolated from soil in Taiwan, China, and Agaricomycotina sp. NRP120 (AB726809) isolated from Japan have the identical D1/D2 sequence with YN25-1, YN33-4, NYNU 224166 and NYNU 224240, which indicated that they may belong to the same species.

New genus Babevozyma in Trimorphomycetaceae (Tremellales)

The genus Saitozyma was created by Liu et al. (2015a) including three species, S. flava, S. paraflava and S. podzolica. The monophyly of Saitozyma was not supported and S. podzolica was separated by the genera Carlosrosaea, Sugitazyma and Trimorphomyces from the subclade represented by S. flava, the type species of Saitozyma in this study (Fig. 1A), which in agreement with our previous study (Li et al. 2020). The ITS+D1/D2 phylogenetic analysis in our study (Fig. 5) and Liu et al. (2015b) also showed that S. podzolica, S. ninhbinhensis and S. wallum which recently reported by Crous et al. (2021) formed a separate subclade from the subclade represented by the type species. The above data analyses indicated that the subclade of S. podzolica should represent a new genus in Trimorphomycetaceae. Strain DBS918-3-4 was related to S. podzolica (Fig. 5) and differed from it by three nt in the D1/D2 domains and 19 nt (~3.7%) in the ITS region.

Carlosrosaea (Trimorphomycetaceae, Tremellales)

Three strains isolated from Yunnan province formed three groups in the Carlosrosaea (Figs 1A and 5). Strain JZXS7-21 has the same sequences in the D1/D2 domains with C. foliicola, but differed from it by 10 nt (~2%) in the ITS region. Strain YN28M1 was closely related to two unpublished strains Carlosrosaea sp. 7 BM 77 and Carlosrosaea sp. 3 BSS 158 with more than 2% and 9% difference in the D1/D2 domains and ITS region, respectively (Fig. 5). Strain YN35-7 has one nt D1/D2 and two nt ITS sequence differences with an unpublished Carlosrosaea sp. NYNU 208206 (Fig. 5), which indicated that they are conspecific. Those two strains differed from the known Carlosrosaea or unpublished species by 17 nt (~3%) in the D1/D2 domains and more than 100 nt (~17%) in the ITS region.

Note: YN28M1 has one to two nt in the ITS2 region with three uncultured fungus clone 2588_218 (KU188658) isolated from the rhizosphere of conifer seedlings and clone 2170_969 (KP891846) isolated from Scolytus multistriatus in Sweden (Menkis et al. 2016a, b) and clone 3980_1319 (MN903603) isolated from Pinus sylvestris in Lithuania (Lynikienė et al. 2020), which indicated that the species represent by strain YN28M1 was distributed worldwide. Three uncultured fungus clones (LS993390, LR887593 and LR781329) isolated from leaf, soil and root in China, clone OTU_540_GW5CJXV07IELAM (JF945390) isolated from Fagus sylvatica phyllosphere in France (Cordier et al. 2012) have identical ITS2 or ITS1 sequences with YN35-7, which indicated that the species represent by YN35-7 is broadly distributed. Carlosrosaea sp. fn_8 (LC333465) isolated from floral nectar in Japan (Tsuji et al. 2018) has one nt in the D1/D2 domains with YN35-7, which indicated that they may be conspecific.

Kockovaella (Cuniculitremaceae, Tremellales)

Two strains isolated from Tibet, MT-2C-31 and RTFM4-1, formed two separate branches in the genus Kockovaella (Figs 1B). They differed from the known and unpublished species of Kockovaella by more than 12 nt (~2%) in the D1/D2 domains and 26 nt (~4%) in the ITS region, respectively.

Papiliotrema (Rhynchogastremaceae, Tremellales)

Three strains motuo45-1, YN83-2 and YN109M3 were located in the genus Papiliotrema (Figs 1B and 6). Strain motuo45-1 has identical D1/D2 sequence and two to four nt ITS differences with ‘Papiliotrema laurentii’ CBS 5297, CBS 7140, IIF4SW-F2, IIF7SW-F3, IF7SW-F4, IF7SW-B5, Kw418bc/19, Kw543bc/19, ‘Cryptococcus laurentii’ IMUFRJ 51940, IMUFRJ 51993, IMUFRJ 51994, IMUFRJ 51996, IMUFRJ 51998, Papiliotrema sp. YP169 and an uncultured fungus ASV 910 (Fig. 6), which indicated that they are conspecific. They differed from the type strains of P. laurentii by one nt in the D1/D2 domains, but by 11 nt (~2%) in the ITS region, which indicated that they should be a different species. Strain YN83-2 closely related to P. aspenensis and YN109M3 closely related to P. bandonii formed two separated branches (Figs 1B and 6), and differed from the other Papiliotrema species by more than 7 nt (~1.2%) in the D1/D2 domains and 31 nt (~5%) in the ITS region, respectively.

Note: ‘Papiliotrema laurentii’ AE7 (MH030123) isolated from soil in Germany, CBS 2174 (KY104468) isolated from a human in Netherlands and ‘Cryptococcus laurentii’ WM 844 (KP131909) isolated from an animal in Australia have 99.21-100% (zero to five nt) similarity with strain motuo45-1 in the ITS region, which indicated that the species represent by motuo45-1 is worldwide distributed and has various ecological niches. Papiliotrema aff. laurentii PYCC 5069 (MZ435815) and A48 (AF485975) isolated from Portugal have the same D1/D2 sequence with YN109M3. ‘Papiliotrema laurentii’ MB2011 (KC798409) isolated from banana nectar in Costa Rica has the identical D1/D2 sequence with the YN83-2 group. The above data indicated that the two species represented by YN83-2 and YN109M3, respectively, may occur worldwide.

Pseudotremella (Bulleraceae, Tremellales)

Four strains, XSR1-5, XSR10-2, XZY97M2 and XZY113-6, formed two groups in the genus Pseudotremella (Figs 1B and 7). Strains XSR1-5, XSR10-2 and XZY113-6 have similar sequences with one and two nt differences in the D1/D2 and ITS regions, respectively, which indicated that they are the same species. Those three strains clustered with ‘Pseudotremella moriformis’ RJB2846-8, Pseudotremella sp. KBP Y-6836, KBP Y-6886 and DBVPG 10729 with two nt D1/D2 sequence difference and 22 nt (~4%) ITS sequence diversity, which indicated that they belong to different taxa. Those strains were closely related to P. lacticolor (Fig. 7), and differed from it by 22 nt (~4%) in the D1/D2 domains and by 92 nt (~17%) in the ITS region. Strain XZY97M2 had a close relationship with P. nivalis, P. moriformis and ‘Tremella’ sp. KBP Y-6774 (Fig. 7), and differed from them by more than 12 nt (~2%) and 48 nt (~8%) in the D1/D2 domains and ITS region, respectively.

New genus Baiozyma in Bulleraceae (Tremellales)

The ITS and D1/D2 sequence BLASTn searches against the GenBank database showed that strain YN107M4 had an affinity to Sirobasidium intermedium, Sirobasidium brefeldianum and Fonsecazyma mujuensis, and had 86.6–90.6% ITS similarity and about 94% D1/D2 similarity with them. Strain YN107M4 was located in a separate branch and closely related to the above three known species in the Bulleraceae in the ITS+D1/D2 and six-gene trees (Figs 1B and 7). The low similarities in the above comparison and the two dataset phylogenetic analyses indicated that YN107M4did not belong to Sirobasidium and Fonsecazyma and represented a new genus in Bulleraceae.

New genus Corticimyces in Sirobasidiaceae (Tremellales)

The D1/D2 sequence BLASTn result showed that strain XSR1-14 had 93% similarity with Fibulobasidium inconspicuum and Bullera unica, however, the ITS BLASTn showed that this strain had about 85% similarity with Tremella lethariae and Naematelia encephala. This conflict BLASTn results indicated that XSR1-14 does not belong to any known genera in the Tremellales. The ITS+D1/D2 and six-gene phylogenetic analyses showed that XSR1-14 was located in a separate branch and had an affinity to Sirobasidium magnum and the genus Fibulobasidium in the Sirobasidiaceae (Figs 1B and 8). The above data analyses indicated that strain XSR1-14 should represent a novel genus.

Note: XSR1-14 had a similar sequence (two to three nt difference in the ITS and one nt in the D1/D2 domain) with two uncultured fungus clones 2_123 (KF274305) isolated wood stump in Finland and clone BOTU002 (KC588703) isolated from the forest floor (leaf litter) in northern hardwoods ecosystem in the USA, which indicated that this new species is worldwide distributed.

Naematelia (Naemateliaceae, Tremellales)

Strain XZY420-1 was closely related to N. encephala, the type species of Naematelia (Figs 1B and 9), and differed from it by 2 nt in the D1/D2 domains and by 11 nt (~1.8%) in the ITS region.

Vishniacozyma (Bulleribasidiaceae, Tremellales)

Nine strains, MLWB11-2, MLWB2-1, NYG10-6, XZY87M1, XZY91-8, XZY496-1, T1-6-1, YN95-2 and YN110-1, formed seven groups in the Vishniacozyma clade (Figs 1B and 10). Strains NYG10-6 and XZY87M1 differed from each other by four nt and 21 nt (~3.4%) in the D1/D2 domains and ITS region, respectively. Those two groups were closely related to V. victoriae (Figs 1B and 10), and differed from it by zero to four nt in the D1/D2 domain and 11-18 nt (~2–3%) in the ITS region. Strains T1-6-1 and XZY91-8 have identical ITS sequence, whereas had one nt difference in the D1/D2 domains, and were closely related to V. tephrensis (Figs 1B and 10). Those two strains differed from the latter by one nt in the D1/D2 domains, but by 16 nt (~3%) in the ITS region. Strain YN110-1 formed a separate branch related to V. victoriae, V. tephrensis, V. carnescens and three new species represented by NYG10-6, XZY87M1 and T1-6-1, and differed from them by 12 nt (~2%) in the D1/D2 domains and by more than 27 nt (~4%) in the ITS region. Strains MLWB2-1 and MLWB11-2 have identical sequences in both the ITS and D1/D2 regions, and clustered with Vishniacozyma sp. strains DMKU-SE39, DMKU4-1, DMKU10-2 and DMKU5-3 with zero to one nt and three to seven nt differences in the D1/D2 and ITS regions, respectively, which indicated that they are conspecific. Strain XZY496-1 formed a separate branch and was closely related to three unpublished strains Anhui D1, D8 and D20 (Fig. 10), and differed them by six nt in the D1/D2 domains and by 25 nt (~4%) in the ITS region. Strain YN95-2 was closely related to V. taibaiensis (Fig. 10), and differed from it by two nt in the D1/D2 domains and 19 nt (~3%) in the ITS region.

Note: Two Japanese strains, ‘Cryptococcus’ sp. KY-763 (AB428344 and AB428345) and Vishniacozyma sp. FCGNE-9 (LC529164) isolated from bed dust have the same D1/D2 sequence with MLWB2-1 and MLWB11-2, however, more than 13 nt（2%）ITS sequence diversity was observed between them, thus they may be different species. ‘Cryptococcus’ sp. BI195 (EU200789) in Brazil and MLWB2-1, MLWB11-2 have identical D1/D2 sequences, but it’s ITS sequence is not available for ‘Cryptococcus’ sp. BI195, which indicated that they may be conspecific. ‘Vishniacozyma taibaiensis’ PYCC 9090 (OM670202) isolated from Rye dough bait in Portugal and ‘Cryptococcus’ sp. KCTC 17070 (AF459686) isolated from Korea have the same D1/D2 sequence with YN95-2, which indicated that they may be conspecific. Vishniacozyma cf. taibaiensis ATT065 (FJ743602) isolated from Atta texana nest in the USA (Rodrigues et al. 2009) has the identical D1D2 sequence with XZY496-1, which indicated that they may belong to the same species. The uncultured fungus clone 3241N21 (KF617423) isolated from Picea mariana forest soil mineral horizon in the USA (Taylor et al. 2014) and clone IU-FSC Fun28b_FuD061 (AB520377) isolated from soil in Japan (Nishizawa et al. 2010) have two to three nt difference from XZY87M1 in the ITS region, which indicated that the species represented by XZY87M1 is worldwide distributed. Vishniacozyma sp. Anhui_B3, Vishniacozyma sp. Anhui_B10, Vishniacozyma sp. Anhui_B13 and an uncultured fungus clone IU-FSC Fun27_FuC053 have three nt in the D1/D2 domains and four nt in the ITS region with T1-6-1 and XZY91-8, which indicated that they are the same species.

Teunia (Cryptococcaceae, Tremellales)

Eight strains, LJGSL37B1, XZY88-7, XZY100M2, XZY129-2, XZY194-1, XZY380-2, XZY380-5 and XZY565-4, formed five groups in the genus Teunia (Figs 1C and 11). Strains XZY380-2 and XZY380-5 with the same D1/D2 sequence and similar ITS (2 nt differences), and differed from XZY100M2 by one nt in the D1/D2 domains and by 20 nt (~3.3%) in the ITS region, which indicated that those three strains belong to two different species. The two groups were closely related to T. korlaensis, T. nitrariae and Fungal sp. ARIZ:P251 (Figs 1C and 11), and differed them by more than four nt in the D1/D2 domains, and by more than 18 nt (~3.1%) in the ITS region. Strain LJGSL37B1 has an identical D1/D2 sequence and one nt ITS difference with Teunia sp. NYUN 2111141, which indicated that they are conspecific. They formed a separate branch closely related to T. korlaensis, T. nitrariae and T. helanensis (Figs 1C and 11), and differed from them by more than 7 nt (~1.4%) in the D1/D2 domains and by more than 29 nt (~6.2%) in the ITS region. Strain XZY194-1 was closely related to T. tronadorensis (Fig. 11), and differed from it by 13 nt (~2%) in the D1/D2 domains and by 35 nt (~5.5%) in the ITS region.

Strains XZY88-7, XZY129-2 and XZY565-4 formed a separate subclade with T. lichenophila (Figs 1C and 11) and three unpublished strains, ‘Kwoniella’ sp. 7, 12A and 23 (Fig. 11). XZY88-7 differed from XZY565-4 and XZY129-2 by 12 nt in the D1/D2 domains and 46 nt (~8%) in the ITS region. Strain XZY565-4 differed from XZY129-2 by one nt in the D1/D2 domain, but by 10 nt (~1.2%) in the ITS region, which indicated that they may belong to different species. Strains XZY88-7 had an affinity to the unpublished species ‘Kwoniella’ sp. 23 (Fig. 11), and differed from it by 6 nt (~1%) in the D1/D2 and by 16 (~2.7%) in the ITS region. Strains XZY88-7 and XZY129-2 were more related to T. lichenophila (Fig. 11), and differed from it by 9–11 nt (~1.6–2%) in the D1/D2 domains and by more than 31 nt (~5%) in the ITS region.

Note: Two Janpanses strains Teunia sp. MUCC2071 (LC715721) isolated from the flower of Nerium oleander and MUCC1912 (LC715712) isolated from the flower of Prunus mume, ‘Kwoniella’ sp. HB31-3 (KJ507251) isolated from the flower of Elaeagnus multiflora in South Korea and ‘Cryptococcus’ sp. BI226 (EU678944) in isolated from Brazil have an identical D1/D2 sequence with LJGSL37B1, which indicated that the species represented by LJGSL37B1 may frequently occur in flowers and is distributed worldwide.

Kwoniella (Cryptococcaceae, Tremellales)

Three strains isolated from Tibet were located in the genus Kwoniella and formed two groups (Figs 1C and 11). Strains XZY162-1 and XZY214M1 with identical ITS sequence and one nt D1/D2 difference were related to K. newhampshirensis (Fig. 11), and differed from it by four nt in the D1/D2 domains and by 13 nt (~2.6%) in the ITS region. Strain XZY364-6 had a close relationship with K. endophytica (Fig. 11), and differed from it by five nt and 27 nt (~4%) in the D1/D2 domains and ITS region, respectively.

Phaeotremella (Phaeotremellaceae, Tremellales)

Strain XZY502-4 was located in the Phaeotremella (Figs 1C and 12), closely related to P. fuscosuccinea (Fig. 12), and differed from it by 4 nt in the D1/D2 domains and by 43 nt (~8%) in the ITS region.

Note: the uncultured fungus clone rcw_144 (KM104080) isolated from red-cockaded woodpecker excavation in living pine in the USA has three nt difference in the ITS region with XZY502-4, which indicated that this new species occurred in the USA.

Vanrija (Trichosporonaceae, Trichosporonales)

Strain WYS 38-3 isolated from Wuying mountain in Fujian province occurred in the Vanrija clade (Figs 1C and 13), and was closely related to V. meifongana (Fig. 13). The former differed from the latter by 9 nt (~1.7%) in the D1/D2 domains and by 22 nt (~4.4%) in the ITS region.

Takashimella (Tetragoniomycetaceae, Trichosporonales)

The genus Takashimella includes four species, namely T. formosensis, T. koratensis, T. lagerstroemiae and T. tepidaria (Wang and Wang 2015). Only five and four nt differences were found in the D1/D2 domains and ITS region, respectively, between T. formosensis and T. koratensis, however, those two species can be distinguished by the difference in GC% content (57.8 mol% for T. formosensis and 49.4 mol% for T. koratensis) (Fungsin et al. 2006). Strain 21812S was located in the Takashimella and closely related to T. formosensis and T. koratensis (Figs 1C and 13), and differed from them by 4 nt in the D1/D2 domains and by 4–7 nt in the ITS region, which indicated that it is a different species from T. formosensis and T. koratensis.

Holtermannia (Holtermanniaceae, Holtermanniales)

The genus Holtermannia includes H. corniformis and H. saccardoi (Li et al. 2020). Strain 21825S-11 was closely related to H. saccardoi (Figs 1D and 13). They differed from each other by zero nt in the D1/D2 domains and by 12 nt (~2%) in the ITS region.

Piskurozyma (Piskurozymaceae, Filobasidiales)

Eight strains, NYG10-7, NYG10-7A, NYG26-1, XZY30M1, XZY134-2, XZY281-2, XZY321-2 and XZY321-4, formed six groups in the Piskurozyma (Figs 1D and 14). Strain XZY321-2 was closely related to two unpublished strains, Piskurozyma sp. KBP Y-4629 isolated from Eriophorum angustifolium and Piskurozyma sp. KBP Y-4633 (ON263269) isolated from soil under Carex ensifolia in Russia and an uncultured fungus clone 91 NA3 P32 N4 (KC965725) isolated from soil in Canada (Fig. 14), and differed from them by three to five nt in the D1/D2 domains and by 6–9 nt (1–1.5%) in the ITS region, which indicated that those two groups may be not conspecific. The BLASTn search showed that strain XZY321-4 had more similar ITS and D1/D2 sequences to P. fildesensis than other species, but this strain was closely related to P. sorana (Fig. 14) and differed from it by 15 nt (~3%) in the D1/D2 domains and by 68 (~11%) in the ITS region. Strains NYG10-7 and NYG10-7A formed a separate branch related to P. arborea, P. silvicultrix and XZY321-2 group (Fig. 14), and differed from them by more than 30 nt (6%) in the D1/D2 domains and by more than 14% in the ITS region. Strain XZY30M1 had an affinity to P. cylindrica (Fig. 14) and differed from it by two nt in the D1/D2 domains and 17 nt (~3%) in the ITS region. Strain NYG26-1 had the same D1/D2 sequence with P. filicatus, however, 12 nt (~2%) ITS sequence difference was found between them. Strain XZY134-2 was closely related to P. capsuligena (Fig. 14), and differed from it by 10 nt (~1.8%) the D1/D2 domains and by 29 nt (~4.6%) in the ITS region.

Note: the uncultured Basidiomycota clone man6_litter_F06 (GU328623) isolated from the Oe layer in the USA (Edwards et al. 2010), clone BTD-A2-ITSFL (GU082960) isolated from raw soil in the USA, clone M1-5-1-20 (LC096584) isolated from the root of Pyrola japonica in Japan have zero to four nt in the ITS region, which indicated that this species represented by XZY30M1 was distributed in the USA and Japan. ‘Cryptococcus cylindricus’ CRUB 1255 (DQ513289) isolated from Argentina has the same D1/D2 sequences with XZY30M1, which indicates that they may be the same species. Agaricomycotina NRS143 (AB726980) isolated from Japan and an uncultured fungus clone B05A10 (KC701830) isolated from forest soil in the USA have zero to two nt with XZY321-4 in the D1/D2 domains, which indicated that they may be conspecific. Tremellomycetes sp. strain bbo2p20 (MW448998) isolated from Vaccinium myrtillus in Finland (Nguyen et al. 2021), the uncultured fungus clone 2170_365 (KP891308) isolated from Scolytus multistriatus in Sweden (Menkis et al. 2016a), clone OTU_943 (KX223240) isolated from Pinus radiata forest roots and soils in New Zealand have zero to three nt in the ITS2 region, which indicated that this species represented by XZY321-2 was worldwide distributed. The uncultured fungus clone 454_1611 (KM494465) isolated from Norway-spruce stump in Sweden has the identical ITS2 sequence with NYG10-7 and NYG10-7A, which indicated that this species occurred in Sweden.

Mrakia (Mrakiaceae, Cystofilobasidiales)

Five strains, XZY289N6, XZY337-1, XZY319-4, XZY347-3 and XZY414-1, were located in the Mrakia clade and formed three separate groups both in ITS+D1/D2 and six-gene trees (Figs 1D and 15). Strains XZY337-1, XZY319-4 and XZY347-3 with the same ITS and D1/D2 sequences had an affinity to M. artica, M. stelviica, M. montana and M. cryoconite (Fig. 15), and differed from them by 5-8 nt in the D1/D2 domains and by 5-7 nt in the ITS region. Strains XZY414-1 and TP-Snow-Y57 have the same sequence both in the ITS region and D1/D2 domains, which indicated that they are conspecific. Strains XZY414-1 and XZY289N6 had the same D1/D2 sequence, but 10 nt differences in the ITS region between them, which indicated that they are different species.

New genus Foliozyma (Mrakiaceae, Cystofilobasidiales)

Strains XZY30-4 and XZY30-6 differed from each other by three and eight nt (~1.3%) in the D1/D2 domains and ITS region, respectively. The BLASTn results showed that XZY30-4 and XZY30-6 had an affinity to Cystofilobasidium species with 90% similarity in the D1/D2 domains and with 70–79% coverage and 82-84% similarity in the ITS region. XZY30-4, XZY30-6 and an uncultured fungus ASV 375 formed a separate branch in Mrakiaceae (Cystofilobasidiales) in the ITS+D1/D2 tree (Fig. 15). Six-gene phylogenetic analysis (Fig. 1D) was in agreement with the result constructed using the ITS+D1/D2 dataset. Two ‘Cystofilobasidium capitatum’ strains HAI-Y-404 and HAI-Y-699, Cystofilobasidium sp. OR 878, CLIB 3011 and four uncultured fungus clones A1b7660, A2e10666 and ASV 375 were clustered together with XZY30-4 and XZY30-6 in the D1/D2 tree (Fig. S1), and were related, whereas separated to the genus Cystofilobasidium. The above different approaches analyses indicated that those two strains represent a new genus in the Cystofilobasidiales.

Note: the uncultured fungus isolate OTU_350 (MK722946) isolated from soil in the USA has two to four nt ITS2 sequences difference with XZY30-6 and XZY30-4, which indicated that this genus represented by XZY30-6 occurs in the USA.

Phaffia (Mrakiaceae, Cystofilobasidiales)

Two strains isolated from Tibet were located in the Phaffia clade (Figs 1D and 15). Strain XZY12-2 clustered with two ‘Xanthophyllomyces dendrorhous’ strains CRUB 0853 and CRUB 1149 with two nt D1/D2 sequence and 10 nt ITS sequence differences, which indicated they may be different species. Strain XZY12-2 was closely related to the known species P. rhodozyma (Fig. 15), and differed from it by one nt in the D1/D2 domains and by 19 nt (~3.2%) in the ITS region. Strain XZY261N1 had an affinity to P. tasmanica (Fig. 15), and differed from it by three nt in the D1/D2 domains and 35 nt (~6.8%) in the ITS region.

New species identification in the Agaricostilbomycetes (Pucciniomycotina)

Kondoa (Kondoaceae, Agaricostilbales)

Strain MT-1B-33 was located in the genus Kondoa and was closely related to K. thailandica (Figs 2A and 16). The former differed from the latter by 14 nt (~2.8%) in the D1/D2 domains and by 36 nt (~7.5%) in the ITS region.

Boekhoutia (Chionosphaeraceae, Agaricostilbales)

The genus Boekhoutia was built by Li et al. (2020) based on only one species B. sterigmata. Strain MT-2C-38 isolated from Motuo in Tibet had an affinity to B. sterigmata (Figs 2A and 16) with four nt D1D2 difference and 58 nt (~9.2%) ITS diversity, which indicated that MT-2C-38 is the second member of the genus Boekhoutia.

New species identification in the Cystobasidiomycetes (Pucciniomycotina)

Buckleyzyma (Buckleyzymaceae, incertae sedis)

Strain YSR26-8 was located in the Buckleyzyma clade (Figs 2B and 17). This strain clustered with Cystobasidiomycetes sp. P44D004 isolated from Vitis vinifera in the USA (Bourret et al. 2013), and differed from it by zero nt in the D1/D2 domains and three nt in the ITS region, which indicated that they are conspecific. They had one nt D1/D2 sequence difference and 98.7% similarity in the ITS region with ‘Buckleyzyma aurantiaca’ RP432 13, ‘Rhodotorula’ sp. HB 1211 and five ‘Rhodotorula’ uncultured clones, KL 2w A02, KL 2d 2G04, KL 2d 2F03, SW 2d G11 and SW 2w B07, which indicated they may be not conspecific. Their taxonomic relationship is need more robust data including genome to resolve in the future.

Microsporomyces (Mycrosporomycetaceae, incertae sedis)

Two strains associated with the plant Betula sp., XZY554-7 and YN35N5, were located in the genus Microsporomyces (Figs 2B and 17). Strain XZY554-7 was isolated from the bark of Betula sp. collected in Tibet, whereas strain YN35N5 was isolated from of Betula sp. leaf in Yunnan province. Strain XZY554-7 clustered with two Cystobasidiomycetes sp. SOR11c5 and SOR12b5 isolated from a soredium of Cladonia fimbriata in the Czech Republic, and they had one nt difference in the D1/D2 domains and 12 nt (~2.5%) difference in the ITS region, which indicated that they may be different species. They were closely related to M. pini and M. orientalis (Fig. 17), and differed from them by 16 nt (~3%) in the D1/D2 domains and by more than 12% in the ITS region. Strain YN35N5 was closely related to an unpublished strain Microsporomyces sp. NYUN 2111170 (Fig. 17) with the same D1/D2 sequence and 97.6% ITS similarity, which indicated that they may be different species. Strain YN35N5 differed from the known Microsporomyces species by about 5% in the D1/D2 domains and by 103 nt (~16%) in the ITS region.

Halobasidium (Cystobasidiaceae, Cystobasidiales)

Strains YN28M2 and YN55-1 clustered with eight strains Cystobasidium sp. A2A4, A14A2, A15D1, A15D4, A15M2, Cystobasidiomycetes sp. JS-40, YM24636 and Halobasidium sp. MT254, and have similar D1/D2 (zero to one difference) and ITS (three to seven nt difference) sequences, which indicated that they may be conspecific. The two Chinese strains differed from H. xiangyangense by 10 nt (~1.8%) in the D1/D2 domains and by 13 nt (~2.3%) in the ITS region. (Figs 2B and 17)

Note: Cystobasidiomycetes sp. isolate R8 (MH237664) isolated from Paeonia lactiflora Pallas in China (Cheng et al. 2018) and BK318 (LR961819) isolated from Fraxinus excelsior in the Czech Republic have four nt difference in ITS with YN28M2 and YN55-1, which indicated that they belong to the same species. ‘Rhodotorula’ sp. YM24691 (JN936880) isolated from China has two nt differences in the D1/D2 domains, which indicated that they may be conspecific.

Cystobasidium (Cystobasidiaceae, Cystobasidiales)

Two strains XSR28-13 isolated from Qinghai province and XZY554-12 isolated from Tibet had similar ITS (one nt) and D1/D2 (one nt) sequences, and related to C. psychroaquaticum (Figs 2B and 17). They differed from the known species by seven nt (~1.3%) in the D1/D2 domains and by 14 nt (~2.3%) in the ITS region.

Note: Cystobasidium sp. AY952 (MG250437) isolated from the inner surface of termite's tapetum samples collected from Namib Desert fairy circles in Namibia has four nt difference from XSR28-13 and XZY554-12 in the ITS region, which indicated that they are conspecific.

New species identification in the Microbotryomycetes (Pucciniomycotina)

Rhodotorula (Sporidiobolaceae, Sporidiobolales)

Strain XZY511-5 was located in the Rhodotorula (Figs 2C and 18A). It had the same D1/D2 sequence and similar ITS sequences (zero to three nt) with four Rhodotorula sp. strains ITS1, LF1M14, LN1M04 and NL1 isolated from Taiwan, China, which indicated that they are conspecific. They differed from the known Rhodotorula species by seven nt (~1.3%) in the D1/D2 domains and by 24 nt (~4%) in the ITS region.

Note: ‘Rhodotorula toruloides’ PV18 (MW895335) isolated from soil in Costa Rica, uncultured fungus clone 3993_1335 (MW215459) isolated from rhizosphere in Lithuania (Marčiulynienė et al. 2021) and uncultured Rhodotorula clone NMF-OTU-85 (MG707557) isolated from roots in China have zero to three nt differences with XZY511-5 in the ITS region, which indicated that the species represented by XZY511-5 was worldwide distributed.

New genus Bauerozyma (Ustilentylomataceae, Microbotryales)

Strain YN25-3 formed a separate branch in the Microbotryales and was closely related to Microbotryozyma swertiae and Ustilentyloma graminis without support (Fig. 2C and 18A). The D1/D2 and ITS BLASTn results showed strain YN25-3 had less than 98% D1/D2 and 85.8% ITS similarity with the known Microbotryales species.

Note: the uncultured fungus clone BS.A2.8 (KY694702) isolated from the rhizosphere in Oman has the same ITS2 sequence as YN25-3, which indicated that this species occurs outside of China.

New genus Andreyozyma (incertae sedis)

The D1/D2 sequence BLASTn showed that strain XZY328-1 was more similar (about 88%) with Chrysozyma than other genera in the Microbotryomycetes, however, ITS BLASTn showed that this strain had an affinity to Phenoliferia and Libkindia with 75–86% coverage and 83–87% similarity. The inconsistant blast results and high diversity in both D1/D2 and region ITS indicated that XZY328-1 did not belong to any known genera in the Microbotryomycetes. (Figs 2C and 18A).

Note: The uncultured fungus clone BOTU112 (KC588812) isolated from the forest floor (leaf litter) in the northern hardwoods ecosystem in the USA (Cline and Zak 2014) has five nt difference in the D1/D2 domains and 12 nt (~3%) diversity in the ITS2 with XZY328-1, which indicated that an undescribed species in this genus represented by XZY328-1 occurred in the USA.

New genus Nakasozyma (incertae sedis)

Strains XZY99M13 and XZY22-1 had five nt in the D1/D2 domains and 29 nt (~5%) in the ITS region, which indicated that they belong to different species. Those two strains have less than 88–94% similarity in the D1/D2 domains with Oberwinklerozyma, Vonarxula javanica and Meredithblackwellia eburnean, however, the ITS BLASTn result showed that those two strains had an affinity to Leucosporidium intermedium and Yunzhangia auriculariae with less than 71–76 coverage and 82–83% similarity. The six-gene phylogenetic tree showed that they were closely related to the newly described genus Andreyozyma and the known genera Pseudoleucosporidium and Curvibasidium (Fig. 2C).

Note: ‘Rhodotorula’ sp. strains KBP:Y-6605 (MT013025) isolated from Cladonia cornuta and KBP:Y-6609 (MZ666410) isolated from Cladonia stellaris in Russia have identical D1/D2 sequence and 2 nt in the ITS region with XZY99M13, which indicated that they are conspecific

New genus Fengyania (incertae sedis)

Strains XZY23-5 and XZY30M2 had three and four nt differences in D1/D2 domains and the ITS region, respectively, which indicated that they are conspecific. The D1/D2 BLASTn showed that those two strains had 92–94% similarity with the genera Curvibasidium and Sampaiozyma, however, the ITS BLASTn showed that they had 80–82% coverage and 86–87% similarity with the genera Leucosporidium and Sampaiozyma. They formed a separate branch in the Microbotryomycetes (Figs 2C and 18A).

Note: ‘Rhodotorula’ sp. strains KBP Y-5569 (MK265711) and KBP Y-5465 (MK265710) isolated from Dryas punctata in Russia, ‘Rhodotorula’ sp. TSN-574 (MK307634) isolated from beech forest soil in Germany formed a well support clade with XZY30M2 and XZY23-5, which indicated that some members of the new genus represented by XZY23-5 are distributed worldwide.

Curvibasidium (incertae sedis)

Two strains XZY26-2 and XZY23-1 with four nt differences in the ITS region were located in the Curvibasidium (Figs 2C and 18B). They had an affinity to C. nothofagi and C. rogersii, and differed them by one nt in the D1/D2 and by 15–16 nt (~2.6%) in the ITS region.

Chrysozyma (Chrysozymaceae, incertae sedis)

Strain XZY20-10 was a member of Chrysozyma (Figs 2C and 18B). It was closely related to three unpublished strains Chrysozyma sp. 3831, 3836 and 3838, and Chrysozyma griseoflava (Fig.18B) by more than six nt in the D1/D2 domains and by more than 48 nt (~8%) in the ITS region.

Yurkovia (Chrysozymaceae, incertae sedis)

The genus Yurkovia include three species, namely Y. longicylindrica Y. mendeliana and Y. nerthusi (Li et al. 2020). Strain YN97-2 and Y. longicylindrica and Y. mendeliana formed a well support clade (Fig. 18B)., and differed from each other by three nt in the D1/D2 domains and by 37 nt (~6%) in the ITS region.

Note: the uncultured fungus clones 5320_639 (ON750211) isolated from Pinus sylvestris needles, 2170_499 (KP891331) isolated from Scolytus multistriatus and 3991_636 (MT266343) isolated from Quercus robur in Sweden (Menkis et al. 2022) and an uncultured Chlamydomonadales isolate soil 2642 (MF483850) isolated from soil in United Kingdom have zero to one nt difference in the ITS2 with YN97-2, which indicated that the species represented by YN97-2 is worldwide distributed.

Pseudohyphozyma (incertae sedis)

Strain XZY290-2 was located in the genus Pseudohyphozyma (Figs 2C and 18B), which differed from the known species by eight nt (~1.4%) in the D1/D2 domains and by 13 nt (~2.9%) in the ITS region.

Slooffia (incertae sedis)

Two strains, NYG09-2B and XZY423-3, have identical ITS and D1/D2 sequences. They were closely related to Slooffia cresolica (Figs 2C and 18B), and differed from it by five nt in the D1/D2 domains and by 12 nt (~2%) in the ITS region.

New genus Litoriozyma (Litoriozymaceae, Litoriozymales)

The BLASTn results showed that strain SHTS17-2 had less 89% and 83% similarity in the D1/D2 domains and ITS region, respectively, with the known species in the Microbotryomycetes. The ITS+D1/D2 and six-gene phylogenetic analyses showed that strain SHTS17-2 was related to the order Rosettozymales with low support value or without support (Figs 2C and 18B).

New taxa identification in the Ustilaginomycetes (Ustialginomycotina)

Ustilago (Ustilaginaceae, Ustilaginales)

Strain 2189S-11 was located in the genus Ustilago, and closely related to U. sparsa and U. cynodontis. The new strain differed from those two species by two to five nt in the D1/D2 domains and more than 39 nt (~94%) in the ITS region (Figs 3 and 19).

New taxa identification in the Exobasidiomycetes(Ustialginomycotina)

Acaromyces (Cryptobasidiaceae, Exobasidiales)

Strain MT-4B-41 had identical with Acaromyces ingoldii in the D1/D2 domains, however, they had ten nt (~1.2%) difference ITS sequence, which indicated that they may be different species.

Exobasidium (Exobasidiaceae, Exobasidiales)

The D1/D2 BLASTn showed that strain LJGSL4-20 had an affinity to E. hemisphaericum and E. rhododendri, and differed from them by five to eithe nt, the ITS BLASTn showed that this strain had less 95% similarity with the other Exobasidium species. The ITS+D1/D2 phylogenetic analysis showed that strain LJGSL4-20 formed a separate branch (Fig. 20).

Meira (Brachybasidiaceae, Exobasidiales)

Two strains 217S13D1 and HJLB11-102 formed two groups in the Meira clade (Figs 3 and 20). Strain 217S13D1 had an affinity with M. plantarum and M. nashicola, and differed from them by zero to one nt in the D1/D2 domains and by 8–24 nt (~4%) ITS region. Strain HJLB11-102 was closely related to M. pileae, and differed from it by two and 22 nt (~3.6%) in the D1/D2 domains and ITS region, respectively.

New genus Vankyiozyma (Vankyiozymaceae, Georgefischeriales)

The ITS BLASTn showed that strain MT-2A-31 had about 30% similarity with all Ustilaginomycotina species, which indicated that ITS1+ITS2 sequences were too diverse to fail to be aligned. The D1/D2 BLASTn showed less 94% similarity with the known species in the Georgefischeriales. This strain was more closely related to four unpublished strains, ‘Tilletiopsis’ sp. TY163, TY235, TY363 and ST105 isolated from phyllosphere in Thailand, with two to eight nt difference (Fig. 21). However, the ITS sequences of Thai strains are not available, therefore their taxonomic relationship with strain MT-2A-31 need to be verified in the future. The other six strains, ‘Tilletiopsis’ sp. TY289 isolated from Thailand, Entylomatales sp. NIP046 isolated from Japan and four Chinese strains Gjaerumia sp. NYNU 208150 Exobasidiomycetes sp. GZ1335, GZ1336 and GZ1871 represent at least two more undescribed species clustered with the clade represented by strain MT-2A-31 with middle statistic support (ML 71%). This newly clade was suggested to be a new family in the Exobasidiomycetes by Richter et al. (2019). The six-gene analysis showed that strain MT-2A-31 formed a separate branch in the Georgefischeriales (Fig. 3), which was in agreement with the suggestion from Richter et al. (2019), therefore a new genus and a new family will be proposed in the taxonomic section for strain MT-2A-31.

Taxonomy

New taxa in the Tremellomycetes (Agaricomycotina)

Derxomyces pseudoclindricus Q.-M. Wang, sp. nov. MycoBank MB846548. Figs 22 A, B.

Etymology: the specific epithet pseudclindricus refers to a similar colony morphology to that of Derxomyces clindricus.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical and ellipsoidal, 2.24–3.87 × 5.67–14.91 µm and single, budding is polar (Fig. 22A), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are subglobosal or ellipsoidal, 2.85–4.46 × 3.18–5.98 µm (Fig. 22B).

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose (weak), melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose (weak), L-rhamnose (weak), D-glucosamine (variable), N-acetyl-D-glucosamine, ethanol (variable), glycerol (variable), erythritol, ribitol (variable), galactitol (variable), D-mannitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid (variable), inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, D-glucitol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, D. pseudoclindricus differs from the closely related species D. cylindricus (Fig. 1A) in its inability to assimilate D-glucitol and sodium nitrite and the ability to assimilate lactose, inulin and erythritol (Table S2.1).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6875^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-3B-3).

Other culture examined: China, Tibet, Motuo county, leaf, Oct. 2021, isolated by Q.M. Wang, W.N. Zhao, culture MT-2A-35.

Derxomyces motuoensis Q.-M. Wang, sp. nov.MycoBank MB846551. Figs 22C, D.

Etymology: the specific epithet motuoensis refers to the geographic origin of the type strain, Motuo county, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.81–6.98 × 6.18–12.34 µm and single, budding is polar (Fig. 22C), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are napiform and reniform, 4.15–7.09 × 6.36–9.41 µm (Fig. 22D).

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose (weak), melibiose, raffinose, inulin (latent), soluble starch (latent), L-arabinose, D-arabinose, L-rhamnose, D-glucosamine (weak), N-acetyl-D-glucosamine (weak), ribitol (latent, weak), galactitol (latent, weak), D-mannitol, α-methyl-D-glucoside, DL-lactic acid, succinic aid, inositol (weak) and D-glucuronic acid (weak) are assimilated as sole carbon sources. Lactose, melezitose, D-xylose, D-ribose, methanol, ethanol, glycerol, erythritol, D-glucitol, salicin, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (latent, weak), ethylamine (weak) and cadaverine (weak) are assimilated as sole nitrogen sources. Sodium nitrite and L-lysine are not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, D. motuoensis differs from the closely related species D. hubeiensis, D. ovatus and D. taiwanicus (Fig. 1A) in its inability to assimilate melezitose, D-xylose, D-glucitol and L-lysine and the ability to assimilate D-glucosamine, N-acetyl-D-glucosamine, ethylamine and cadaverine (Table S2.1).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6874^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-5C-1).

Derxomyces orientalis Q.-M. Wang, sp. nov.MycoBank MB846552. Figs 22E, F.

Etymology: the specific epithet orientalis refers to the geographic origin of the type strain, China, eastern hemisphere.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, oval and subglobose, 2.39–4.84 × 3.46–6.66 µm and single, budding is polar (Fig. 22E), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are subglobosal to napiform, 2.53–6.90 × 3.86–9.30 µm (Fig. 22F).

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose (variable), sucrose (variable), maltose, cellobiose, trehalose (variable), lactose (variable), melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose (variable), L-arabinose, D-arabinose (latent), D-ribose (latent), L-rhamnose, D-glucosamine (variable), N-acetyl-D-glucosamine (variable), glycerol (variable), erythritol, ribitol, galactitol (variable), D-mannitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, inositol (variable) and D-glucuronic acid are assimilated as sole carbon sources. Methanol, ethanol, D-glucitol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine (latent, weak) and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, D. orientalis differs from the closely related species D. longicylindricus (Fig. 1A) in its inability to assimilate D-glucitol and the ability to assimilate D-arabinose, D-ribose, erythritol, ribitol, D-mannitol, salicin, DL-lactic acid, potassium nitrate and sodium nitrite (Table S2.1).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6871^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-1B-10).

Other culture examined: China, Tibet, Motuo county, leaf, Oct. 2021, isolated by Q.M. Wang, W.N. Zhao, culture MT-4A-7.

Derxomyces foliicola Q.-M. Wang, sp. nov.MycoBank MB846553. Figs 22G, H.

Etymology: the specific epithet foliicola refers to the type strain isolated from a leaf.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical and ellipsoidal, 2.65–4.32 × 3.46–7.63 µm and single, budding is polar (Fig. 22G), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are ellipsoidal, 2.24–3.99 × 3.63–6.50 µm (Fig. 22H).

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, melibiose, raffinose, melezitose, inulin (weak), soluble starch (weak), D-xylose, L-arabinose, D-arabinose (weak), L-rhamnose, D-glucosamine (latent, weak), N-acetyl-D-glucosamine, glycerol (latent, weak), ribitol (weak), D-mannitol, α-methyl-D-glucoside, salicin (latent, weak), DL-lactic acid, succinic aid, inositol (weak) and D-glucuronic acid (weak) are assimilated as sole carbon sources. Lactose, D-ribose, methanol, ethanol, erythritol, galactitol, D-glucitol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine (weak), ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, D. foliicola differs from the closely related species D. anomala (Fig. 1A) in its inability to assimilate lactose, D-ribose, L-sorbose, galactitol and D-glucitol and the ability to assimilate cellobiose, inulin, D-glucosamine, glycerol, ribitol, salicin, inositol and potassium nitrate (Table S2.1).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6872^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-3B-4).

Hannaella pteridophytae Q.-M. Wang, sp. nov.MycoBank MB846554. Fig. 22I.

Etymology: the specific epithet pteridophytae refers to Pteridophyta, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and subglobosal, 3.10–5.88 × 4.20–8.85 µm and single, budding is polar (Fig. 22I), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose (latent), sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, D-arabinose (latent), D-ribose, L-rhamnose (latent, weak), D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol (latent), erythritol (latent), ribitol, galactitol (latent), D-glucitol, α-methyl-D-glucoside (latent), salicin (latent, weak), DL-lactic acid, succinic aid, inositol (latent) and D-glucuronic acid (latent) are assimilated as sole carbon sources. L-arabinose, methanol, D-mannitol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (latent), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, H. pteridophytae differs from the closely related species H. taiwanensis (Fig. 4) in its inability to assimilate L-arabinose, D-mannitol and citric acid and the ability to assimilate inulin and sodium nitrite (Table S2.2).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Pteridophyta sp., Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6903^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN28-3).

Hannaella artemisiae Q.-M. Wang, sp. nov.MycoBank MB846555. Fig. 22J.

Etymology: the specific epithet artemisiae refers to Artemisia, the plant genus from which the strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.91–5.49 × 3.09–6.98 µm and single, budding is polar (Fig. 22J), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, glycerol (latent, weak), erythritol, ribitol (weak), galactitol (latent), D-mannitol, D-glucitol (latent), α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid (latent), inositol (latent) and D-glucuronic acid (latent) are assimilated as sole carbon sources. N-acetyl-D-glucosamine, methanol, ethanol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, H. artemisiae differs from the closely related species H. pagnoccae and H. dianchiensis (Fig. 4) in its ability to assimilate D-glucosamine and potassium nitrate (Table S2.2).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Bambusoideae, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6904^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN33-4).

Other culture examined: China, Yunnan province, Zhenxiong county, Dawan town, Shili village, leaf, Oct. 2019, isolated by Q.M. Wang, G.S. Wang, culture YN25-1.

Hannaella urticae Q.-M. Wang, sp. nov.MycoBank MB846556. Fig. 22K.

Etymology: the specific epithet urticae refers to Urtica, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.59–6.71 × 3.85–7.37 µm and single, budding is polar (Fig. 22K), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose (latent), maltose, cellobiose (latent), trehalose (latent), lactose (weak), melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose (latent), D-arabinose (latent), D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol, erythritol, ribitol, galactitol (latent), D-mannitol, D-glucitol, salicin (latent), DL-lactic acid, succinic aid, inositol (latent) and D-glucuronic acid (latent) are assimilated as sole carbon sources. Methanol, α-methyl-D-glucoside, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (latent, weak), L-lysine, ethylamine (latent) and cadaverine (latent) are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, H. urticae differs from the closely related species H. surugaensis (Fig. 4) in its inability to assimilate methyl-α-D-glucoside and citric acid and the ability to assimilate lactose, inulin, soluble starch, N-acetyl-D-glucosamine, ethanol, potassium nitrate and sodium nitrite (Table S2.2).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Urtica fissa, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6905^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN14M1).

Babevozyma Q.-M. Wang gen. nov.MycoBank MB846557.

Etymology: the genus is named in honour of IP Babeva for his proposal of the species of Candida podzolica.

This genus is proposed for the separate clade represented by Babevozyma podzolica. Member of the Trimorphomycetaceae (Tremellales,Tremellomycetes). The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate clade within the Trimorphomycetaceae (Fig. 1A).

Sexual reproduction not known. Colonies cream to yellowish-cream, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Babevozyma podzolica (Babeva & Reshetova) Q.-M. Wang.

Babevozyma podzolica (Babeva & Reshetova) Q.-M. Wang, comb. nov.MycoBank MB846625.

Basionym: Candida podzolica Babeva & Reshetova, Mikrobiologiya, Seriya B 44(2): 333 (1975).

Synonym: Saitozyma podzolica (Babeva & Reshetova) Xin Zhan Liu, F.Y. Bai, M. Groenew. & Boekhout, Stud. Mycol. 81: 128 (2015).

Babevozyma ninhbinhensis (D.T. Luong, M. Takash., Ty, Dung & Nakase) Q.-M. Wang, comb. nov.MycoBank MB846626.

Basionym: Bullera ninhbinhensis D.T. Luong, M. Takash., Ty, Dung & Nakase, Journal of Genetics and Applications, Vietnam (Special Issue) Biotechnology: 39 (2002).

Synonym: Saitozyma ninhbinhensis (D.T. Luong, M. Takash., Ty, Dung & Nakase) Yurkov, Stud. Mycol. 96: 137 (2020).

Babevozyma wallum (Gogorza Gondra, J. Kruse, McTaggart, Boekhout & R.G. Shivas) Q.-M. Wang, comb. nov.MycoBank MB846627.

Basionym: Saitozyma wallum Gogorza Gondra, J. Kruse, McTaggart, Boekhout & R.G. Shivas, Persoonia 42: 459 (2019).

Babevozyma hericia Q.-M. Wang, sp. nov.MycoBank MB846560. Fig. 22L.

Etymology: the specific epithet hericia refers to Hericium, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal, ellipsoidal and oval, 2.30–5.19 × 3.91–7.65 µm and single, budding is polar (Fig. 22L), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose (latent), D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol (latent, weak), galactitol, D-mannitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, glycerol, erythritol, ribitol, D-glucitol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (weak), sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 33 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, B. hericia differs from the closely related species B. podzolica (Fig. 5) in its inability to assimilate D-glucitol and the ability to assimilate inulin and potassium nitrate (Table S2.3).

Typus: China, Taohuachong mountain, Yingshan county, Hubei province, obtained from a Hericium erinaceus. Sep. 2021, Q.-M. Wang (holotype CGMCC 2.6869^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = DBS918-3-4).

Carlosrosaea rhododendri Q.-M. Wang, sp. nov.MycoBank MB846561. Fig. 22M.

Etymology: the specific epithet rhododendri refers to Rhododendron, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 1.53–4.58 × 2.67–5.29 µm and single, budding is polar (Fig. 22M), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is tawny, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose (latent), galactose, sucrose (latent), maltose (latent, weak), trehalose, lactose (latent, weak), melibiose, raffinose (latent), melezitose, inulin, D-xylose, L-arabinose, D-ribose, L-rhamnose (latent), D-glucosamine, N-acetyl-D-glucosamine (weak), erythritol (weak), ribitol, D-mannitol, D-glucitol, α-methyl-D-glucoside (weak), DL-lactic acid, succinic aid, inositol and D-glucuronic acid (latent) are assimilated as sole carbon sources. L-sorbose, cellobiose, soluble starch, D-arabinose, methanol, ethanol, glycerol, galactitol, salicin, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite and ethylamine are not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, C. rhododendri differs from the closely related species C. follicola (Fig. 5) in its inability to assimilate cellobiose, soluble starch, D-arabinose, ethanol, glycerol, galactitol, salicin and citric acid and the ability to assimilate inulin, potassium nitrate, L-lysine and cadaverine (Table S2.4).

Typus: China, Jiaozi snow mountain, Kunming, Yunnan province, obtained from a leaf of Rhododendron simsii. Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6895^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = JZXS7-21).

Carlosrosaea yunnanensis Q.-M. Wang, sp. nov.MycoBank MB846562. Fig. 22N.

Etymology: the specific epithet yunnanensis refers to the geographic origin of the type strain, Yunnan province.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 1.69–4.87 × 3.60–6.79 µm and single, budding is polar (Fig. 22N), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale white, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose (latent), galactose, sucrose (latent), maltose (latent, weak), trehalose, lactose (latent, weak), melibiose, raffinose (latent), melezitose, inulin, D-xylose, L-arabinose, D-ribose, L-rhamnose (latent), D-glucosamine, N-acetyl-D-glucosamine (weak), erythritol (weak), ribitol, D-mannitol, D-glucitol, α-methyl-D-glucoside (weak), DL-lactic acid, succinic aid, inositol and D-glucuronic acid (latent) are assimilated as sole carbon sources. L-sorbose, cellobiose, soluble starch, D-arabinose, methanol, ethanol, glycerol, galactitol, salicin, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite and ethylamine are not assimilated. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, C. yunnanensis differs from the all known species of Carlosrosaea in its inability to assimilate melibiose, D-xylose, L-arabinose and D-mannitol (Table S2.4).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a Pteridophyta sp., Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6913^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN28M1).

Carlosrosaea betulae Q.-M. Wang, sp. nov.MycoBank MB846563. Fig. 22O.

Etymology: the specific epithet betulae refers to Betula, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal, oval and ellipsoidal, 2.05–4.91 × 2.73–5.68 µm and single, budding is polar (Fig. 22O), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose (latent, weak), lactose, melibiose, melezitose, inulin (latent, weak), soluble starch, D-xylose (weak), L-arabinose (weak), D-arabinose (weak), D-ribose, L-rhamnose (weak), D-glucosamine, N-acetyl-D-glucosamine (latent), D-mannitol, α-methyl-D-glucoside (weak), succinic aid, citric acid and D-glucuronic acid are assimilated as sole carbon sources. Raffinose, methanol, ethanol, glycerol, erythritol, ribitol, galactitol, D-glucitol, salicin, DL-lactic acid, inositol and hexdecane are not assimilated. Ammonium sulfate (latent), potassium nitrate, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, C. betulae differs from the closely related species C. follicola and C. rhododendri (Fig. 1A) in its inability to assimilate raffinose, erythritol, ribitol, D-glucitol, DL-lactic acid and inositol and the ability to assimilate L-sorbose and ethylamine (Table S2.4).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Betula sp. Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6900^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN35-7).

Kockovaella foliicola Q.-M. Wang, sp. nov.MycoBank MB846564. Figs 22P, 23A.

Etymology: the specific epithet foliicola refers to the substrate origin of the type strain, leaves.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and ellipsoidal, 1.55–3.67 × 2.94–11.32 µm and single, budding is polar (Fig. 22P), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale-white, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are subglobosal and ellipsoidal, 2.21–3.36 × 3.28–4.23 µm (Fig. 23A)

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, D-glucitol (weak), α-methyl-D-glucoside, succinic aid, inositol (weak) and D-glucuronic a cid are assimilated as sole carbon sources. L-sorbose, D-arabinose, D-ribose, methanol, ethanol, glycerol, erythritol, ribitol, galactitol, D-mannitol, salicin, DL-lactic acid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, K. foliicola differs from the closely related species K. barringtoniae (Fig. 1B) in its inability to assimilate L-sorbose, D-arabinose, D-ribose, ethanol, glycerol, erythritol, ribitol, galactitol, D-mannitol, salicin and citric acid and the ability to assimilate inulin, potassium nitrate, ethylamine and cadaverine (Table S2.5).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6867^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-2C-31).

Kockovaella silvicultrix Q.-M. Wang, sp. nov.MycoBank MB846565. Figs 23B, C.

Etymology: the specific epithet silvicultrix refers to the substrate origin of the type strain, rotten wood.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and oval, 2.40–4.94 × 3.56–7.03 µm and single, budding is polar (Fig. 23B), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is whiteish-cream, butyrous, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are subglobosal, 2.11–3.32 × 3.16–4.24 µm (Fig. 23C).

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-Sorbose, sucrose, maltose, cellobiose, trehalose, raffinose (weak), melezitose, inulin, soluble starch, D-Xylose, L-Arabinose, D-Arabinose, D-Ribose, L-Rhamnose, D-Glucosamine, N-Acetyl-D-glucosamine, erythritol, ribitol, galactitol (latent, weak), D-Mannitol, D-Glucitol, Methyl-α-D-glucoside, salicin (latent), succinic acid (latent, weak), inositol and D-Glucuronic acid (latent) are assimilated as sole carbon sources. Lactose, melibiose, methanol, ethanol, glycerol, DL-Lactic acid, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, ethylamine (latent, weak) and cadaverine (latent) are assimilated as sole nitrogen sources. Sodium nitrite and L-lysine are not assimilated. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, K. silvicultrix differs from the closely related species K. Mexicana, K. lichenicola, K. chinensis, K. sichuanensis, K. calophylli, K. cucphuongensis, K. vietnamensis, K. litseae, K. schimae and K. ogasawarensis (Fig. 1B) in its inability to assimilate lactose and citric acid and the ability to assimilate methyl-α-D-glucoside and potassium nitrate (Table S2.5).

Typus: China, Naqu area, Tibet, obtained from a rotten wood, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6892^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = RTFM4-1).

Papiliotrema catalpae Q.-M. Wang, sp. nov.MycoBank MB846566. Fig. 23D.

Etymology: the specific epithet catalpae refers to Catalpa, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and oval, 2.74–5.34 × 3.18–6.61 µm and single, budding is polar (Fig. 23D), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale-yellow, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose (latent), sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose (latent), L-arabinose, D-arabinose, D-ribose (latent), L-rhamnose, D-glucosamine (latent), N-acetyl-D-glucosamine, ethanol (latent, weak), ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside (latent, weak), salicin (latent, weak), DL-lactic acid, succinic aid, inositol (latent, weak) and D-glucuronic acid are assimilated as sole carbon sources. Methanol, glycerol, erythritol, citric acid and hexdecane are not assimilated. Ammonium sulfate (latent), potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine (latent) are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. catalpae differs from the closely related species P. bandonii, P. siamensis, P. perniciosus and P. nemorosa (Fig. 1B) in its inability to assimilate citric acid and the ability to assimilate inulin and potassium nitrate (Table S2.6).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Catalpa ovata, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6897^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN109M3).

Papiliotrema pseudolaurentii Q.-M. Wang, sp. nov.MycoBank MB846567. Fig. 23E.

Etymology: the specific epithet pseudolaurentii refers to the similar colony morphology to that of Papiliotrema laurentii.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal, ellipsoidal and oval, 2.24–5.40 × 2.98–6.39 µm and single, budding is polar (Fig. 23E), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, smooth and dimmed. The margin is wavy. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol, ribitol, galactitol, D-mannitol, D-glucitol (weak), α-methyl-D-glucoside, salicin, DL-lactic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, erythritol, succinic aid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 37 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. pseudolaurentii differs from the closely related species P. laurentii (Fig. 1B) in its inability to assimilate erythritol, succinic acid and citric acid and the ability to assimilate inulin, potassium nitrate (Table S2.6).

Typus: China, Motuo county, Tibet, obtained from a bark of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6866^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = motuo45-1).

Papiliotrema castaneae Q.-M. Wang, sp. nov.MycoBank MB846568. Fig. 23F.

Etymology: the specific epithet castaneae refers to Castanea, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 1.79–5.54 × 2.83–6.44 µm and single, budding is polar (Fig. 23F), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose (latent), cellobiose (latent), trehalose, lactose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose (latent), D-arabinose (latent), D-ribose, ethanol (latent), glycerol (latent, weak), erythritol, ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside (latent), salicin, DL-lactic acid, inositol (latent) and D-glucuronic acid are assimilated as sole carbon sources. Melibiose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, succinic aid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine (latent), ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. castaneae differs from the closely related species P. aspenensis, P. pseudolaurentii, P. laurentii, P. aspenensis, P. phichitensis, and P. rajasthanensis (Fig. 6) in its inability to assimilate melibiose, L-rhamnose, D-glucosamine and N-acetyl-D-glucosamine (Table S2.6).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Castanea mollissima, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6898^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN83-2).

Pseudotremella hippophae Q.-M. Wang, sp. nov.MycoBank MB846569. Fig. 23G.

Etymology: the specific epithet hippophae refers to Hippophae, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal, ellipsoidal and oval, 1.92–5.43 × 3.51–6.95 µm and single, budding is polar (Fig. 23G), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellowish-cream, butyrous, crease and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose (latent), raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol, erythritol (latent, weak), galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, succinic aid, inositol (latent) and D-glucuronic acid are assimilated as sole carbon sources. Melibiose, methanol, ribitol, DL-lactic acid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (variable), L-lysine, ethylamine (weak) and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. hippophae differs from the closely related species P. lacticolor (Fig. 7) in its inability to assimilate melibiose, ribitol and citric acid and the ability to assimilate potassium nitrate and cadaverine (Table S2.7).

Typus: China, Qilian mountain, Qinghai province, obtained from a bark of Hippophae rhamnoides, Jun. 2019, Q.-M. Wang (holotype CGMCC 2.6838^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XSR10-2).

Other culture examined: China, Qinghai province, Qilian mountain, bark of Quercus acutissima, Jun. 2019, isolated by Q.M. Wang, G.S. Wang, culture XSR1-5; Tibet, Lhasa, Norbu Lingka, leaf of Tacamahaca, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY113-6.

Pseudotremella rhododendri Q.-M. Wang, sp. nov.MycoBank MB846570. Fig. 23H.

Etymology: the specific epithet rhododendri refers to Rhododendron, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 1.83–4.51 × 3.01–6.97 µm and single, budding is polar (Fig. 23H), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, smooth and faint glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melezitose, inulin, soluble starch, D-xylose, L-arabinose (latent), D-arabinose (latent), D-ribose, L-rhamnose, D-glucosamine (latent), ethanol, glycerol, galactitol (latent), D-mannitol, salicin (latent), DL-lactic acid, succinic aid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Melibiose, raffinose, N-acetyl-D-glucosamine, methanol, erythritol, ribitol, D-glucitol, α-methyl-D-glucoside, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (latent, weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is negative. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. rhododendri differs from the closely related species P. moriformi and P. nivalis (Fig. 7) in its inability to assimilate raffinose, ribitol, D-glucitol and citric acid and the ability to assimilate inulin, ethanol and potassium nitrate (Table S2.7).

Typus: China, Baiba county, Tibet, obtained from a leaf of Rhododendron simsii, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6854^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY97M2).

Baiozyma Q.-M. Wang gen. nov.MycoBank MB846571.

Etymology: the genus is named in honour of Feng-Yan Bai for his research contributions to yeast taxonomy.

This genus is proposed for the separate branch represented by strain YN107M4. Member of the Tremellales (Tremellomycetes). The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate branch closely relate to the genus Fonsecazyma and Sirobasidium intermedium within the Tremellales (Fig. 1B).

Sexual reproduction not known. Colonies yellowish-cream, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Baiozyma castanea Q.-M. Wang.

Baiozyma castanea Q.-M. Wang, sp. nov.MycoBank MB846572. Fig. 23I.

Etymology: the specific epithet castanea refers to Castanea, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal and oval, 1.96–5.27 × 2.57–5.65 µm and single, budding is polar (Fig. 23I), sediment and pellicle are formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth, bulging and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (latent), L-sorbose (latent), sucrose, maltose, cellobiose (latent), trehalose, lactose (latent), raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, ethanol, glycerol (latent), erythritol (latent), ribitol, galactitol (latent), D-mannitol, D-glucitol (latent), α-methyl-D-glucoside (latent), salicin (latent), DL-lactic acid, succinic aid, inositol and D-glucuronic acid (latent) are assimilated as sole carbon sources. Melibiose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine (latent) and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, leaf of Castanea mollissima, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6899^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN107M4).

Corticimyces Q.-M. Wang gen. nov.MycoBank B846573.

Etymology: the specific epithet Corticimyces refers to the origin of the substrate of the species, cortex.

This genus is proposed for the separate branch represented by the strain XSR1-14. Member of the Sirobasidiaceae (Tremellales,Tremellomycetes). The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate branch closely related to the genus Fibulobasidium and Sirobasidium magnum within the Sirobasidiaceae (Fig. 1B).

Sexual reproduction not known. Colonies cream to cream, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Corticimyces cunninghamiae Q.-M. Wang.

Corticimyces cunninghamiae Q.-M. Wang, sp. nov.MycoBank MB846574. Fig. 23J.

Etymology: the specific epithet cunninghamiae refers to Cunninghamia, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval, ellipsoidal and subglobosal 2.57–4.94 × 3.91–5.65 µm and single, budding is polar (Fig. 23J), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is whiteish-cream, butyrous, the surface of microbiota has streak and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol, erythritol (latent, weak), ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid (latent, weak), succinic aid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Galactose, melezitose, methanol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (latent, weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Qilia mountain, Qinghai province, obtained from a bark of Cunninghamia sp., Jun. 2019, Q.-M. Wang (holotype CGMCC 2.6840^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XSR1-14).

Naematelia pseudoencephala Q.-M. Wang, sp. nov.MycoBank MB846575. Fig. 23K.

Etymology: the specific epithet pseudoencephala refers to the similar colony morphology to that of Naematelia encephala.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal, oval and globosal, 3.15–6.57 × 4.06–7.63 µm and single, budding is polar (Fig. 23K), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale white, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose (latent), raffinose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol (latent), glycerol (weak), erythritol, ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid (latent, weak), succinic aid (latent), citric acid (latent, weak), inositol (latent, weak) and D-glucuronic acid are assimilated as sole carbon sources. Melezitose, D-ribose, methanol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, N. pseudoencephala differs from the closely related species N. encephala (Fig. 9) in its inability to assimilate melezitose and D-ribose and the ability to assimilate inulin, erythritol, potassium nitrate and sodium nitrite (Table S2.8).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from the bark of Huodendron tibeticum, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6844^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY420-1).

Vishniacozyma sinopodophylli Q.-M. Wang, sp. nov.MycoBank MB846576. Fig. 23L.

Etymology: the specific epithet sinopodophylli refers to Sinopodophyllum, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and subglobosal, 2.22–5.06 × 3.49–7.53 µm and single, budding is polar (Fig. 23L), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale pink, butyrous, smooth and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-Acetyl-D-glucosamine, glycerol, erythritol, ribitol, galactitol, D-mannitol, Methyl-α-D-glucoside, salicin, DL-lactic acid, succinic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, ethanol, D-glucitol, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. sinopodophylli differs from the closely related species V. victoriae and V. carnescens in its inability to assimilate D-glucitol and citric acid, and the ability to assimilate inulin and potassium nitrate.

V. sinopodophylli differs from the closely related species V. paravictoriae (Fig. 1B) in its inability to assimilate ethanol, and the ability to assimilate sucrose, lactose, melibiose, raffinose, D-glucosamine, N-acetyl-D-glucosamine and galactitol (Table S2.9).

Typus: China, Baiba county, Tibet, obtained from a leaf of Sinopodophyllum hexandrum, Aug.2019, Q.-M. Wang (holotype CGMCC 2.6857^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY87M1).

Vishniacozyma paravictoriae Q.-M. Wang, sp. nov.MycoBank MB846577. Fig. 23M.

Etymology: the specific epithet paravictoriae refers to the similar colony morphology to that of Vishniacozyma victoriae.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, 1.92–4.46 × 3.34–6.41 µm and single, budding is polar (Fig. 23M), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale amber, mucilaginous, smooth and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, maltose, cellobiose, trehalose, melezitose (latent, weak), inulin, soluble starch, D-xylose, L-arabinose (weak), D-arabinose, D-ribose, L-rhamnose (latent weak), ethanol (weak), glycerol, erythritol, ribitol (weak), D-mannitol, Methyl-α-D-glucoside (weak), salicin, DL-lactic acid, succinic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Sucrose, lactose, melibiose, raffinose, D-glucosamine, N-Acetyl-D-glucosamine, methanol, galactitol, D-Glucitol, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 17ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. paravictoriae differs from the closely related species V. victoriae (Fig. 1B) in its inability to assimilate sucrose, lactose, melibiose, raffinose, D-glucosamine, galactitol, D-glucitol and citric acid and the ability to assimilate inulin, ethanol, DL-lactic acid and potassium nitrate (Table S2.9).

Typus: China, Nanyigou, Linzhi, Tibet, obtained from soil, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6918^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = NYG10-6).

Vishniacozyma pyri Q.-M. Wang, sp. nov.MycoBank MB846578. Fig. 23N.

Etymology: the specific epithet pyri refers to Pyrus, the plant genus from which the strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal, subglobosal and ellipsoidal, 2.57–4.32 × 2.94–6.49 µm and single, budding is polar (Fig. 23N), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, smooth, protruding and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-Acetyl-D-glucosamine (variable), glycerol (variable), erythritol, ribitol, galactitol, D-mannitol, D-glucitol (variable, weak), Methyl-α-D-glucoside, salicin, DL-lactic acid, succinic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, ethanol, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. pyri differs from the closely related species V. tephrensis (Fig. 1B) in its inability to assimilate citric acid and the ability to assimilate inulin and potassium nitrate (Table S2.9).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6870^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = T1-6-1).

Other culture examined: China, Tibet, Baiba county, leaf of Pyrus sorotina, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY91-8.

Vishniacozyma catalpae Q.-M. Wang, sp. nov.MycoBank MB846579. Fig. 23O.

Etymology: the specific epithet catalpae refers to Catalpa, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 1.76–4.52 × 2.98–5.88 µm and single, budding is polar (Fig. 23O), a sediment is formed. After 1 month at 17 ºC, pellicle and sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale pink, butyrous, smooth and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-ribose, L-rhamnose, N-acetyl-D-glucosamine, glycerol (latent, weak), erythritol, ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. D-arabinose, D-glucosamine, methanol, ethanol, succinic aid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. catalpae differs from the closely related species V. tephrensis, V. victoriae, V. paravictoriae, V. sinopodophylli, V. pyri and V. carnescens (Fig. 1B) in its inability to assimilate D-arabinose and succinic acid (Table S2.9).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Catalpa ovata, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6902^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN110-1).

Vishniacozyma marinae Q.-M. Wang, sp. nov.MycoBank MB846580. Fig. 23P.

Etymology: the specific epithet marinae refers to seawater, the substrate origin of the type strain.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, 2.17–3.58 × 3.28–5.23 µm and single, budding is polar (Fig. 23P), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, smooth and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose (latent), D-ribose, L-rhamnose, D-glucosamine, N-Acetyl-D-glucosamine, glycerol, erythritol, ribitol (latent, weak), D-mannitol, D-glucitol, Methyl-α-D-glucoside, salicin, DL-lactic acid, succinic acid, inositol (latent) and D-glucuronic acid (latent) are assimilated as sole carbon sources. Methanol, ethanol, galactitol, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. marinae differs from the closely related species V. pseudopenaeus (Fig. 1B) in its inability to assimilate galactitol and citric acid and the ability to assimilate inulin (Table S2.9).

Typus: China, Mulongwan of Fuoluo town, Ledong county, Hainan province, obtained from seawater, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6837^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MLWB11-2).

Other culture examined: China, Hainan province, Ledong county, Fuoluo town, Mulongwan, seawater, Feb. 2022, isolated by Q.M. Wang, W.N. Zhao, culture MLWB2-1.

Vishniacozyma pini Q.-M. Wang, sp. nov.MycoBank MB846581. Fig. 24A.

Etymology: the specific epithet pini refers to Pinus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.62–5.12 × 3.35–6.57 µm and single, budding is polar (Fig. 24A), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellow, butyrous, smooth and pale glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose (latent), trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, L-arabinose, D-arabinose, L-rhamnose, D-glucosamine, N-Acetyl-D-glucosamine, erythritol, ribitol, galactitol, D-mannitol, D-glucitol (latent, weak), Methyl-α-D-glucoside, salicin, DL-lactic acid, succinic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Galactose, D-xylose, D-ribose, methanol, ethanol, glycerol, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. pini differs from the closely related species V. peneaus (Fig. 1B) in its inability to assimilate galactose, D-xylose, D-ribose, glycerol and citric acid and the ability to assimilate inulin, DL-lactic acid and potassium nitrate (Table S2.9).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a bark of Pinus sp., Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6849^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY496-1).

Vishniacozyma zhenxiongensis Q.-M. Wang, sp. nov.MycoBank MB846582. Fig. 24B.

Etymology: the specific epithet zhenxiongensis refers to the geographic origin of the type strain, zhenxiong county, Yunnan province.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal, oval and ellipsoidal, 1.99–5.18 × 2.81–5.92 µm and single, budding is polar (Fig. 24B), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is amber, butyrous, smooth and light glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-Acetyl-D-glucosamine, erythritol, ribitol, galactitol, D-mannitol, Methyl-α-D-glucoside, salicin, DL-lactic acid, succinic acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, ethanol, glycerol, D-glucitol, citric acid and hexadecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. zhenxiongensis differs from the closely related species V. taibaiensis (Fig. 10) in its inability to assimilate D-glucitol and the ability to assimilate L-sorbose, soluble starch and potassium nitrate (Table S2.9).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Pteridophyta sp., Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6901^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN95-2).

Teunia myricariae Q.-M. Wang, sp. nov.MycoBank MB846583. Fig. 24C.

Etymology: the specific epithet myricariae refers to Myricaria, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.75–5.77 × 4.86–9.06 µm and single, budding is polar (Fig. 24C), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is amber, butyrous, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, maltose, cellobiose (latent), trehalose, inulin, soluble starch (variable), D-xylose, L-arabinose, D-ribose (variable), L-rhamnose (variable), ethanol (variable), ribitol (variable), galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin (latent, weak), succinic aid (variable), inositol (variable) and D-glucuronic acid (weak) are assimilated as sole carbon sources. Sucrose, lactose, melibiose, raffinose, melezitose, D-arabinose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, DL-lactic acid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (weak), L-lysine (variable) and cadaverine (variable) are assimilated as sole nitrogen sources. Sodium nitrite and ethylamine are not assimilated. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. myricariae differs from the closely related species T. chimonanthi (Fig. 1C) in its inability to sucrose, lactose, melezitose and ethylamine and the ability to assimilate D-glucuronic acid and potassium nitrate (Table S2.10).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a leaf of Myricaria laxiflora, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6846^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY380-2).

Other culture examined: China, Tibet, Linzhi, Zhenxiong county, New basom lake, leaf of Myricaria laxiflora, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY380-5.

Teunia chimonanthi Q.-M. Wang, sp. nov.MycoBank MB846584. Fig. 24D.

Etymology: the specific epithet chimonanthi refers to Chimonanthus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, subglobosal and oval, 2.81–5.47 × 4.76–7.76 µm and single, budding is polar (Fig. 24D), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is amber, butyrous, smooth and dimmed. The margin is wavy. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, L-rhamnose, galactitol, D-mannitol, D-glucitol (latent, weak), α-methyl-D-glucoside, salicin (weak) and succinic aid are assimilated as sole carbon sources. Melibiose, raffinose, D-arabinose, D-ribose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, glycerol, erythritol, ribitol, DL-lactic acid, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, ethylamine and cadaverine (latent, weak) are assimilated as sole nitrogen sources. Potassium nitrate, sodium nitrite and L-lysine are not assimilated. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. chimonanthi differs from the closely related species T. myricariae (Fig. 1C) in its inability to assimilate D-glucuronic acid and potassium nitrate and the ability to assimilate sucrose, lactose, melezitose and ethylamine (Table S2.10).

Typus: China, Baiba county, Tibet, obtained from a leaf of Chimonanthus praecox, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6916^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY100M2).

Teunia rhododendri Q.-M. Wang, sp. nov.MycoBank MB846585. Fig. 24E.

Etymology: the specific epithet rhododendri refers to Rhododendron, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal, subglobosal and oval, 2.28–5.72 × 3.35–7.63 µm and single, budding is polar (Fig. 24E), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellowish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, raffinose, melezitose, inulin, soluble starch, D-xylose, D-ribose, L-rhamnose, ethanol, ribitol, galactitol, D-mannitol, D-glucitol, DL-lactic acid, inositol and D-glucuronic acid (latent, weak) are assimilated as sole carbon sources. Melibiose, L-arabinose, D-arabinose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, α-methyl-D-glucoside, salicin, succinic aid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. rhododendri differs from the closely related species T. myricariae, T. chimonanthi, T. helanensis, T. korlaensis and T. nitrariae (Fig. 11) in its inability to assimilate L-arabinose and salicin and the ability to assimilate raffinose (Table S2.10).

Typus: China, Lijiang Alpine Botanical Garden, Yunnan province, obtained from a leaf of Rhododendron simsii, Feb, 2022, Q.-M. Wang (holotype CGMCC 2.6896^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = LJGSL37B1).

Teunia querci Q.-M. Wang, sp. nov.MycoBank MB846586. Fig. 24F.

Etymology: the specific epithet querci refers to Quercus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and oval, 2.28–6.59 × 4.04–8.73 µm and single, budding is polar (Fig. 24F), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale amber, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, maltose, cellobiose, trehalose, lactose, melezitose, inulin, soluble starch, D-xylose, L-arabinose (latent, weak), L-rhamnose, ethanol, galactitol, D-mannitol, α-methyl-D-glucoside, salicin (latent, weak), succinic aid and D-glucuronic acid are assimilated as sole carbon sources. Sucrose, melibiose, raffinose, D-arabinose, D-ribose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, ribitol, D-glucitol, DL-lactic acid, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (weak), L-lysine (latent, weak) and ethylamine are assimilated as sole nitrogen sources. Sodium nitrite and cadaverine are not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. querci differs from the closely related species T. tronadorensis (Fig. 11) in its inability to assimilate sucrose, melibiose, raffinose, D-arabinose, D-ribose, D-glucosamine, glycerol, ribitol, D-glucitol, Inositol and cadaverine and the ability to assimilate soluble starch and potassium nitrate (Table S2.10).

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Quercus semicarpifolia, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6959^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY194-1).

Teunia heritierae Q.-M. Wang, sp. nov.MycoBank MB846587. Fig. 24G.

Etymology: the specific epithet heritierae refers to Heritiera, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal, ellipsoidal and globosal, 2.78–6.29 × 4.05–7.83 µm and single, budding is polar (Fig. 24G), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose (latent), maltose, cellobiose, trehalose, lactose, melezitose, inulin, soluble starch (latent), D-xylose (latent), L-arabinose (latent), D-arabinose, D-ribose, L-rhamnose (latent), D-glucosamine (latent), N-acetyl-D-glucosamine, ethanol (latent), glycerol (latent), D-mannitol, D-glucitol (latent), α-methyl-D-glucoside, salicin, succinic aid (latent) and D-glucuronic acid are assimilated as sole carbon sources. Galactose, sucrose, melibiose, raffinose, methanol, erythritol, ribitol, galactitol, DL-lactic acid, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (latent, weak), L-lysine (latent), ethylamine (latent) and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. heritierae differs from the closely related species T. lichenophila and T. parabetulicola (Fig.11) in its inability to assimilate sucrose and galactitol and the ability to assimilate D-ribose (Table S2.10).

T. heritierae differs from the closely related species T. betulicola (Fig.11) in its inability to assimilate inositol and the ability to assimilate melezitose, D-xylose, methyl-α-D-glucoside and potassium nitrate (Table S2.10).

Typus: China, Baiba county, Tibet, obtained from a leaf of Heritiera littoralis, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6856^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY88-7).

Teunia betulicola Q.-M. Wang, sp. nov.MycoBank MB846588. Fig. 24H.

Etymology: the specific epithet betulicola refers to the substrate origin of the type strain, Betula.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal and oval, 3.42–5.23 × 4.42–6.35 µm and single, budding is polar (Fig. 24H), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is amber, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, maltose (latent), cellobiose, trehalose, lactose (latent), inulin, soluble starch (latent), L-arabinose (latent) D-arabinose (latent), D-ribose (latent), L-rhamnose (latent), D-glucosamine (latent), N-acetyl-D-glucosamine, ethanol (latent), glycerol (latent), D-mannitol, D-glucitol (latent), salicin (latent, weak), succinic aid (latent), inositol (latent, weak) and D-glucuronic acid are assimilated as sole carbon sources. Galactose, sucrose, melibiose, raffinose, melezitose, D-xylose, methanol, erythritol, ribitol, galactitol, α-methyl-D-glucoside, DL-lactic acid, citric acid and hexdecane are not assimilated. Ammonium sulfate, L-lysine (latent), ethylamine and cadaverine (latent, weak) are assimilated as sole nitrogen sources. Potassium nitrate and sodium nitrite are not assimilated. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. betulicola differs from the closely related species T. parabetulicola (Fig. 11) in its inability to assimilate sucrose, melezitose, galactitol, methyl-α-D-glucoside and potassium nitrate and the ability to assimilate D-ribose (Table S2.10).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a leaf of Betula platyphylla, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.7195^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY565-4).

Teunia parabetulicola Q.-M. Wang, sp. nov.MycoBank MB846589. Fig. 24I.

Etymology: the specific epithet parabetulicola refers to the similar colony morphology to that of Teunia betulicola.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal and oval, 2.73–5.83 × 2.99–6.90 µm and single, budding is polar (Fig. 24I), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melezitose, inulin, soluble starch, L-arabinose (latent), D-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol (latent), galactitol (latent, weak), D-mannitol, D-glucitol (latent), α-methyl-D-glucoside (latent), salicin, succinic aid (latent), inositol (latent) and D-glucuronic acid are assimilated as sole carbon sources. galactose, melibiose, raffinose, D-xylose, D-ribose, methanol, erythritol, ribitol, DL-lactic acid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. parabetulicola differs from the closely related species T. betulicola (Fig. 11) in its inability to assimilate D-ribose and the ability to assimilate sucrose, melezitose, galactitol, methyl-α-D-glucoside and potassium nitrate (Table S2.10).

Typus: China, Basom lake, Linzhi, Tibet, obtained from a leaf of Betula platyphylla, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6852^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY129-2).

Kwoniella hippophae Q.-M. Wang, sp. nov.MycoBank MB846590. Fig. 24J.

Etymology: the specific epithet hippophae refers to Hippophae, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are and globosal and subglobosal, 2.76–7.46 × 2.90–7.76 µm and single, budding is polar (Fig. 24J), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, raffinose, melezitose, inulin, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, ethanol, glycerol, ribitol (weak), galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside (weak), salicin, inositol and D-glucuronic acid are assimilated as sole carbon sources. Melibiose, soluble starch, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, DL-lactic acid, succinic aid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, K. hippophae differs from the closely related species K. endophytica (Fig .11) in its inability to assimilate L-sorbose, methyl-α-D-glucoside and salicin and the ability to assimilate DL-lactic acid and succinic acid (Table S2.11).

Typus: China, Naidong county, Tibet, obtained from a leaf of Hippophae rhamnoides, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6893^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY364-6).

Kwoniella lonicerae Q.-M. Wang, sp. nov.MycoBank MB846591. Fig. 24K.

Etymology: the specific epithet lonicerae refers to Lonicera, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and subglobosal, 3.05–7.95 × 4.05–9.65 µm and single, budding is polar (Fig. 24K), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle is present. On YM agar, after 1 month at 17 ºC, the streak culture is pale amber, butyrous, warty and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, raffinose, melezitose (variable), inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose (variable), L-rhamnose, N-acetyl-D-glucosamine, ethanol, glycerol (latent, weak), ribitol (latent, weak), galactitol (latent, weak), D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin (latent, weak), succinic aid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Melibiose, D-glucosamine, methanol, erythritol, DL-lactic acid, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine, ethylamine and cadaverine (variable) are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, K. lonicerae differs from the closely related species K. newhampshirensis (Fig. 11) in its ability to assimilate inositol and potassium nitrate (Table S2.11).

Typus: China, Sejila mountain, Linzhi, Tibet, obtained from a leaf of Lonicera japonica, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6883^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY162-1).

Other culture examined: China, Tibet, Baiba county, leaf of Lonicera japonica, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY214M1.

Phaeotremella pini Q.-M. Wang, sp. nov.MycoBank MB846592. Fig. 24L.

Etymology: the specific epithet pini refers to Pinus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 1.79–4.04 × 3.26–6.94 µm and single, budding is polar (Fig. 24L), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is saffron yellowish-cream, butyrous, smooth and faint glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose (latent), cellobiose, trehalose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-ribose (latent, weak), L-rhamnose, D-glucosamine (latent), N-acetyl-D-glucosamine, ethanol (latent, weak), erythritol (latent), ribitol, D-mannitol, D-glucitol (latent), α-methyl-D-glucoside, salicin (latent), DL-lactic acid, succinic aid (latent), citric acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Lactose, melibiose, raffinose, D-arabinose, methanol, glycerol, galactitol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. pini differs from the closely related species P. foliacea, P. ovata, P. lacteal, P. skinneri, P. fagi and P. neofoliacea (Fig. 1C) in its inability to assimilate D-arabinose (Table S2.12).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a rotten wood of Pinus sp., Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6848^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY502-4).

Vanrija silvicola Q.-M. Wang, sp. nov.MycoBank MB846593. Fig. 24M.

Etymology: the specific epithet silvicola refers to the type strain isolated from rotten wood.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical and elongate fusiform, 2.39–6.35 × 3.71–18.07 µm and single, budding is polar (Fig. 24M), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol, erythritol, ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 33 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, V. silvicola differs from the closely related species V. meifongana, V. meifongana, V. humicola, V. albida, V. music, V. pseudolongus and V. longus (Fig. 1C) in its inability to assimilate hexadecane and the ability assimilate potassium nitrate (Table S2.13).

Typus: China, Wuyi mountain, Fujian province, obtained from a rotten wood, Q.-M. Wang (holotype CGMCC 2.4340^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = WYS 38-3).

Takashimella corticis Q.-M. Wang, sp. nov.MycoBank MB846594. Fig. 24N.

Etymology: the specific epithet corticis refers to the type strain isolated from bark.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical, elongate oval and ellipsoidal, 1.68–4.04 × 4.5–8.19 µm and single, budding is polar (Fig. 24N), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, glossy and the surface of the colony has hair-like protrusions. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, sucrose, maltose, cellobiose, trehalose, lactose (latent), raffinose (latent), inulin, soluble starch, D-xylose, L-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, erythritol (latent, weak), ribitol (latent, weak), D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid (latent), succinic aid, citric acid and D-glucuronic acid are assimilated as sole carbon sources. L-sorbose, melibiose, melezitose, D-arabinose, D-ribose, methanol, ethanol, glycerol, galactitol, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate (weak), L-lysine, ethylamine and cadaverine (latent) are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, T. corticis differs from the closely related species T. formosensis and T. koratensis (Fig. 13) in its inability to assimilate melezitose, D-arabinose, D-ribose and inositol and the ability to assimilate inulin and potassium nitrate (Table S2.14).

Typus: China, Bawangling mountain, Changjiang, Hainan province, obtained from a bark an unidentified plant, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6836^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = 21812S).

Other culture examined: China, Tibet, Motuo county, leaf, Oct. 2021, isolated by Q.M. Wang, W.N. Zhao, culture MT-3C-2.

Holtermannia pseudosaccardoi Q.-M. Wang, sp. nov.MycoBank MB846595. Figs 24O, P.

Etymology: the specific epithet pseudosaccardoi refers to the similar colony morphology to that of Holtermannia saccardoi.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal, 2.55–7.03 × 3.39–7.30 µm and single, budding is polar (Fig. 24O), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale-yellow, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are oval and subglobosal, 3.69–5.76 × 6.23–6.78 µm (Fig. 24P).

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, L-rhamnose, ethanol, erythritol, ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, inositol and D-glucuronic acid (latent, weak) are assimilated as sole carbon sources. Galactose, lactose, D-ribose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, H. pseudosaccardoi differs from the closely related species H. saccardoi (Fig.13) in its inability to assimilate galactose and citric acid and the ability to assimilate melibiose, raffinose, inulin, erythritol, potassium nitrate and sodium nitrite (Table S2.15).

Typus: China, Bawangling mountain, Changjiang, Hainan province, obtained from a bark an unidentified plant, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6835^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = 21825S-11).

Piskurozyma terricola Q.-M. Wang, sp. nov.MycoBank MB846596. Fig. 25A.

Etymology: the specific epithet terricola refers to the type strain isolated from soil.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.44–4.76 × 5.61–7.83 µm and single, budding is polar (Fig. 25A), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol (weak), ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, erythritol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. terricola differs from the closely related species P. arborea and P. silvicultrix (Fig. 14) in its ability to assimilate lactose, D-ribose, D-glucosamine, N-acetyl-D-glucosamine and sodium nitrite (Table S2.16).

Typus: China, Sejila mountain, Linzhi, Tibet, obtained from soil, Aug, 2019, Q.-M. Wang (holotype CGMCC 2.6888^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY321-2).

Piskurozyma linzhiensis Q.-M. Wang, sp. nov.MycoBank MB846597. Fig. 25B.

Etymology: the specific epithet linzhiensis s refers to the geographic origin of the type strain, Linzhi, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and subglobosal, 4.13–6.35 × 4.73–6.87 µm and single, budding is polar (Fig. 25B), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, cellobiose, trehalose, lactose, raffinose (weak), melezitose, inulin, soluble starch (latent, weak), D-xylose, L-arabinose, D-arabinose, D-ribose (latent), L-rhamnose, ethanol, glycerol (latent, weak), ribitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, DL-lactic acid, succinic aid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Maltose, melibiose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, salicin, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. linzhiensis differs from the closely related species P. humicola, P. terricola, P. cuscutae, P. capsuligena, P. yama, P. fildesensis, P. arborea and P. silvicultrix (Fig. 14) in its inability to assimilate maltose (Table S2.16).

Typus: China, Sejila mountain, Linzhi, Tibet, obtained from soil, Aug, 2019, Q.-M. Wang (holotype CGMCC 2.6919^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY321-4).

Piskurozyma humicola Q.-M. Wang, sp. nov.MycoBank MB846598. Fig. 25C.

Etymology: the specific epithet humicola refers to the type strain isolated from soil.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.27–5.43 × 4.12–7.20 µm and single, budding is polar (Fig. 25C), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale-yellow, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose (latent, weak), melibiose, raffinose, melezitose, inulin, soluble starch (weak), D-xylose, L-arabinose, D-arabinose, D-ribose (latent, weak), L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine (latent, weak), ethanol (latent, weak), glycerol (latent, weak), ribitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, DL-lactic acid, succinic aid, citric acid and inositol are assimilated as sole carbon sources. Methanol, erythritol, galactitol, salicin, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. humicola differs from the closely related species P. arborea, P. silvicultrix and P. terricola (Fig. 14) in its inability to assimilate D-glucuronic acid (Table S2.16).

Typus: China, Nanyigou, Linzhi, Tibet, obtained from soil, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6879^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = NYG10-7).

Piskurozyma planta Q.-M. Wang, sp. nov.MycoBank MB846599. Fig. 25D.

Etymology: the specific epithet planta refers to the type strain isolated from plant (a leaf of Liliaceae).

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical and ellipsoidal, 2.81–5.23 × 5.98–10.37 µm and single, budding is polar (Fig. 25D), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, rough and dimmed. The margin is incised. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol, ribitol, galactitol, D-mannitol, D-glucitol (latent, weak), α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid, inositol and D-glucuronic acid (weak) are assimilated as sole carbon sources. Methanol, erythritol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. planta differs from the closely related species P. cylindrica (Fig. 14) in its ability to assimilate lactose, inulin, D-ribose, glycerol, galactitol and citric acid (Table S2.16).

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Liliaceae, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6862^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY30M1).

Piskurozyma nanyiensis Q.-M. Wang, sp. nov.MycoBank MB846600. Fig. 25E.

Etymology: the specific epithet nanyiensis s refers to the geographic origin of the type strain, Nanyi county, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, oval and subglobosal, 3.03–6.28 × 4.52–8.12 µm and single, budding is polar (Fig. 25E), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale-yellow, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, erythritol, ribitol, galactitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid, citric acid, inositol and D-glucuronic acid are assimilated as sole carbon sources. Methanol, glycerol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. nanyiensis differs from the closely related species P. filicatus and P. taiwanensis (Fig. 14) in its inability to assimilate hexadecane and the ability to assimilate inulin (Table S2.16).

Typus: China, Nanyigou, Linzhi, Tibet, obtained from soil, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6880^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = NYG26-1).

Piskurozyma cuscutae Q.-M. Wang, sp. nov.MycoBank MB846601. Fig. 25F.

Etymology: the specific epithet cuscutae refers to Cuscuta, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and oval, 4.23–7.43 × 4.64–8.68 µm and single, budding is polar (Fig. 25F), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, maltose (weak), cellobiose, trehalose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose, D-ribose (latent, weak), ethanol, glycerol, ribitol, D-mannitol, salicin, DL-lactic acid, succinic aid and D-glucuronic acidare assimilated as sole carbon sources. Galactose, sucrose, lactose, melibiose, raffinose, melezitose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, D-glucitol, α-methyl-D-glucoside, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. cuscutae differs from the closely related species P. capsuligena in its inability to assimilate galactose, sucrose, raffinose, D-glucitol, methyl-α-D-glucoside, citric acid and inositol and the ability to assimilate L-orbose, DL-lactic acid, potassium nitrate and sodium nitrite (Table S2.16).

Typus: China, Basom lake, Linzhi, Tibet, obtained from a leaf of Cuscuta chinensis, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6853^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY134-2).

Mrakia rhododendri Q.-M. Wang, sp. nov.MycoBank MB846602. Fig. 25G.

Etymology: the specific epithet rhododendri refers to Rhododendron, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 2.15–6.00 × 4.10–8.92 µm and single, budding is polar (Fig. 25G), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is present. Glucose, galactose, L-sorbose, sucrose (variable), maltose, cellobiose, trehalose, lactose (latent), melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose (latent), L-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, glycerol (latent, weak), erythritol (latent, weak), ribitol, D-mannitol, D-glucitol (variable), α-methyl-D-glucoside (variable), salicin, DL-lactic acid, succinic aid and D-glucuronic acid are assimilated as sole carbon sources. D-arabinose, methanol, galactitol, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. rhododendri differs from the closely related species M. arctica, M. stelviica, M. montana and M. cryoconite (Fig. 15) in its inability to assimilate D-arabinose and Inositol (Table S2.17).

Typus: China, Sejila mountain, Linzhi, Tibet, obtained from a leaf of Rhododendron parvifolium, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6914^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY337-1).

Other culture examined: China, Tibet, Linzhi, Sejila mountain, leaves, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, cultures XZY319-4, XZY347-3.

Mrakia pini Q.-M. Wang, sp. nov.MycoBank MB846603. Fig. 25H.

Etymology: the specific epithet pini refers to Pinus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 2.22–6.25 × 3.05–8.46 µm and single, budding is polar (Fig. 25H), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellowish-cream, butyrous, smooth and faint glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is present. Glucose, galactose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose (latent), melibiose, raffinose, melezitose, inulin, soluble starch, D-xylose (weak), L-arabinose, D-arabinose (weak), D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, ethanol, ribitol, D-mannitol, D-glucitol (weak), α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid and D-glucuronic acid are assimilated as sole carbon sources. Methanol, glycerol, erythritol, galactitol, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. pini differs from the closely related species M. floricola (Fig. 15) in its inability to assimilate galactitol and the ability to assimilate D-arabinose. In addition, the glucose fermentation of M. floricola is present (Table S2.17).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a bark of Pinus sp., Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6847^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY414-1).

Mrakia floricola Q.-M. Wang, sp. nov.MycoBank MB846604. Fig. 25I.

Etymology: the specific epithet floricola refers to the type of strain isolated from a flower (a flower of Rhododendron simsii).

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval, subglobosal and ellipsoidal, 2.25–5.55 × 3.43–7.76 µm and single, budding is polar (Fig. 25I), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale whiteish-cream, butyrous, bulge, smooth and glistening. The margin is wavy. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (latent), L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose (latent), melibiose (latent), raffinose, melezitose, inulin, soluble starch, D-xylose (latent, weak), L-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine (weak), ethanol, ribitol, galactitol (latent, weak), D-mannitol, D-glucito, α-methyl-D-glucoside (weak), salicin (weak), DL-lactic acid, succinic aid and D-glucuronic acid (latent, weak) are assimilated as sole carbon sources. D-arabinose, methanol, glycerol, erythritol, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. floricola differs from the closely related species M. pini (Fig. 15) in its inability to assimilate D-arabinose and the ability to assimilate galactitol. In addition, the glucose fermentation of M. floricola is absent (Table S2.17).

Typus: China, Sejila Mountain, Linzhi, Tibet, obtained from a flower of Rhododendron simsii, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6886^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY289N6).

Foliozyma Q.-M. Wang gen. nov.MycoBank MB846605.

Etymology: the specific epithet Foliozyma refers to the origin of the substrate of the species, leaf.

This genus is proposed for the separate branch represented by strains XZY30-4 and XZY30-6. Member of the Mrakiaceae (Cystofilobasidiales,Tremellomycetes). The genus is mainly circumscribed by the phylogenetic analysis of the six genes dataset, in which it occurred as a separate branch closely related to the genus Cystofilobasidium within the Cystofilobasidiales (Fig. 1D).

Sexual reproduction not known. Colonies cream to cream, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Foliozyma lilicola Q.-M. Wang.

Foliozyma lilicola Q.-M. Wang, sp. nov.MycoBank MB846606. Fig. 25J.

Etymology: the specific epithet lilicola refers to the substrate origin of the type strain, a leaf of Liliacea.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and cylindrical, 2.05–6.55 × 3.56–9.99 µm and single, budding is polar (Fig. 25J), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, rough and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, maltose, cellobiose (latent), trehalose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose (variable), L-rhamnose, D-glucosamine (variable), N-acetyl-D-glucosamine (variable), glycerol (variable), ribitol, D-mannitol, D-glucitol (variable), salicin, DL-lactic acid, succinic aid (variable), inositol and D-glucuronic acid are assimilated as sole carbon sources. Galactose, sucrose, lactose, melibiose, raffinose, D-ribose, methanol, ethanol, erythritol, galactitol, α-methyl-D-glucoside, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (variable), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Liliaceae, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6861^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY30-6).

Other culture examined: China, Tibet, Linzhi, Kadinggou, leaf of Liliaceae, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY30-4.

Phaffia (Mrakiaceae, Cystofilobasidiales)

Phaffia rhododendri Q.-M. Wang, sp. nov.MycoBank MB846608. Fig. 25K.

Etymology: the specific epithet rhododendri refers to Rhododendron, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal, ellipsoidal and oval, 2.89–7.66 × 3.75–9.47 µm and single, budding is polar (Fig. 25K), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is salmon, butyrous, smooth and pale glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is present. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose, raffinose, melezitose, inulin, soluble starch, D-xylose (weak), L-arabinose, D-ribose (latent, weak), L-rhamnose (weak), ethanol, D-mannitol, α-methyl-D-glucoside, salicin (weak), DL-lactic acid (weak) and succinic aid are assimilated as sole carbon sources. galactose, lactose, melibiose, D-arabinose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, ribitol, galactitol, D-glucitol, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. rhododendri differs from the closely related species P. rhodozyma (Fig. 1D) in its inability to assimilate glycerol and D-glucuronic acid and the ability to assimilate L-rhamnose, potassium nitrate, sodium nitrite and ethylamine (Table S2.18).

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Rhododendron capitatum, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6920 ^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY12-2).

Phaffia paratasmanica Q.-M. Wang, sp. nov.MycoBank MB846609. Fig. 25L.

Etymology: the specific epithet paratasmanica refers to the similar colony morphology to that of Phaffia tasmanica.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are subglobosal, oval and ellipsoidal, 3.04–8.28 × 6.05–11.12 µm and single, budding is polar (Fig. 25L), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale-pink, mucilaginous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is present. Glucose, sucrose, maltose, trehalose, melibiose, raffinose, melezitose, inulin, soluble starch, L-arabinose, glycerol, D-mannitol, DL-lactic acid, succinic aid (latent, weak), inositol (latent, weak) and D-glucuronic acid are assimilated as sole carbon sources. galactose, L-sorbose, cellobiose, lactose, D-xylose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, erythritol, ribitol, galactitol, D-glucitol, α-methyl-D-glucoside, salicin, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. paratasmanica differs from the closely related species P. tasmanica (Fig. 15) in its inability to assimilate galactose, cellobiose, ethanol, methyl-α-D-glucoside, salicin and citric acid and the ability to assimilate inulin, potassium nitrate, sodium nitrite and ethylamine (Table S2.18).

Typus: China, Sejila mountain of Linzhi, Tibet, obtained from a rotten wood, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.7159^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY261N1).

New taxa in the Agaricostilbomycetes (Pucciniomycotina)

Kondoa tibetensis Q.-M. Wang, sp. nov.MycoBank MB846610. Fig. 25M.

Etymology: the specific epithet tibetensis refers to the geographic origin of the type strain, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical and elongate oval, 1.97–3.50 × 6.97–12.55 µm and single, budding is polar (Fig. 25M), a sediment is formed. After 1 month at 17 ºC, pellicle and sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale white, butyrous, rough, bulging and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, sucrose (latent), maltose, melibiose, raffinose, melezitose (latent, weak), inulin (latent, weak), soluble starch, ethanol (weak), glycerol (weak), ribitol (weak), galactitol, D-mannitol (latent, weak), salicin, DL-lactic acid, hexdecane (weak) and D-glucuronic acid are assimilated as sole carbon sources. Galactose, L-sorbose, cellobiose, trehalose, lactose, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, D-glucitol, α-methyl-D-glucoside, succinic aid, citric acid and inositol are not assimilated. Ammonium sulfate, potassium nitrate (weak), sodium nitrite (weak) and cadaverine (weak) are assimilated as sole nitrogen sources. L-lysine and ethylamine are not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, K. tibetensis differs from the closely related species K. thailandica (Fig. 16) in its inability to assimilate cellobiose, trehalose, succinic acid and citric acid and the ability to assimilate inulin, ethanol, DL-lactic acid, hexadecane, D-glucuronic acid and cadaverine (Table S2.19).

Typus: China, Motuo county, Tibet, obtained from a leaf an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6877^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-1B-33).

Boekhoutia foliicola Q.-M. Wang, sp. nov.MycoBank MB846611. Fig. 25N.

Etymology: the specific epithet foliicola refers to the substrate origin of the type strain, leaves.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical and ellipsoidal, 1.72–3.78 × 2.31–7.44 µm and single, budding is polar (Fig. 25N), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale red, butyrous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, sucrose, maltose, cellobiose, trehalose, raffinose, melezitose, inulin, soluble starch, D-xylose (weak), L-arabinose, ethanol (latent, weak), ribitol, D-mannitol, α-methyl-D-glucoside (weak), salicin (weak) and DL-lactic acid (weak) are assimilated as sole carbon sources. L-sorbose, lactose, melibiose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, galactitol, D-glucitol, succinic aid, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine (latent), ethylamine (weak) and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, B. foliicola differs from the closely related species B. sterigmata (Fig. 16) in its inability to assimilate D-glucuronic acid and the ability to assimilate raffinose, soluble starch, D-xylose, L-arabinose, ethanol, ribitol, D-mannitol, methyl-α-D-glucoside, salicin, DL-lactic acid, potassium nitrate and sodium nitrite (Table S2.20).

Typus: China, Motuo county, Tibet, obtained from a leaf an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6878^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-2C-38).

Buckleyzyma pseudoaurantiaca Q.-M. Wang, sp. nov.MycoBank MB846612. Fig. 25O.

Etymology: the specific epithet pseudoaurantiaca refers to the similar colony morphology to that of Buckleyzyma aurantiaca.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, 3.38–6.53 × 4.84–8.49 µm and single, budding is polar (Fig. 25O), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale orangish, butyrous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (weak), L-sorbose (weak), sucrose (latent, weak), trehalose (latent), D-xylose, N-acetyl-D-glucosamine (weak), glycerol (weak), ribitol (weak), galactitol, D-mannitol, α-methyl-D-glucoside (latent, weak), DL-lactic acid, succinic aid (latent), citric acid (latent) and D-glucuronic acid are assimilated as sole carbon sources. Maltose, cellobiose, lactose, melibiose, raffinose, melezitose, inulin, soluble starch, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, methanol, ethanol, erythritol, D-glucitol, salicin, inositol and hexdecane are not assimilated. Ammonium sulfate (latent), potassium nitrate (latent, weak), sodium nitrite (latent), L-lysine (weak) and cadaverine are assimilated as sole nitrogen sources. Ethylamine is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, B. pseudoaurantiaca differs from the closely related species B. aurantiaca (Fig. 2A) in its inability to assimilate maltose, cellobiose, melezitose, L-arabinose, D-arabinose, ethanol and D-glucitol and the ability to assimilate galactitol (Table S2.21).

Typus: China, Chengguan county, Lhasa, Tibet, obtained from a rotten wood, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6834^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YSR26-8).

Microsporomyces betulae Q.-M. Wang, sp. nov.MycoBank MB846613. Fig. 25P.

Etymology: the specific epithet betulae refers to Betula, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are long oval and ellipsoidal, 2.00–4.31 × 4.02–6.11 µm and single, budding is polar (Fig. 25P), a sediment is formed. After 1 month at 17 ºC, a sediment is present,a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale orange, butyrous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose (latent), sucrose (latent), cellobiose, trehalose, melezitose, inulin, soluble starch, L-arabinose, glycerol, D-mannitol, salicin, succinic aid and D-glucuronic acid are assimilated as sole carbon sources. Galactose, maltose, lactose, melibiose, raffinose, D-xylose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, erythritol, ribitol, galactitol, D-glucitol, α-methyl-D-glucoside, DL-lactic acid, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate and L-lysine (latent, weak) are assimilated as sole nitrogen sources. Sodium nitrite, ethylamine and cadaverine are not assimilated. Maximum growth temperature is 20 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. betulae differs from the closely related species M. pini and M. orientalis (Fig. 2A) in its inability to assimilate N-acetyl-D-glucosamine, erythritol, DL-lactic acid, citric acid and cadaverine and the ability to assimilate inulin (Table S2.22).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a bark of Betula sp., Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6850^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY554-7).

Microsporomyces foliicola Q.-M. Wang, sp. nov.MycoBank MB846614. Fig. 26A.

Etymology: the specific epithet foliicola refers to the type strain isolated from a leaf.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.12–4.99 × 3.14–6.10 µm and single, budding is polar (Fig. 26A), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale orangish, butyrous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, sucrose, cellobiose (latent), melezitose (latent, weak), inulin (latent, weak), D-xylose (latent, weak), L-arabinose, glycerol, α-methyl-D-glucoside and D-glucuronic acid are assimilated as sole carbon sources. Galactose, L-sorbose, maltose, trehalose, lactose, melibiose, raffinose, soluble starch, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, erythritol, ribitol, galactitol, D-mannitol, D-glucitol, salicin, DL-lactic acid, succinic aid, citric acid, inositol and hexdecane are not assimilated. L-lysine (latent, weak) and ethylamine (latent, weak) are assimilated as sole nitrogen sources. Ammonium sulfate, potassium nitrate, sodium nitrite and cadaverine are not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. foliicola differs from the closely related species M. pini, M. orientalis, M. betulae, M. magnisporus, M. bloemfonteinensis, M. pseudomagnisporus, M. rubellus and M. ellipsoideus (Fig. 2A) in its inability to assimilate trehalose (Table S2.22).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Betula sp., Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6907^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN35N5).

Halobasidium phyllophila Q.-M. Wang, sp. nov.MycoBank MB846615. Fig. 26B.

Etymology: the specific epithet phyllophila refers to the type strain isolated from a leaf.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and subglobosal, 2.87–5.76 × 4.04–7.07 µm and single, budding is polar (Fig. 26B), a sediment is formed. After 1 month at 17 ºC, sediment and pellicle are present. On YM agar, after 1 month at 17 ºC, the streak culture is amber, butyrous, smooth and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (weak), L-sorbose (variable), sucrose, maltose, cellobiose (variable), trehalose, lactose (variable), raffinose (variable), melezitose, inulin, soluble starch (variable), D-xylose, N-acetyl-D-glucosamine, ethanol (variable), glycerol (latent, weak), ribitol, D-mannitol, α-methyl-D-glucoside (variable), salicin (latent, weak), DL-lactic acid, succinic aid, inositol (variable) and D-glucuronic acid are assimilated as sole carbon sources. Melibiose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, methanol, erythritol, galactitol, D-glucitol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (latent, weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, H. phyllophila differs from the closely related species H. xiangyangense (Fig. 2A) in its inability to assimilate melibiose, L-arabinose, D-arabinose, D-ribose, ethanol and citric acid and the ability to assimilate DL-lactic acid, sodium nitrite and cadaverine (Table S2.23).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Pteridophyta sp., Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6906^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN28M2).

Other culture examined: China, Yunnan province, Zhenxiong county, Dawan town, Shili village, leaf, Oct. 2019, isolated by Q.M. Wang, G.S. Wang, culture YN55-1.

Cystobasidium cunninghamiae Q.-M. Wang, sp. nov.MycoBank MB846616. Fig. 26C.

Etymology: the specific epithet cunninghamiae refers to Cunninghamia, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and oval, 2.42–4.86 × 3.34–6.10 µm and single, budding is polar (Fig. 26C), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is amber, mucilaginous, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, trehalose, melezitose, inulin, D-xylose (latent), L-arabinose (variable), D-arabinose (weak), D-ribose (variable), N-acetyl-D-glucosamine (latent), glycerol, ribitol, D-mannitol, salicin, DL-lactic acid, succinic aid (variable) and D-glucuronic acid (variable) are assimilated as sole carbon sources. Galactose, maltose, cellobiose, lactose, melibiose, raffinose, soluble starch, L-rhamnose, D-glucosamine, methanol, ethanol, erythritol, galactitol, D-glucitol, α-methyl-D-glucoside, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine (latent), ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, C. cunninghamiae differs from the closely related species C. psychroaquaticum (Fig. 17) in its ability to assimilate inulin, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine (Table S2.24).

Typus: China, Qilian mountain, Qinghai province, obtained from a bark of Cunninghamia sp., Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6841^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XSR28-13).

Other culture examined: China, Tibet, Linzhi, New basom lake, the bark of Pinus sp., Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY554-12.

Rhodotorula linzhiensis Q.-M. Wang, sp. nov.MycoBank MB846617. Fig. 26D.

Etymology: the specific epithet linzhiensis refers to the geographic origin of the type strain, Linzhi, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells, ellipsoidal and oval, 2.18–4.33 × 3.07–6.74 µm and single, budding is polar (Fig. 26D), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale pink, butyrous, smooth and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (weak), L-sorbose, cellobiose, trehalose, lactose (latent, weak), inulin, D-xylose, D-arabinose (latent, weak), D-ribose (latent), ethanol (latent), glycerol, ribitol, D-mannitol, α-methyl-D-glucoside, DL-lactic acid and succinic aid are assimilated as sole carbon sources. Sucrose, maltose, melibiose, raffinose, melezitose, soluble starch, L-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, D-glucitol, salicin, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 33 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, R. pinicola differs from the closely related species R. toruloides (Fig. 18A) in its inability to assimilate sucrose, maltose, raffinose, melezitose, L-arabinose, D-glucitol, salicin, citric acid and D-glucuronic acid and the ability to assimilate L-sorbose, lactose and inulin (Table S2.25).

Typus: China, New Basom lake, Linzhi, Tibet, obtained from a rotten wood of Pinus sp. Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6911^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY511-5).

Bauerozyma Q.-M. Wang gen. nov.MycoBank MB846618.

Etymology: the genus is named in honour of Robert Bauer for his contributions to the taxonomy of smuts.

This genus is proposed for the separate branch represented by strain YN25-3. Member of the Microbotryales (Microbotryomycetes). The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate branch within the Microbotryales (Fig. 2B).

Sexual reproduction not known. Colonies yellowish, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Bauerozyma artemisiae Q.-M. Wang.

Bauerozyma artemisiae Q.-M. Wang, sp. nov.MycoBank MB846619. Fig. 26E.

Etymology: the specific epithet artemisiae refers to Artemisia, the plant genus from which the strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical, ellipsoidal and botuliform, 2.28–5.55 × 3.25–6.05 µm and single, budding is polar (Fig. 26E), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, rough, crease and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose (weak), sucrose, maltose, cellobiose, trehalose, raffinose (latent, weak), melezitose, inulin, soluble starch (weak), glycerol (weak), ribitol (latent, weak), D-mannitol, methyl-α-D-glucoside, DL-lactic acid (weak), succinic acid and citric acid are assimilated as sole carbon sources. Galactose, lactose, melibiose, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine N-acetyl-D-glucosamine, methanol, ethanol, erythritol, galactitol, D-glucitol, salicin, inositol, hexadecane and D-glucuronic acid not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 32 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Artemisia carvifolia, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6908^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN25-3).

Andreyozyma Q.-M. Wang gen. nov.MycoBank MB846620.

Etymology: the genus is named in honour of Dr. Andrey Yurkov for his contributions to yeast taxonomy.

This genus is proposed for the separate branch represented by strain XZY328-1. Member of the Microbotryomycetes. The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate branch within the Microbotryomycetes (Fig. 2B).

Sexual reproduction not known. Colonies yellowish, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Andreyozyma sejilaensis Q.-M. Wang.

Andreyozyma sejilaensis Q.-M. Wang, sp. nov.MycoBank MB846621. Fig. 26F.

Etymology: the specific epithet sejilaensis refers to the geographic origin of the type strain, Sejila mountain, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical, oval and ellipsoidal, 2.03–4.06 × 4.21–9.26 µm and single, budding is polar (Fig. 26F), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, cellobiose, trehalose, melezitose, inulin, soluble starch, L-arabinose, ethanol, glycerol (latent, weak), ribitol, D-mannitol, D-glucitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid and citric acid are assimilated as sole carbon sources. Galactose, maltose, lactose, melibiose, raffinose, D-xylose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, ethylamine and cadaverine are assimilated as sole nitrogen sources. L-lysine is not assimilated. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Sejila mountain, Linzhi, Tibet, China, obtained from a fallen leaf, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6923^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY328-1).

Nakasozyma Q.-M. Wang gen. nov.MycoBank MB846622.

Etymology: the genus is named in honour of Takashi Nakase for his contributions to yeast taxonomy.

This genus is proposed for the separate clade represented by XZY22-1 and XZY99M13. Member of the Microbotryomycetes. The genus is mainly circumscribed by the phylogenetic analysis of the six genes dataset, in which it occurred as a separate branch within the Microbotryomycetes (Fig. 2B).

Sexual reproduction not known. Colonies cream to yellowish, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Nakasozyma lonicerae Q.-M. Wang.

Nakasozyma lonicerae Q.-M. Wang, sp. nov.MycoBank MB846623. Fig. 26G.

Etymology: the specific epithet lonicerae refers to Lonicera, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical, fusiformis and ellipsoidal, 1.82–3.49 × 3.59–9.20 µm and single, budding is polar (Fig. 26G), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, verrucous and glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, sucrose, maltose, cellobiose, trehalose, lactose (latent, weak), melezitose, inulin, soluble starch, L-arabinose, D-arabinose (latent, weak), D-ribose (weak), ethanol, ribitol, D-mannitol, α-methyl-D-glucoside (latent, weak), salicin, DL-lactic acid and succinic aid are assimilated as sole carbon sources. Galactose, L-sorbose, melibiose, raffinose, D-xylose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, galactitol, D-glucitol, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, N. lonicerae differs from the closely related species N. junca (Fig. 18A) in its inability to assimilate D-xylose, inositol and D-glucuronic acid and the ability to assimilate sodium nitrite (Table S2.26).

Typus: China, Baiba county, Tibet, obtained from a leaf of Lonicera japonica, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6922^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY99M13).

Nakasozyma junca Q.-M. Wang, sp. nov.MycoBank MB846624. Fig. 26H.

Etymology: the specific epithet junca refers to Juncus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells fusiformis, ellipsoidal and oval, 2.12–3.39 × 3.47–7.24 µm and single, budding is polar (Fig. 26H), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, smooth and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, sucrose, maltose, cellobiose, trehalose, lactose, melezitose, inulin, soluble starch (weak), D-xylose, L-arabinose, D-arabinose, D-ribose, ethanol, ribitol, D-mannitol, α-methyl-D-glucoside (weak), salicin, DL-lactic acid, succinic aid, inositol (weak) and D-glucuronic acid (weak) are assimilated as sole carbon sources. galactose, L-sorbose, melibiose, raffinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, erythritol, galactitol, D-glucitol, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, L-lysine, ethylamine and cadaverine (weak) are assimilated as sole nitrogen sources. Sodium nitrite is not assimilated. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, N. junca differs from the closely related species N. lonicerae (Fig. 18A) in its inability to assimilate sodium nitrite and the ability to assimilate D-xylose, inositol and D-glucuronic acid (Table S2.26).

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Juncus effusus, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.7158^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY22-1).

Fengyania Q.-M. Wang gen. nov.MycoBank MB846628.

Etymology: the genus is named in honour of Feng-Yan Bai for his for his contributions to yeast taxonomy.

This genus is proposed for the separate clade represented by XZY30M2 and XZY23-5. Member of the Microbotryomycetes. The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate branch within the Microbotryomycetes (Fig. 2B).

Sexual reproduction not known. Colonies yellowish-cream, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Fengyania pteridophytae Q.-M. Wang.

Fengyania pteridophytae Q.-M. Wang, sp. nov.MycoBank MB846629. Fig. 26I.

Etymology: the specific epithet pteridophytae refers to Pteridophyta, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, cylindrical and oval, 2.28–5.55 × 3.25–6.05 µm and single, budding is polar (Fig. 26I), a sediment is formed. After 1 month at 17 ºC, a sediment is present,a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, rough, crease and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose (variable), lactose (variable), melezitose, inulin, soluble starch (weak), D-xylose, ethanol, glycerol, ribitol (variable), D-mannitol, α-methyl-D-glucoside, DL-lactic acid and succinic aid are assimilated as sole carbon sources. Galactose, melibiose, raffinose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, D-glucitol, salicin, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (weak), L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Pteridophyta sp., Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6915^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY23-5).

Other culture examined: China, Tibet, Linzhi, Kadinggou, leaf of Liliaceae, Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY30M2.

Curvibasidium pinum Q.-M. Wang, sp. nov.MycoBank MB846630. Fig. 26J.

Etymology: the specific epithet pinum refers to Pinus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 2.83–5.88 × 3.48–7.54. µm and single, budding is polar (Fig. 26J), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, smooth and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (latent, weak), L-sorbose, cellobiose, inulin, D-xylose (weak), L-arabinose (latent, weak), ethanol, glycerol (latent, weak), ribitol, galactitol, D-mannitol, α-methyl-D-glucoside, salicin, DL-lactic acid and succinic aid are assimilated as sole carbon sources. Sucrose, maltose, trehalose, lactose, melibiose, raffinose, melezitose, soluble starch, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, D-glucitol, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, C. pinum differs from the closely related species C. nothofagi and C. rogersii (Fig. 18B) in its inability to assimilate D-arabinose and D-glucitol, and the ability to assimilate inulin, methyl-α-D-glucoside and potassium nitrate (Table S2.27).

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a Parasite of Pinus massoniana, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6865^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY26-2).

Chrysozyma quercicola Q.-M. Wang, sp. nov.MycoBank MB846631. Fig. 26K.

Etymology: the specific epithet quericola refers to the substrate origin of the type strain, a leaf of Quercus semicarpifolia.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal and cylindrical, 2.10–4.82 × 5.47–8.95 µm and single, budding is polar (Fig. 26K), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, smooth and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose (latent, weak), sucrose, maltose, cellobiose, trehalose, lactose (latent, weak), melezitose, inulin, soluble starch (weak), ethanol (weak), glycerol (weak), ribitol, D-mannitol, α-methyl-D-glucoside (weak), salicin, DL-lactic acid (latent, weak), succinic aid and D-glucuronic acid (weak) are assimilated as sole carbon sources. Galactose, melibiose, raffinose, D-xylose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, D-glucitol, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, C. quercicola differs from the closely related species C. cylindrica (Fig. 2B) in its inability to assimilate galactose and D-glucitol and the ability to assimilate L-sorbose, maltose, cellobiose, lactose, inulin, soluble starch, ethanol, glycerol, ribitol, methyl-α-D-glucoside, salicin, succinic acid, D-glucuronic acid, sodium nitrite and L-lysine (Table S2.28).

Typus: China, Kadinggou, Linzhi, Tibet, obtained from a leaf of Quercus semicarpifolia, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6863^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY20-10).

Yurkovia castanea Q.-M. Wang, sp. nov.MycoBank MB846632. Fig. 26L.

Etymology: the specific epithet castanea refers to Castanea, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are fusiformis and ellipsoidal to elongate, 1.53–3.48 × 3.88–8.73 µm and single, budding is polar (Fig. 26L), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, butyrous, smooth and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose (weak), sucrose, maltose, cellobiose, trehalose, lactose (weak), melezitose, inulin, soluble starch (weak), D-xylose (latent, weak), D-glucosamine (weak), N-acetyl-D-glucosamine, ethanol, glycerol, ribitol, D-mannitol, α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid and D-glucuronic acid are assimilated as sole carbon sources. Galactose, melibiose, raffinose, L-arabinose, D-arabinose, D-ribose, L-rhamnose, methanol, erythritol, galactitol, D-glucitol, citric acid, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, Y. castanea differs from the closely related species Y. longicylindrica Y. mendeliana and Y. nerthusi (Fig. 18B) in its inability to assimilate galactose and D-glucitol and the ability to assimilate lactose and salicin (Table S2.29).

Typus: China, Shili village of Dawan town, Zhenxiong county, Yunnan province, obtained from a leaf of Castanea mollissima, Oct. 2019, Q.-M. Wang (holotype CGMCC 2.6909^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = YN97-2).

Pseudohyphozyma sanghuangpori Q.-M. Wang, sp. nov.MycoBank MB846633. Fig. 26M.

Etymology: the specific epithet sanghuangpori refers to Sanghuangporus, the plant genus from which the type strain was isolated.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are fusiformis, cylindrical and ellipsoidal, 2.03–4.23 × 3.42–8.41 µm and single, budding is polar (Fig. 26M), a sediment is formed. After 1 month at 17 ºC, a sediment is present,a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is whiteish-cream, butyrous smooth bulging and glossy and dimmed. The margin is wavy. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (latent, weak), L-sorbose, maltose, cellobiose, trehalose, melezitose, inulin, soluble starch, D-xylose, L-arabinose, D-arabinose (latent, weak), D-glucosamine, ethanol, glycerol, ribitol, D-mannitol, α-methyl-D-glucoside, DL-lactic acid, succinic aid, citric acid and D-glucuronic acid are assimilated as sole carbon sources. Sucrose, lactose, melibiose, raffinose, D-ribose, L-rhamnose, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, D-glucitol, salicin, inositol and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 17 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, P. sanghuangpori differs from the closely related species P. bogoriensis and P. lulangensis (Fig. 18B) in its inability to assimilate D-ribose, D-glucitol and salicin and the ability to assimilate methyl-α-D-glucoside, potassium nitrate and sodium nitrite (Table S2.30).

Typus: China, Sejila mountain, Linzhi, Tibet, obtained from a Sanghuangporus lonicericola, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6891^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = XZY290-2).

Slooffia terricola Q.-M. Wang, sp. nov.MycoBank MB846634. Fig. 26N.

Etymology: the specific epithet terricola refers to the origin of the substrate of the type strain, soil.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval and ellipsoidal, 2.93–4.09 × 3.98–5.66. µm and single, budding is polar (Fig. 26N), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is amber, butyrous, smooth and faint glossy. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose, melezitose, inulin, soluble starch (weak), D-xylose (variable), L-rhamnose (variable), ethanol, glycerol (weak), ribitol (latent, weak), D-mannitol, D-glucitol, α-methyl-D-glucoside, DL-lactic acid, succinic aid, inositol (variable) and D-glucuronic acid are assimilated as sole carbon sources. Galactose, lactose, melibiose, raffinose, L-arabinose, D-arabinose, D-ribose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, salicin, citric acid and hexdecane are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 25 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, S. terricola differs from the closely related species S. cresolica (Fig. 18B) in its inability to assimilate galactose, D-arabinose, D-ribose, D-glucosamine and Citric acid and the ability to assimilate L-sorbose, inulin and soluble starch (Table S2.31).

Typus: China, Nanyigou, Linzhi, Tibet, obtained from soil, Aug. 2019, Q.-M. Wang (holotype CGMCC 2.6882^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = NYG09-2B).

Other culture examined: China, Tibet, Linzhi, Zhenxiong county, New basom lake, leaf of Polyporus sp., Aug. 2019, isolated by Q.M. Wang, G.S. Wang, culture XZY423-3.

Litoriozymales Q.-M. Wang ord. nov. MycoBank MB846635.

Member of the Microbotryomycetes (Pucciniomycotina). The diagnosis of the order Litoriozymales is based on the genus Litoriozyma. The nomenclature of the order is based on the genus Litoriozyma.

Type family: Litoriozymaceae Q.-M. Wang.

Litoriozymaceae Q.-M. Wang fam. nov. MycoBank MB846636.

Member of the Microbotryomycete (Pucciniomycotina). The diagnosis of the family Litoriozymaaceae is based on the genus Litoriozyma. The nomenclature of the family is based on the genus Litoriozyma.

Type genus: Litoriozyma Q.-M. Wang.

Genus accepted: Litoriozyma Q.-M. Wang.

Litoriozyma Q.-M. Wang gen. nov.MycoBank MB846637.

Etymology: the specific epithet Litoriozyma refers to the origin of the substrate of the species, intertidal zone.

This genus is proposed for the separate branch represented by SHTS17-2. Member of the Microbotryomycetes. The genus is mainly circumscribed by the phylogenetic analysis of the six-gene dataset, in which it occurred as a separate branch within the Microbotryomycetes (Fig. 2B).

Sexual reproduction not known. Colonies cream to yellowish-cream, butyrous. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Litoriozyma hainanensis Q.-M. Wang.

Litoriozyma hainanensis Q.-M. Wang, sp. nov.MycoBank MB846638. Fig. 26O.

Etymology: the specific epithet hainanensis refers to the geographic origin of the type strain, Hainan province.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are oval, ellipsoidal and subglobosal, 2.82–5.74 × 4.48–7.34 µm and single, budding is polar (Fig. 26O), a sediment is formed. After 1 month at 17 ºC, a sediment is present, a pellicle does not appear. On YM agar, after 1 month at 17 ºC, the streak culture is yellowish-cream, butyrous, smooth and glistening. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, L-sorbose, sucrose, maltose, cellobiose, trehalose, lactose, melibiose, raffinose, melezitose (latent), inulin, D-xylose, L-arabinose, D-arabinose, D-ribose (latent, weak), ethanol, glycerol, ribitol (latent, weak), D-mannitol, D-glucitol (latent, weak), α-methyl-D-glucoside, salicin, DL-lactic acid, succinic aid (latent) and hexdecane are assimilated as sole carbon sources. Galactose, soluble starch, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, erythritol, galactitol, citric acid, inositol and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 32 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Shanhaitian of Longlou town, Wenchang, Hainan province, obtained from an Intertidal zone, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6843^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = SHTS17-2).

New taxa in the Ustilaginomycetes (Ustialginomycotina)

Ustilago foliicola Q.-M. Wang, sp. nov.MycoBank MB846639. Fig. 26P.

Etymology: the specific epithet foliicola refers to the type strain isolated from a leaf.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are globosal and oval, 2.51–5.65 × 3.10–7.03 µm and single, budding is polar (Fig. 26P), a sediment is formed. After 1 month at 17 ºC, a pellicle is present and the sediment disappears. On YM agar, after 1 month at 17 ºC, the streak culture is pale white, mealiness, rough and dimmed. The margin is erose. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose (latent), sucrose, maltose, cellobiose, trehalose, lactose (latent), raffinose, melezitose, inulin, soluble starch, D-xylose (latent), L-arabinose, D-arabinose (latent), D-ribose (latent), L-rhamnose (latent, weak), glycerol, erythritol, D-mannitol, D-glucitol, α-methyl-D-glucoside (latent), salicin (latent), succinic aid and inositol are assimilated as sole carbon sources. L-sorbose, melibiose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, ribitol, galactitol, DL-lactic acid, citric acid, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate (latent, weak), sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 33 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Bawangling mountain, Changjiang, Hainan province, obtained from a leaf of an unidentified plant, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6890^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = 2189S-11).

New taxa in the Exobasidiomycetes (Ustialginomycotina)

Acaromyces pseudoingoldii Q.-M. Wang, sp. nov.MycoBank MB846640. Fig. 27A.

Etymology: the specific epithet pseudoingoldii refers to the similar colony morphology to that of Acaromyces ingoldii.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are ellipsoidal, oval and globosal, 2.93–6.08 × 3.11–7.77 µm and moniliform, budding is polar (Fig. 27A), a sediment is formed. After 1 month at 17 ºC, a pellicle and a sediment are present. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, dried, rough and dimmed. The margin is entire. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose (weak), sucrose, maltose, cellobiose, trehalose, lactose (latent), raffinose (weak), melezitose, inulin, soluble starch (latent, weak), D-xylose, L-arabinose, D-ribose, ethanol (latent, weak), erythritol (latent, weak), ribitol (latent, weak), D-mannitol, D-glucitol (latent, weak), succinic aid (latent, weak) and inositol (latent, weak) are assimilated as sole carbon sources. Melibiose, D-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, galactitol, α-methyl-D-glucoside, salicin, DL-lactic acid, citric acid, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite, L-lysine, ethylamine and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, A. pseudoingoldii differs from the closely related species A. ingoldii (Fig. 20) in its inability to assimilate D-arabinose, salicin, and D-glucuronic acid and the ability to assimilate L-sorbose, inulin and soluble starch (Table S2.32).

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6881^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-4B-41).

Exobasidium lijiangensis Q.-M. Wang, sp. nov.MycoBank MB846641. Fig. 27B.

Etymology: the specific epithet lijiangensis refers to the geographic origin of the type strain, Lijiang county, Yunnan province.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are fusiformis, cylindrical and ellipsoidal, 1.45–2.68 × 3.09–10.75 µm and single, budding is polar (Fig. 27B), a sediment is formed. After 1 month at 17 ºC, a pellicle is present and the sediment disappears. On YM agar, after 1 month at 17 ºC, the streak culture is white, leathery, smooth and faint glistening. The margin is erose. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, sucrose (latent), maltose (latent, weak), melezitose, inulin (latent, weak), D-xylose, L-arabinose, D-ribose (latent, weak), glycerol (latent), erythritol (latent), D-mannitol, D-glucitol and succinic aid (latent, weak) are assimilated as sole carbon sources. Galactose, L-sorbose, cellobiose, trehalose, lactose, melibiose, raffinose, soluble starch, D-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, ribitol, galactitol, α-methyl-D-glucoside, salicin, DL-lactic acid, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate, sodium nitrite (latent), ethylamine and cadaverine are assimilated as sole nitrogen sources. L-lysine is not assimilated. Maximum growth temperature is 33 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Lijiang Alpine Botanical Garden, Yunnan province, obtained from a leaf of Rhododendron decorum, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6921^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = LJGSL4-20)

Meira pseudoplantarum Q.-M. Wang, sp. nov.MycoBank MB846642. Fig. 27C.

Etymology: the specific epithet pseudoplantarum refers to the similar colony morphology to that of Meira plantarum.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are fusiformis and cylindrical, 1.04–2.09 × 4.49–15.94 µm and single, budding is polar (Fig. 27C), a sediment is formed. After 1 month at 17 ºC, a pellicle is present and the sediment disappears. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, rough and dimmed. The margin is erose. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose (latent), sucrose, maltose, cellobiose (latent), trehalose, melibiose, raffinose, melezitose, inulin (latent), D-xylose, L-arabinose (latent), D-ribose, ethanol (latent), erythritol, D-mannitol, D-glucitol, salicin (latent), DL-lactic acid (latent), succinic aid and citric acid (latent, weak) are assimilated as sole carbon sources. lactose, soluble starch, D-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, ribitol, galactitol, α-methyl-D-glucoside, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, sodium nitrite (weak), ethylamine and cadaverine are assimilated as sole nitrogen sources. potassium nitrate and L-lysine are not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. pseudoplantarum differs from the closely related species M. plantarum and M. nashicola (Fig. 20) in its inability to assimilate potassium nitrate and L-lysine and the ability to assimilate L-sorbose, inulin, ethanol, DL-lactic acid and citric acid (Table S2.33).

Typus: China, Jianfeng mountain, Jianfeng town, Ledong Li Autonomou county , Hainan province, obtained from a leaf of an unidentified plant, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6889^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = 217S13D1).

Meira marina Q.-M. Wang, sp. nov.MycoBank MB846643. Fig. 27D.

Etymology: the specific epithet marina refers to the origin of the substrate of the type strain, seawater.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are fusiformis and cylindrical, 1.17–2.59 × 3.13–6.63 µm and single, budding is polar (Fig. 27D), a sediment is formed. After 1 month at 17 ºC, a pellicle is present and the sediment disappears. On YM agar, after 1 month at 17 ºC, the streak culture is pale white, leathery, rough and dimmed. The margin is erose. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, galactose, L-sorbose (latent, weak), sucrose (latent), maltose, cellobiose (latent), trehalose, melibiose (latent), raffinose, melezitose, inulin (latent), soluble starch (latent, weak), D-xylose, L-arabinose, D-ribose (latent, weak), ethanol, erythritol (latent), D-mannitol, D-glucitol, salicin (latent, weak), DL-lactic acid (latent), succinic aid (latent) and citric acid (latent, weak) are assimilated as sole carbon sources. Lactose, D-arabinose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, glycerol, ribitol, galactitol, α-methyl-D-glucoside, inositol, hexdecane and D-glucuronic acid are not assimilated. Ammonium sulfate, potassium nitrate (latent), sodium nitrite (latent), L-lysine (latent), ethylamine (latent) and cadaverine are assimilated as sole nitrogen sources. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Physiologically, M. marina differs from the closely related species M. pileae (Fig. 20) in its inability to assimilate D-arabinose, glycerol and ribitol and the ability to assimilate L-sorbose, soluble starch, DL-lactic acid and citric acid (Table S2.33).

Typus: China, Haijing road of Basuo town, Dongfang, Hainan province, obtained from seawater, Feb. 2022, Q.-M. Wang (holotype CGMCC 2.6912^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = HJLB11-102).

Vankyiozymaceae Q.-M. Wang fam. nov. MycoBank MB846644.

Member of the Georgefischeriales (Exobasidiomycetes, Ustilaginomycotina). The diagnosis of the family Vankyiozymaceae is based on the genus Vankyiozyma. The nomenclature of the family is based on the genus Vankyiozyma.

Type genus: Vankyiozyma Q.-M. Wang.

Genus accepted: Vankyiozyma Q.-M. Wang.

Vankyiozyma Q.-M. Wang gen. nov.MycoBank MB846645.

Etymology: the genus is named in honour of Kálmán Vánky for his for his for his contributions to smut taxonomy.

This genus is proposed for the separate clade represented by MT-2A-31. Member of the Georgefischeriales. The genus is mainly circumscribed by the phylogenetic analysis of the six genes dataset, in which it occurred as a separate branch within the Georgefischeriales (Fig. 3).

Sexual reproduction not known. Colonies yellowish, rough and dimmed. Budding cells present. Pseudohyphae and hyphae not produced. Ballistoconidia not formed.

Type species: Vankyiozyma motuoensis Q.-M. Wang.

Vankyiozyma motuoensis Q.-M. Wang, sp. nov.MycoBank MB846646. Fig. 27E.

Etymology: the specific epithet motuoensis refers to the geographic origin of the type strain, Motuo, Tibet.

Culture characteristics: In YM broth, after 7 days at 17 ºC, cells are cylindrical, fusiformis and ellipsoidal, 1.71–2.59 × 5.55–12.04 µm and single, budding is polar (Fig. 27E), a sediment is formed. After 1 month at 17 ºC, a pellicle is present and the sediment disappears. On YM agar, after 1 month at 17 ºC, the streak culture is pale yellow, leathery, rough and dimmed. The margin is fimbricate. In Dalmau plate culture on corn meal agar, pseudohyphae are not formed. Sexual structures are not observed on YM, PDA, V8 and CM agar. Ballistoconidia are not produced.

Physiological and biochemical characteristics: Glucose fermentation is absent. Glucose, maltose, cellobiose, raffinose, melezitose, soluble starch, D-xylose, L-arabinose (weak), D-arabinose, D-mannitol, α-methyl-D-glucoside, are assimilated as sole carbon sources. Galactose, L-sorbose, sucrose, trehalose, lactose, melibiose, inulin, D-ribose, L-rhamnose, D-glucosamine, N-acetyl-D-glucosamine, methanol, ethanol, glycerol, erythritol, ribitol, galactitol, D-glucitol, salicin, DL-lactic acid, succinic aid, citric acid, inositol, hexdecane and D-glucuronic acid are not assimilated. Ethylamine and cadaverine assimilated as sole nitrogen sources. Ammonium sulfate, potassium nitrate, sodium nitrite and L-lysine are not assimilated. Maximum growth temperature is 30 ºC. Growth in vitamin-free medium is positive. Starch-like substances are not produced. Growth on 50% (w/w) glucose-yeast extract agar is negative. Urease activity is positive. Diazonium Blue B reaction is positive.

Typus: China, Motuo county, Tibet, obtained from a leaf of an unidentified plant, Oct. 2021, Q.-M. Wang (holotype CGMCC 2.6855^T preserved in a metabolically inactive state, ex-type JCM xxx = TISTR xxx = MT-2A-31).

Validated taxa

Guomyces Q.-M. Wang, E. Tanaka, M. Groenew. & D. Begerow, gen. nov. Mycobank MB846647.

For a description see Frontiers in Microbiology 12 (no. 777338): 19 (2022).

Type species: Guomyces nicotianae H.K. Wang & F.C. Lin ex Q.-M. Wang, E. Tanaka, M. Groenew. & D. Begerow.

Synonym: Guomyces Q.-M. Wang, E. Tanaka, M. Groenew. & D. Begerow, Frontiers in Microbiology 12 (no. 777338): 19 (2022), nom. inval., Art. 40.1 (Shenzhen).

Guomyces nicotianae H.K. Wang & F.C. Lin ex Q.-M. Wang, E. Tanaka, M. Groenew. & D. Begerow, sp. nov. Mycobank MB846648.

For a description see Frontiers in Microbiology 12 (no. 777338): 19 (2022).

Synonyms: Meira nicotianae H.K. Wang & F.C. Lin ex Denchev & T. Denchev, Mycobiota 11: 3 (2021)

= Meira nicotianae H.K. Wang & F.C. Lin, Phytotaxa 365 (2): 176 (2018), nom. inval., Art. 35.1 (Shenzhen).

Contributions

Q.-M.W. conceived and designed the project. Q.-M.W., G.-S.W. and W.-N.Z. performed sampling and yeast isolation., Y.-L.J., W.-J.B., G.-S.W., M.-M.W., Y.-Y.L., F.L. and X.-H.C. performed phenotypic characterization and analyzed the molecular data. A.-H. Li deposited the new taxa. Q.-M.W. and M.-M.W. wrote the paper.

Acknowledgements

This study was supported by grants No. 319611330200 and No. 31770018 from the National Natural Science Foundation of China (NSFC), No. 2021FY100905 and No. 2019FY100700 from the Ministry of Science and Technology of China, No. 521000981388 from Advanced Talents Incubation Program of the Hebei University. The authors are solely responsible for the content of this work.

Conflict of interest

The authors declare that there are no conflicts of interest

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Tables 1 is available in the Supplementary Files section.

TableS1.xlsx
TableS2.1Derxomyces.xlsx
TableS2.2Hannaella.xlsx
TableS2.5Kockovaella.xlsx
TableS2.6Papiliotrema.xlsx
TableS2.7Pseudotremella.xlsx
TableS2.8Naematelia.xlsx
TableS2.9Vishniacozyma.xlsx
TableS2.11Kwoniella.xlsx
TableS2.12Phaeotremella.xlsx
TableS2.13Vanrija.xlsx
TableS2.14Takashimella.xlsx
TableS2.15Holtermannia.xlsx
TableS2.16Piskurozyma.xlsx
TableS2.17Mrakia.xlsx
TableS2.18Phaffia.xlsx
TableS2.19Kondoa.xlsx
TableS2.20Boekhoutia.xlsx
TableS2.21Buckleyzyma.xlsx
TableS2.22Microsporomyces.xlsx
TableS2.23Halobasidium.xlsx
TableS2.24Cystobasidium.xlsx
TableS2.25Rhodotorula.xlsx
TableS2.26Nakasozyma.xlsx
TableS2.27Curvibasidium.xlsx
TableS2.28Chrysozyma.xlsx
TableS2.29Yurkovia.xlsx
TableS2.30Pseudohyphozyma.xlsx
TableS2.31Slooffia.xlsx
TableS2.32Acaromyces.xlsx
TableS2.33Meria.xlsx
s1.jpg
Fig. S1.Phylogenetic tree inferred using the LSU rDNA D1/D2 domains, depicting the phylogenetic positions of the new genus Foliozyma (Mrakiaceae, Cystofilobasidiales, Tremellomycetes, Agaricomycotina) by maximum likelihood analysis and over 50% from 1000 bootstrap replicates is shown. Bar = 0.05 substitutions per nucleotide position.
Table1.docx
Supportingtablecaptions.docx

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Proposal of one new order, two new families, nine new genera and seventy-eight new basidiomycetous yeast species isolated from China

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