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Research Article
Five new epiphytic species of Vishniacozyma (Bulleribasidiaceae, Tremellales) from China
expand article infoShan Liu‡, Dan-Yang Cai‡, Chun-Yue Chai‡, Feng-Li Hui‡
‡ Nanyang Normal University, Nanyang, China

Corresponding author: Feng-Li Hui ( fenglihui@yeah.net )

Academic editor: Margarita Dueñas
© 2025 Shan Liu, Dan-Yang Cai, Chun-Yue Chai, Feng-Li Hui.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Liu S, Cai D-Y, Chai C-Y, Hui F-L (2025) Five new epiphytic species of Vishniacozyma (Bulleribasidiaceae, Tremellales) from China. MycoKeys 113: 321-336. https://doi.org/10.3897/mycokeys.113.140598
Open Access

Abstract

The genus Vishniacozyma, globally distributed, encompasses numerous epiphytic and endophytic species. In this study, five new species are proposed to accommodate eleven yeast strains isolated from leaves of different plants: V. diospyri sp. nov. (holotype CICC 33574T), V. guiyangensis sp. nov. (holotype CICC 33569T), V. pingtangensis sp. nov. (holotype CICC 33596T), V. eriobotryae sp. nov. (holotype GDMCC 2.312T), and V. tianchiensis sp. nov. (holotype CICC 33617T) using phenotypic and phylogenetic characters. Phylogenetic analysis was based on the internal transcribed spacer (ITS) region and the D1/D2 domain of the large subunit (LSU) rRNA gene. Illustrations and descriptions of these five taxa are provided, along with comparative analyses with closely related species within the genus. This research highlights the considerable diversity of Vishniacozyma species in China and contributes valuable data for future investigations in fungal systematics and evolution.

Key words:

Basidiomycota, new species, phylloplane, phylogenetic analysis, taxonomy

Introduction

The genus Vishniacozyma, typified by V. carnescens (Verona & Luchetti) X.Z. Liu, F.Y. Bai, M. Groenew. & Boekhout, was established by Liu et al. (2015) based on results of their seven-marker phylogeny. In this study, 11 previously described species in the genera Bullera, Cryptococcus, and Trimorphomyces were transferred to Vishniacozyma. Subsequently, 21 new species, including V. ellesmerensis M. Tsuji, Y. Tanabe, W.F. Vincent & Mas (Tsuji et al. 2019), V. changhuana C.F. Lee & Chin F. Chang, V. taiwanica C.F. Lee & Chin F. Chang (Chang et al. 2019), V. kurtzmanii Yurkov (Yurkov and Kurtzman 2019), V. alagoana C.R. Félix, D.A. Andrade, J.H. Almeida, H.M. Navarro, Fell & Landell (Félix et al. 2020), V. phoenicis Kachalkin, A.S. Venzhik & Tomash (Crous et al. 2020), V. europaea Q.M. Wang, F.Y. Bai & A.H. Li, V. melezitolytica Q.M. Wang, F.Y. Bai & A.H. Li, V. pseudopenaeus Q.M. Wang, F.Y. Bai & A.H. Li (Li et al. 2020), V. insularis Y.P. Tan, Marney & R.G. Shivas (Tan et al. 2021), V. pseudodimennae X.Z. Liu, F.Y. Bai & X.Y. Wei (Wei et al. 2022), V. terrae Y. Park, Maeng & S. Sriniv (Maeng et al. 2023), V. pseudocarnescens H.Y. Zhu, X.Z. Liu & F.Y. Bai (Zhu et al. 2023), V. catalpae Q.M. Wang, V. marinae Q.M. Wang, V. paravictoriae Q.M. Wang, V. pini Q.M. Wang, V. pyri Q.M. Wang, V. sinopodophylli Q.M. Wang, V. zhenxiongensis Q.M. Wang (Jiang et al. 2024), and V. floricola Péter, Álvarez-Pérez & Dlauchy (Dlauchy et al. 2024) have been accepted in the genus.

To date, the sexual state of Vishniacozyma is only known in V. nebularis (Vishniac) A.M. Yurkov, which was found on a dead branch from Taiwan (Kirschner and Chen 2008). The remaining representatives of the genus are asexual morphs resembling yeast stages from the genus Cryptococcus and reproduce via budding. Some species may form ballistoconidia and poorly developed pseudohyphae (Liu et al. 2015). Physiologically, the members of the genus lack fermentative ability, possess CoQ-9 or Q-10 as a predominant ubiquinone, and assimilate various carbon sources, but not methanol and hexadecane (Liu et al. 2015; Zhu et al. 2023).

Vishniacozyma has a crucial role in maintaining forest biodiversity, so some species, such as V. changhuana and V. taiwanica, are abundant in the mangrove ecosystem (Chang et al. 2019; Nimsi et al. 2023). Besides ecological functions, biocontrol, biotechnological, and medicinal values of this genus are also notable. For instance, V. foliicola (Q.M. Wang & F.Y. Bai) A.M. Yurkov and V. victoriae (M.J. Montes, Belloch, Galiana, M.D. García, C. Andrés, S. Ferrer, Torr.-Rodr. & J. Guinea) X.Z. Liu, F.Y. Bai, M. Groenew. & Boekhout, as endophytes, inhibit the growth of phytopathogenic fungi to control postharvest diseases in stored fruit (Bilański and Kowalski 2022; Gorordo et al. 2022; Nian et al. 2023). V. psychrotolerans (V. de García, Zalar, Brizzio, Gunde-Cim. & van Broock) A.M. Yurkov can accumulate lipids for biodiesel production (Deeba et al. 2017, 2018). V. victoriae is a suitable candidate for the production of fatty acids and ergosterol at the industrial scale (Villarreal et al. 2018).

Vishniacozyma is a ubiquitous genus, widely distributed in subtropical, tropical, temperate, and even cold regions, including America, Asia, Australia, and Europe (Montes et al. 1999; Vishniac 2002; de Garcia et al. 2012; Tsuji et al. 2019; Yurkov and Kurtzman 2019; Crous et al. 2020; Félix et al. 2020; Li et al. 2020; Tan et al. 2021; Zhu et al. 2023; Jiang et al. 2024). Vishniacozyma species are mainly endophytes, epiphytes, and saprophytes of plants, especially of the leaves (Wang et al. 2011; Félix et al. 2020; Li et al. 2020; Tan et al. 2021; Jiang et al. 2024), but they can also grow across diverse terrestrial environments, including soil (Li et al. 2020; Maeng et al. 2023; Jiang et al. 2024), air inside caves (Ogórek et al. 2021), and dogs (Harvey et al. 2023).

Vishniacozyma seems to be a very diversified genus in China, since of the 32 species included in them, 15 have been described from this country (Kirschner and Chen 2008; Wang et al. 2011; Chang et al. 2019; Li et al. 2020; Wei et al. 2022; Zhu et al. 2023; Jiang et al. 2024), but to know the true richness of Vishniacozyma species, more studies are necessary in different areas. The aim of this study is to contribute to knowledge of this genus, based on eleven strains of tremellomycetous fungi isolated from different areas across China. According to molecular (ITS and LSU) and phenotypic analyses, they were identified as five new species of Vishniacozyma that are described and illustrated in this paper.

Materials and methods

Sample collection and yeast isolation

Samples were collected from Guizhou and Henan Provinces of China. Yeast strains were isolated from leaf surfaces using the improved ballistospore-fall method as described by Nakase and Takashima (1993). Specifically, the fresh and healthy leaves were sectioned into small pieces and attached to the inner lid of a Petri dish using a thin layer of petroleum jelly. The dish contained yeast extract-malt extract (YM) agar (0.3% yeast extract, 0.3% malt extract, 0.5% peptone, 1% glucose, and 2% agar) with supplemented 0.01% chloramphenicol to limit bacterial growth. Plates were incubated at 20 °C and monitored daily for the presence of colonies, which were selected and purified by streaking them on separate YM agar plates. Following purification, yeast strains were suspended in 20% (v/v) glycerol and stored at −80 °C. Cultures of all isolates were preserved at the Microbiology Lab, Nanyang Normal University, Henan, China, cultures type are preserved as a metabolically inactive state in the CICC (China Centre of Industrial Culture Collection, Beijing, PR China) and cultures ex-type in the PYCC (Portuguese Yeast Culture Collection, Caparica, Portugal).

Phenotypic examination

Morphological characterization and physiological and biochemical tests were carried out according to standard methods described by Kurtzman et al. (2011). The potential sexual reproduction in new species was assessed on corn meal agar (CMA: 2.5% corn starch and 2% agar), potato dextrose agar (PDA: 20% potato infusion, 2% glucose, and 2% agar), and V8 agar (10% V8 juice and 2% agar) (Li et al. 2020). A loopful of cells of each test strain is mixed on an agar plate incubated at 17 °C for up to eight weeks. The ballistoconidium-forming activity of all new species was observed by the inverted-plate method (do Carmo-Sousa and Phaff 1962) using CMA at 17 °C. Glucose fermentation tests were carried out in a liquid medium using Durham fermentation tubes. Assimilation of carbon and nitrogen compounds was examined in a liquid medium, and starved inoculum was employed for the nitrogen test (Kurtzman et al. 2011). Growth at different temperatures (15, 20, 25, 30, 35, and 37 °C) was determined by cultivation on YM agar. Cell morphology was examined using LEICA DM2500 cameras (LEICA, Wetzlar, Germany) and LAS V4.13 software. All novel taxonomic descriptions and proposed names were deposited in the MycoBank database (http://www.mycobank.org).

DNA extraction, PCR, and sequencing

The genomic DNA was extracted from yeast strains using the Ezup Column Yeast Genomic DNA Purification Kit, according to the manufacturer’s directions (Sangon Biotech Co., Shanghai, China). The ITS region and the D1/D2 domain of the LSU rRNA gene were amplified using the ITS1/ITS4 (White et al. 1990) and NL1/NL4 (Kurtzman and Robnett 1998) primers, respectively. The amplifications were conducted in a 25 µL reaction-volume tube containing 9.5 µL of ddH2O, 12.5 µL of 2× Taq PCR Master Mix with blue dye (Sangon Biotech Co., Shanghai, China), 1 µL of DNA template, and 1 µL of each primer. The following parameters were used to amplify the ITS and D1/D2 regions: an initial denaturation step of 2 min at 95 °C, followed by 35 cycles of 30 s at 95 °C, 30 s at 51 °C, 40 s at 72 °C, and a final extension of 10 min at 72 °C (Toome et al. 2013). The PCR products were purified and sequenced by Sangon Biotech Co., Ltd. (Shanghai, China) using the same primers. All newly generated sequences were deposited in the GenBank database (https://www.ncbi.nlm.nih.gov/genbank/; Table 1).

Table 1.
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Species name, strain numbers, and GenBank accession numbers for the phylogenetic analyses performed in the present study. Entries in bold were newly generated for this study.

Species name Strain number Locality GenBank accession number
ITS LSU D1/D2
Vishniacozyma alagoana CBS 15966T Brazil MH885328 MH909005
Vishniacozyma carnescens CBS 973T Italy NR_130695 NG_058430
Vishniacozyma catalpae CGMCC 2.6902T China OP470302 OP470206
Vishniacozyma changhuana CBS 16556T Taiwan NR_182838 MT906468
Vishniacozyma dimennae CBS 5770T New Zealand NR_144808 NG_058431
Vishniacozyma diospyri NYNU 221044T China OP954624 OP954569
Vishniacozyma diospyri NYNU 2211329 China PQ496711 PQ496709
Vishniacozyma ellesmerensis JCM 32573T Canadian Arctic NR_173768 LC335797
Vishniacozyma eriobotryae NYNU 229203T China OP566897 OP566895
Vishniacozyma eriobotryae NYNU 229144 China PP580371 PP580369
Vishniacozyma europaea CGMCC 2.3099T China NR_174757 MK050335
Vishniacozyma floricola NCAIM Y.02320T Hungary PP337022 PP261370
Vishniacozyma foliicola CGMCC 2.2471T China NR_144809 NG_067769
Vishniacozyma globispora CBS 6981T Canada NR_073235 NG_070507
Vishniacozyma guiyangensis NYNU 22831T China OP566869 OP566870
Vishniacozyma guiyangensis NYNU 236231 China PP580372 PP580370
Vishniacozyma guiyangensis NYNU 236232 China PP580374 PP580373
Vishniacozyma heimaeyensis CBS 8933T Iceland NR_077070 NG_058432
Vishniacozyma insularis BRIP 28256T Australia NR_175761 –
Vishniacozyma kurtzmanii CBS 12229T USA NR_168771 FR820582
Vishniacozyma marinae CGMCC 2.6837T China OP470294 OP470198
Vishniacozyma melezitolytica CGMCC 2.3472T China NR_174755 MK050330
Vishniacozyma nebularis CBS 12283T Taiwan – EU266921
Vishniacozyma paravictoriae CGMCC 2.6918T China OP470300 OP470204
Vishniacozyma peneaus CBS 2409T USA NR_165987 AB035051
Vishniacozyma phoenicis KBP Y-6564T Russia MN449981 MN449981
Vishniacozyma pingtangensis NYNU 23281T China OQ851896 OQ851894
Vishniacozyma pingtangensis NYNU 236258 China PQ496708 PQ496707
Vishniacozyma pini CGMCC 2.6849T China OP470296 OP470200
Vishniacozyma pseudocarnescens CGMCC 2.6457T China OR077051 OR077057
Vishniacozyma pseudodimennae CGMCC 2.6790T China OM417179 OM417179
Vishniacozyma pseudopenaeus CGMCC 2.3165T China NR_174756 MK050333
Vishniacozyma psychrotolerans CBS 12690T Portugal NR_111656 JN193445
Vishniacozyma pyri CGMCC 2.6870T China OP470298 OP470202
Vishniacozyma sinopodophylli CGMCC 2.6857T China OP470297 OP470201
Vishniacozyma taibaiensis CBS 9919T Taiwan NR_144810 NG_058434
Vishniacozyma taiwanica BCRC 23477 Taiwan NR_182839 MT906477
Vishniacozyma tephrensis CBS 8935T Iceland NR_144812 KX507032
Vishniacozyma terrae KCTC 27988T Korea MZ734447 NG_241953
Vishniacozyma tianchiensis NYNU 236163T China OR426458 OR426457
Vishniacozyma tianchiensis NYNU 2311236 China PQ496716 PQ496715
Vishniacozyma victoriae CBS 8685T Antarctica NR_073260 AF363647
Vishniacozyma zhenxiongensis CGMCC 2.6901T China OP470301 OP470205
Tremella globispora CBS 6972T Canada AF444432 AF189869
Tremella flava CBS 8471T Taiwan NR_155935 AF042221
Tremella taiwanensis CBS 8479T Taiwan AF042412 AF042230
Tremella resupinata CBS 8488T Taiwan AF042421 AF042239
Tremella brasiliensis CBS 6966T Costa Rica AF444429 AF189864
Apiotrichum porosum CBS 2040T Germany AF414694 AF189833

T, type strain.

Phylogenetic analysis

Sequences from 49 strains were employed for phylogenetic analysis. Apiotrichum porosum Stautz, CBS2040, was chosen as the outgroup. Phylogenetic analyses were based on a combined ITS and LSU dataset. Aside from the newly generated sequences, additional related sequences were obtained from GenBank (Table 1). Sequences for the individual loci were aligned using MAFFT v.7.110 (Katoh and Standley 2013) with the G-INI-I option. Poorly aligned regions were removed and manually adjusted using MEGA v.11 (Tamura et al. 2021). PhyloSuite v.1.2.2 (Zhang et al. 2020) was employed to concatenate the aligned sequences of the different loci.

Maximum likelihood (ML) analyses were performed using RAxML v.8.2.3 (Stamatakis 2014) under a GTRGAMMA model with one thousand rapid bootstrap (BS) replicates. Bayesian inference (BI) analyses were conducted using MrBayes v.3.2.2 with a GTR + I + G model of DNA substitution and a gamma distribution rate variation across locations (Ronquist et al. 2012). Two independent runs were employed, and each run had four chains and began from random trees. Trees were sampled every 1000th generation, and the first 25% of trees were removed, while the other 75% of trees were retained to construct a 50% majority consensus tree and calculated Bayesian posterior probabilities (BPP). Each tree was visualized with its BS and BPP bootstrap values using Figtree v.1.4.3 (Andrew 2016).

Results

Molecular phylogeny

The total length of the concatenated dataset of two loci across the 49 samples was 1098 bp, including 493 bp for ITS and 605 bp for LSU. ML and BI methods generated similar topologies in main lineages, and only the topology generated by the ML method was presented along with BS values and BPP above 50% and 0.95, respectively, at the nodes (Fig. 1).

Figure 1. 

Maximum likelihood (ML) phylogenetic tree of Vishniacozyma founded on combined ITS and LSU sequence data. The tree was rooted with Apiotrichum porosum CBS 2040. The bootstrap values and Bayesian posterior probabilities over 50%/0.95 (BS/BPP) are indicated at the nodes. Sequences from type strains are marked with (T), and the new species are indicated in bold.

In the phylogenetic tree (Fig. 1), Vishniacozyma was monophyletic with well statistical support (BS/95, BPP/1.0). Eleven newly isolated strains formed five distinct and well-supported lineages distant from other Vishniacozyma species.

Taxonomy

According to the phylogenetic and phenotypic analysis, five new species of Vishniacozyma are described.

Vishniacozyma diospyri C.Y. Chai & F.L. Hui, sp. nov.

MycoBank No: 853420
Figs 2A, F

Etymology.

The specific epithet diospyri refers to Diospyros, the name of the genus from which the type strain was isolated.

Typus.

China. • Henan Prov.: Neixiang Co., Baotianman Nature Reserve (33°29'07"N, 111°52'51"E), Sep 2022, J.Z. Li, in the phylloplane of Diospyros lotus, NYNU 221044 (holotype CICC 33574T, GenBank: OP954624, OP954569); culture ex-type PYCC 9955.

Description.

On YM agar after seven days at 20 °C, the streak culture is yellowish-cream, butyrous, and smooth, with an entire margin. After three days in YM broth at 20 °C, cells are ovoid, ellipsoidal, and cylindrical, 2.5–4.4 × 4.1–13.7 μm, and single; budding is polar. After one month at 20 °C, a ring and sediment are present. In Dalmau plate culture on CMA, pseudohyphae and hyphae are not formed. Sexual structures are not observed on PDA, CMA or V8 agar. On corn meal agar, ballistoconidia are not produced. Glucose fermentation is absent. Glucose, sucrose, raffinose, melibiose, galactose, lactose, trehalose, maltose, melezitose, methyl-α-D-glucoside, cellobiose, salicin (weak), L-sorbose, L-rhamnose, D-xylose, L-arabinose, D-arabinose, 5-keto-D-gluconate, D-ribose, glycerol (delayed), erythritol, ribitol (delayed), galactitol, D-mannitol, D-glucitol, myo-inositol, DL-lactate, succinate, D-gluconate, D-glucosamine, N-acetyl-D-glucosamine, 2-keto-D-gluconate, D-glucuronate, and glucono-1,5-lactone are assimilated as sole carbon sources. Inulin, methanol, and ethanol are not assimilated. Nitrite and L-lysine are assimilated as sole nitrogen sources. Nitrate, ethylamine, and cadaverine are not assimilated. Maximum growth temperature is 25 °C. Growth in vitamin-free medium is positive. Growth on 50% (w/w) glucose-yeast extract agar is negative. Starch-like substances are not produced. Urease activity is positive. Diazonium Blue B reaction is positive.

Figure 2. 

Budding cells of A Vishniacozyma diospyri sp. nov. NYNU 221044T B V. guiyangensis sp. nov. NYNU 22831T C V. pingtangensis sp. nov. NYNU 23281T D V. eriobotryae sp. nov. NYNU 229203T E V. tianchiensis sp. nov. NYNU 236163T following growth in YM broth for three days at 20 °C F Elongate budding cells of V. diospyri sp. nov. NYNU 221044T following growth on CMA for seven days at 20 °C. Scale bars: 10 μm.

Additional strain examined.

China. • Henan Prov.: Neixiang Co., Baotianman Nature Reserve (33°29'07"N, 111°52'51"E), in the phylloplane of Cornus officinalis, Sep 2022, J.Z. Li, NYNU 2211329 (GenBank: PQ496711, PQ496709).

Note.

In the phylogenetic analyses, V. diospyri formed a separate branch that clustered with V. catalpae with high support (BS/99, BPP/1.0; Fig. 1). However, V. diospyri differs from V. catalpae by 20 nucleotides (12/486 in ITS and 8/559 in LSU) and by its ability to assimilate D-arabinose, succinate, and D-glucosamine and its inability to assimilate inulin, ethylamine, and cadaverine.

Vishniacozyma guiyangensis C.Y. Chai & F.L. Hui, sp. nov.

MycoBank No: 853421
Fig. 2B

Etymology.

The specific epithet guiyangensis refers to the geographic origin of the type strain, Guiyang city, Guizhou Province.

Typus.

China. • Guizhou Prov.: Guiyang City, Guiyang Medicinal Botanical Garden (26°34'51"N, 106°42'36"E), Aug 2022, in the phylloplane of Distylium racemosum, L. Zhang & F.L. Hui, NYNU 22831 (holotype CICC 33569T, GenBank: OP566869, OP566870); culture ex-type PYCC 9934.

Description.

On YM agar after seven days at 20 °C, the streak culture is white-cream, mucoid, glistening, and smooth, with an entire margin. After three days in YM broth at 20 °C, cells are ovoid to ellipsoidal, 2.5–3.9 × 4.1–7.4 μm, and single; budding is polar. After one month at 20 °C, sediment is present. In Dalmau plate culture on CMA, pseudohyphae and hyphae are not formed. Sexual structures are not observed on PDA, CMA or V8 agar. On corn meal agar, ballistoconidia are not produced. Glucose fermentation is absent. Glucose, sucrose, raffinose, melibiose, galactose, lactose, trehalose, maltose, melezitose, methyl-α-D-glucoside, cellobiose, L-sorbose (delayed and weak), L-rhamnose, D-xylose, L-arabinose, D-ribose, glycerol, erythritol, galactitol (delayed), D-mannitol, D-glucitol, myo-inositol, DL-lactate, D-gluconate, D-glucosamine, N-acetyl-D-glucosamine, 2-keto-D-gluconate, and D-glucuronate are assimilated as sole carbon sources. Inulin, salicin, D-arabinose, 5-keto-D-gluconate, methanol, ethanol, ribitol, succinate, and glucono-1,5-lactone are not assimilated. Nitrate, nitrite, ethylamine, L-lysine, and cadaverine are not assimilated as sole nitrogen sources. Maximum growth temperature is 25 °C. Growth in vitamin-free medium is negative. Growth on 50% (w/w) glucose-yeast extract agar is positive. Starch-like substances are not produced. Urease activity is positive. Diazonium Blue B reaction is positive.

Additional strain examined.

China. • Henan Prov.: Songxian Co., Tianchi mountain (34°32'27"N, 112°16'39"E), Jun 2023, in the phylloplane of Morus alba, J.Z. Li, NYNU 236231 (GenBank: PP580372, PP580370), NYUN 236232 (GenBank: PP580374, PP580373).

Note.

In the phylogenetic analyses, three strains of V. guiyangensis clustered in a single clade with full support values (BS/100, BPP/1.0; Fig. 1), and V. guiyangensis was close to a clade formed by V. pingtangensis and V. eriobotryae, described in this study with significant support (BS/97, BPP/1.0; Fig. 1). V. guiyangensis differs from V. pingtangensis by 92 nucleotides (65/501 in ITS and 27/600 in LSU) and from V. eriobotryae by 64 nucleotides (17/596 in ITS and 47/596 in LSU). Physiologically, V. guiyangensis can be differentiated from these species by its inability to assimilate salicin, D-arabinose, 5-keto-D-gluconate, ribitol, succinate, and glucono-1,5-lactone. In addition, V. guiyangensis can grow on 50% (w/w) glucose-yeast extract agar, while the other two species cannot.

Vishniacozyma pingtangensis C.Y. Chai & F.L. Hui, sp. nov.

MycoBank No: 853422
Fig. 2C

Etymology.

The specific epithet pingtangensis refers to the geographic origin of the type strain, Pingtang County, Guizhou Province.

Typus.

China. • Guizhou Prov.: Pingtang Co., Sifangjing Vil. (25°7'45'N, 107°2'54"E), Feb 2023, in the phylloplane of Acer saccharum, D. Lu, NYNU 23281 (holotype CICC 33596T, GenBank: OQ851896, OQ851894); culture ex-type PYCC 9976.

Description.

On YM agar after seven days at 20 °C, the streak culture is white-cream, mucoid, glistening, and smooth, with an entire margin. After three days in YM broth at 20 °C, cells are ellipsoida, 2.4–3.7 × 3.7–6.4 μm, and single; budding is polar. After one month at 20 °C, a ring and sediment are present. In Dalmau plate culture on CMA, pseudohyphae and hyphae are not formed. Sexual structures are not observed on PDA, CMA or V8 agar. On corn meal agar, ballistoconidia are not produced. Glucose fermentation is absent. Glucose, sucrose, raffinose, melibiose, galactose, lactose, trehalose, maltose, melezitose, methyl-α-D-glucoside, cellobiose, salicin, L-sorbose (weak), L-rhamnose, D-xylose, L-arabinose, D-arabinose, 5-keto-D-gluconate, D-ribose, ethanol (weak), glycerol (delayed and weak), ribitol, galactitol, D-mannitol, D-glucitol, myo-inositol, DL-lactate (delayed), succinate (weak), D-gluconate, D-glucosamine (weak), N-acetyl-D-glucosamine, 2-keto-D-gluconate, D-glucuronate (weak), and glucono-1,5-lactone are assimilated as sole carbon sources. Inulin, methanol, and erythritol are not assimilated. Nitrate, nitrite (weak), ethylamine, L-lysine (weak), and cadaverine (delayed) are assimilated as sole nitrogen sources. Maximum growth temperature is 25 °C. Growth in vitamin-free medium is positive. Growth on 50% (w/w) glucose-yeast extract agar is negative. Starch-like substances are not produced. Urease activity is positive. Diazonium Blue B reaction is positive.

Additional strain examined.

China. • Henan Prov.: Songxian Co., Tianchi mountain (34°32'27"N, 112°16'39"E), Jun 2023, in the phylloplane of Morus alba, J.Z. Li, NYNU 236258 (GenBank: PQ496708, PQ496707).

Note.

In the phylogenetic analyses, V. pingtangensis was closely related to V. eriobotryae; both differ by 102 nucleotides (52/487 in ITS and 50/600 in LSU). Physiologically, they can be differentiated because V. eriobotryae has the ability to assimilate glycerol and the inability to assimilate inulin and erythritol, in addition to being able to grow at 30 °C, while V. pingtangensis cannot.

Vishniacozyma eriobotryae C.Y. Chai & F.L. Hui, sp. nov.

MycoBank No: 853423
Fig. 2D

Etymology.

The specific epithet eriobotryae refers to Eriobotrya, the plant genus from which the type strain was isolated.

Typus.

China. • Guizhou Prov.: Guiyang City, Guiyang Medicinal Botanical Garden (26°34'51"N, 106°42'36"E), Aug 2022, in the phylloplane of Eriobotrya japonica, L. Zhang & F.L. Hui, NYNU 229203 (holotype GDMCC 2.312T, GenBank: OP566897, OP566895); culture ex-type PYCC 9940.

Description.

On YM agar after seven days at 20 °C, the streak culture is yellowish cream, mucoid, glistening, and smooth, with an entire margin. After three days in YM broth at 20 °C, cells are ovoid and ellipsoidal, 2.2–3.2 × 3.7–5.5 μm, and single; budding is polar. After one month at 20 °C, a ring and sediment are present. In Dalmau plate culture on CMA, pseudohyphae and hyphae are not formed. Sexual structures are not observed on PDA, CMA or V8 agar. On corn meal agar, ballistoconidia are not produced. Glucose fermentation is absent. Glucose, inulin (weak), sucrose, raffinose, melibiose, galactose, lactose, trehalose, maltose, melezitose, methyl-α-D-glucoside, cellobiose, salicin, L-sorbose (weak), L-rhamnose (weak), D-xylose, L-arabinose, D-arabinose, 5-keto-D-gluconate, ethanol (delayed and weak), erythritol, ribitol, galactitol, D-mannitol, D-glucitol, myo-inositol, DL-lactate (delayed and weak), succinate, D-gluconate, D-glucosamine (delayed and weak), N-acetyl-D-glucosamine, 2-keto-D-gluconate, D-glucuronate, and glucono-1,5-lactone are assimilated as sole carbon sources. Methanol and glycerol are not assimilated. Nitrate (delayed and weak), nitrite (delayed and weak), ethylamine (delayed and weak), and cadaverine (delayed and weak) are assimilated as sole nitrogen sources. Maximum growth temperature is 30 °C. Growth in vitamin-free medium is positive. Growth on 50% (w/w) glucose-yeast extract agar is negative. Starch-like substances are not produced. Urease activity is positive. Diazonium Blue B reaction is positive.

Additional strain examined.

China. • Guizhou Prov.: Guiyang City, Guiyang Medicinal Botanical Garden (26°34'51"N, 106°42'36"E), Aug 2022, in the phylloplane of Buddleja davidii, L. Zhang & F.L. Hui, NYNU 229144 (GenBank: PP580371, PP580369).

Note.

V. eriobotryae assimilated nitrate as a sole source of nitrogen in a liquid medium, but not nitrite. If a yeast strain assimilates nitrate, it is expected to assimilate nitrite as well (Kurtzman et al. 2011). Nitrite is rather toxic, therefore, the auxanographic method was used to test the ability to utilize nitrite. On solid media, V. eriobotryae can weakly assimilate nitrate as a sole nitrogen source.

Vishniacozyma tianchiensis C.Y. Chai & F.L. Hui, sp. nov.

MycoBank No: 853424
Fig. 2E

Etymology.

The specific epithet tianchiensis refers to the geographic origin of the type strain, Tianchi Mountain, Songxian County, Henan Province.

Typus.

China. • Henan Prov.: Songxian Co., Tianchi mountain (34°32'27"N, 112°16'39"E), Jun 2023, in the phylloplane of Salix matsudana, J.Z. Li, NYNU 236163 (holotype CICC 33617T, GenBank: OR426458, OR426457); culture ex-type PYCC 9988.

Description.

On YM agar after seven days at 20 °C, the streak culture is yellowish cream, mucoid, glistening, and smooth, with an entire margin. After three days in YM broth at 20 °C, cells are globose, 3.7–5.4 × 4–6.4 μm, and single; budding is polar. After one month at 20 °C, a ring and sediment are present. In Dalmau plate culture on CMA, pseudohyphae and hyphae are not formed. Sexual structures are not observed on PDA, CMA or V8 agar. On corn meal agar, ballistoconidia are not produced. Glucose fermentation is absent. Glucose, inulin, sucrose, raffinose, melibiose, galactose, lactose, trehalose, maltose, methyl-α-D-glucoside (delayed), cellobiose, salicin (weak), L-sorbose, L-rhamnose, D-xylose, L-arabinose, D-arabinose (weak), 5-keto-D-gluconate, D-ribose, glycerol (weak), erythritol, ribitol, galactitol, D-mannitol, D-glucitol, myo-inositol, DL-lactate, succinate, D-gluconate, D-glucosamine (delayed and weak), N-acetyl-D-glucosamine (weak), 2-keto-D-gluconate, D-glucuronate, and glucono-1,5-lactone are assimilated as sole carbon sources. Melezitose, methanol, ethanol, and citrate are not assimilated. Nitrate, nitrite, ethylamine, L-lysine, and cadaverine (weak) are assimilated as sole nitrogen sources. Maximum growth temperature is 25 °C. Growth in vitamin-free medium is positive. Growth on 50% (w/w) glucose-yeast extract agar is positive. Starch-like substances are not produced. Urease activity is positive. Diazonium Blue B reaction is positive.

Additional strain examined.

China. Henan Prov.: Xixia Co., Funiu Mountain (33°20'45"N, 111°47'37"E), Oct 2023, in the phylloplane of Diospyros lotus, S. Liu & Y.Z. Qiao, NYNU 2311236 (GenBank: PQ496716, PQ496715).

Note.

In the phylogenetic analyses, V. tianchiensis was grouped with V. dimennae (Fell & Phaff) X.Z. Liu, F.Y. Bai, M. Groenew. & Boekhout, V. globispora (B.N. Johri & Bandoni) X.Z. Liu, F.Y. Bai, M. Groenew. & Boekhout, and V. pseudodimennae in a clade with high support values (BS /100, BPP /1.0; Fig. 1). V. tianchiensis differs from V. dimennae by 21 nucleotides (14/479 in ITS and 7/596 in LSU), from V. globispora by 40 nucleotides (28/478 in ITS and 12/596 in LSU), and from V. pseudodimennae by 37 nucleotides (22/437 in ITS and 15/566 in LSU). Physiologically, V. tianchiensis can be differentiated from three closest known species, by the ability to assimilate inulin, melibiose, methyl-α-D-glucoside, and erythritol and the inability to grow at 30 °C.

Discussion

The Vishniacozyma species share several phenotypic similarities, making it difficult to classify them accurately using phenotypic data alone. In this study, five new species were identified in Vishniacozyma according to the polyphasic approach recommended by Li et al. (2020) and Zhu et al. (2023). Based on both phenotypic and phylogenetic analyses, our study identified five novel species in which only asexual morphs were found. V. diospyri, V. guiyangensis, V. pingtangensis, V. eriobotryae, and V. tianchiensis are morphologically similar to their sister taxa, with significant differences regarding molecular data and physiochemical features.

Of all reported Vishniacozyma species, over 50% are linked to plant materials, of which approximately half are from plant leaves. The five new species described in this paper were also isolated from the surface of plant leaves, further enriching the diversity of the phylloplane fungi. However, considering their low abundance and rare occurrence, they could be classified as transient species. More importantly, most of these transient species colonized in plants, giving us an apparent indication of their role in plant growth in arid areas (Wei et al. 2022). For example, V. pseudodimennae and V. victoriae are transient species that have been frequently found in plants inhabiting dry environments (Buzzini et al. 2018; Wei et al. 2022), implying that these species may help plants survive in these areas. Similarly, five new Vishniacozyma species here described, V. diospyri, V. guiyangensis, V. pingtangensis, V. eriobotryae, and V. tianchiensis, were also isolated from plants. It is possible that these new species provide similar ecological roles as do V. pseudodimennae and V. victoriae.

To date, studies on Vishniacozyma have primarily focused on their taxonomy and ecosystem function (Chang et al. 2019; Nimsi et al. 2023). Consequently, interest in these fungi is not limited to their biodiversity and ecological role but encompasses potential agricultural, industrial, and medical uses with associated economic value (Bilański and Kowalski 2022; Nimsi et al. 2023).

Acknowledgments

The authors are very grateful to their colleagues at the School of Life Science and Agricultural Engineering, Nanyang Normal University, including Dr. Jing-Zhao Li for providing specimens; Ting Lei for help with phylogenetic analysis; Wen-Ting Hu and Ya-Zhuo Qiao for help with morphological observations.

References

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was funded by the National Natural Science Foundation of China (Project No. 31570021).

Author contributions

Data curation: S.L.; Methodology: S.L. and D.Y.C.; Molecular phylogeny: C.Y.C.; Writing—original draft: S.L.; Writing—review and editing: FLH. All authors read and approved the final manuscript.

Author ORCIDs

Shan Liu https://orcid.org/0009-0003-2845-1495

Dan-Yang Cai https://orcid.org/0009-0002-6866-7322

Chun-Yue Chai https://orcid.org/0000-0003-0284-5560

Feng-Li Hui https://orcid.org/0000-0001-7928-3055

Data availability

All of the data that support the findings of this study are available in the main text.

Acknowledgments

The authors are very grateful to their colleagues at the School of Life Science and Agricultural Engineering, Nanyang Normal University, including Dr. Jing-Zhao Li for providing specimens; Ting Lei for help with phylogenetic analysis; Wen-Ting Hu and Ya-Zhuo Qiao for help with morphological observations.

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