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Research Article
Two new species of Penicillium (Eurotiales, Aspergillaceae) from China based on morphological and molecular analyses
expand article infoRui-Na Liang‡, Xiang-Hao Lin‡, Miao-Miao An‡, Guo-Zhu Zhao‡
‡ Beijing Forestry University, Beijing, China

Corresponding author: Guo-Zhu Zhao ( zhaogz@bjfu.edu.cn )

Academic editor: Ning Jiang
© 2025 Rui-Na Liang, Xiang-Hao Lin, Miao-Miao An, Guo-Zhu Zhao.
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: Liang R-N, Lin X-H, An M-M, Zhao G-Z (2025) Two new species of Penicillium (Eurotiales, Aspergillaceae) from China based on morphological and molecular analyses. MycoKeys 116: 255-274. https://doi.org/10.3897/mycokeys.116.149376
Open Access

Abstract

Penicillium is a large and significant genus of fungi, exhibiting widespread distribution across diverse substrates. Ongoing taxonomic and nomenclatural revisions have led to an annual increase in the number of newly described species. This study described two new Penicillium species, i.e., P. lentum and P. tibetense, discovered in China. They have been identified and characterized through morphological examination and both single gene and multigene phylogenetic analyses. Based on these analyses, P. lentum was classified within the section Brevicompacta, while P. tibetense was placed in the section Lanata-Divaricata. Both species exhibited the morphological features typical of their respective sections. Penicillium lentum is characterized by restricted growth with dense colonies on agar media and predominantly generates terverticillate conidiophores. Penicillium tibetense demonstrates rapid growth on media and has vigorous growth on CYA at 30 °C, producing biverticillate conidiophores. Comprehensive descriptions and detailed illustrations of these new species were presented. A morphological comparison between the new species and their closely related taxa was provided.

Key words:

Aspergillaceae, DNA barcodes, section Brevicompacta, section Lanata-Divaricata, taxonomy

Introduction

Penicillium is widely distributed across various substrates, primarily in soil, as well as in the atmosphere, food, plant tissues, and other environments. Several species possess considerable value for human applications in food production, biocontrol, and biotechnology. For instance, P. sclerotiorum exhibits antagonistic activity against certain plant pathogens, demonstrating potential as a biocontrol agent (Jahan et al. 2024). The food industry utilizes P. nalgiovense as starter cultures for dry-fermented sausages (Ludemann et al. 2010). The capability of certain species to synthesize pigments has prompted the evaluation of these species for the production of highly stable and safe natural pigments (Morales-Oyervides et al. 2020). Nevertheless, mycotoxins generated by specific species present a significant risk to human and animal health (Nielsen et al. 2017). Notably, patulin exhibits multiple toxicities, including genotoxicity and immunotoxicity, and is predominantly produced by P. expansum and P. griseofulvum (Bandoh et al. 2009; Puel et al. 2010; Tannous et al. 2014).

Link (1809) introduced the generic name Penicillium, which is classified in the family Aspergillaceae. Traditional taxonomy of Penicillium primarily relied on morphological characters, including colony diameter, texture, conidial color, and conidiophore branching patterns. However, the variability in morphology has presented substantial challenges in accurately identifying novel species, frequently resulting in the erroneous classification of new isolates under known species (Visagie et al. 2016). Conversely, contemporary taxonomy adopts a polyphasic strategy that incorporates morphological, extrolite, genetic, and multigene phylogenetic data (Visagie et al. 2014). Houbraken et al. (2020) delivered the most comprehensive update on the genus Penicillium based on a phylogenetic approach combined with phenotypic, physiologic, and extrolite data. This study recognized 483 species and introduced a novel series classification, which is deemed highly predictive of potential functional traits (Houbraken et al. 2020). Subsequently, Visagie et al. (2024b) applied GCPSR (Genealogical Concordance Phylogenetic Species Recognition) and phylogenetic analyses to reassess the list of Penicillium species published up to 31 December 2022, resulting in an updated count of 535 species. An additional 100 species of this genus were described from 1 January 2023 to 31 December 2024 (Ansari et al. 2023; Crous et al. 2023; da Silva et al. 2023; Khuna et al. 2023; Li et al. 2023; Liu et al. 2023; Tan 2023; Tan and Shivas 2023, 2024; Tan et al. 2023, 2024a, 2024b; Wang et al. 2023; Zhang et al. 2023; Araújo et al. 2024; Crous et al. 2024; Liang et al. 2024; Lima et al. 2024; Nóbrega et al. 2024; Song et al. 2024; Visagie et al. 2024a, 2024c; Zhang et al. 2024). The increase in species numbers in recent years indicates the possibility of numerous undiscovered Penicillium species, and their biodiversity, ecological functions, and potential for resource development warrant further investigation.

During a comprehensive survey of Penicillium biodiversity in China, we found two isolates that could not be classified within existing species. In this paper, we compare these isolates with related species using multi-locus phylogenetic analyses and morphological character assessments. As a result, the isolates are described as species new to science. This study is expected to offer new perspectives on the diversity, function, ecology, and distribution of Penicillium members.

Materials and methods

Isolates

Soil samples were collected from the rhizosphere of plants in the Kangyu Tunnel, Tibet, China, while indoor dust samples were sourced from Beijing Forestry University, Beijing, China. To isolate the fungus, the samples were suspended in sterile water at a ratio of 1:10, vortexed to ensure homogeneity, and then diluted to 10-4 concentrations. Each of 100 μL from 10-2, 10-3, and 10-4 dilutions was spread on potato dextrose agar (PDA) and Martin medium with 50 ppm penicillin and 50 ppm streptomycin. The cultures were incubated at 25 °C for 5–7 days. Individual colonies were then picked from the plates and transferred to fresh PDA plates until pure cultures were obtained. Type specimens, preserved as dry cultures, were deposited in the Fungarium (HMAS), Institute of Microbiology, Chinese Academy of Sciences, while ex-type strains, maintained as living cultures, were stored at the China General Microbiological Culture Collection Centre (CGMCC).

Morphological studies

Morphological observations of colonies were conducted under strictly standardized conditions, encompassing media preparation, inoculation technique, incubation parameters, and description methods (Visagie et al. 2014). Colony characters and diameters were recorded from cultures grown on Czapek yeast autolysate agar (CYA), malt extract agar (MEA), yeast extract sucrose agar (YES), dichloran 18% glycerol agar (DG18), and creatine sucrose agar (CREA) at 25 °C for 7 days. Additional CYA plates were incubated at 30 and 37 °C. Color names and codes adhered to the book “Color Standards and Color Nomenclature” (Ridgway 1912). Ehrlich reaction was employed to assess the production of indole metabolites; a violet ring observed within ten min was deemed a positive result, while other color changes were interpreted as negative (Lund 1995; Houbraken et al. 2016).

For light microscopic observations, slides were prepared from cultures grown on MEA, and phenol glycerin solution was used as mounting fluid, with cotton blue staining if necessary. In addition, a field emission scanning electron microscope (Hitachi SU8010, Japan) was employed to examine microstructural characteristics. Agar blocks (3–4 mm × 3–4 mm) were fixed in 2.5% v/v glutaraldehyde at 4 °C for 8–12 hr, then washed three times for 10 min each with 0.1M phosphate buffer. Dehydration was performed with a gradient of ethanol (30, 50, 70, 95, and 100% v/v) for 10–20 min per step, followed by replacement with tert-butanol and ultimate vacuum freeze-dried and gold-sprayed for observation (Wei et al. 2024).

DNA extraction, sequencing, and phylogenetic analyses

Colonies were cultivated on MEA plates for 5–7 days, and DNA extraction was conducted using the E.Z.N.A.® Fungal DNA Mini Kit (Omega Bio-Tek, Inc., United States). The internal transcribed spacer (ITS), beta-tubulin (BenA), calmodulin (CaM), and RNA polymerase II second largest subunit (RPB2) genes were amplified using primer pairs ITS1/ITS4 (White et al. 1990), Bt2a/Bt2b (Glass and Donaldson 1995), CMD5/CMD6 (Hong et al. 2006), and RPB2-5F/RPB2-7CR (Liu et al. 1999), respectively. Polymerase chain reaction (PCR) amplification followed Visagie et al. (2014). Sequencing reactions were performed by Sangon Biotech (Shanghai) Company Limited, China. DNAMAN software (Lynnon Biosoft) was used for the assembly and trimming of the Sanger chromatograms. Sequences were submitted to GenBank (www.ncbi.nlm.nih.gov).

Sequence similarity searches were conducted using the mega BLAST program of basic local alignment search tool (BLAST) within the NCBI core nucleotide database (core_nt). Comprehensive sequence datasets were compiled containing newly generated sequences alongside reference sequences sourced from GenBank (Table 1). Sequence alignments were performed using the ClustalW algorithm and subsequently manually edited using MEGA 11 (Tamura et al. 2021). The resulting multiple sequence alignments have been deposited in TreeBASE (submission number: 31847) (www.treebase.org). Phylogenetic trees were constructed based on the ITS, BenA, CaM, and RPB2 genes as well as the concatenated sequences of the latter three genes. Phylogenetic analyses were conducted using both maximum likelihood (ML) and Bayesian Inference (BI). ML phylogenies were performed using IQtree v. 1.6.12 (Nguyen et al. 2015), including 1000 standard non-parametric bootstrap replicates with the best partition scheme and substitution model selected using ModelFinder (Kalyaanamoorthy et al. 2017). BI phylogenies were run in MrBayes v. 3.2.7 (Ronquist et al. 2012). Best fit models were selected according to the Akaike information criterion (AIC) using MrModeltest v. 2.4 (Nylander 2004). Posterior probabilities (PP) were estimated using Markov Chain Monte Carlo (MCMC) sampling, set to run for 1,000,000 generations with the average standard deviation of split frequencies less than 0.01 as the stopping criterion. In cases where this threshold was not achieved, the run was continued until the condition was met. Additionally, the initial 25% of the generated trees were discarded as burn-in.

Table 1.
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Strains of Penicillium used for phylogenetic analyses.

Species Strain Substrate and origin GenBank accession numbers
ITS BenA CaM RPB2
P. abidjanum CBS 246.67T Soil, Ivory Coast GU981582 GU981650 MN969234 JN121469
P. alagoense URM 8086T Leaves of Miconia sp., Brazil MK804503 MK802333 MK802336 MK802338
P. amphipolaria CBS 140997T Soil, Antarctica KT887872 KT887833 KT887794 MN969177
P. annulatum CBS 135126T Air sample, South Africa JX091426 JX091514 JX141545 KF296410
P. araracuaraense CBS 113149T Leaf litter, Colombia GU981597 GU981642 MN969237 KF296414
P. astrolabium CBS 122427T Grapes, Portugal DQ645804 DQ645793 DQ645808 JN406634
P. ausonanum CBS 148237T Sediment of freshwater stream, Spain LR655808 LR655809 LR655810 LR655811
P. austrosinense CGMCC 3.18797T Acidic soil, China KY495007 KY495116 MN969328 KY495061
P. bialowiezense CBS 227.28T Soil under conifers, Poland EU587315 AY674439 AY484828 JN406604
P. bissettii CBS 140972T Soil from spruce forest, Canada KT887845 KT887806 KT887767 MN969178
P. brasilianum CBS 253.55T Herbarium exsiccata, Brazil GU981577 GU981629 MN969239 KF296420
P. brevicompactum NRRL 28139 Stroma of a wood decay fungus, USA AY484917 DQ645795 AY484825 –
CV1492 Unknown, South Africa JX091398 JX091533 JX141574 –
CBS 257.29T Unknown, Belgium AY484912 AY674437 AY484813 JN406594
P. buchwaldii CBS 116980 Wheat, United Kingdom JX313163 JX313181 JX313147 –
CBS 116935 Wheat, United Kingdom JX313156 JX313174 JX313140 –
CBS 116929 Wheat flour, Denmark JX313152 JX313170 JX313136 –
CBS 117181T Hordeum vulgare, Denmark JX313164 MN969374 JX313148 JN406637
P. camponoti CBS 140982T Carpenter ants, Canada KT887855 KT887816 KT887777 MN969179
P. cataractarum CBS 140974T Fallen nuts of Carya cordiformis, Canada KT887847 KT887808 KT887769 MN969180
P. coffeatum CGMCC 3.25152T Soil, China OQ870815 OR051121 OR051298 OR051466
P. daleae CBS 211.28T Soil under conifer, Poland GU981583 GU981649 MN969251 KF296427
P. echinulonalgiovense CBS 328.59T Unknown, Japan GU981587 GU981631 KX961269 KX961301
P. excelsum DTO 357-D7T Brazil nut shell, Brazil KR815341 KP691061 KR815342 MN969166
ITAL 7804 Flowers, Brazil KT749963 KT749959 KT749962 –
P. expansum CBS 325.48T Malus sylvestris, USA AY373912 AY674400 DQ911134 JF417427
P. fengjieense CGMCC 3.25157T Soil, China OQ870765 OR051156 OR051333 OR051489
P. fennelliae CBS 711.68T Soil, Congo JX313169 MN969382 JX313151 JN406536
P. flaviroseum CGMCC 3.18805T Acidic soil, China KY495032 KY495141 MN969329 KY495083
P. fructuariae-cellae CBS 145110T Dried fruit of Vitis vinifera, Italy MK039434 KU554679 MK045337 –
P. globosum CBS 144639T Acidic soil, China KY495014 KY495123 MN969330 KY495067
P. griseoflavum CGMCC 3.18799T Acidic soil, China KY495011 KY495120 MN969331 KY495064
P. griseopurpureum CBS 406.65T Soil under Pinus sp., United Kingdom KF296408 KF296467 MN969261 KF296431
P. guaibinense CCDCA 11512T Soil, Brazil MH674389 MH674391 MH674393 –
P. guangxiense CBS 144526T Soil, China KY494986 KY495095 MN969332 KY495045
P. hainanense CGMCC 3.18798T Acidic soil, China KY495009 KY495118 MN969333 KY495062
P. infrabuccalum CBS 140983T Camponotus pennsylvanicus, Canada KT887856 KT887817 KT887778 MN969181
P. jianfenglingense CGMCC 3.18802T Acidic soil, China KY495016 KY495125 MN969334 KY495069
P. jinyunshanicum CGMCC 3.25162T Soil, China OQ870766 OR051157 OR051334 OR051490
P. kongii AS3.15329T leaf sample of Cotoneaster sp., China KC427191 KC427171 KC427151 –
P. laevigatum CGMCC 3.18801T Acidic soil, China KY495015 KY495124 MN969335 KY495068
P. lentum CGMCC 3.28596T = B24 Indoor dust, Beijing, China PQ643282 PQ519854 PQ519855 PQ519856
P. mariae-crucis CBS 271.83T Secale cereale, Spain GU981593 GU981630 MN969275 KF296439
P. marykayhuntiae BRIP 74934aT Soil, Australia OR271913 OR269446 – OR269440
P. neocrassum CBS 122428T Grapes, Madeira DQ645805 DQ645794 DQ645809 JN406633
P. newtonturnerae BRIP 74909aT Soil, Australia OP903478 OP921964 OP921962 OP921963
P. ochrochloron CBS 357.48T Copper sulphate solution, USA GU981604 GU981672 MN969280 KF296445
DTO 189-A6 Unknown, Japan KC346347 KC346324 KC346341 KC346318
P. olsonii CBS 232.60T Musa, France EU587341 AY674445 DQ658165 JN121464
P. onobense CBS 174.81T Soil, andosol, Spain GU981575 GU981627 MN969281 KF296447
P. panissanguineum CBS 140989T Soil near termite mound, Tanzania KT887862 KT887823 KT887784 MN969182
P. paraherquei CBS 338.59T Soil, Japan AF178511 KF296465 MN969285 KF296449
P. pauciramulum CGMCC 3.25164T Soil, associated with nest of Formicidae, China OQ870726 OR051111 OR051288 OR051457
P. pedernalense CBS 140770T Litopenaeus vannamei, Ecuador KU255398 KU255396 MN969322 MN969184
P. penarojense CBS 113178T Leaf litter, Colombia GU981570 GU981646 MN969287 KF296450
P. piscarium CBS 362.48T Cod-liver oil emulsion, Germany GU981600 GU981668 MN969288 KF296451
P. pulvillorum CBS 280.39T Acidic soil, United Kingdom AF178517 GU981670 MN969289 KF296452
CBS 275.83 Rye grain, Spain GU981601 GU981671 KC346336 KF296423
P. rolfsii CBS 368.48T Fruit of Ananas sativus, USA JN617705 GU981667 MN969294 KF296455
P. roodeplaatense DTO 444-C8 Soil, South Africa OR819195 OR820176 OR820180 OR820186
P. rotoruae CBS 145838T Pinus radiata timber stake in ground contact, New Zealand MN315103 MN315104 MN315102 MT240842
P. rubriannulatum CGMCC 3.18804T Acidic soil, China KY495029 KY495138 MN969336 KY495080
P. salamii CBS 135391T Salami, Italy HG514431 HG514437 HG514432 MN969160
P. simplicissimum CBS 372.48T Secale cereale, Spain GU981588 GU981632 MN969297 JN121507
P. singorense CBS 138214T House dust, Thailand KJ775674 KJ775167 KJ775403 MN969138
P. skrjabinii CBS 439.75T Soil, Russia GU981576 GU981626 MN969299 EU427252
P. soliforme CGMCC 3.18806T Acidic soil, China KY495038 KY495147 MN969337 KY495047
NN072390 Acidic soil, China KY495019 KY495128 KY494959 KY495072
NN072399 Acidic soil, China KY495022 KY495131 KY494962 KY495074
P. spathulatum CBS 117192T Mouldy chestnut (Castanea sp.), France JX313165 MN969400 JX313149 JN406636
P. spinuliferum CBS 144483T Acidic soil, associated with Litchi chinensis, China KY495040 KY495149 MN969338 KY495090
P. stangiae URM 8347T Soil, Brazil MW648590 MW646388 MW646390 MW646392
P. stolkiae CBS 315.67T Soil, South Africa AF033444 JN617717 AF481135 JN121488
P. subfuscum CBS 147455T Soil, South Africa MT949907 MT957412 MT957454 MT957480
P. subrubescens CBS 132785T Soil of Helianthus tuberosus field, Finland KC346350 KC346327 KC346330 KC346306
P. subrutilans CGMCC 3.25174T Soil, China OQ870816 OR051137 OR051314 OR051479
P. svalbardense CBS 122416T Glacial ice, Svalbard GU981603 DQ486644 KC346338 KF296457
P. taii CGMCC 3.25176T Soil, China OQ870778 OR051170 OR051347 OR051496
P. tanzanicum CBS 140968T Soil near termite mound, Tanzania KT887841 KT887802 KT887763 MN969183
P. terrarumae CBS 131811T Soil contaminated by heavy metals, China MN431397 KX650295 MN969323 MN969185
CS23-08 Unknown, China OQ870751 OR051141 OR051318 OR051481
P. tularense CBS 430.69T Soil under Pinus ponderosa and Quercus kelloggii, USA AF033487 KC427175 JX313135 JN121516
CBS 431.69 Soil under Pinus ponderosa and Quercus kelloggii, USA JX313167 AY674433 JX313134 –
P. vanderhammenii CBS 126216T Leaf litter, Colombia GU981574 GU981647 MN969308 KF296458
P. vasconiae CBS 339.79T Soil, Spain GU981599 GU981653 MN969309 MN969144
P. vickeryae BRIP 72552aT Soil, Australia OP903479 OP921966 – OP921965
P. viridissimum CGMCC 3.18796T Acidic soil, China KY495004 KY495113 MN969339 KY495059
P. wotroi CBS 118171T Leaf litter, Colombia GU981591 GU981637 MN969313 KF296460
P. tibetense CGMCC 3.28597T = XZ5-3 Rhizosphere soil, Tibet, China PQ643284 PQ519857 PQ519858 PQ519859
P. yuyongnianii CGMCC 3.25187T Soil, China OQ870820 OR051175 OR051352 OR051499
P. zonatum CBS 992.72T Soil, USA GU981581 GU981651 MN969315 KF296461

Results

Morphology

Two novel species, Penicillium lentum and P. tibetense, were introduced within the sections Brevicompacta and Lanata-Divaricata, respectively, based on comprehensive phylogenetic analyses. General morphological characteristics and ecological information for the species included in these sections are provided in Table 2. Both newly described species exhibited morphological traits consistent with their respective sections. Specifically, P. lentum displayed limited growth with dense colonies on agar media and primarily produced terverticillate conidiophores. In contrast, P. tibetense demonstrated rapid growth on agar media, particularly exhibiting robust development on CYA at 30 °C, and predominantly formed biverticillate conidiophores. The morphological features of the new species and their closely related species are summarized in Table 3.

Table 2.
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Morphological and ecological data pertaining to the sections of the new species in this study.

Section Morphology Ecology References
Brevicompacta Colonies restricted (occasionally moderately fast), texture velutinous; conidiophores terverticillate or multiramulate branched with wide stipes, smooth-walled. Mainly soil and foods, also on plant leaves and rotting wood. (Houbraken and Samson 2011; Frisvad et al. 2013; Wang and Wang 2013; Houbraken et al. 2020)
Lanata-Divaricata Colonies grow rapidly, occasionally moderately fast; conidiophores monoverticillate, biverticillate or divaricate, occasionally terverticillate. Commonly found in soil, also on rotting leaf litter and vegetable. (Houbraken and Samson 2011; Houbraken et al. 2020)
Table 3.
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Morphological features of new species and their closely related taxa.

Species Growth rates (mm) Conidiophores branching Cleistothecia /sclerotia Conidia Acid production on CREA
CYA CYA 30 °C CYA 37 °C Size Shape Roughening
P. lentum 7–10 No growth No growth Terverticillate, sometimes biverticillate Absent 2–3 × 1.5–2.5 μm Broadly ellipsoidal Smooth Absent
P. tularense a n.a. n.a. n.a. Asymmetric and divaricate Cleistothecia 2.2–2.6 μm Globose to subglobose Smooth n.a.
P. tibetense 42–50 42–52 21–27 Biverticillate Absent 1.5–3 μm Globose to subglobose Finely rough Absent
P. excelsum b 35–50 n.a. 8–22 Biverticillate, sometimes terverticillate Absent 4–5 × 2–3.2 μm Ellipsoidal Smooth Absent

aDescription based on Paden (1971), bDescription based on Taniwaki et al. (2016).

Phylogenetic analyses

A BLAST search revealed that strain CGMCC 3.28596 is most closely related to Penicillium tularense (Identities: ITS: 97.52%, BenA: 81.13%, CaM: 84.91%, RPB2: 91.00%) within section Brevicompacta, and strain CGMCC 3.28597 exhibits the highest similarity to P. excelsum (Identities: ITS: 98.64%, BenA: 94.37%, CaM: 89.66%, RPB2: 94.84%) within section Lanata-Divaricata.

Section Brevicompacta

The analyses of the concatenated dataset (BenA, CaM, and RPB2) comprised 20 predominantly ex-type strains, each with a total sequence length of 1876 bp (BenA: 469 bp, CaM: 512 bp, RPB2: 895 bp). Phylogenetic analyses divided section Brevicompacta into four distinct clades, with the new species Penicillium lentum forming a robustly supported clade alongside P. tularense (100% bs, 1.00 pp) (Fig. 1). In the phylogenetic analyses of individual genes, the new species, together with P. tularense, consistently formed a well-supported clade, mostly with high support values (>97% bs, 1.00 pp), except for ITS (Fig. 2).

Figure 1. 

ML tree based on the concatenated data set (BenA, CaM, and RPB2) of section Brevicompacta. Penicillium expansum CBS 325.48T was designated as the outgroup. Nodes display bootstrap values (bs) exceeding 70% or posterior probabilities (pp) greater than 0.95. Branches with bs of 95% or higher and pp of 1.00 are depicted in bold. The strain described as the new species P. lentum is indicated with blue text. * Indicates bs = 100% or pp = 1.00, T = ex-type strain.

Figure 2. 

ML trees for section Brevicompacta based on ITS, BenA, CaM, and RPB2. Penicillium expansum CBS 325.48T was designated as the outgroup. Nodes display bootstrap values (bs) exceeding 70% or posterior probabilities (pp) greater than 0.95. Branches with bs of 95% or higher and pp of 1.00 are depicted in bold. The strain described as the new species P. lentum is indicated with blue text. * Indicates bs = 100% or pp = 1.00, T = ex-type strain.

Section Lanata-Divaricata

In this section, we selected the series Simplicissima, Dalearum, and Rolfsiorum, comprising 71 predominantly ex-type strains, for phylogenetic analyses based on the concatenated dataset totaling 1876 bp (BenA: 504 bp, CaM: 617 bp, RPB2: 755 bp). The resulting phylogenies revealed that Penicillium tibetense is closely related to P. excelsum (64% bs, 0.97 pp; not depicted in Fig. 3). However, the significant evolutionary divergence observed supports the recognition of P. tibetense as a distinct species (Fig. 3). Phylogenetic analyses of individual genes within series Rolfsiorum demonstrated generally weak clustering support, with variations observed among the ITS, BenA, CaM, and RPB2 datasets. Furthermore, P. ochrochloron and P. rotoruae share identical ITS sequences, making them indistinguishable through ITS phylogeny alone (Fig. 4).

Figure 3. 

ML tree based on the concatenated data set (BenA, CaM, and RPB2) of section Lanata-Divaricata (series Simplicissima, Dalearum, and Rolfsiorum). Penicillium stolkiae CBS 315.67T was designated as the outgroup. Nodes display bootstrap values (bs) exceeding 70% or posterior probabilities (pp) greater than 0.95. Branches with bs of 95% or higher and pp of 1.00 are depicted in bold. The strain described as the new species P. tibetense is indicated with blue text. * Indicates bs = 100% or pp = 1.00, T = ex-type strain.

Figure 4. 

ML trees for section Lanata-Divaricata series Rolfsiorum based on ITS, BenA, CaM, and RPB2. Penicillium stolkiae CBS 315.67T was designated as the outgroup. Nodes display bootstrap values (bs) exceeding 70% or posterior probabilities (pp) greater than 0.95. Branches with bs of 95% or higher and pp of 1.00 are depicted in bold. The strain described as the new species P. tibetense is indicated with blue text. * Indicates bs = 100% or pp = 1.00, T = ex-type strain.

Taxonomy

Penicillium lentum R.N. Liang & G.Z. Zhao, sp. nov.

MycoBank No: 857346
Fig. 5

Infrageneric classification.

Subgenus Penicillium, section Brevicompacta, series Tularensia.

Figure 5. 

Penicillium lentum CGMCC 3.28596. A Colonies on medium at 25 °C for 7d (left to right, top row: CYA, YES, DG18, MEA obverse; second row: CYA reverse, YES reverse, DG18 reverse, CREA obverse) B–E conidiophores F conidia G–I SEM micrograph of conidiophores J SEM micrograph of conidia. Scale bars: 10 μm (B–I); 2 μm (J).

Etymology.

The specific epithet “lentum” is derived from lentus (Latin), reflecting the slow growth rate characteristic of this species.

Type.

China • Beijing, Haidian District, Beijing Forestry University, 40°0'20"N, 116°20'51"E, from indoor dust, 1 February 2024, collected by G.Z. Zhao, B24 (holotype HMAS 353385, dried culture; culture ex-type CGMCC 3.28596).

Colony diameter after 7 d (mm).

CYA 7–10; CYA 30 °C, 37 °C no growth; MEA 6–9; YES 9–13; DG18 7–11; CREA 3.5–5.

Colony characteristics (7 d).

CYA at 25 °C: Colonies deep, raised at center, margins low, narrow, irregular; mycelium white; texture velutinous, floccose areas present; sporulation moderate to good, conidia antique green (R. Pl. VI); exudate clear; reverse capucine buff (R. Pl. III); soluble pigment absent. MEA at 25 °C: Colonies deep, raised at center, margins low, narrow, entire; mycelium white; texture velutinous, floccose areas present; sporulation moderate to good, conidia celandine green (R. Pl. XLVII) to deep turtle green (R. Pl. XXXII); exudate clear; reverse light orange-yellow (R. Pl. III); soluble pigment absent. YES at 25 °C: Colonies deep, radially and concentrically sulcate, raised at center, margins low, narrow, entire; mycelium white; texture velutinous and fasciculate; sporulation good to strong, conidia glaucous-green (R. Pl. XXXIII); exudate absent; reverse cinnamon (R. Pl. XXIX); soluble pigment absent. DG18 at 25 °C: Colonies low, plane, margins low, wide, entire; mycelium white; texture velutinous and fasciculate; sporulation good, conidia bluish gray-green (R. Pl. XLII); exudate absent; reverse antimony yellow (R. Pl. XV); soluble pigment absent. CREA at 25 °C: Weak growth, no acid production. Ehrlich reaction negative.

Micromorphology.

Conidiophores biverticillate to terverticillate; stipes smooth-walled, 70–236.5 × 2.5–4.5 μm; rami two when present, 6.5–18 × 2–4 μm; metulae divergent, 2–4 per branch/ramus, 4.0–13.0 × 2.5–4.5 μm; phialides ampulliform, 3–8 per metula, 4.5–8.0 × 2–3 μm; conidia broadly ellipsoidal, smooth-walled, 2–3 × 1.5–2.5 μm.

Notes.

Penicillium lentum belongs to section Brevicompacta and is most closely related to P. tularense (Fig. 1). Penicillium tularense produces light brown to pale tan cleistothecia, which are not found in the new species (Paden 1971). Additionally, P. lentum has broadly ellipsoidal conidia, while P. tularense produces globose to subglobose conidia (Table 3).

Penicillium tibetense R.N. Liang & G.Z. Zhao, sp. nov.

MycoBank No: 857347
Fig. 6

Infrageneric classification.

Subgenus Aspergilloides, section Lanata-Divaricata, series Rolfsiorum.

Figure 6. 

Penicillium tibetense CGMCC 3.28597. A Colonies on medium at 25 °C for 7d (left to right, top row: CYA, YES, DG18, MEA obverse; second row: CYA reverse, YES reverse, DG18 reverse, CREA obverse) B–D conidiophores E conidia F, G SEM micrograph of conidiophores H SEM micrograph of conidia. Scale bars: 10 μm (B–G); 2 μm (H).

Etymology.

The specific epithet “tibetense” denotes the geographical origin of the species, indicating its discovery in Tibet.

Type.

China • Tibet, Changdu City, Basu County, Kangyu Tunnel, 30°33'53"N, 96°15'25"E, from rhizosphere soil of grasses, 19 July 2023, collected by X.W. Peng, XZ5-3 (holotype HMAS 353386, dried culture; culture ex-type CGMCC 3.28597).

Colony diameter after 7 d (mm).

CYA 42–50; CYA 30 °C 42–52; CYA 37 °C 21–27; MEA 48–52; YES 46–52; DG18 20–26; CREA 24–26.

Colony characteristics (7 d).

CYA at 25 °C: Colonies low to moderately deep, radially sulcate, margins low, narrow, entire; mycelium white; texture floccose; sporulation moderate, conidia livid pink (R. Pl. XXVII); exudate clear; reverse light purple-drab (R. Pl. XLV) to avellaneous (R. Pl. XL); soluble pigment absent. CYA at 30 °C: Colonies low to moderately deep, radially sulcate, margins low, narrow, entire; mycelium white; texture floccose; sporulation moderate, conidia livid pink (R. Pl. XXVII); exudate clear; reverse brownish vinaceous (R. Pl. XXXIX); soluble pigment absent. CYA at 37 °C: Colonies moderately deep, radially sulcate, margins low, narrow, entire; mycelium white; texture floccose; sporulation sparse, conidia livid pink (R. Pl. XXVII); exudate clear; reverse light buff (R. Pl. XV); soluble pigment absent. MEA at 25 °C: Colonies low to moderately deep, radially sulcate, margins low, narrow, entire; mycelium white; texture floccose; sporulation sparse to moderate, conidia pale brownish vinaceous (R. Pl. XXXIX); exudate clear; reverse antimony yellow (R. Pl. XV); soluble pigment absent. YES at 25 °C: Colonies moderately deep, randomly sulcate, margins low, wide, entire; mycelium white; texture floccose; sporulation moderate, conidia antique green (R. Pl. VI); exudate clear; reverse antimony yellow (R. Pl. XV); soluble pigment absent. DG18 at 25 °C: Colonies low, radially sulcate, margins low, wide, entire; mycelium white; texture floccose; sporulation sparse, conidia ecru-drab (R. Pl. XLVI); exudate absent; reverse orange-pink (R. Pl. II); soluble pigment absent. CREA at 25 °C: Strong growth, no acid production. Ehrlich reaction negative.

Micromorphology.

Conidiophores biverticillate; stipes finely rough-walled, 27–364.5 × 2–3 μm; metulae appressed to divergent, 2–4 per stipe, 8–15 × 1.5–3 μm; phialides ampulliform to cylindrical, 2–6 per metula, 5–10.5 × 1.5–3 μm; conidia globose to subglobose, finely rough-walled, 1.5–3 μm diam.

Notes.

Penicillium tibetense is classified in section Lanata-Divaricata and exhibits a close phylogenetic relationship to P. excelsum (Fig. 3). This novel species generates globose to subglobose, finely rough-walled conidia that distinguish it from P. excelsum (Table 3). Additionally, P. tibetense demonstrates more robust growth on CYA at 37 °C compared to P. excelsum (21–27 mm vs. 8–22 mm) (Taniwaki et al. 2016).

Discussion

Penicillium, a ubiquitous and diverse fungal genus, plays pivotal roles in natural ecosystems while maintaining substantial economic importance and significant relevance to human affairs. The recent rapid increase in newly described species within this genus suggests that numerous taxa remain undiscovered. Given the extensive biotechnological applications of Penicillium species, accurate taxonomic identification is paramount, necessitating comprehensive species delineation through polyphasic approaches. In the present study, we introduced two novel species: one belonging to section Brevicompacta and the other to section Lanata-Divaricata.

Section Brevicompacta currently comprises 15 species distributed across four series (Visagie et al. 2024a, 2024b) and is represented in our findings by the newly described P. lentum sp. nov. This species, classified within series Tularensia, is characterized by predominantly terverticillate conidiophores and demonstrates a close relationship with other members of section Brevicompacta (Table 2). Section Lanata-Divaricata is characterized by its remarkable species diversity and rapid colony growth, primarily comprising soil-inhabiting species, with over 90 taxa currently recognized across five series (Houbraken et al. 2020; Visagie et al. 2024b). The newly identified Penicillium tibetense assigned to series Rolfsiorum exhibits characteristic rapidly expanding colonies and produces biverticillate conidiophores, consistent with the morphological features typical of this series. Members of section Lanata-Divaricata are ecologically significant as decomposers of organic matter (Lichtner et al. 2022), with notable biotechnological potential exemplified by P. subrubescens, which has demonstrated efficient inulinase production (Mansouri et al. 2013).

Phylogenetic analyses of section Brevicompacta demonstrated that our strain P. lentum formed a well-supported clade with its closest relative, P. tularense (Fig. 1), a relationship corroborated by shared morphological characteristics such as conidiophore branching patterns and growth rates. However, our strain could be clearly distinguished from P. tularense based on distinct phenotypic features, including the presence or absence of cleistothecia and differences in conidial morphology (Table 3). The phylogenetic relationships within section Lanata-Divaricata remain unresolved, primarily due to the poor support values in certain clades (Houbraken et al. 2020), exemplified by a clade comprising P. camponoti, P. piscarium, P. rolfsii, P. subrutilans, and P. terrarumae (Fig. 3), which highlights the persistent challenges in resolving certain taxonomic groups even with multigene phylogenetic approaches.

To address these limitations, we recommend expanding the taxonomic sampling to include strains from diverse geographical origins and ecological niches. This strategy would not only generate additional reference sequences but also facilitate the discovery of novel species and the detection of infraspecific variation (Visagie and Houbraken 2020). Furthermore, sequencing additional gene regions represents a promising approach to enhance phylogenetic resolution (Visagie et al. 2021). The rapid development of high-throughput sequencing technologies has resulted in an accelerated increase in genomic data availability (Kapli et al. 2020), positioning phylogenomics as an essential tool in modern fungal taxonomy and systematics. By leveraging genome-scale data, phylogenomics is poised to overcome the limitations of single gene or multigene analyses, providing robust statistical support for clade resolution and enabling the reconstruction of a highly resolved fungal tree of life (Burki et al. 2020; Zhou and May 2023). These advancements underscore the transformative potential of phylogenomics in addressing long-standing taxonomic challenges and refining our understanding of fungal evolutionary relationships.

Acknowledgements

We express our sincere gratitude to Professor Xia-Wei Peng for her invaluable assistance in the collection of soil samples from Tibet.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was funded by the National Natural Science Foundation of China (No. 31570019, 31093440) and the Survey Project of Alien Invasive Species and Grassland Pest in Mentougou District (2022HXFWSWXY038).

Author contributions

Rui-Na Liang: Formal analysis, investigation, data curation, writing – original draft preparation, visualization; Xiang-Hao Lin and Miao-Miao An: Investigation, visualization; Guo-Zhu Zhao: Conceptualization, methodology, validation, resources, writing – review and editing, supervision, funding acquisition. All authors have read and agreed to the published version of the manuscript.

Data availability

All of the data that support the findings of this study are available in the main text. All sequences generated in this study have been submitted to GenBank.

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