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Article

Seven New Species of Eurotiales (Ascomycota) Isolated from Tidal Flat Sediments in China

1
State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
2
College of Life Sciences, Yangtze University, Jingzhou 434025, China
*
Author to whom correspondence should be addressed.
J. Fungi 2023, 9(10), 960; https://doi.org/10.3390/jof9100960
Submission received: 4 August 2023 / Revised: 19 September 2023 / Accepted: 23 September 2023 / Published: 25 September 2023

Abstract

:
Tidal flats have been reported to contain many microorganisms and play a critical role in maintaining biodiversity. In surveys of filamentous fungi from tidal flat sediments in China, seven new species of Eurotiales were discovered and described. Morphological characteristics and DNA sequence analyses of combined datasets of the BenA, CaM, and RPB2 regions support their placements and recognition as new species. Aspergillus liaoningensis sp. nov. and A. plumeriae sp. nov. belong to sections Candidi and Flavipedes of subgenus Circumdati, and A. subinflatus sp. nov. is a member of section Cremei of subgenus Cremei. Penicillium danzhouense sp. nov., P. tenue sp. nov., and P. zhanjiangense sp. nov. are attributed to sections Exilicaulis and Lanata-Divaricata of subgenus Aspergilloides. Talaromyces virens sp. nov. is in section Talaromyces. Detailed descriptions and illustrations of these novel taxa are provided. Their differences from close relatives were compared and discussed.

1. Introduction

Tidal flats, which link the ocean and the land, contain plentiful microorganisms [1]. Filamentous fungi were reported to be dominant in the intertidal fungal ecosystem [2,3]. The order Eurotiales is one of the most abundant groups, which contains five families with about 28 genera, including Aspergillus P. Micheli, Penicillium Link, and Talaromyces C.R. Benj. [4]. These genera are economically important in the fields of human health, agriculture, industry, and pharmaceutics [2,4,5,6,7]. For example, A. fumigatus Fresen. and T. marneffei (Segretain, Capponi & Sureau) Samson, N. Yilmaz, Frisvad & Seifert are two well-known human pathogens [8,9]. Penicillin, an effective anti-infective drug [10], was produced by P. chrysogenum Thom. Aspergillus oryzae (Ahlb.) Cohn can be used in food fermentation and was reported as a producer of enzymes [11,12]. Therefore, the discovery of these fungi is of theoretical and practical importance.
Due to the sophisticated classification, infrageneric taxonomy has commonly been used for Aspergillus, Penicillium, and Talaromyces [4]. With the application of multiple loci phylogeny, the inter-specific relationships within these genera have become more clear [13,14,15]. Currently, the genus Aspergillus comprises 483 species belonging to 27 sections [11,16,17,18,19,20,21,22], Penicillium contains 530 species in 33 sections [20,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38], and Talaromyces includes 204 species in 7 sections [6,18,32,39,40,41,42]. They show a broad range of habitats, such as woody substratum, sandy soil, tidal flats, water, and indoor air [43,44,45,46,47].
During the examinations of filamentous fungi isolated from tidal flat sediments in different provinces in China, seven undescribed taxa were encountered. Judging by the cultural and microscopic characteristics, they belong to Aspergillus, Penicillium, and Talaromyces. Their taxonomic placements were further confirmed by carrying out multilocus phylogenetic analyses of β-tubulin (BenA), calmodulin (CaM), and the second-largest subunit of RNA polymerase II (RPB2). The distinctions between the novel species and their close relatives were compared.

2. Materials and Methods

2.1. Sampling and Fungal Isolation

Tidal flat sediment samples were collected from Guangdong, Hainan, and Liaoning provinces from August to October 2020. Sediment samples were kept at 4 °C until used. Strains were isolated using a 3% sea salt medium with the dilution plate method and were preserved in the China General Microbiological Culture Collection Center (CGMCC). Dried cultures were deposited in the Herbarium Mycologicum Academiae Sinicae (HMAS).

2.2. Morphological Observations

Colony characteristics were observed and described following the method of Visagie et al. [14]. Four standard growth media were used: Czapek yeast autolysate agar (CYA, yeast extract Oxoid), malt extract agar (MEA, Amresco), yeast extract agar (YES), and potato dextrose agar (PDA) [48,49,50]. A twenty-five percent lactic acid (LA) solution was used as the mounting medium for the microscopic examinations of structures and measurements of the conidial head, stipe, phialide, vesicle, and conidia. The methods for inoculation, morphological observation, and digital recordings were performed following previous studies [51,52].

2.3. DNA Extraction, PCR Amplification, and Sequencing

Genomic DNA was extracted from fungal mycelium with the Plant Genomic DNA Kit (Tiangen Biotech (Beijing) Co., Ltd., Beijing, China). The sequences of nuclear the ribosomal DNA ITS1-5.8S-ITS2 (ITS), BenA, CaM, and RPB2 regions were amplified on an ABI 2720 Thermal Cycler (Applied Biosciences, Foster City, CA, USA) with the primer pairs ITS5 and ITS4 [53], T1 and Bt2a (or Bt2b) [54,55], CMD5 and CMD6 [56], and fRPB2-5F and fRPB2-7cR [57], respectively. PCR products were sequenced in both directions on an ABI 3730 DNA Sequencer (Applied Biosciences, Foster City, CA, USA).

2.4. Phylogenetic Analyses

The newly generated sequences and those retrieved from GenBank are listed in Table 1. They were assembled and aligned with BioEdit 7.0.5 [58] and manually edited. To evaluate statistical congruence amongst the loci BenA, CaM, and RPB2, the partition homogeneity test (PHT) was performed in PAUP*4.0b10 [59] with 1000 replicates. To determine the positions of these strains, the datasets of these regions belonging to Aspergillus sect. Candidi, Cremei and Flavipedes, Penicillium sect. Exilicaulis and Lanata-Divaricata, and Talaromyces sect. Talaromyces were compiled and analyzed by the maximum likelihood (ML) and Bayesian inference (BI) methods. ML analysis was performed with the default GTRCAT model using RAxML [60]. The BI analysis was conducted by MrBayes 3.2.5 [61]. Nucleotide substitution models were determined by MrModeltest 2.3 [62]. Dendrogram trees were visualized and edited using TreeView v. 1.6.6 [63] and FigTree v. 1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/ (accessed on 25 November 2018)). A Bayesian inference posterior probability (BIPP) greater than 90% and a maximum likelihood bootstrap support (MLBS) greater than 70% were shown at the nodes.

3. Results

3.1. Phylogenetic Analyses

To determine the positions of the Aspergillus strains, Hamigera avellanea Stolk & Samson and Penicillium expansum Link were used as outgroup taxa. The partition homogeneity tests (p = 0.01 and 0.25, respectively) indicated that the individual partitions were not highly incongruent [64]; thus, these three loci were combined for the phylogenetic analyses. The phylogenetic trees showed that strains CGMCC 3.25201 and 3.25202 were located in sect. Candidi and Flavipedes, respectively (Figure 1). The strain CGMCC 3.25201 shared a close relationship with A. subalbidus Visagie, Hirooka & Samson (BIPP/MLBS = 91%/89%), while CGMCC 3.25202 clustered with A. movilensis A. Nováková, Hubka, S.W. Peterson & M. Kolařík, receiving high supporting values (BIPP/MLBS = 100%/100%). The strain CGMCC 3.25203 grouped with other members of sect. Cremei ser. Inflati, receiving high statistic values (BIPP/MLBS = 100%/100%) (Figure 2).
In the phylogenetic analyses of Penicillium sect. Exilicaulis, H. avellanea and A. glaucus (L.) Link served as outgroup taxa. The partition homogeneity test (p = 0.01) indicated that the individual partitions were not highly incongruent [64]; thus, these three loci were combined for the phylogenetic analyses. The phylogenetic trees showed that strains CGMCC 3.25204 and 3.25205 were located in ser. Erubescentia (BIPP/MLBS = 100%/93%) (Figure 3). The strain CGMCC 3.25204 was clustered with P. canis S.W. Peterson, while strain CGMCC 3.25205 was grouped with P. striatisporum Stolk, both receiving full support.
In the phylogenetic analyses of Penicillium sect. Lanata-Divaricata, H. avellanea and P. glabrum (Wehmer) Westling were used as outgroup taxa. The partition homogeneity test (p = 0.01) indicated that the individual partitions were not highly incongruent [64]; thus, these three loci were combined for the phylogenetic analyses. The strain CGMCC 3.25206 was placed in ser. Janthinella and clustered with P. janthinellum Biourge with high supporting values (BIPP/MLBS = 100%/100%) (Figure 4).
To determine the position of the Talaromyces strain, T. trachyspermus (Shear) Stolk & Samson and T. chongqingensis X.C. Wang & W.Y. Zhuang were used as outgroup taxa. The partition homogeneity test (p = 0.01) indicated that the individual partitions were not highly incongruent [64]; thus, these three loci were combined for the phylogenetic analyses. The phylogenetic tree showed that strain CGMCC 3.25207 was grouped with the others of sect. Talaromyces (BIPP/MLBS = 100%/100%) and was closely related to T. xishaensis X.C. Wang, L. Wang & W.Y. Zhuang (BIPP/MLBS = 100%/98%) (Figure 5).

3.2. Taxonomy

  • Aspergillus liaoningensis C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 6.
Fungal Names: FN 571613.
Etymology: The specific epithet refers to the type locality “Liaoning Province” of the fungus.
In: Aspergillus subgen. Circumdati sect. Candidi ser. Candidi.
Typification: CHINA, Liaoning Province, Donggang City, Zhongshan District, Yalu River Wetland Park, 39°49′0″ N 124°3′20″ E, in fluvial sediments, 13 October 2020, Chang Liu, tt32414 (holotype HMAS 247877, ex-type strain CGMCC 3.25201).
DNA barcodes: ITS ON563148, BenA ON231293, CaM ON470836, RPB2 ON470844.
Colony diam.: 7 days, 25 °C (unless stated otherwise): CYA 20–23 mm; CYA 37 °C, 8–9 mm; CYA 5 °C no growth; MEA 13–14 mm; PDA 16–22 mm; YES 18–20 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies irregular, plain, cracked; margins narrow, nearly entire; mycelia white to cream; texture velutinous to floccose; sporulation dense; conidia en masse cream; no exudate, no soluble pigments; reverse cream to light yellow, white at periphery. On CYA 37 °C, 7 days: Colonies irregular, slightly protuberant in centers; margins narrow to moderately wide; mycelia cream; texture velutinous to floccose; sporulation dense; conidia en masse white to cream; no exudate, no soluble pigments; reverse dark cream, brown at centers. On MEA 25 °C, 7 days: Colonies nearly circular, slightly protuberant in centers; margins narrow to moderately wide, nearly entire; mycelia white to light cream; texture velutinous to floccose; sporulation dense in center, light cream; no exudate, no soluble pigments; reverse cream, brown at centers, white at periphery. On PDA 25 °C, 7 days: Colonies irregular, plain, cracked; margins narrow, nearly entire; mycelia white to grey; texture velutinous to floccose; sporulation dense; conidia en masse grey; no exudate, no soluble pigments; reverse cream, brown at centers, white at periphery. On YES 25 °C, 7 days: Colonies nearly circular, protuberant in centers, edges irregular; margins narrow, nearly entire; mycelium light grey at center, white at margin; texture velutinous to floccose; sporulation moderately dense; no exudate, no soluble pigments; reverse white, light yellow at centers.
Micromorphology: Conidial heads radiate; stipes thick walls, smooth, hyaline, not septate, 50–260 × 5.0–7.5 µm; vesicles globose to broad ellipsoidal, 5.5–17.3 × 5.3–17.1 µm; biseriate; metulae cylindrical to obovate, 5.2–8.0 × 3.3–5.1 µm, covering two-thirds to almost the entire surface of the vesicle; phialides flask-shaped to acerose, 4.8–8.4 × 2.3–3.1 µm; conidia globose to subglobose, smooth, 2.6–4.2 µm in diam.
Note: This species is phylogenetically related to A. subalbidus (Figure 1), but the latter differs in that it lacks growth on CYA at 37 °C, has faster growth rates on MEA (17–19 mm) and YES (25–30 mm) at 25 °C, and its colonies do not crack on CYA and PDA [14].
  • Aspergillus plumeriae C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 7.
Fungal Names: FN 571614.
Etymology: The specific epithet refers to the yellowish-white colony on PDA.
In: Aspergillus subgen. Circumdati sect. Flavipedes ser. Spelaei.
Typification: CHINA, Liaoning Province, Dalian City, Zhongshan District, Binhai East Road, 38°52′1″ N 121°41′20″ E, in tidal flat sediments, 12 October 2020, Chang Liu, tt30226 (holotype HMAS 247878, ex-type strain CGMCC 3.25202).
DNA barcodes: ITS ON563147, BenA ON231292, CaM ON470835, RPB2 ON470843.
Colony diameter: 7 days, 25 °C (unless stated otherwise): CYA 24–26 mm; CYA 37 °C no growth; CYA 5 °C no growth; MEA 18–22 mm; PDA 17–19 mm; YES 25–26 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies nearly circular, plain, slightly protuberant in centers; margins narrow to moderately wide, nearly entire; mycelia white to cream; texture velutinous; sporulation dense, conidia en masse white; no exudate, no soluble pigments; reverse light yellow. On MEA 25 °C, 7 days: Colonies nearly circular, wrinkled, slightly protuberant in centers, radially sulcate; margins moderately wide, nearly entire; mycelia white to cream; texture velutinous; sporulation dense in center, cream to light yellow; no exudate, no soluble pigments; reverse yellowish-brown. On PDA 25 °C, 7 days: Colonies nearly circular, wrinkled, slightly concave at centers, radially sulcate; margins moderately wide, nearly entire; mycelia white, bright yellow at center; texture velutinous to floccose; sporulation dense in center, bright yellow; no exudate, no soluble pigments; reverse light–brown at centers, white at periphery. On YES 25 °C, 7 days: Colonies nearly circular, wrinkled, slightly concave at centers, radially sulcate; margins moderately wide, nearly entire; mycelium white to cream at center, white at margin; texture velutinous; sporulation moderately dense, white to cream; no exudate, no soluble pigments; reverse white to light yellow.
Micromorphology: Conidial heads radiate; stipes thick walls, smooth, hyaline or blackish, not septate, longer than 340 µm; vesicles globose to subglobose, 13–22.7 µm in diam.; biseriate; metulae cylindrical, 4.5–7.8 × 2.6–4.9 µm, covering two-thirds to almost the entire surface of the vesicle; phialides flask-shaped to acerose, slightly curved at the mouth, 5.4–8.4 × 1.7–2.9 µm; conidia globose to subglobose, smooth, 2.6–3.3 µm in diam.
Note: Among the known species of Aspergillus, A. plumeriae is distinct because of its yellowish-white colony on PDA. It is phylogenetically related to A. movilensis (Figure 1), but the latter differs in its ability to grow at 37 °C and production of pyriform vesicles with smaller sizes (5.0–16 µm in diam.) [65].
  • Aspergillus subinflatus C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 8.
Fungal Names: FN 571615.
Etymology: The specific epithet refers to the similarity of the fungus to A. inflatus.
In: Aspergillus subgen. Cremei sect. Cremei. Ser. Inflati.
Typification: CHINA, Hainan Province, Ledong Li Autonomous County, Liguo Town, 108°56′22″ N 18°24′38″ E, in mangrove sediments, 3 September 2020, Hai-Yan Zhu, tt14122 (holotype HMAS 247879, ex-type strain CGMCC 3.25203).
DNA barcodes: ITS ON563146, BenA ON231291, CaM ON470834, RPB2 ON470845.
Colony diameter: 7 days, 25 °C (unless stated otherwise): CYA 16–17 mm; CYA 37 °C 4–5 mm; CYA 5 °C no growth; MEA 13–14 mm; PDA 17–18 mm; YES 14–16 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies nearly circular, wrinkled, protuberant in centers, radially sulcate; margins narrow to moderately wide, nearly entire; mycelia white to grey; texture velutinous; sporulation dense; conidia en masse grey; no exudate, no soluble pigments; reverse light yellow. On CYA 37 °C, 7 days: Colonies nearly circular, lain; mycelia light pink to white; texture velutinous; no sporulation; no exudate, no soluble pigments; reverse light pink to white. On MEA 25 °C, 7 days: Colonies nearly circular, slightly wrinkled, slightly protuberant in centers; margins wide to moderately wide, nearly entire; mycelia white to grey; texture velutinous to floccose; sporulation dense in center, grey; no exudate, no soluble pigments; reverse yellowish-brown, yellow to white at centers. On PDA 25 °C, 7 days: Colonies nearly circular, slightly wrinkled, slightly protuberant in centers; margins moderately wide, nearly entire; mycelia white to light grey; texture velutinous to floccose; sporulation dense in center, light grey; no exudate, no soluble pigments; reverse pale yellow. On YES 25 °C, 7 days: Colonies nearly circular, wrinkled, protuberant in centers; margins narrow, nearly entire; mycelium white, light grey at center; texture velutinous to floccose; sporulation moderately dense, white; no exudate, no soluble pigments; reverse yellow, white at periphery.
Micromorphology: Conidial heads radiate; stipes smooth-walled, slightly swollen at the apex, 12–318 × 3.5–5.0 µm; vesicle ellipsoidal, 5.3–17 × 3.1–6.4 μm; metulae developing successively on the vesicle and also occurring on its subterminal and terminal portion, swollen, cylindrical to obovate, 3.8–9.4 × 2.5–4.1 µm; phialides flask-shaped to acerose, tapering into thin neck, 2.5–7.9 × 1.9–2.6 µm; conidia globose to subglobose, smooth to finely rough, 1.6–2.3 μm in diam.
Note: This species is morphologically similar and phylogenetically related to A. inflatus (Figure 2) but differs in that its metulae occur at the subterminal and terminal positions [13,66]. Moreover, there are 69 bp, 74 bp, and 88 bp divergences in the BenA, CaM, and RPB2 regions between the ex-type cultures of the two taxa (CGMCC 3.25203 and CBS 682.70).
  • Penicillium danzhouense C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 9.
Fungal Names: FN 571616.
Etymology: The specific epithet refers to the type locality “Danzhou City” of the fungus.
In: Penicillium subgen. Aspergilloides sect. Exilicaulis ser. Erubescentia.
Typification: CHINA, Hainan Province, Danzhou City, Eman Town, 19°51′24″ N 109°13′54″ E, in tidal flat sediments, 2 September 2020, Hai-Yan Zhu, tt13610 (holotype HMAS 247880, ex-type strain CGMCC 3.25204).
DNA barcodes: ITS ON563150, BenA ON231295, CaM ON470838.
Colony diameter: 7 days, 25 °C (unless stated otherwise): CYA 21–24 mm; CYA 37 °C 6–7 mm; CYA 5 °C no growth; MEA 14–17 mm; PDA 17–19 mm; YES 12–15 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies nearly circular, deep, wrinkled, protuberant at centers, radially sulcate; margins narrow, entire; mycelia white, light pink at center; texture velutinous to floccose; no sporulation; exudate colorless, no soluble pigments; reverse yellow. On CYA 37 °C, 7 days: Colonies nearly circular, plain, strongly wrinkled; margins narrow; mycelia white; texture velutinous to floccose; no sporulation; no exudate, no soluble pigments; reverse light yellow. On MEA 25 °C, 7 days: Colonies nearly circular, concave at centers, protuberant at margins; margins narrow to moderately wide, nearly entire; mycelia white, light grey at center; texture velutinous to floccose; sporulation sparse, light grey; no exudate, no soluble pigments; reverse yellow, white at periphery. On PDA 25 °C, 7 days: Colonies nearly circular, slightly protuberant at centers, edges irregular; margins narrow to moderately wide, nearly entire; mycelia white to cream, grey at center; texture velutinous to floccose; sporulation moderately dense, grey; no exudate, no soluble pigments; reverse light yellow to cream. On YES 25 °C, 7 days: Colonies nearly circular, deep, protuberant at centers; margins narrow, entire; mycelium white; texture velutinous to floccose; sporulation sparse; no exudate, no soluble pigments; reverse yellow, white at periphery.
Micromorphology: Conidiophores monoverticillate, rarely biverticillate; stipes smooth-walled, 12–40 × 2.0–3.0 µm; phialides flask-shaped to acerose, 2–5 per metula, 4.6–8.7 × 1.7–2.2 µm; conidia globose to subglobose, smooth to finely roughened, 2.2–3.0 µm in diam.
Note: This species is phylogenetically related to P. catenatum (Figure 3), but the latter differs in its larger phialides (8.0–12 × 2.5–3.0 µm) [67]. Moreover, the growth rates of P. catenatum were relatively slower than P. danzhouense, and the former inhabited the desert rather than tidal flat sediments.
  • Penicillium tenue C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 10.
Fungal Names: FN 571617.
Etymology: The specific epithet refers to the slender phialides.
In: Penicillium subgen. Aspergilloides sect. Exilicaulis ser. Erubescentia.
Typification: CHINA, Hainan Province, Danzhou City, Duntou Town, 19°09′06″ N 108°40′19″ E, in tidal flat sediments, 2 September 2020, Hai-Yan Zhu, tt13918 (holotype HMAS 247881, ex-type strain CGMCC 3.25205).
DNA barcodes: ITS ON563151, BenA ON231296, CaM ON470839, RPB2 ON470842.
Colony diameter: 7 days, 25 °C (unless stated otherwise): CYA 11–12 mm; CYA 37 °C 12–13 mm; CYA 5 °C no growth; MEA 11–14 mm; PDA 12–13 mm; YES 11–12 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies nearly circular, deep, protuberant at centers, radially sulcate; margins narrow, entire; mycelia white, light yellow at center; texture velutinous to floccose; no sporulation; exudate yellow, no soluble pigments; reverse yellow, deepens in the center. On CYA 37 °C, 7 days: Colonies nearly circular, concave at centers, protuberant at margins, slightly wrinkled, radially sulcate; margins narrow; mycelia white, light yellow at centers; texture velutinous to floccose; sporulation sparse; exudate yellow to yellowish-brown, no soluble pigments; reverse yellow, yellowish-brown in centers. On MEA 25 °C, 7 days: Colonies nearly circular, protuberant at centers; margins narrow to moderately wide, nearly entire; mycelia white, light yellow at center; texture velutinous to floccose; sporulation sparse; no exudate, no soluble pigments; reverse yellowish-brown, white at periphery. On PDA 25 °C, 7 days: Colonies nearly circular, protuberant at centers; margins narrow, nearly entire; mycelia white to yellow; texture velutinous to floccose; sporulation sparse; exudate yellow to yellowish-brown, no soluble pigments; reverse light yellow. On YES 25 °C, 7 days: Colonies nearly circular, deep, concave at centers, protuberant at margins; margins narrow, entire; mycelium white; texture velutinous to floccose; no sporulation; no exudate, no soluble pigments; reverse yellowish-brown, white at center and periphery.
Micromorphology: Conidiophores monoverticillate; stipes smooth-walled, 5.0–24 × 2.0–3.0 µm; phialides flask-shaped to acerose, 2–4 metula, 3.1–6.1 × 1.6–2.1 µm; conidia globose to subglobose, spinulose, 2.1–2.7 µm in diam.
Note: This species is phylogenetically related to P. striatisporum (Figure 3), but the latter differs in its ellipsoidal to ovoid and striate conidia [68]. Sequence comparisons between the ex-type cultures of the two species revealed that 8 bp, 12 bp, and 3 bp divergences were detected for the BenA, CaM, and RPB2 regions.
  • Penicillium zhanjiangense C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 11.
Fungal Names: FN 571618.
Etymology: The specific epithet refers to the type locality “Zhanjiang City“ of the fungus.
In: Penicillium subgen. Aspergilloides section sect. Lanata-Divaricata ser. Janthinella.
Typification: CHINA, Guangdong Province, Zhanjiang City, Xuwen County, Southeast Village of China Mainland, 20°59′09″ N 109°40′53″ E, in tidal flat sediments, 30 August 2020, Hai-Yan Zhu, tt12003 (holotype HMAS 247882, ex-type strain CGMCC 3.25206).
DNA barcodes: ITS ON563149, BenA ON231294, CaM ON470837.
Colony diameter: 7 days, 25 °C (unless stated otherwise): CYA 28–30 mm; CYA 37 °C 42–43 mm; CYA 5 °C no growth; MEA 41–48 mm; PDA 36–37 mm; YES 22–24 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies nearly circular, concave at centers, protuberant at margins, wrinkled, radially sulcate; margins narrow to moderately wide, nearly entire; mycelia white, light grey at center; texture velutinous to floccose; sporulation sparse; conidia en masse cream to light grey; no exudate, no soluble pigments; reverse yellowish-brown, light green to blackish-green at centers, white at periphery. On CYA 37 °C, 7 days: Colonies nearly circular, concave at centers, protuberant at margins, wrinkled, radially sulcate; margins narrow; mycelia pink, white to yellowish-grey at centers; texture velutinous; texture velutinous to floccose; sporulation sparse; exudate pink to pinkish-brown, no soluble pigments; reverse pink to dark pink, yellow at periphery. On MEA 25 °C, 7 days: Colonies nearly circular, concave at centers, protuberant at margins, wrinkled, radially sulcate; margins narrow to moderately wide, nearly entire; mycelia cream, brown at center; texture velutinous to floccose; sporulation dense; conidia en masse brownish-yellow; no exudate, no soluble pigments; reverse yellowish-brown, deepens in the center. On PDA 25 °C, 7 days: Colonies nearly circular, concave at centers, protuberant at margins, slightly wrinkled, radially sulcate; margins narrow to moderately wide, nearly entire; mycelia light yellow, brown at center; texture velutinous to floccose; sporulation dense; conidia en masse brownish-yellow; no exudate, no soluble pigments; reverse light green to dark green, deepens in the center. On YES 25 °C, 7 days: Colonies nearly circular, deep, concave at centers, protuberant at margins, slightly wrinkled, radially sulcate; margins narrow, nearly entire; mycelium white to cream; texture velutinous to floccose; sporulation sparse; no exudate, no soluble pigments; reverse yellow, white at periphery.
Micromorphology: Conidiophores monoverticillate to biverticillate; stipes smooth-walled, 24–170 × 2.4–3.7 µm; metulae cylindrical, 8.1–28.3 × 2.2–3.1 µm; phialides flask-shaped to acerose, tapering into thin neck, 2–3 per metula, 4.9–15.5 × 1.9–2.9 µm; conidia globose to subglobose, smooth to finely roughened, 2.0–3.3 µm in diam.
Note: This species is phylogenetically related to P. janthinellum (Figure 4), but the latter differs in its faster growth rate on YES (44–46 mm) at 25 °C, while a slower growth rate was observed on CYA at 37 °C (20–30 mm) [69].
  • Talaromyces virens C. Liu, Z.Q. Zeng & W.Y. Zhuang, sp. nov. Figure 12.
Fungal Names: FN 571619.
Etymology: The specific epithet refers to the green conidia.
In: Talaromyces section Talaromyces.
Typification: CHINA, Hainan Province, Wenchang City, Dongjiao Town, 110°50′40″ N 19°32′27″ E, in tidal flat sediments, 1 September 2020, Hai-Yan Zhu, tt13401 (holotype HMAS 247883, ex-type strain CGMCC 3.25207).
DNA barcodes: ITS ON563152, BenA ON231297, CaM ON470840, RPB2 ON470841.
Colony diameter: 7 days, 25 °C (unless stated otherwise): CYA 19–20 mm; CYA 37 °C 6–7 mm; CYA 5 °C no growth; MEA 23–26 mm; PDA 22–23 mm; YES 18–19 mm.
Colony characteristics: On CYA 25 °C, 7 days: Colonies nearly circular, protuberant in centers; margins narrow to moderately wide, nearly entire; mycelia white; texture velutinous; sporulation dense; conidia en masse dark olive green; no exudate, no soluble pigments; reverse light khaki, light brown at centers, light yellow and white at periphery. On CYA 37 °C, 7 days: Colonies nearly circular, deep, wrinkled, deeply concave in centers; margins narrow; mycelia white; texture velutinous; sporulation dense; conidia en masse grey; no exudate, no soluble pigments; reverse grey or white. On MEA 25 °C, 7 days: Colonies nearly circular, slightly protuberant in centers; margins wide, nearly entire; mycelia white; texture velutinous; sporulation dense in center, olive-drab; no exudate, no soluble pigments; reverse light yellow, light orange at centers. On PDA 25 °C, 7 days: Colonies nearly circular, protuberant in centers; margins wide, nearly entire; mycelia white; texture velutinous to floccose; sporulation dense in center, deep green; no exudate, no soluble pigments; reverse white, light coral to red at centers, white at periphery. On YES 25 °C, 7 days: Colonies nearly circular, protuberant in centers; margins narrow to moderately wide, nearly entire; mycelium gray-green to yellowish-green at center, white at margin; texture velutinous to floccose; sporulation moderately dense, yellow to green; no exudate, no soluble pigments; reverse white, dark yellow at centers.
Micromorphology: Conidiophores biverticillate; stipes smooth-walled, 147–220 × 2.4–4.0 µm; metulae cylindrical, 9.1–14.1 × 2.5–3.4 µm; phialides flask-shaped to acerose, tapering into thin neck, 2–6 per metula, 7.7–11.2 × 2.2–3.3 µm; conidia globose to subglobose, green, 2.4–4.0 µm in diam.
Note: This species is morphologically and phylogenetically related to T. xishaensis (Figure 5). However, the latter has greyish-green colonies on CYA, yellowish-green colonies on MEA, and grey to bluish-green colonies on YES [70].

4. Discussion

The genus Aspergillus is divided into six subgenera with 27 sections [13,22]. Our new species A. liaoningensis was well-located in the ser. Candidi of sect. Candidi (BIPP/MLBS = 100%/100%) (Figure 1). Many species within this section have been reported as producers of secondary metabolites, such as taichunamides, shikimic acid derivatives, and terpene-derived taichunins [71,72,73]. Studies on the metabolic application of A. liaoningensis are surely our future goal. Aspergillus plumeriae belongs to ser. Spelaei of sect. Flavipedes, which is in accordance with the growth rate at 37 °C [65]. Aspergillus subinflatus was classified as a member of ser. Inflati, sect. Cremei of subgen. Cremei, and is most related to A. inflatus in both phylogeny and morphology. However, sequence comparisons revealed that there were 69 bp, 74 bp, and 88 bp unmatched loci detected in the BenA, CaM, and RPB2 regions between them.
Both P. danzhouense and P. tenue form monoverticillate conidiophores, consistent with the other members of ser. Erubescentia in sect. Exilicaulis [7,48,74,75]. The phylogenetic results indicate that P. danzhouense is closely related to P. catenatum with high statistical support values (Figure 3); however, the latter differs in having larger phialides (8.0–12 × 2.5–3.0 µm vs. 4.6–8.7 × 1.7–2.2 µm) [67]. Moreover, there were 20 bp and 56 bp divergences in the BenA and CaM regions between them. Our results also showed that P. tenue was grouped with P. striatisporum, receiving full support (Figure 3), but the latter possesses ellipsoidal to ovoid and striate conidia [68]. Penicillium dravuni, a marine-derived species belonging to this section, was not included in our phylogenetic analysis because no sequence data are available at present. However, it can be easily distinguished from P. danzhouense (white to light pink colonies) and P. tenue (white to light yellow colonies) because it forms yellow-gray colonies and has faster growth rates (25–35 mm vs. 21–24 mm, and 11–12 mm in CYA at 25 °C) [76].
Sect. Lanata-Divaricata is a species-rich section in Penicillium, and about 85 species have been described hitherto [4,20,27,28,31,33]. Our phylogenetic tree indicated that P. zhanjiangense was well-located among other species of sect. Lanata-Divaricata, with high supporting values (BIPP/MLBS = 97%/100%), and it clustered with P. janthinellum (BIPP/MLBS = 100%/100%) (Figure 4). A new species of the section was added in this study.
Since the establishment of Talaromyces, eight sections have been proposed: Bacillispori, Helici, Islandici, Purpurei, Subinflati, Talaromyces, Tenues, and Trachyspermi [40]. The phylogenetic overview of the sect. Talaromyces was conducted by Wang and Zhuang [40], and about 88 species are currently known in this section [40,41,42]. The three-locus phylogeny formed a well-supported monophyletic group (BIPP/MLBS = 100%/100%) and indicates that T. virens is related to but distinct from T. xishaensis (Figure 5). They can be easily distinguished by their different colony features on different media [70]. Moreover, there are 11 bp, 34 bp, and 32 bp unmatched loci detected in the BenA, CaM, and RPB2 regions between the ex-type cultures (CGMCC 3.25207 and CGMCC 3.17995).
Species of the genera Penicillium, Aspergillus, and Talaromyces have been isolated from various substrates, including dust, soil, dung, cloth, human tissue, plants, and insects [4,11]. Due to the special ecological habitat, the fungi of these groups within tidal flats have high biodiversity and are an important source of active natural products [77,78,79,80]. Recently, three novel taxa of Penicillium were reported in tidal flats [28]. Similarly, the present study introduces three species of Aspergillus as well as three taxa of Penicillium and one of Talaromyces derived from tidal flat sediments. With the extensive use of molecular approaches, large-scale surveys in these unexplored tidal flats regions will significantly improve our knowledge of fungal species diversity in special ecological environments.

5. Conclusions

The filamentous fungi from tidal flat sediments in China were surveyed, and seven novel taxa of the genera Aspergillus, Penicillium, and Talaromyces were discovered. With the joining of the new species, the phylogenetic relationships among species of these three genera were updated.

Author Contributions

Conceptualization, Z.-Q.Z.; methodology, formal analysis, and investigation, C.L. and Z.-Q.Z.; resources and data curation, W.-Y.Z. and Z.-Q.Z.; original draft preparation and writing, C.L., X.-C.W., Z.-H.Y., Z.-Q.Z. and W.-Y.Z.; supervision and project administration, Z.-Q.Z. All authors have read and agreed to the published version of the manuscript.

Funding

This project was supported by the National Natural Science Foundation of China (32270009, 31870012, 31750001); the Biological Resources Programme, Chinese Academy of Sciences (KFJ-BRP-017-082); the National Project on Scientific Groundwork, Ministry of Science and Technology of the People’s Republic of China (2019FY100700); and the Frontier Key Program of Chinese Academy of Sciences (QYZDY-SSW-SMC029).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The names of the new species were formally registered in the database Fungal Names (https://nmdc.cn/fungalnames (accessed on 10 July 2023)). Specimens were deposited in the Herbarium Mycologicum Academiae Sinicae (https://nmdc.cn/fungarium/ (accessed on 18 February 2023)). Cultures were deposited in the China General Microbiological Culture Collection Center (https://cgmcc.net/ (accessed on 18 July 2023)). The newly generated sequences were deposited in GenBank (https://www.ncbi.nlm.nih.gov/genbank (accessed on 20 May 2022)).

Acknowledgments

The authors would like to thank Feng-Yan Bai and Hai-Yan Zhu for collecting samples jointly for this study.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Aspergillus sect. Candidi and Flavipedes. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
Figure 1. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Aspergillus sect. Candidi and Flavipedes. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
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Figure 2. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Aspergillus sect. Cremei species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
Figure 2. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Aspergillus sect. Cremei species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
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Figure 3. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Penicillium sect. Exilicaulis species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
Figure 3. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Penicillium sect. Exilicaulis species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
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Figure 4. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Penicillium sect. Lanata-Divaricata species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
Figure 4. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Penicillium sect. Lanata-Divaricata species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
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Figure 5. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Talaromyces sect. Talaromyces species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
Figure 5. BI tree generated from analyses of combined BenA, CaM, and RPB2 sequences of Talaromyces sect. Talaromyces species. BIPP ≥ 90% (left) and MLBS ≥ 70% (right) are indicated at nodes.
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Figure 6. Colonial and microscopic morphology of Aspergillus liaoningensis (CGMCC 3.25201). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bf) Conidiophores; (g) Conidia. Scale bars: (bg) = 10 μm.
Figure 6. Colonial and microscopic morphology of Aspergillus liaoningensis (CGMCC 3.25201). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bf) Conidiophores; (g) Conidia. Scale bars: (bg) = 10 μm.
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Figure 7. Colonial and microscopic morphology of Aspergillus plumeriae (CGMCC 3.25202). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
Figure 7. Colonial and microscopic morphology of Aspergillus plumeriae (CGMCC 3.25202). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
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Figure 8. Colonial and microscopic morphology of Aspergillus subinflatus (CGMCC 3.25203). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
Figure 8. Colonial and microscopic morphology of Aspergillus subinflatus (CGMCC 3.25203). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
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Figure 9. Colonial and microscopic morphology of Penicillium danzhouense (CGMCC 3.25204). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bg) Conidiophores; (h,i) Conidia. Scale bars: (bi) = 10 μm.
Figure 9. Colonial and microscopic morphology of Penicillium danzhouense (CGMCC 3.25204). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bg) Conidiophores; (h,i) Conidia. Scale bars: (bi) = 10 μm.
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Figure 10. Colonial and microscopic morphology of Penicillium tenue (CGMCC 3.25205). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bd) Conidiophores; (e) Conidia. Scale bars: (be) = 10 μm.
Figure 10. Colonial and microscopic morphology of Penicillium tenue (CGMCC 3.25205). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bd) Conidiophores; (e) Conidia. Scale bars: (be) = 10 μm.
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Figure 11. Colonial and microscopic morphology of Penicillium zhanjiangense (CGMCC 3.25206). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
Figure 11. Colonial and microscopic morphology of Penicillium zhanjiangense (CGMCC 3.25206). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
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Figure 12. Colonial and microscopic morphology of Talaromyces virens (CGMCC 3.25207). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
Figure 12. Colonial and microscopic morphology of Talaromyces virens (CGMCC 3.25207). (a) Colonies after 7 days at 25 °C; top row left to right: obverse CYA, MEA, PDA, and YES; bottom row left to right: reverse CYA, MEA, PDA, and YES; (bh) Conidiophores; (i) Conidia. Scale bars: (bi) = 10 μm.
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Table 1. Names, strain numbers, and corresponding GenBank accession numbers of the taxa used in this study.
Table 1. Names, strain numbers, and corresponding GenBank accession numbers of the taxa used in this study.
SpeciesStrain NumbersITSBenACaMRPB2
sect. Candidi
A. campestrisCBS 348.81 TEF669577EU014091EF669535EF669619
A. candidusCBS 566.65 TEF669592EU014089EF669550EF669634
A. dobrogensisCBS 143370 TLT626959LT627027LT558722LT627028
A. liaoningensisCGMCC 3.25201 TON563148ON231293ON470836ON470844
A. magnusUAMH 1324 TON156376ON164570ON164619ON164517
A. neotriticiCCF 3853 TFR727136FR775327HE661598LT627021
A. pragensisCBS 135591 TFR727138HE661604FR751452LN849445
A. subalbidusCBS 567.65 TEF669593KP987050EF669551EF669635
A. taichungensisCCF 5597 TLT626957EU076297HG916679LT627016
A. tenebricusCBS 147048 TON156389ON164584ON164623ON164532
sect. Flavipedes
A. ardalensisCBS 134372 TFR733808HG916683HG916725HG916704
A. capensisCBS 138188 TKJ775550KJ775072KJ775279KP987020
A. flavipesATCC 24487 TEF669591EU014085EF669549EF669633
A. iizukaeCBS 541.69 TEF669597EU014086EF669555EF669639
A. luppiiCBS 653.74 TEF669617EU014079EF669575EF669659
A. micronesiensisCBS 138183 TKJ775548KJ775085KP987067KP987023
A. movilensisCBS 134395 TKP987089HG916697HG916740HG916718
A. neoflavipesCBS 260.73 TEF669614EU014084EF669572EF669656
A. neoniveusCBS 261.73 TEF669612EU014098EF669570KP987024
A. olivimuriaeNRRL 66783MH298877MH492010MH492011MH492012
A. plumeriaeCGMCC 3.25202 TON563147ON231292ON470835ON470843
A. polyporicolaNRRL 32683 TEF669595EU014088EF669553EF669637
A. spelaeusCBS 134371 THG915905HG916698HG916741HG916719
A. suttoniaeUTHSCSA DI14-215LT899487LT899536LT899589LT899644
A. templicolaCBS 138181 TKJ775545KJ775092KJ775394KP987017
A. urmiensisCBS 139558 TKP987073KP987041KP987056KP987030
sect. Cremei
A. arxiiCBS 525.83 TMN431361MN969365 MN969223JN121529
A. brunneouniseriatusNRRL 4273 TEF652141EF652123EF652138EF652089
A. chaetosartoryaeNRRL 5501 TEF652144EF652117EF652129EF652099
A. chrysellusNRRL 5084 TEF652155EF652109EF652136EF652090
A. citocrescensCBS 140566 TFR727121FR775317LN878969MN969163
A. cremeusNRRL 5081 TEF652149EF652120EF652125EF652101
A. dimorphicusNRRL 3650 TEF652154EF652111EF652135EF652096
A. europaeusCBS 134393TLN908996LN909006LN909007LT548274
A. flaschentraegeriNRRL 5042 TEF652150EF652113EF652130EF652102
A. gorakhpurensisNRRL 3649 TEF652145EF652114EF652126EF652097
A. inflatusCBS 682.70 TFJ531054FJ531008FJ531090JN406529
A. itaconicusNRRL 161 TEF652147EF652118EF652140EF652103
A. koreanusNIBR EML-GSNP1-1KX216525KX216530KX216528KX216531
A. lebretiiURM 8451 TON862928OP672381OP290539OP290510
A. pulvinusNRRL 5078 TEF652159EF652121EF652139EF652104
A. stromatoidesCBS 500.65 TEF652146FJ531038EF652127EF652098
A. subinflatusCGMCC 3.25203 TON563146ON231291ON470834ON470845
A. tardusCBS 433.93 TFJ531045FJ531001FJ531084n.a.
A. wentiiNRRL 375 TEF652151EF652106EF652131EF652092
Sect. Lanata-divaricata
P. abidjanumCBS 246.67 TGU981582GU981650KF296383JN121469
P. alagoenseURM 8086 TMK804503MK802333MK802336MK802338
P. amphipolariaDAOMC 250551 TKT887872KT887833KT887794n.a.
P. annulatumCBS 135126 TJX091426JX091514JX141545KF296410
P. araracuaraenseCBS 113149 TGU981597GU981642KF296373KF296414
P. ausonanumFMR 16948 TLR655808LR655809LR655810LR655811
P. austrosinenseCGMCC 3.18797 TKY495007KY495116KY494947KY495061
P. bissettiiDAOMC 167011 TKT887845KT887806KT887767MN969107
P. brasilianumCBS 253.55 TGU981577GU981629MN969239KF296420
P. brefeldianumCBS 235.81 TAF033435GU981623EU021683KF296421
P. camponotumKAS 2177 TKT887855KT887816KT887777MN969179
P. caperatumCBS 443.75 TKC411761GU981660KF296392KF296422
P. cataractumDAOMC 250534 TKT887847KT887808KT887769n.a.
P. cluniaeCBS 326.89 TKF296406KF296471KF296402KF296424
P. coeruleumCBS 141.45 TGU981606GU981655KF296393KF296425
P. cremeogriseumCBS 223.66 TGU981586GU981624KF296403KF296426
P. curticauleCBS 135127 TFJ231021JX091526JX141536KF296417
P. daleaeCBS 211.28 TGU981583GU981649KF296385KF296427
P. diatomitisCCF 3904 TFJ430748HE651133LT970912LT797560
P. donggangicumAS3.15900 TMW946996MZ004914MZ004918MW979253
P. echinulonalgiovenseCBS 328.59 TGU981587GU981631KX961269KX961301
P. ehrlichiiCBS 324.48 TAF033432GU981652KF296395KF296428
P. elleniaeCBS 118135 TGU981612GU981663KF296389KF296429
P. excelsumITAL7572 TKR815341KP691061KR815342MN969166
P. flaviroseumCGMCC 3.18805 TKY495032KY495141KY494972KY495083
P. fructuariae-cellaeCBS 145110 TMK039434KU554679MK045337n.a.
P. glaucoroseumCBS 138908 TMN431390MN969383MN969257MN969119
P. globosumCGMCC 3.18800 TKY495014 KY495123KY494954KY495067
P. griseoflavumCGMCC 3.18799 TKY495011 KY495120KY494951KY495064
P. griseopurpureumCBS 406.65 TKF296408 KF296467KF296384KF296431
P. guaibinenseCCDCA 11512 TMH674389MH674391MH674393n.a.
P. guangxienseCGMCC 3.18793 TKY494986 KY495095KY494926n.a.
P. hainanenseCGMCC 3.18798 TKY495009KY495118KY494949n.a.
P. hepuenseAS3.16039 TMW946994MZ004912MZ004916MW979254
P. infrabuccalumDAOMC 250537 TKT887856 KT887817KT887778n.a.
P. janthinellumCBS 340.48 TGU981585 GU981625KF296401JN121497
P. javanicumCBS 341.48 TGU981613 GU981657KF296387JN121498
P. jianfenglingenseCGMCC 3.18802 TKY495016 KY495125KY494956KY495069
P. jiaozhouwanicumAS3.16038 TMW946993MZ004911MZ004915MW979252
P. koreenseKACC 47721 TKJ801939 KM000846MN969317MN969159
P. laevigatumCGMCC 3.18801 TKY495015 KY495124KY494955KY495068
P. levitumCBS 345.48 TGU981607 GU981654KF296394KF296432
P. limosumCBS 339.97 TGU981568 GU981621KF296398KF296433
P. lineolatumCBS 188.77 TGU981579 GU981620KF296397KF296434
P. ludwigiiCBS 417.68 TKF296409 KF296468KF296404KF296435
P. malacosphaerulumCBS 135120 TFJ231026 JX091524JX141542KF296438
P. mariae-crucisCBS 271.83 TGU981593 GU981630KF296374KF296439
P. meloformeCBS 445.74 TKC411762 GU981656KF296396KF296440
P. newtonturneraeBRIP74909a TOP903478OP921964OP921962OP921963
P. nordestinenseURM 8423 TOV265270OV265324OV265272OM927721
P. ochrochloronCBS 357.48 TGU981604 GU981672KF296378KF296445
P. onobenseCBS 174.81 TGU981575 GU981627KF296371KF296447
P. ortumCBS 135669 TJX091427 JX091520JX141551KF296443
P. oxalicumCBS 219.30 TAF033438 KF296462KF296367JN121456
P. panissanguineumDAOMC 250562 TKT887862 KT887823KT887784n.a.
P. paraherqueiCBS 338.59 TAF178511 KF296465KF296372KF296449
P. pedernalenseCBS 140770 TKU255398KU255396MN969322MN969184
P. penarojenseCBS 113178 TGU981570 GU981646KF296381KF296450
P. piscariumCBS 362.48 TGU981600 GU981668KF296379KF296451
P. pulvillorumCBS 280.39 TAF178517 GU981670KF296377KF296452
P. raperiCBS 281.58 TAF033433 GU981622KF296399KF296453
P. reticulisporumCBS 122.68 TAF033437 GU981665KF296391KF296454
P. rolfsiiCBS 368.48 TJN617705 GU981667KF296375KF296455
P. rubriannulatumCGMCC 3.18804 TKY495029 KY495138KY494969KY495080
P. setosumCBS 144865 TKT852579MF184995MH105905MH016196
P. simplicissimumCBS 372.48 TGU981588 GU981632KF296368JN121507
P. singorenseCBS 138214 TKJ775674 KJ775167KJ775403n.a.
P. skrjabiniiCBS 439.75 TGU981576 GU981626KF296370EU427252
P. soliformeCGMCC 3.18806 TKY495038 KY495147KY494978n.a.
P. soosanumCCF 3778 TFJ430745FM865811LT970913LT797561
P. spinuliferumCGMCC 3.18807 TKY495040 KY495149KY494980KY495090
P. subfuscumCMW56196 TMT949907MT957412 MT957454MT957480
P. subrubescensCBS 132785 TKC346350 KC346327KC346330KC346306
P. svalbardenseCBS 122416 TGU981603 KC346325KC346338KF296457
P. tanzanicumCBS 140968 TKT887841KT887802KT887763MN969183
P. terrarumaeCBS 131811 TMN431397KX650295MN969323MN969185
P. ucsense2HH TOM914583ON024157ON024158ON024159
P. uruguayenseCBS 143247 TLT904729LT904699LT904698MN969200
P. vanderhammeniiCBS 126216 TGU981574 GU981647KF296382KF296458
P. vasconiaeCBS 339.79 TGU981599 GU981653KF296386KF296459
P. vickeryaeBRIP72552a TOP903479OP921966n.a.OP921965
P. viridissimumCGMCC 3.18796 TKY495004 KY495113KY494944KY495059
P. wotroiCBS 118171 TGU981591GU981637KF296369KF296460
P. yunnanenseCGMCC 3.18794 TKY494990KY495099KY494930KY495048
P. zhanjiangenseCGMCC 3.25206 TON563149ON231294ON470837n.a.
P. zonatumCBS 992.72 TGU981581GU981651KF296380KF296461
sect. Exilicaulis
P. allaniaeBRIP 74886a TOP903475OP921956OP921954OP921955
P. alutaceumCBS 317.67 TAF033454KJ834430KP016768JN121489
P. amapaenseURM 8639 TOL764382OL782590OL782584ON854925
P. aotearoaePDD 107543 TKT887874KT887835KT887796MN969174
P. arabicumCBS 414.69 TKC411758KP016750KP016770KP064574
P. archeraeBRIP 72549c TOP903477OP921961n.a.OP921960
P. atrolazulinumCBS 139136 TJX140913JX141077JX157416KP064575
P. atrosanguineumCBS 380.75 TJN617706KJ834435KP016771JN406557
P. burgenseCBS 325.89 TKC411736KJ834437KP016772JN406572
P. canisNRRL 62798 TKJ511291KF900167KF900177KF900196
P. catenatumCBS 352.67 TKC411754KJ834438KP016774JN121504
P. chalabudaeCBS 219.66 TKP016811KP016748KP016767KP064572
P. cinerascensNRRL 748 TAF033455JX141041JX157405MN969112
P. cinereoatrumCBS 222.66 TKC411700KJ834442KP125335JN406608
P. citreonigrumCBS 258.29 TAF033456EF198621EF198628JN121474
P. citreosulfuratumIMI 92228 TKP016814KP016753KP016777KP064615
P. consobrinumCBS 139144 TJX140888JX141135JX157453KP064619
P. corylophilumCBS 312.48 TAF033450JX141042KP016780KP064631
P. cravenianumCBS 139138 TJX140900JX141076JX157418KP064636
P. danzhouenseCGMCC 3.25204 TON563150ON231295ON470838n.a.
P. decumbensCBS 230.81 TAY157490KJ834446KP016782JN406601
P. diabolicalicenseCBS 140967 TKT887840KT887801KT887762MN969175
P. dimorphosporumCBS 456.70 TAF081804KJ834448KP016783JN121517
P. dravuniF01V25 TAY494856n.a.n.a.n.a.
P. erubescensCBS 318.67 TAF033464HQ646566EU427281JN121490
P. fagiCBS 689.77 TAF481124KJ834449KP016784JN406540
P. fundyenseCBS 140980 TKT887853KT887814KT887775MN969176
P. guttulosumNRRL 907 THQ646592HQ646576HQ646587MG386247
P. hemitrachumCBS 139134 TFJ231003JX141048JX157526KP064642
P. hermansiiCBS 124296 TMG333472MG386214MG386229MG386242
P. heteromorphumCBS 226.89 TKC411702KJ834455KP016786JN406605
P. katangenseCBS 247.67 TAF033458KP016757KP016788KP064646
P. kurssanoviiCBS 625.67 TEF422849KP016758KP016789KP064647
P. labradorumCBS 145775 TMK881918 MK887898MK887899MK887900
P. laeveCBS 136665 TKF667369KF667365KF667367KF667371
P. lapidosumCBS 343.48 TMN431392KJ834465FJ530984JN121500
P. maclennaniaeCBS 198.81 TKC411689KJ834468KP016791KP064648
P. meliniiCBS 218.30 TAF033449KJ834471KP016792JN406613
P. menonorumNRRL 50410 THQ646591HQ646573HQ646584KF900194
P. meridianumCBS 314.67 TAF033451KJ834472KP016794JN406576
P. momoiiCBS 139157 TJX140895JX141073JX157479KP064673
P. namyslowskiiCBS 353.48 TAF033463JX141067KP016795JF417430
P. nepalenseCBS 203.84 TKC411692KJ834474KP016796JN121453
P. ovatumCBS 136664 TKF667370KF667366KF667368KF667372
P. pagulumCBS 139166 TJX140898JX141070JX157519KP064655
P. parvofructumCBS 141690 TLT559091LT627645LT627646MN969197
P. parvumCBS 359.48 TAF033460HQ646568KF900173JN406559
P. philippinenseCBS 623.72 TKC411770KJ834482KP016799JN406543
P. pimiteouienseCBS 102479 TAF037431HQ646569HQ646580JN406650
P. punicaeJMRC:SF:12421 Tn.a.KX839673KX839671KX839675
P. raciborskiiCBS 224.28 TAF033447JX141069KP016800JN406607
P. repensicolaCBS 139160 TJX140893JX141150JX157490KP064660
P. restrictumCBS 367.48 TAF033457KJ834486KP016803JN121506
P. rubefaciensCBS 145.83 TKC411677 KJ834487KP016804JN406627
P. rubidurumCBS 609.73 TAF033462HQ646574HQ646585JN406545
P. smithiiCBS 276.83 TKC411723KJ834492KP016806JN406589
P. striatisporumCBS 705.68 TAF038938MN969401KP016807JN406538
P. subturcoseumCBS 139132 TFJ231006JX141161JX157532KP064674
P. tenueCGMCC 3.25205 TON563151ON231296ON470839ON470842
P. terrenumCBS 313.67 TAM992111KJ834496KP016808JN406577
P. velutinumCBS 250.32 TAF033448JX141170MT478037KP064682
P. vinaceumCBS 389.48 TAF033461HQ646575HQ646586JN406555
P. xanthomeliniiCBS 139163 TJX140921JX141120JX157495KP064683
sect. Talaromyces
T. aculeatusCBS 289.48 TKF741995KF741929KF741975MH793099
T. adpressusCGMCC 3.18211 TKU866657KU866844KU866741KU867001
T. alveolarisCBS 142379 TLT558969LT559086LT795596LT795597
T. amazonensisCBS 140373 TKX011509KX011490KX011502MN969186
T. amestolkiaeCBS 132696 TJX315660JX315623KF741937JX315698
T. angelicaeKACC 46611 TKF183638KF183640KJ885259KX961275
T. annesophieaeCBS 142939 TMF574592MF590098MF590104MN969199
T. apiculatusCBS 312.59 TJN899375KF741916KF741950KM023287
T. argentinensisNRRL 28750 TMH793045MH792917MH792981MH793108
T. aspriconidiusCBS 141835 TMN864274MN863343MN863320MN863332
T. atkinsoniaeBRIP 72528a TOP059084OP087524n.a.OP087523
T. aurantiacusCBS 314.59 TJN899380KF741917KF741951KX961285
T. aureolinusCGMCC 3.15865 TMK837953MK837937MK837945MK837961
T. australisCBS 137102 TKF741991KF741922KF741971KX961284
T. bannicusCGMCC 3.15862 TMK837955MK837939MK837947MK837963
T. beijingensisCGMCC 3.18200 TKU866649KU866837KU866733KU866993
T. brevisCBS 141833 TMN864269MN863338MN863315MN863328
T. calidicaniusCBS 112002 TJN899319HQ156944KF741934KM023311
T. californicusNRRL 58168 TMH793056MH792928MH792992MH793119
T. cnidiiKACC 46617 TKF183639KF183641KJ885266KM023299
T. coprophilusCBS 142756 TLT899794LT898319LT899776LT899812
T. cucurbitiradicusACCC 39155 TKY053254KY053228KY053246n.a.
T. derxiiCBS 412.89 TJN899327JX494306KF741959KM023282
T. dimorphusCGMCC 3.15692 TKY007095KY007111KY007103KY112593
T. domesticusNRRL 58121 TMH793055MH792927MH792991MH793118
T. duclauxiiCBS 322.48 TJN899342JX091384KF741955JN121491
T. euchlorocarpiusCBM PF1203 TAB176617KJ865733KJ885271KM023303
T. flavovirensCBS 102801 TJN899392JX091376KF741933KX961283
T. flavusCBS 310.38 TJN899360JX494302KF741949JF417426
T. francoaeCBS 113134 TKX011510KX011489KX011501MN969188
T. funiculosusCBS 272.86 TJN899377MN969408KF741945KM023293
T. fuscoviridisCBS 193.69 TKF741979KF741912KF741942MN969156
T. fusiformisCGMCC 3.18210 TKU866656KU866843KU866740KU867000
T. galapagensisCBS 751.74 TJN899358JX091388KF741966KX961280
T. ginkgonis10725 TOL638158OL689844OL689846OL689848
T. haitouensisHR1-7MZ045695MZ054634MZ054637MZ054631
T. indigoticusCBS 100534 TJN899331JX494308KF741931KX961278
T. intermediusCBS 152.65 TJN899332JX091387KJ885290KX961282
T. kabodanensisCBS 139564 TKP851981KP851986KP851995MN969190
T. kendrickiiCBS 136666 TKF741987KF741921KF741967MN969158
T. lentulusCGMCC 3.15689 TKY007088KY007104KY007096KY112586
T. lianiCBS 225.66 TJN899395JX091380KJ885257KX961277
T. louisianensisNRRL 35823 TMH793052MH792924MH792988MH793115
T. macrosporusCBS 317.63 TJN899333JX091382KF741952KM023292
T. maeCGMCC 3.15690 TKY007090KY007106KY007098KY112588
T. malicolaNRRL 3724 TMH909513MH909406MH909459MH909567
T. mangshanicusCGMCC 3.18013 TKX447531KX447530KX447528KX447527
T. marneffeiCBS 388.87 TJN899344JX091389KF741958KM023283
T. muroiiCBS 756.96 TMN431394KJ865727KJ885274KX961276
T. mycothecaeCBS 142494 TMF278326LT855561LT855564LT855567
T. nanjingensisCCTCCM 2012167 TMW130720MW147759MW147760MW147762
T. neofusisporusCGMCC 3.15415 TKP765385KP765381KP765383MN969165
T. oumae-annaeCBS 138208 TKJ775720KJ775213KJ775425KX961281
T. panamensisCBS 128.89 TJN899362HQ156948KF741936KM023284
T. penicillioidesCGMCC 3.15822 TMK837956MK837940MK837948MK837964
T. pinophilusCBS 631.66 TJN899382JX091381KF741964KM023291
T. pratensisNRRL 62170 TMH793075MH792948MH793012MH793139
T. primulinusCBS 321.48 TJN899317JX494305KF741954KM023294
T. pseudofuniculosusCBS 143041 TLT899796LT898323LT899778LT899814
T. purgamentorumCBS 113145 TKX011504KX011487KX011500MN969189
T. purpureogenusCBS 286.36 TJN899372JX315639KF741947JX315709
T. qiiCGMCC 3.15414 TKP765384KP765380KP765382MN969164
T. rapidusCBS 142382 TLT558970LT559087LT795600LT795601
T. rosarhizaGUCC 190040.1 TMZ221603MZ333143MZ333137MZ333141
T. ruberCBS 132704 TJX315662JX315629KF741938JX315700
T. rubicundusCBS 342.59 TJN899384JX494309KF741956KM023296
T. rufusCBS 141834 TMN864272MN863341MN863318MN863331
T. santanderensisHF05 TOP082331OP067657OP067656OP067655
T. sayulitensisCBS 138204 TKJ775713KJ775206KJ775422MN969146
T. shilinensisXCW_SN259 TOL638159OL689845OL689847OL689849
T. siamensisCBS 475.88 TJN899385JX091379KF741960KM023279
T. soliNRRL 62165 TMH793074MH792947MH793011MH793138
T. sparsusCGMCC 3.16003 TMT077182MT083924MT083925MT083926
T. stellenboschiensisCBS 135665 TJX091471JX091605JX140683MN969157
T. stipitatusCBS 375.48 TJN899348KM111288KF741957KM023280
T. stolliiCBS 408.93 TJX315674JX315633JX315646JX315712
T. striatoconidiumCBS 550.89 TMN431418MN969441MN969360MT156347
T. thailandensisCBS 133147 TJX898041JX494294KF741940KM023307
T. tumuliNRRL 62151 TMH793071MH792944MH793008MH793135
T. veerkampiiCBS 500.78 TKF741984KF741918KF741961KX961279
T. verruculosusNRRL 1050 TKF741994KF741928KF741944KM023306
T. versatilisIMI 134755 TMN431395MN969412MN969319MN969161
T. virensCGMCC 3.25207 TON563150ON231297ON470840ON470841
T. viridisCBS 114.72 TAF285782JX494310KF741935JN121430
T. viridulusCBS 252.87 TJN899314JX091385KF741943JF417422
T. wushanicusCS17.05 TMZ356356MZ361347MZ361354MZ361361
T. xishaensisCGMCC 3.17995 TKU644580KU644581KU644582MZ361364
T. yunnanensisKUMCC 18-0208 TMT152339MT161683MT178251n.a.
T. zhenhaiensisZH3-18 TMZ045697MZ054636MZ054639MZ054633
Outgroup
A. glaucusCBS 516.65 TEF652052EF651887EF651989EF651934
Hamigera avellaneaCBS 295.48TAF454075EU021664EU021682EU021627
P. expansumCBS 325.48TAY373912AY674400DQ911134JF417427
P. glabrumCBS 125543 TGU981567GU981619KM089152JF417447
T. chongqingensisCS26-67 TMZ358001MZ361343MZ361350MZ361357
T. trachyspermusCBS 373.48 TJN899354KF114803KJ885281JF417432
Note: Numbers in boldface indicate newly submitted sequences. T means type strain.
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MDPI and ACS Style

Liu, C.; Wang, X.-C.; Yu, Z.-H.; Zhuang, W.-Y.; Zeng, Z.-Q. Seven New Species of Eurotiales (Ascomycota) Isolated from Tidal Flat Sediments in China. J. Fungi 2023, 9, 960. https://doi.org/10.3390/jof9100960

AMA Style

Liu C, Wang X-C, Yu Z-H, Zhuang W-Y, Zeng Z-Q. Seven New Species of Eurotiales (Ascomycota) Isolated from Tidal Flat Sediments in China. Journal of Fungi. 2023; 9(10):960. https://doi.org/10.3390/jof9100960

Chicago/Turabian Style

Liu, Chang, Xin-Cun Wang, Zhi-He Yu, Wen-Ying Zhuang, and Zhao-Qing Zeng. 2023. "Seven New Species of Eurotiales (Ascomycota) Isolated from Tidal Flat Sediments in China" Journal of Fungi 9, no. 10: 960. https://doi.org/10.3390/jof9100960

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