Research Article
Research Article
Morphological characteristics and phylogenetic evidence reveal two new species of Acremonium (Hypocreales, Sordariomycetes)
expand article infoXin Li, Zhi-Yuan Zhang, Yu-Lian Ren, Wan-Hao Chen§, Jian-Dong Liang§, Ji-Mei Pan§, Jian-Zhong Huang|, Zong-Qi Liang, Yan-Feng Han
‡ Guizhou University, Guiyang, China
§ Guizhou University of Traditional Chinese Medicine, Guiyang, China
| Fujian Normal University, Fuzhou, China
Open Access


Using chicken feathers as bait, Acremonium globosisporum sp. nov. and Acremonium curvum sp. nov. were collected from the soil of Yuncheng East Garden Wildlife Zoo and Zhengzhou Zoo in China. They were identified by combining the morphological characteristics and the two-locus DNA sequence (LSU and ITS) analyses. In the phylogenetic tree, both new species clustered into separate subclades, respectively. They were different from their allied species in their morphology. The description, illustrations, and phylogenetic tree of the two new species were provided.


Acremonium, filamentous fungi, phylogeny, taxonomy


The genus Acremonium Link, established in 1929, with A. alternatum Link as the type species, is one of the largest and most complex genera of asexually typified. The morphological characteristics consist of hyphae septate, mostly tapered and lateral phialides, produced singly or in small groups, and unicellular conidia produced in mucoid heads or unconnected chains (Summerbell et al. 2011). Nowadays, Acremonium has 217 records in the Index Fungorum (, retrieval on 30 Jun. 2022). The traditionally circumscribed Acremonium is polyphyletic, which explains why many Acremonium species were transferred to other genera and families (Yang et al. 2019). Thus, there are still many unidentified, suspect or misidentified taxa that require taxonomic investigation.

Due to the poor differentiation of asexual forms of the genus Acremonium, it is difficult to identify species only by morphological differences. To address this issue, there are many unidentified and suspicious species that require further phylogenetic analysis. To date, many isolates of Acremonium spp. lack the gene loci such as SSU, TEF 1-α and RPB2 (Table 1), therefore, phylogenetic analyses of this genus are generally performed based on the single locus sequences, especially LSU (Hyde et al. 2020).

In the present study, two new species of Acremonium were identified in a survey of keratinolytic fungi from China, which were enriched by the baiting technique. We provided a description, illustrations, and phylogenetic tree for the two new species.

Materials and methods

Fungal isolation and morphology

Soil samples were collected from Yuncheng East Garden Wildlife Zoo (35°6'26"N, 111°4'24"E) (three isolates), Yuncheng City, Shanxi Province and Zhengzhou Zoo (34°47'20"N, 113°40'41"E) (one isolate), Zhengzhou City, Henan Province, China by Yu-Lian Ren on July 2021. We collected 3–10 cm below the soil surface, placed the samples in sterile Ziploc plastic bags (Kaixin Biotechnology, Guizhou, China), and transported them to the laboratory (Zhang et al. 2019a, b). Then, they were treated and isolated according to the baiting method (using chicken feathers as bait: a method specifically designed for isolating keratinophilic microbes) of Zhang et al. (2020a, b; 2021). We washed the chicken feathers, sterilized them in an autoclave for 30 minutes at 121 °C, and dried them in an oven at 50 °C. The sterile and dried chicken feathers were mixed with soil samples and then wet with sterile distilled water and cultured at darkroom temperature for 1 month (Li et al. 2022).

Then, the 2 g samples were weighed in a conical flask with glass beads containing 20 mL sterile water and mixed evenly by eddy shock for 10 min. Next, 1 mL samples were mixed evenly in 9 mL sterile water in a sterile environment and diluted to 10-3. Then, 1 mL 10-3 samples were put into a sterile petri dish, and SDA medium containing 50 mg/L penicillin and 50 mg/L streptomycin was added and mixed. The target strains were isolated. The purified strains were transferred to PDA, OA, and MEA plates for dark culture at 25 °C for 7 days. Microscopic features were examined by making direct wet mounts with 25% lactic acid on PDA, with a light microscope.

The cultures were placed to slowly dry at 50 °C to produce the dried holotype. The dried holotype was deposited in the Mycological Herbarium of the Institute of Microbiology, Chinese Academy of Sciences, Beijing, China (HMAS), while ex-type living culture was stored in PDA test tubes which were deposited in the China General Microbiological Culture Collection Center (CGMCC), and the Institute of Fungus Resources, Guizhou University, Guiyang City, Guizhou, China (GZUIFR).

DNA extraction, PCR amplification, and sequencing

We used a 5% chelex-100 solution for total genomic DNA extraction. ITS1/ITS4 (White et al. 1990), LROR/LR7 (Vilgalys and Hester 1990), EF1-983F/ EF1-2218R (Rehner and Buckley 2005), fRPB2-5f/ fRPB2-7cR (Liu et al. 1999), and NS1 and NS4 (White et al. 1990) primers were used for amplification of the internal transcribed spacers (ITS), the 28S nrRNA locus (LSU), translation elongation factor 1-alpha gene region (TEF 1-α), RNA polymerase II second largest subunit gene (RPB2), and small subunit rDNA (SSU), respectively. Purification and sequencing were performed by Quintarabio (Wuhan, China). The new sequences were submitted to GenBank (Table 1).

Table 1.

Strains included in the present study.

Species Strains LSU ITS SSU TEF 1 RPB2
Acremonium alcalophilum CBS 114.92T JX158443 DQ825967 JX158486 JX158399 JX158465
Acremonium alternatum CBS 407.66T HQ231988 HE798150
Acremonium alternatum CBS 831.97 HQ231989
Acremonium arthrinii MFLU 18-1225T MN036334 MN036335 MN038169
Acremonium behniae CBS 146824T MW175400 MW175360
Acremonium biseptum CBS 750.69T HQ231998
Acremonium blochii CBS 993.69 HQ232002 HE608636
Acremonium borodinense CBS 101148T HQ232003 HE608635
Acremonium brachypenium CBS 866.73T HQ232004 AB540570
Acremonium camptosporum CBS 756.69T HQ232008 HQ232186
Acremonium cavaraeanum CBS 101149T HF680202 HF680220
Acremonium cavaraeanum CBS 111656 HF680203 HF680221
Acremonium cavaraeanum CBS 758.69 HQ232012 HF680222
Acremonium cerealis CBS 207.65 HQ232013
Acremonium cerealis CBS 215.69 HQ232014
Acremonium chiangraiense MFLUCC 14-0397T MN648329 MN648324
Acremonium chrysogenum CBS 144.62T HQ232017 HQ232187
Acremonium chrysogenum CBS 401.65 MH870276 MH858636
Acremonium citrinum CBS 384.96T HF680217 HF680236
Acremonium dimorphosporum CBS 139050T LN810506 LN810515
Acremonium exiguum CBS 587.73T HQ232035
Acremonium exuviarum UAMH 9995T HQ232036 AY882946
Acremonium felinum CBS 147.81T AB540488 AB540562
Acremonium flavum CBS 596.70T HQ232037 HQ232191
Acremonium flavum CBS 316.72 MH872204 MH860487
Acremonium fuci CBS 112868T AY632653
Acremonium fuci CBS 113889 AY632652
Acremonium fusidioides CBS 109069 HF680204 HF680223
Acremonium fusidioides CBS 991.69 HF680211 HF680230
Acremonium fusidioides CBS 840.68T HQ232039 FN706542
Acremonium hansfordii CBS 390.73 HQ232043 AB540578
Acremonium hennebertii CBS 768.69T HQ232044 HF680238
Acremonium inflatum CBS 212.69T HQ232050
Acremonium mali ACCC 39305T MF993114 MF987658
Acremonium moniliforme CBS 139051T LN810507 LN810516
Acremonium moniliforme FMR 10363 LN810508 LN810517
Acremonium parvum CBS 381.70A HQ231986 HF680219
Acremonium persicinum CBS 310.59T HQ232077
Acremonium persicinum CBS 101694 HQ232085
Acremonium pinkertoniae CBS 157.70T HQ232089 HQ232202
Acremonium polychroma CBS 181.27T HQ232091 AB540567
Acremonium potronii CBS 189.70 HQ232094
Acremonium pseudozeylanicum CBS 560.73T HQ232101
Acremonium pteridii CBS 782.69T HQ232102
Acremonium pteridii CBS 784.69 HQ232103
Acremonium sclerotigenum CBS 124.42T HQ232126 FN706552 HQ232209
Acremonium sclerotigenum A101 KC987215 KC987139 KC987177 KC998961
Acremonium sclerotigenum A130 KC987242 KC987166 KC987204 KC998988
Acremonium sp. E102 KC987248 KC987172 KC987210 KC998994 KC999030
Acremonium spinosum CBS 136.33T HQ232137 HE608637 HQ232210
Acremonium stroudii CBS 138820T KM225291
Acremonium tumulicola CBS 127532T AB540478 AB540552
Acremonium variecolor CBS 130360T HE608651 HE608647
Acremonium variecolor CBS 130361 HE608652 HE608648
Acremonium verruculosum CBS 989.69T HQ232150
Acrophialophora hechuanensis GZUIFR-H08-1T MK926789 DQ185070 EU053286
Brunneomyces brunnescens CBS 559.73T HQ231966 LN810520 HQ232184 LN810534
Brunneomyces hominis UTHSC 06-415T LN810509 KP131517 LN810535
Bryocentria brongniartii M139 EU940105 EU940052
Bryocentria brongniartii M190 EU940125 EU940052
Bryocentria metzgeriae M140 EU940106
Bulbithecium hyalosporum CBS 318.91T AF096187 HE608634
Cephalosporium purpurascens CBS 149.62T HQ232071
Cosmospora lavitskiae CBS 530.68T HQ231997
Emericellopsis alkalina CBS 127350T KC987247 KC987171 KC987209 KC998993 KC999029
Emericellopsis terricola CBS 120.40T U57082 U57676 U44112
Gliomastix roseogrisea CBS 134.56T HQ232121
Hapsidospora irregularis ATCC 22087T AF096192 AF096177
Kiflimonium curvulum CBS 430.66T HQ232026 HE608638 HQ232188
Lanatonectria flavolanata CBS 230.31 HQ232157
Leucosphaerina arxii CBS 737.84T HE608662 HE608640
Nigrosabulum globosum ATCC 22102T AF096195
Paracremonium contagium CBS 110348T HQ232118 KM231831 KM231966
Parasarocladium breve CBS 150.62T HQ232005
Parasarocladium radiatum CBS 142.62T HQ232104 HQ232205
Pestalotiopsis hawaiiensis CBS 114491T KM116239 KM199339 KM199514
Pestalotiopsis spathulata CBS 356.86T KM116236 KM199338 KM199513
Phialemonium atrogriseum CBS 604.67T HQ231981 HE610367 FJ176825
Pseudoacremonium sacchari CBS 137990T KJ869201 KJ869144
Sarcopodium vanillae CBS 100582 HQ232174 KM231780 KM231911
Sarocladium bacillisporum CBS 425.67T HQ231992 HE608639 HQ232179
Sarocladium bactrocephalum CBS 749.69T HQ231994 HG965006 HQ232180
Sarocladium strictum CBS 346.70T HQ232141 AY214439 HQ232211
Sarocladium terricola CBS 243.59T HQ232046 HQ232196
Selinia pulchra AR 2812 GQ505992 HM484859 HM484841
Trichothecium crotocinigenum CBS 129.64T HQ232018 AJ621773
Trichothecium indicum CBS 123.78T AF096194 AF096179
Trichothecium roseum DAOM 208997 U69891 U69892
Trichothecium sympodiale ATCC 36477 U69889 U69890
Acremonium curvum CGMCC 3.20954 = GZUIFR 22.035T ON041050 ON041034 ON876754 ON494579 ON494583
Acremonium globosisporum CGMCC 3.20955 = GZUIFR 22.036T ON041051 ON041035 ON876755 ON494580 ON494584
Acremonium globosisporum GZUIFR 22.037 ON041052 ON041036 ON876756 ON494581 ON494585
Acremonium globosisporum GZUIFR 22.038 ON041053 ON041037 ON876757 ON494582 ON494586

Phylogenetic analyses

The ITS and LSU sequences of Acremonium were downloaded from GenBank (Table 1). Two strains of Pestalotiopsis spathulata (CBS 356.86) and P. hawaiiensis (CBS 114491) were chosen as the outgroup taxa. The TBtools were used for name simplification and renaming (Chen et al. 2020). Sequences were aligned by MAFFT v7.037 (Katoh and Standley 2013). Multi-locus was concatenated by PhyloSuite v1.16 (Zhang et al. 2020a).

Bayesian inference (BI) and maximum likelihood (ML) methods were used in the analysis. For BI analysis was conducted with MrBayes v3.2 (Ronquist et al. 2012) and Markov chain Monte Carlo (MCMC) simulations; ML analysis was performed using IQ-TREE v1.6.11 (Nguyen et al. 2015), as outlined in Li et al (2022). All analyses were performed in PhyloSuite V1.16 (Zhang et al. 2020b).



Based on a BLAST search ( using the LSU sequences, our isolates were identified as belonging to the genus Acremonium. To further determine the phylogenetic position of these strains, we performed a multi-locus phylogenetic analysis. The dataset was composed of LSU (1–430 bp) and ITS (431–1005 bp) gene, comprising a total of 1005 characters (including gaps). The best-fit partition model for ML analysis and BI analysis is shown in Table 2. The results showed that the CGMCC 3.20955, GZUIFR 22.037, and GZUIFR 22.038 are still grouped in the Pinkertoniae-clade (Fig. 1). The CGMCC 3.20954 is still grouped in the Chrysogenum-clade (Fig. 1).

Table 2.

The best-fit substitution models are used in multi-locus phylogenetic construction.

ML analysis TN+F+R5 GTR+F+R4
BI analysis GTR+F+I+G4 GTR+F+I+G4
Figure 1. 

Phylogenetic tree of the genus Acremonium constructed from LSU and ITS. Bayesian posterior probability (≥ 0.95) and ML bootstrap values (≥ 70%) are indicated along branches (BPP/ML).


Acremonium globosisporum Xin Li, Y.F. Han & Z.Q. Liang, sp. nov.

MycoBank No: 843765
Fig. 2


Yuncheng East Garden Wildlife Zoo, Yuncheng City, Shanxi Province, China N35°6'26", E111°4'24", isolated from green belt soil, July 2021, Yu-Lian Ren (dried holotype culture HMAS 351939, ex-holotype culture CGMCC 3.20955 = GZUIFR 22.036). ITS sequences, GenBank ON041035; LSU sequences, GenBank ON041051; SSU sequences, GenBank ON876755; TEF 1-α sequences, GenBank ON494580; RPB2 sequences, GenBank ON494584.

Figure 2. 

Morphology of Acremonium globosisporum sp. nov. a–f colony on PDA, OA and MEA after 7 d at 25 °C (upper surface and lower surface) g–j conidia are borne on the phialides k–l winding hyphae and inflate hyphae. Scale bars: 4 mm (a–f); 10 μm (g–l).


Colonies on PDA and OA at 25 °C attaining 11–13 mm and 9–11 mm diam respectively after 7 d, white, flat or raised, velvety to slightly cottony. On MEA at 25 °C, reaching 8–10 mm after 7 d, white to yellowish white, raised, slimy. Hyphae hyaline, septate, sometimes winding and inflate, 1.5–11.0 µm wide. Sporulation abundant. Phialides are mostly borne singly, hyaline, erect to slightly curved, sometimes forming a collarette, 9.0–22.0 µm long, tapering from 1.5–3.5 µm near the base to 0.5–1.5 µm. Conidia cohering in long chains, with minutely truncate ends, up to 27.5 µm long, globose or subglobose, 2.5–4.5 × 2.5–4.5 µm (x– ± SD = 3.4 ± 0.77 × 3.6 ± 0.52, n = 50) diam. Chlamydospores and teleomorph stage were not observed.


globosisporum. A reference to the global conidia.

Additional specimens examined

Yuncheng East Garden Wildlife Zoo, Yuncheng City, Shanxi Province, China N35°6'26", E111°4'24", isolated from green belt soil, July 2021, Yu-Lian Ren, GZUIFR 22.037, ITS, LSU, SSU, TEF 1-α, RPB2 sequences GenBank ON041036, ON041052, ON876756, ON494581, ON494585; GZUIFR 22.038, ITS, LSU, SSU, TEF 1-α, RPB2 sequences GenBank ON041037, ON041053, ON876757, ON494582, ON494586.

Known distribution

Yuncheng City, Shanxi Province, China.


The phylogeny results showed that the CGMCC 3.20955, GZUIFR 22.037 and GZUIFR 22.038 still nested in the Pinkertoniae-clade. The morphological characteristics of Acremonium globosisporum were similar to other species of the Pinkertoniae-clade in that phialides were erect on the hyphae; sporulation was abundant, and conidia were subglobose (Ito et al. 2000). However, Acremonium globosisporum hyphae were sometimes winding and inflated, with conidia cohering in long chains, unlike other species.

Acremonium curvum Xin Li, Y.F. Han & Z.Q. Liang, sp. nov.

MycoBank No: 843766
Fig. 3


Zhengzhou Zoo, Zhengzhou City, Henan Province, China N34°47'20", E113°40'41", isolated from green belt soil, July 2021, Yu-Lian Ren (dried holotype culture HMAS 351938, ex-holotype culture CGMCC 3.20954 = GZUIFR 22.035). ITS sequences, GenBank ON041034, LSU sequences, GenBank ON041050; SSU sequences, GenBank ON876754; TEF 1-α sequences, GenBank ON494579; RPB2 sequences, GenBank ON494583.

Figure 3. 

Morphology of Acremonium curvum sp. nov. a–f colony on PDA, OA and MEA after 7 d at 25 °C (upper surface and lower surface) g, I, j conidia are borne on the phialides h Winding hyphae. Scale bars: 4 mm (a–f); 10 μm (g–j).


Colonies on PDA and OA at 25 °C attaining 11–14 mm and 7–9 mm diam respectively after 7 d, white, flat, radially folded or rugose. On MEA at 25 °C, reaching 6–8 mm after 7 d, white to yellowish-white, slimy. Hyphae hyaline, septate, sometimes winding, 1.5–2.5 µm wide. Sporulation abundant. Phialides are mostly borne singly, curved, slightly inflated at the base, tapered at the tip, up to 38.0 µm long. tapering from 1.5–3.5 µm near the base to 0.5–1.5 µm. Conidia cohering together on the top of phialides, one-celled, solitary, or several fascicled, ovoid or subglobose, 3.0–7.0 × 2.5–3.5 µm (x– ± SD = 4.1 ± 1.18 ×3.2 ± 0.77, n = 50) diam. Chlamydospores and teleomorph stage were not observed.


curvum. Referring to the curved Phialides.

Known distribution

Henan Province, China.


Based on the multi-locus analysis we found that Acremonium curvum had close phylogenetic affinities to other taxa of the Chrysogenum-clade. Morphologically, A. curvum was similar to other taxa of the Chrysogenum-clade in having simple or rarely branched conidiophores, slightly inflated at the base and tapered at tip phialides, and ovoid to subglobose conidia (Yang et al. 2019). Conidia of Hapsidospora irregularis and A. curvum had several fascicled at the tips of the conidiophores (Malloch and Cain 1970). However, A. curvum was differentiated by having mostly curved phialides and the conidia were several fascicled at the tips of the phialides.


In the present study, four strains of Acremonium fungi were isolated from soil in the Shanxi and Henan Province, China. Two-locus (LSU and ITS) phylogenetic analyses in combination with morphological characteristics were used for identification. As a result, two new species of A. curvum (one isolate) and A. globosisporum (three isolates) were proposed.

With the development of biotechnology, a growing number of studies have combined morphological and phylogenetic features to distinguish between species. This provides the basis for more precise species naming. Generally, the fungal ITS marker includes considerably more sequence variability, and consequently provides high interspecific resolution, and also some degree of intraspecific variability (Nilsson et al. 2008). Therefore, ITS has been widely used in studies of fungal inter- and intraspecific relationships (Dai et al. 2020; Szczepańska et al. 2021). There are numerous ITS sequences stored in public databases, which are incomparable to other molecular markers (Zhang et al. 2022). In addition, according to Vu et al. (2019), combining ITS and LSU can improve the accuracy of fungal species discrimination with high generality. They think that fungi commonly present in clinical, environmental, or economically relevant communities can often be identified to species level by their ITS and LSU barcodes (Vu et al. 2019).

Although, Yang et al. (2019) used a multi-locus phylogenetic analysis in introducing the new species Acremonium arthrinii, lacking loci such as SSU, TEF 1-α and RPB2 (Table 1) in the isolates of Acremonium spp. were relatively serious, so it is not difficult to find that the strains with SSU and TEF 1-α in this analysis were not yet 50% or even 30% of the total number of strains. Therefore, although we sequenced these above loci in the new isolates, they were not included in the phylogenetic analysis. In the future, the phylogeny relationships of Acremonium members will undoubtedly vary and become clearer with the increase of the number and type of molecular used.

In recent years, Acremonium spp. has been reported to cause immunocompetent and immunocompromised individual diseases, such as brain abscess (Anis et al. 2021), fungal keratitis (Liu et al. 2021), fungal osteomyelitis (Jalan et al. 2021), and fungal maxillary sinusitis (Durbec et al. 2011). In the present study, all strains were isolated by a method specifically designed for the isolation of keratinophilic microbes. Therefore, more studies are necessary to confirm whether A. curvum and A. globosisporum are opportunistic infectious pathogens that infect the skin and cause skin infection, as well as their potential application in the degradation of keratin-rich matrices.


This work was financially supported by the National Natural Science Foundation of China (no. 32060011, 32160007, 31860002), “Hundred” Talent Projects of Guizhou Province (Qian Ke He [2020] 6005), and the Key Areas of Research and Development Program of Guangdong Province (no. 2018B020205003), and Construction Program of Biology First-class Discipline in Guizhou (GNYL [2017] 009). We appreciate MDPI for the English-language editing of the whole manuscript.


  • Chen CJ, Chen H, Zhang Y, Thomas HR, Frank MH, He YH, Xia R (2020) TBtools: An integrative toolkit developed for interactive analyses of big biological data. Molecular Plant 13(8): 1194–1202.
  • Dai YD, Wu CK, Yuan F, Wang YB, Huang LD, Chen ZH, Zeng WB, Wang Y, Yang ZL, Zeng PS, Lemetti P, Mo XX, Yu H (2020) Evolutionary biogeography on Ophiocordyceps sinensis: An indicator of molecular phylogeny to geochronological and ecological exchanges. Geoscience Frontiers 11(3): 807–820.
  • Durbec M, Bienvenu AL, Picot S, Dubreuil C, Cosmidis A, Tringali S (2011) Maxillary sinus fungal infection by Acremonium. European Annals of Otorhinolaryngology, Head and Neck Diseases 128(1): 41–43.
  • Hyde KD, Dong Y, Phookamsak R, Jeewon R, Bhat DJ, Jones EBG, Liu NG, Abeywickrama PD, Mapook A, Wei D, Perera RH, Manawasinghe IS, Pem D, Bundhun D, Karunarathna A, Ekanayaka AH, Bao DF, Li JF, Samarakoon MC, Chaiwan N, Lin CG, Phutthacharoen K, Zhang SN, Senanayake IC, Goonasekara ID, Thambugala KM, Phukhamsakda C, Tennakoon DS, Jiang HB, Yang J, Zeng M, Huanraluek N, Liu JK, Wijesinghe SN, Tian Q, Tibpromma S, Brahmanage RS, Boonmee S, Huang SK, Thiyagaraja V, Lu YZ, Jayawardena RS, Dong W, Yang EF, Singh SK, Singh SM, Rana S, Lad SS, Anand G, Devadatha B, Niranjan M, Sarma VV, Liimatainen K, Aguirre-Hudson B, Niskanen T, Overall A, Alvarenga RLM, Gibertoni TB, Pfliegler WP, Horváth E, Imre A, Alves AL, Bahkali AH, Doilom M, Elgorban AM, Maharachchikumbura SSN, Rajeshkumar KC, Haelewaters D, Mortimer PE, Zhao Q, Lumyong S, Xu JC, Sheng J (2020) Fungal diversity notes 1151–1276: Taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 100(1): 5–277.
  • Ito T, Okane I, Nakagiri A, Gams W (2000) Two species of Acremonium section Acremonium: A. borodinense sp. nov. and A. cavaraeanum rediscovered. Mycological Research 104(1): 77–80.
  • Jalan D, Saini MK, Elhence P, Elhence A, Jain P (2021) Calcaneal osteomyelitis caused by Acremonium sp. in an immunocompetent adult – A case report. The Foot 47: е101781.
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780.
  • Li X, Zhang ZY, Chen WH, Liang JD, Huang JZ, Han YF, Liang ZQ (2022) A new species of Arthrographis (Eremomycetaceae, Dothideomycetes), from the soil in Guizhou, China. Phytotaxa 538(3): 175–181.
  • Liu J, Freiberg FJ, Yeung SN, Iovieno A (2021) Late-onset recurrent Acremonium fungal keratitis after therapeutic penetrating keratoplasty. Canadian Journal of Ophthalmology 56(4): e135–e137.
  • Malloch D, Cain RF (1970) Five new genera in the new family Pseudeurotiaceae. Canadian Journal of Botany 48(10): 1815–1825.
  • Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274.
  • Nilsson RH, Kristiansson E, Ryberg M, Hallenberg N, Larsson KH (2008) Intraspecific ITS variability in the kingdom fungi as ex-pressed in the international sequence databases and its implications for molecular species identification. Evolutionary Bioinformatics Online 4: 193–201.
  • Rehner SA, Buckley E (2005) A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: Evidence for cryptic diversification and links to Cordyceps teleomorphs. Mycologia 97(1): 84–98.
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542.
  • Summerbell RC, Gueidan C, Schroers HJ, de Hoog GS, Starink M, Arocha Rosete Y, Guarro J, Scott JA (2011) Acremonium phylogenetic overview and revision of Gliomastix, Sarocladium, and Trichothecium. Studies in Mycology 68(1): 139–162.
  • Szczepańska K, Guzow-Krzemińska B, Urbaniak J (2021) Infraspecific variation of some brown Parmeliae (in Poland) – a comparison of ITS rDNA and non-molecular characters. MycoKeys 85: 127–160.
  • Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172(8): 4238–4246.
  • Vu D, Groenewald M, de Vries M, Gehrmann T, Stielow B, Eberhardt U, Al-Hatmi A, Groenewald JZ, Cardinali G, Houbraken J, Boekhout T, Crous PW, Robert V, Verkley GJM (2019) Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Studies in Mycology 92(1): 135–154.
  • White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (Eds) PCR protocols: a guide to methods and applications, Academic Press, San Diego, California, 315–322.
  • Yang CL, Xu XL, Jeewon R, Boonmee S, Liu YG, Hyde KD (2019) Acremonium arthrinii sp. nov., a mycopathogenic fungus on Arthrinium yunnanum. Phytotaxa 420(4): 283–299.
  • Zhang ZY, Chen WH, Zou X, Han YF, Huang JZ, Liang ZQ, Deshmukh SK (2019a) Phylogeny and taxonomy of two new Plectosphaerella (Plectosphaerellaceae, Glomerellales) species from China. MycoKeys 57: 47–60.
  • Zhang ZY, Han YF, Chen WH, Liang ZQ (2019b) Gongronella sichuanensis (Cunninghamellaceae, Mucorales), a new species isolated from soil in China. Phytotaxa 416(2): 167–174.
  • Zhang ZY, Dong CB, Chen WH, Mou QR, Lu XX, Han YF, Huang JZ, Liang ZQ (2020a) The enigmatic Thelebolaceae (Thelebolales, Leotiomycetes): One new genus Solomyces and five new species. Frontiers in Microbiology 11: е572596.
  • Zhang ZY, Zhao YX, Shen X, Chen WH, Han YF, Huang JZ, Liang ZQ (2020b) Molecular phylogeny and morphology of Cunninghamella guizhouensis sp. nov. (Cunninghamellaceae, Mucorales), from soil in Guizhou, China. Phytotaxa 455(1): 31–39.
  • Zhang ZY, Shao QY, Li X, Chen WH, Liang JD, Han YF, Huang JZ, Liang ZQ (2021) Culturable fungi from urban soils in China I: Description of 10 new taxa. Microbiology Spectrum 9(2): e00867–e21.
  • Zhang Z, Chen W, Liang Z, Zhang L, Han Y, Huang J, Liang Z (2022) Revealing the non-overlapping characteristics between original centers and genetic diversity of Purpureocillium lilacinum. Fungal Ecology [accepted].