Morphological and phylogenetic analyses reveal novel entomopathogenic fungi infecting scale insects and aphids in China

1 Morphological and phylogenetic analyses reveal novel entomopathogenic fungi infecting scale insects and aphids in China Chunlin Yang1,2 , Xiulan Xu3, Xinyue Li1,2, Feng Liu1,2, Zhen Zeng3, Qiangang Xiao3, Yinggao Liu1,2 1 College of Forestry, Sichuan Agricultural University, Chengdu 611130, China 2 National Forestry and Grassland Administration Key Laboratory of Forest Resources Conservation and Ecological Safety on the Upper Reaches of the Yangtze River, College of Forestry, Sichuan Agricultural University, Chengdu 611130, China 3 Forestry Research Institute, Chengdu Academy of Agricultural and Forestry Sciences, Chengdu 611130, China Corresponding authors: Chunlin Yang ([email protected]); Xiulan Xu ([email protected]) Copyright: © Chunlin Yang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Entomopathogenic fungi exhibit a cosmopolitan distribution across diverse ecosystems, with their ubiquitous presence intrinsically linked to insect habitats—essentially occurring wherever insect populations exist. These fungi represent a vital biological resource, particularly in agriculture and forestry. They serve as a crucial repository of fungal strains for biological pest control. This investigation identified seven species from southwest China based on multi-gene (ITS, LSU, SSU, act, rpb1, rpb2, and tef1-α) phylogenetic analyses and morphological characteristics, including one new species and one newly recorded species in Cladosporium (Cladosporiaceae); four new species in Moelleriella (Clavicipitaceae); and one new species in Podonectria (Podonectriaceae). All seven fungi are in their asexual morphs and were discovered on aphids or scale insects, which are common and significant pests. These include Cladosporium kuwanaspidis, Cladosporium guizhouense, Moelleriella eucalypti, Moelleriella boehmeriae, Moelleriella cinnamomum, Moelleriella citrus, and Podonectria multiarmata. Descriptions and illustrations for all seven taxa are provided. Six of these species were collected from scale insects, specifically those found in bamboo forests, broad-leaved forests, or understory vegetation, and one was collected from aphids, primarily distributed on the underside of night-blooming jasmine leaves. This work reveals the rich diversity of entomopathogenic fungi in southwestern China, not only on larger insects such as Lepidoptera and Hymenoptera but also on smaller Hemiptera, many of which are significant agricultural and forestry pests. This study contributes fungal resources that may support the development of innovative pest control methods in the future. Key words: 6 new taxa, Cladosporiaceae, Clavicipitaceae, fungal entomopathogens, Podonectriaceae Introduction To ensure environmental safety, fungal agents are considered key biological regulators in the application of insecticides for pest control (Roy and Cottrell 2008; Sharma and Sharma 2021). The initial recommendation to use microbial insect pathogens for controlling insects was proposed by LeConte (1874) and Pasteur (1874). Subsequently, the first successful application of Metarhizium anisopliae was conducted in Russia against Bothynoderes punctiventris Academic editor: João Araújo Received: 27 August 2025 Accepted: 9 September 2025 Published: 8 October 2025 Citation: Yang C, Xu X, Li X, Liu F, Zeng Z, Xiao Q, Liu Y (2025) Morphological and phylogenetic analyses reveal novel entomopathogenic fungi infecting scale insects and aphids in China. IMA Fungus 16: e170123. https://doi.org/10.3897/ imafungus.16.170123 IMA Fungus 16: e170123 (2025) DOI: 10.3897/imafungus.16.170123 2 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China (Coleoptera: Curculionidae) by Krassilstschik (1888). This achievement built upon the earlier identification of the microbial agent, initially named Entomophthora anisopliae, which targeted Anisoplia austriaca (Coleoptera: Scarabaeidae). To date, more than 1,000 species of entomopathogenic fungi have been documented across over 100 genera (Hywel-Jones 1993; Vega et al. 2012; Hawksworth and Lücking 2017; Corallo et al. 2019; Wu et al. 2024). Among these species, such as Akanthomyces Lebert, Beauveria Vuill., Cordyceps Fr. (=Isaria Pers.), Lecanicillium W. Gams & Zare, and Metarhizium Sorokin, are widely recognized for their potential in biological control. Notably, species like A. aculeatus, B. bassiana, B. brongniartii, C. fumosorosea, L. lecanii (≡Verticillium lecanii), L. muscarium, L. longisporum, and M. anisopliae are commonly employed in pest management strategies (Faria and Wraight 2007; McKinnon et al. 2017; Corallo et al. 2019; Stone and Bidochka 2020; Nicoletti and Becchimanzi 2020; Nishi et al. 2021; Khonsanit et al. 2024). For example, 171 products have been developed as biocontrol agents, primarily based on B. bassiana, B. brongniartii, C. fumosorosea, and M. anisopliae (Faria and Wraight 2007). Beauveria bassiana, marketed as “Mycotrol”, has been utilized as a commercial mycoinsecticide to target a broad spectrum of insect pests in North America (Wraight et al. 2021). Similarly, the commercial product “Green Muscle,” derived from M. acridum (≡M. anisopliae var. acridum), has proven effective against locust and grasshopper pests in Africa (Niassy and Diarra 2011). In addition, entomopathogenic fungi exhibit diverse nutritional behaviors—functioning as biotrophs, necrotrophs, or hemibiotrophs (Vega et al. 2009)—and demonstrate additional ecological roles, including rhizosphere colonization, fungal endophytism, plant disease antagonism, and enhancement of plant growth or stress resistance (Kabaluk and Ericsson 2007; Kim et al. 2008; Ownley et al. 2008; Vega 2008; Pava-Ripoll et al. 2011; Akutse et al. 2013; Jaber and Enkerli 2017; Jaber and Ownley 2017; Mantzoukas and Eliopoulos 2020; Sharma and Sharma 2021; Muola et al. 2023). Insect-associated fungi have been widely reported across the extensive regions south of the Yellow River basin, with most species displaying distribution patterns characteristic of tropical, subtropical, and temperate zones (Liang et al. 2016; Chen et al. 2021; Wei et al. 2022; Xiao et al. 2023; Chuang et al. 2024; Wang et al. 2024, 2025), but have been rarely documented in the Sichuan region. The development of entomogenous fungi is closely linked to vegetation and insect populations, as well as climatic and soil conditions (Hajek and Leger 1994; Quesada-Moraga et al. 2007, 2024; Vega et al. 2009; Medo and Cagáň 2011), although experimental validation in this area remains limited. Scattered studies have indicated that entomogenous fungi are more commonly found in areas with rich vegetation, relatively high humidity, suitable temperatures, and shaded environments. Chuang et al. (2024) found that in Taiwan, China, entomogenous fungi belonging to the Cordycipitaceae are more readily collected in regions with an average temperature of 19–20.5 °C, rainfall ranging from 2,300 to 3,150 mm, and relative humidity between 81% and 82.5%. Wang et al. (2025) collected and described two new genera and 13 new species within the Clavicipitaceae, associated with whiteflies and scale insects on various host plants in Yunnan and Hainan, China. Sichuan is recognized as one of the key biodiversity research hotspots, owing to its diverse ecosystems, complex habitat conditions, and rich plant species. While the region supports a wide variety of insect fauna, entomogenous fungi have not been systematically studied, and their species diversity is likely 3 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China significantly underestimated. In our ongoing search for entomogenous fungi associated with miniature insects, several new taxa of entomopathogenic fungi (Nectriaceae; Podonectriaceae) were isolated from scale insects collected in Sichuan Province, primarily from bamboo and walnut hosts (Xu et al. 2021a; Liu et al. 2023). To further explore the diversity of entomogenous fungi, we conducted field collections at multiple sites across Sichuan. Detailed morphological descriptions are provided along with relevant ecological data. Comprehensive phylogenetic analyses were performed using a seven-locus dataset (ITS, LSU, SSU, tef1-α, rpb1, rpb2, and act) to ensure accurate taxonomic position. This study enriches the entomogenous fungi database in Sichuan and offers valuable strains for sustainable pest control in agriculture and forestry. Materials and methods Fungal collection and isolation Specimens consisting of whole leaves or branches bearing fruiting bodies were collected from Chengdu, Dazhou, Guangan, Leshan, and Meishan in Sichuan Province, China, placed in sterilized plastic bags, and transported to the laboratory for analysis. The procedure for obtaining axenic cultures, as described by Chomnunti et al. (2014), involved isolating pure cultures, transferring them to PDA slants for 10 days of cultivation, and subsequently storing them at 4 °C. The specimens were deposited in the Herbarium of Sichuan Agricultural University (SICAU), Chengdu, China, while the strains were stored in the Culture Collection of Sichuan Agricultural University (SICAUCC), China. Morphological observations Fruiting bodies were observed and photographed using a dissecting microscope NVTGG (Shanghai Advanced Photoelectric Technology Co. Ltd., Shanghai, China) fitted with a VS-800C micro-digital camera (Shenzhen Weishen Times Technology Co. Ltd., Shenzhen, China). The dimensions of conidiomata, paraphyses, conidiophores, conidiogenous cells, and conidia were measured from field samples and photographed using an Olympus BX43 compound microscope equipped with an Olympus DP22 digital camera, in conjunction with ACDSee v3.1 software. Measurements were made using Tarosoft® Image Frame Work v.0.9.7 (Tarosoft (R), Nontha Buri, Thailand). Lactophenol cotton blue reagent was used to observe the conidiogenous structures and determine the number of septa. To observe and document the color and texture of the colonies, fresh plates were prepared from purified colonies and incubated at 25 °C for 1 to 3 weeks. DNA extraction, amplification, and sequencing Total genomic DNA was extracted from mycelia grown on PDA or single fruiting bodies using the Plant Genomic DNA extraction kit (Tiangen, China). Primer pairs ITS5 and ITS4 (White et al. 1990), NS1 and NS4 (White et al. 1990), LR0R and LR5 (Vilgalys and Hester 1990), EF1-983F and EF1-2218R (Rehner and Buckley 2005), RPB1-Ac and RPB1-Cr (Lombard et al. 2015), fRPB2-5F and fRPB2-7cR (Liu et al. 2000), and ACT512F/ACT-783R (Carbone and Kohn 4 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China 1999) were used to amplify the internal transcribed spacer (ITS), the partial small subunit nuclear rDNA (SSU), the nuclear ribosomal large subunit (LSU), the translation elongation factor tef1-α, the largest subunit of RNA polymerase II (rpb1), the second largest subunit of RNA polymerase II (rpb2), and the actin gene (act), respectively. Additionally, for Cladosporium species, EF1728F and EF2 (Lombard et al. 2015) were used to amplify tef1-α. Polymerase chain reaction (PCR) was performed in a 25 μL reaction mixture containing 22 μL Master Mix (Beijing TsingKe Biotech Co. Ltd., Beijing, China), 1 μL DNA template (10–30 ng/μL), and 1 μL of each primer (120–150 ng/μL). The amplification reactions were performed as described by Lombard et al. (2015), Dai et al. (2016), and Wang et al. (2022a). PCR products were sequenced at TsingKe Biological Technology Co. Ltd., Chengdu, China. The newly generated sequences were deposited in GenBank. Phylogenetic analyses Phylogenetic analyses were conducted using sequences from Cladosporium (ITS, act, and tef1-α), Moelleriella (LSU, rpb1, and tef1-α), and Podonectria (ITS, LSU, SSU, tef1-α, and rpb2). Multigene sequences from various species (see Tables 1–3) were obtained from GenBank, along with additional sequences generated in this study. DNA alignments were carried out using the MAFFT v.7.429 online service (Katoh et al. 2019), and ambiguous regions were excluded using BioEdit version 7.0.5.3 (Hall 1999). Multigene sequences were concatenated using Mesquite software (Maddison and Maddison 2019). Multigene phylogenetic analyses were conducted using maximum likelihood (ML) and Bayesian inference (BI) methods. The optimal nucleotide substitution model was identified with MrModeltest v.2.2 (Nylander 2004). ML and BI analyses were performed using the CIPRES Science Gateway web server (Miller et al. 2010). For the ML analysis, RAxML-HPC2 on XSEDE (v.8.2.10) (Stamatakis 2014) was employed with the GTR+GAMMA substitution model and 1,000 bootstrap iterations. For BI analyses, the best-fit models were selected using MrModeltest v.2.2 for each dataset: Cladosporium (ITS: SYM+I+G, act: GTR+G, tef1-α: GTR+I+G), Moelleriella (LSU: GTR+I+G, rpb1: GTR+I+G, tef1-α: GTR+I+G), and Podonectriaceae (ITS: GTR+I+G, LSU: GTR+I+G, SSU: GTR+I+G, tef1-α: GTR+I+G, rpb2: GTR+I+G). The analyses were computed with six simultaneous Markov Chain Monte Carlo (MCMC) chains, run for 1,000,000 to 10,000,000 generations with a sampling frequency of every 100 generations. The burn-in fraction was set to 0.25, and the run was automatically terminated when the average standard deviation of split frequencies fell below 0.01. Phylogenetic trees were visualized using FigTree v.1.4.3 (Rambaut and Drummond 2016) and further edited with Adobe Illustrator CS6 (Adobe Systems Inc., United States). Maximum likelihood bootstrap values (MLBS) of 60% or greater and Bayesian posterior probabilities (BIPP) of 0.95 or higher were indicated on the trees. Abbreviations act The actin gene BI Bayesian inference 5 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China bp Base pair BIPP Bayesian inference posterior probabilities DNA Deoxyribonucleic acid ITS internal transcribed spacer LSU The nuclear ribosomal large subunit MEA Malt extract agar medium ML Maximum likelihood MLBS Maximum likelihood bootstrap proportions PCR Polymerase chain reaction PDA Potato dextrose agar medium rpb1 The largest subunits of RNA polymerase II rpb2 The RNA polymerase II second largest subunit SICAU The herbarium of Sichuan Agricultural University SICAUCC The culture collection in Sichuan Agricultural University SNA Synthetic low nutrient agar medium SSU The partial small subunit nuclear rDNA tef1-α The translation elongation factor 1α Table 1. Voucher information and GenBank accession numbers of the taxa used in the Cladosporium. Species Voucher information ITS act tef1-α References Cladosporium acalyphae CBS 125982 T HM147994 HM148481 HM148235 Bensch et al. (2010) Cl. alboflavescens CBS 140690 T LN834420 LN834604 LN834516 Sandoval-Denis et al. (2016) Cl. angulosum CBS 140692 T LN834425 LN834609 LN834521 Sandoval-Denis et al. (2016) Cl. angustisporum CBS 125983 T HM147995 HM148482 HM148236 Bensch et al. (2010) Cl. angustiterminale CBS 140480 T KT600379 KT600575 KT600476 Bensch et al. (2015) Cl. anthropophilum CBS 140685 T LN834437 LN834621 LN834533 Sandoval-Denis et al. (2016) Cl. arenosum CHFC-EA 566 T MN879328 MN890008 MN890011 Crous et al. (2020a) Cl. armandiae CBS 153756 T PQ066521 PQ067354 PQ067355 Tan et al. (2024) Cl. asperulatum CBS 126340 T HM147998 HM148485 HM148239 Bensch et al. (2010) Cl. aulonemiae COAD 2269 T MZ318427 MT373119 MT680198 Costa et al. (2022) Cl. aulonemiae COAD 2270 MZ318428 MT373120 MT680199 Costa et al. (2022) Cl. australiense CBS 125984 T HM147999 HM148486 HM148240 Bensch et al. (2010) Cl. austroafricanum CBS 140481 T KT600381 KT600577 KT600478 Bensch et al. (2015) Cl. austrolitorale CBS 148321 T MN879327 MN890007 MN890010 Crous et al. (2021a) Cl. bambusicola COAD 2256 T MZ318433 MT373125 MT680204 Costa et al. (2022) Cl. bambusicola COAD 2565 OP535372 OP598124 OP676083 Costa et al. (2022) Cl. benschiae COAD 2263 T MZ318436 MT373128 MT680207 Costa et al. (2022) Cl. benschiae COAD 2265 MZ318437 MT373129 MT680208 Costa et al. (2022) Cl. bentivoglioae BRIP 74745a T OR947065 OR964960 OR964961 Tan and Shivas (2023a) Cl. brigadeirense COAD 2257 T MZ318435 MT373127 MT680206 Costa et al. (2022) Cl. caprifimosum FMR 16532 T LR813198 LR813205 LR813210 Iturrieta-González et al. (2021) Cl. cavernicola URM 8389 T MZ518829 MZ555746 MZ555733 Pereira et al. (2022) Cl. chalastosporoides CBS 125985 HM148001 HM148488 HM148242 Bensch et al. (2010) Cl. chasmanthicola CPC 21300 T KY646221 KY646224 KY646227 Marin-Felix et al. (2017) Cl. chlamydosporigenum AUMC 11340 T MN826919 OL514009 Moharram et al. (2022) Cl. chlamydosporiformans COAD 2571 T OP535374 OP598126 OP676085 Pereira et al. (2024) Cl. chlamydosporiformans COAD 2568 OP535378 OP598130 OP676089 Pereira et al. (2024) 6 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Species Voucher information ITS act tef1-α References Cl. chusqueae COAD 2258 T MZ318430 MT373122 MT680201 Costa et al. (2022) Cl. chusqueae COAD 2261 MZ318431 MT373124 MT680203 Costa et al. (2022) Cl. chubutense CBS 124457 T FJ936158 FJ936165 FJ936161 Schubert et al. (2009) Cl. cladosporioides CBS 112388 T HM148003 HM148490 HM148244 Bensch et al. (2010) Cl. compactisporum AUMC 11366 T MN826822 OL514010 Moharram et al. (2022) Cl. colocasiae CBS 386.64 T HM148067 HM148555 HM148310 Bensch et al. (2010) Cl. colombiae CBS 274.80B T FJ936159 FJ936166 FJ936163 Schubert et al. (2009) Cl. congjiangense GUCC 21208.3 T OP852675 OP863094 OP859042 Yang et al. (2023) Cl. congjiangense GUCC 21208.5 OP852676 OP863095 OP859043 Yang et al. (2023) Cl. coprophilum FMR 16164 T LR813201 LR813207 LR813213 Iturrieta-González et al. (2021) Cl. corticola BRIP 74385a T OP256851 OP288997 OP288998 Crous et al. (2023) Cl. crousii CBS 140686 T LN834431 LN834615 LN834527 Sandoval-Denis et al. (2016) Cl. cucumerinum CBS 171.52 T HM148072 HM148561 HM148316 Bensch et al. (2010) Cl. delicatulum CBS 126344 HM148081 HM148570 HM148325 Bensch et al. (2010) Cl. devikae BRIP 72278a T MZ303808 MZ344212 MZ344193 Prasannath et al. (2021a, 2021b) Cl. diamantinense COAD 3108 T ON062328 ON141933 ON982817 Dutra et al. (2023) Cl. endoviticola JZBH 390018 T MN654960 MN984220 MN984228 Manawasinghe et al. (2020) Cl. eucommiae GUCC 401.1 T OL587465 OL519775 OL504966 Wang et al. (2022b) Cl. eucommiae GUCC 401.9 ON334729 ON383337 Wang et al. (2022b) Cl. europaeum CBS 134914 T HM148056 HM148543 HM148298 Bensch et al. (2018) Cl. europaeum CBS 116744 HM148053 HM148540 HM148294 Bensch et al. (2018) Cl. exasperatum CBS 125986 HM148090 HM148579 HM148334 Bensch et al. (2010) Cl. exile CBS 125987 HM148091 HM148580 HM148335 Bensch et al. (2010) Cl. flabelliforme CBS 126345 HM148092 HM148581 HM148336 Cl. flavovirens CBS 140462 T LN834440 LN834624 LN834536 Sandoval-Denis et al. (2016) Cl. funiculosum CBS 122129 T HM148094 HM148583 HM148338 Bensch et al. (2010) Cl. fuscoviride FMR 16385 T LR813200 LR813206 LR813212 Iturrieta-González et al. (2021) Cl. gamsianum CBS 125989 T HM148095 HM148584 HM148339 Bensch et al. (2010) Cl. globisporum CBS 812.96 T HM148096 HM148585 HM148340 Crous et al. (2011) Cl. grevilleae CBS 114271 T JF770450 JF770473 JF770472 Crous et al. (2011) Cl. guizhouense GUCC 401.7 T OL579741 OL519780 OL504965 Wang et al. (2022b) Cl. guizhouense GUCC 401.8 ON334728 ON383338 ON383470 Wang et al. (2022b) Cl. guizhouense SICAUCC 25-0057 PV156487 PV153466 PV153474 In this study Cl. guizhouense SICAUCC 25-0058 PV156488 PV153467 PV153475 In this study Cl. hemileiicola COAD 2567 T OP535376 OP598128 OP676087 Pereira et al. (2024) Cl. hemileiicola COAD 3350 OP535381 OP598133 OP676092 Pereira et al. (2024) Cl. heteropogonicola BRIP 72465a T OL307932 OL332743 OL332742 Tan et al. (2022) Cl. hillianum CBS 125988 T HM148097 HM148586 HM148341 Bensch et al. (2010) Cl. inversicolor CBS 401.80 T HM148101 HM148590 HM148345 Bensch et al. (2010) Cl. ipereniae CBS 140483 T KT600394 KT600589 KT600491 Bensch et al. (2015) Cl. iranicum CBS 126346 T HM148110 HM148599 HM148354 Bensch et al. (2010) Cl. kaiyangense GUCC 21265.2 T OP852665 OP863097 OP859045 Yang et al. (2023) Cl. kenpeggii CPC 19248 T KY646222 KY646225 KY646228 Marin-Felix et al. (2017) Cl. kuwanaspidis SICAUCC 25-0063 T PV156489 PV153468 PV153480 In this study Cl. kuwanaspidis SICAUCC 25-0064 PV156490 PV153469 PV153481 In this study Cl. lagenariiforme SFC20230103-M23 T OQ186119 OQ185167 OQ185128 Lee et al. (2023) 7 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Species Voucher information ITS act tef1-α References Cl. lentulum FMR 16288 T LR813203 LR813209 LR813215 Iturrieta-González et al. (2021) Cl. licheniphilum CBS 125990 HM148111 HM148600 HM148355 Bensch et al. (2010) Cl. longicatenatum CBS 140485 KT600403 KT600598 KT600500 Bensch et al. (2015) Cl. longissimum CBS 300.96 DQ780352 EF101385 EU570259 Cl. lycoperdinum CBS 574.78C HM148115 HM148604 HM148359 Prasannath et al. (2021a, 2021b) Cl. macadamiae BRIP 72269a T MZ303810 MZ344214 MZ344195 Jayasiri et al. (2019) Cl. magnoliigena MFLUCC 18-1559 T MK347813 MK340864 Lee et al. (2023) Cl. maltirimosum SFC20230103-M51 T OQ186147 OQ185195 OQ185155 Lee et al. (2023) Cl. marinum SFC20230103-M33 T OQ186129 OQ185177 OQ185137 Bensch et al. (2015) Cl. montecillanum CBS 140486 T KT600406 KT600602 KT600504 Bensch et al. (2010) Cl. myrtacearum CBS 126350 T HM148117 HM148606 HM148361 Yang et al. (2023) Cl. nayongense GUCC 21260.3 T OP852669 OP863106 OP859054 Zimowska et al. (2021) Cl. neapolitanum MgPo1 T MK387890 MK416051 MK416094 Bensch et al. (2018) Cl. needhamense CBS 143359 T MF473142 MF473991 MF473570 Bensch et al. (2018) Cl. neerlandicum CBS 143360 T KP701887 KP702010 KP701764 Bensch et al. (2018) Cl. neopsychrotolerans CGMCC 3.18031 T KX938383 KX938366 KX938400 Bensch et al. (2010) Cl. oxysporum CBS 125991 HM148118 HM148607 HM148362 Bensch et al. (2010) Cl. paracladosporioides CBS 171.54 T HM148120 HM148609 HM148364 Bensch et al. (2015) Cl. parapenidielloides CBS 140487 T KT600410 KT600606 KT600508 Rosado et al. (2019) Cl. passiflorae COAD 2135 T MH682175 MH729795 MH724943 Rosado et al. (2019) Cl. passifloricola COAD 2140 T MH729800 MH724948 Bensch et al. (2010) Cl. perangustum CBS 125996 T HM148121 HM148610 HM148365 Bensch et al. (2018) Cl. perangustum CBS 126365 HM148123 HM148612 HM148367 Bensch et al. (2018) Cl. perangustum CPC 11663 HM148128 HM148617 HM148372 Sandoval-Denis et al. (2016) Cl. perangustum CPC 13870 HM148142 HM148631 HM148386 Bensch et al. (2010) Cl. perangustum FMR 13321 LN834380 LN834564 LN834476 Sandoval-Denis et al. (2016) Cl. pernambucoense URM 8390 T MZ518830 MZ555747 MZ555734 Pereira et al. (2022) Cl. pernambucoense URM 8391 MZ518828 MZ555745 MZ555732 Pereira et al. (2022) Cl. phaenocomae CBS 128769 JF499837 JF499881 JF499875 Crous and Groenewald (2011) Cl. phyllactiniicola CBS 126352 HM148150 HM148639 HM148394 Cl. phyllophilum CBS 125992 T HM148154 HM148643 HM148398 Bensch et al. (2010) Cl. pini-ponderosae CBS 124456 T FJ936160 FJ936167 FJ936164 Schubert et al. (2009) Cl. polonicum MgPo1 T MK387894 MK416055 MK416098 Zimowska et al. (2021) Cl. proteacearum BRIP 72301a T MZ303809 MZ344213 MZ344194 Prasannath et al. (2021a, 2021b) Cl. pruni-salicinae GUCC 21206.1 T OP852683 OP863092 OP859041 Yang et al. (2023) Cl. pseudocladosporioides CBS 125993 T HM148158 HM148647 HM148402 Bensch et al. (2010) Cl. pseudochalastoporoides CBS 140490 T KT600415 KT600611 KT600513 Bensch et al. (2015) Cl. pseudotenuissimum COAD 2266 T MZ318439 MT373132 MT680211 Costa et al. (2022) Cl. punicae GUCC 21271.5 T OP852672 OP863108 OP859056 Yang et al. (2023) Cl. puris COAD 2487 T MK253337 MK249980 MK293777 Freitas et al. (2021) Cl. queenslandicum BRIP 72447a T OL307928 OL332736 OL332735 Tan et al. (2022) Cl. queenslandicum BRIP 72452a OL307929 OL332738 OL332737 Tan et al. (2022) Cl. queenslandicum BRIP 72455a OL307931 OL332741 OL332740 Tan et al. (2022) Cl. rectoides CBS 125994 T HM148193 HM148683 HM148438 Bensch et al. (2010) Cl. ribis GUCC 21244.1 T OP852666 OP863098 OP859046 Yang et al. (2023) Cl. rubrum CMG 28 T MN053018 MN066639 MN066644 Vicente et al. (2021) 8 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Species Voucher information ITS act tef1-α References Cl. rugulovarians CBS 140495 T KT600459 KT600656 KT600558 Bensch et al. (2015) Cl. ruthsangerae BRIP 75808a T OR290122 OR335736 OR335741 Tan and Shivas (2023b) Cl. scabrellum CBS 126358 T HM148195 HM148685 HM148440 Bensch et al. (2010) Cl. setoides COAD 3470 T OP535379 OP598131 OP676090 Pereira et al. (2024) Cl. setoides COAD 2576 OP535380 OP598132 OP676091 Pereira et al. (2024) Cl. silenes CBS 109082 T EF679354 EF679506 EF679429 Schubert et al. (2007) Cl. sinuatum CGMCC 3.18096 T KX938385 KX938368 KX938402 Bensch et al. (2018) Cl. speluncae COAD 3116 T ON062329 ON141934 ON982818 Dutra et al. (2023) Cl. splattiae BRIP 75807a OR290120 OR335734 OR335739 Tan and Shivas (2023b) Cl. sphaerospermum CBS 193.54 DQ780343 EF101380 EU570261 Crous et al. (2021b) Cl. stipagrostidicola CBS 146978 T MZ064420 MZ078146 MZ078223 Bensch et al. (2010) Cl. subuliforme CBS 126500 T HM148196 HM148686 HM148441 Bensch et al. (2010) Cl. tenuissimum CBS 125995 T HM148197 HM148687 HM148442 Bensch et al. (2018) Cl. tianshanense CGMCC 3.18033 T KX938381 KX938364 KX938398 Braun et al. (2003); Bensch et al. (2010) Cl. uredinicola ACC 46649 AY251071 HM148712 HM148467 Bensch et al. (2018) Cl. uwebraunianum CBS 143365 T MF473306 MF474156 MF473729 Bensch et al. (2010) Cl. varians CBS 126362 T HM148224 HM148715 HM148470 Bensch et al. (2010) Cl. verrucocladosporioides CBS 126363 T HM148226 HM148717 HM148472 Bensch et al. (2018) Cl. vicinum CBS 143366 T MF473311 MF474161 MF473734 Bensch et al. (2018) Cl. vignae CBS 121.25 HM148227 HM148718 HM148473 Marin-Felix et al. (2017) Cl. welwitschiicola CPC 18648 T KY646223 KY646226 KY646229 Yang et al. (2023) Cl. wenganense GUCC 21220.1 T OP852682 OP863101 OP859049 Bensch et al. (2018) Cl. westerdijkiae CBS 113746 T HM148061 HM148548 HM148303 Sandoval-Denis et al. (2016) Cl. xanthochromaticum CBS 140691 T LN834415 LN834599 LN834511 Bensch et al. (2010) Cl. xylophilum CBS 125997 T HM148230 HM148721 HM148476 Xu et al. (2021b) Cl. yunnanensis KUN-HKAS 121704 T OK338502 OL466937 OL825680 Zalar et al. (2007); Dugan et al. (2008) Cl. longissimum CBS 300.96 T DQ780352 EF101385 EU570259 Zalar et al. (2007); Dugan et al. (2008) T Type material. New species were shown in bold. Table 2. Voucher information and GenBank accession numbers of the taxa used in the Moelleriella. Species Voucher information LSU rpb1tef1-α References Hypocrella citrina P.C. 606 EU392556 EU392640 EU392694 Chaverri et al. (2008) H. citrina P.C. 597 AY986905 AY986930 DQ000331 Chaverri et al. (2005a) H. discoidea BCC2097 AY986945 DQ000346 Chaverri et al. (2005a) H. discoidea I93-901d EU392567 EU392646 EU392700 Chaverri et al. (2008) H. disciformis P.C. 655 EU392560 EU392643 EU392697 Chaverri et al. (2008) H. disciformis P.C. 676 EU392566 EU392645 EU392699 Chaverri et al. (2008) H. hirsuta P.C. 436.2 AY986922 AY986949 DQ000350 Chaverri et al. (2005a) H. hirsuta P.C. 543 EU392569 EU392648 EU392702 Chaverri et al. (2008) H. viridans P.C. 635 EU392572 EU392651 EU392705 Chaverri et al. (2008) H. viridans I89-490 EU392570 EU392649 EU392703 Chaverri et al. (2008) Moelleriella africana P.C. 736 AY986917 AY986943 DQ000344 Chaverri et al. (2005a) M. alba BCC49492 JQ269645 KX254424 JQ256905 Mongkolsamrit et al. (2015) M. alba BCC49409 T JQ269646 KX254423 JQ256906 Mongkolsamrit et al. (2015) M. basicystis P.C. 374 AY986903 AY986928 DQ000329 Chaverri et al. (2005a) 9 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Species Voucher information LSU rpb1tef1-α References M. basicystis F183147 EU392577 EU392653 Chaverri et al. (2008) M. boehmeriae SICAUCC 25-0061 T PV124784 PV153478 PV153492 In this study M. boehmeriae SICAUCC 25-0062 PV124785 PV153479 PV153493 In this study M. boliviensis P.C. 603 AY986923 AY986950 DQ000351 Chaverri et al. (2005a) M. chiangmaiensis BCC18029 T MT659360 MW091560 Khonsanit et al. (2021) M. chiangmaiensis BBH33051 MT659362 MT672277 MT672269 Khonsanit et al. (2021) M. chiangmaiensis BCC60941 MT659361 MT672278 MT672270 Khonsanit et al. (2021) M. chumphoensis BCC47575 JQ269648 KX254422 JQ256908 Mongkolsamrit et al. (2015) M. chumphoensis BCC47574 T JQ269647 KX254421 JQ256907 Mongkolsamrit et al. (2015) M. cinnamomum SICAUCC 25-0067 T PV124788 PV153484 PV153494 In this study M. cinnamomum SICAUCC 25-0068 PV124789 PV153485 PV153495 In this study M. citrus SICAUCC 25-0059 T PV124782 PV153476 PV153490 In this study M. citrus SICAUCC 25-0060 PV124783 PV153477 PV153491 In this study M. disjuncta J.B. 205 EU392578 EU392654 Chaverri et al. (2008) M. epiphylla P.C. 545 EU392585 EU392660 EU392711 Chaverri et al. (2008) M. epiphylla I93-813 EU392583 EU392656 EU392707 Chaverri et al. (2008) M. eucalypti SICAU 25-0072 T PV124778 PV153470 PV153486 In this study M. eucalypti SICAU 25-0073 PV124779 PV153471 PV153487 In this study M. eucalypti SICAU 25-0074 PV124780 PV153472 PV153488 In this study M. eucalypti SICAU 25-0075 PV124781 PV153473 PV153489 In this study M. evansii P.C. 627 T AY986916 AY986942 DQ000343 Chaverri et al. (2005a) M. flava BCC60924 T KF951146 KX254430 MT672271 Khonsanit et al. (2021) M. flava BCC60925 KF951147 KX254431 MT672272 Khonsanit et al. (2021) M. flava BCC60929 KX298238 KX254432 MT672273 Khonsanit et al. (2021) M. flava BCC60930 KX298237 KX254429 Khonsanit et al. (2021) M. globostromata YFCC 22109275 T OR828408 OR831942 OR831952 Wang et al. (2025) M. globostromata YHH 221010 OR828403 OR831940 OR831950 Wang et al. (2025) M. gracilispora CGMCC 3.18989 T KC964202 KC964191 KC964179 Yuan et al. (2020) M. gracilispora CGMCC 3.18990 KC964203 KC964192 KC964180 Yuan et al. (2020) M. hainanensis YHH 2303020 OR828400 OR831938 OR831948 Wang et al. (2025) M. hainanensis YFCC 23039277 T OR831939 OR831949 Wang et al. (2025) M. insperata ARSEF 2396 T AY518374 DQ070029 EU392713 Chaverri et al. (2008) M. jinghongensis YFCC 23089312 T OR854253 OR837093 Wang et al. (2025) M. jinghongensis YHH 2308025 OR828411 OR854254 OR837094 Wang et al. (2025) M. jinuoana YHH MJBP2309031 T PP178643 PP776170 PP776160 Wang et al. (2024) M. jinuoana YHH MJBP2309032 PP178644 PP776171 PP776161 Wang et al. (2024) M. jinuoana YFCC MJBP23099451 PP178645 PP776172 PP776162 Wang et al. (2024) M. kanchanaburiensis BCC75979 MT659363 MT672279 MT843900 Khonsanit et al. (2021) M. kanchanaburiensis BCC75980 MT659364 MT672280 MT843901 Khonsanit et al. (2021) M. kanchanaburiensis BCC75981 T MT659365 MT672281 Khonsanit et al. (2021) M. libera P.C. 444 EU392591 EU392662 EU392714 Chaverri et al. (2008) M. libera P.C. 445 AY986900 AY986925 DQ000326 Chaverri et al. (2005a) M. longzhuensis YHH MLFSL2310012 T PP178646 PP776173 PP776163 Wang et al. (2024) M. longzhuensis YHH MLFSL2310013 PP178647 PP776174 PP776164 Wang et al. (2024) M. longzhuensis YFCC-MLFSL23109453 PP776175 PP776165 Wang et al. (2024) M. macrostroma P.C. 605 T AY986919 AY986946 DQ000347 Chaverri et al. (2005a) 16 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Figure 2. Phylogenetic tree based on maximum likelihood analysis of the combined LSU, rpb1, and tef1-α sequences within the Clavicipitaceae. Numbers above the nodes indicate MLBS values (≥ 60%, left) and BIPP values (≥ 0.95, right), with lower values denoted as “–”. The tree is rooted to Purpureocillium lilacinum (CBS 431.87, CBS 284.36). Ex-type strains are highlighted in bold, and isolates from the present study are shown in red. 17 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Figure 3. Phylogenetic tree based on maximum likelihood analysis of the combined ITS, LSU, SSU, tef1-α, and rpb2 sequences within the Pleosporales. Numbers above the nodes represent MLBS values (≥ 60%, left) and BIPP values (≥ 0.95, right), with lower values denoted as “–”. The tree is rooted to Tubeufia javanica (MFLUCC 12-0545) and T. chiangmaiensis (MFLUCC 11-0514). Ex-type strains are highlighted in bold, and isolates from the present study are shown in red. 18 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Taxonomy Cladosporium kuwanaspidis X.L. Xu & C.L. Yang, sp. nov. MycoBank No: MB 858369 Fig. 4 Etymology. In reference to the generic name for the associated scale insect (Kuwanaspis howardi). Diagnosis. Similar to Cladosporium perangustum in having conidiophores of comparable size, but C. kuwanaspidis differs by its unbranched conidiophores and larger ramoconidia. Type. CHINA • Sichuan Province, Meishan City, Hongya County. Infected scale insects (Kuwanaspis howardi) were found on the culms of bamboo (Pleioblastus amarus (Keng) P. C. Keng), 29°41.88'N, 103°14.04'E, alt. 540 m, 13 Mar. 2021, C.L. Yang, YCL202103004 (SICAU 25-0082 – holotype preserved in the Herbarium of Sichuan Agricultural University; living culture SICAUCC 25-0063 – ex-holotype stored in the Culture Collection in Sichuan Agricultural University). Description. Parasitic on scale insect from Pleioblastus amarus (Poaceae). Sexual morph: Not observed. Asexual morph: Hyphomycetous. Mycelium superficial and immersed, with abundant sporulation on the surface of scale insect. Conidiophores erect, fasciculate, usually macronematous, cylindrical, subnodulose or nodulose, geniculate, septate, unbranched, pale brown to brown, slightly roughened to verruculose, thick-walled, and pronounced loci, 45–120 × 3.5–6 μm. Conidiogenous cells terminal or intercalary, cylindrical, sometimes sinuous, proliferation sympodial, 14–25(–80) × 2.5–6.5 μm, conidiogenous loci at the apex (2–5) or in lateral shoulders (0–2). Ramoconidia olive to brown, septate or aseptate, ellipsoidal to subcylindrical, smoothand thickwalled, 6–13.5 × 3–5.5 μm. Secondary ramoconidia oblong, pale brown, 0–1 septate, 2–4 distal hila, 5.5–9 × 2.5–5 μm. Conidia numerous, catenate, forming short branched chains, aseptate, olive to brown, smoothand thinwalled, ellipsoid-ovoid, obovoid, 2–7.5 × 2–4 μm. Intercalary conidia limoniform, oval to ellipsoid, with hila protuberant, 3.5–7.5 × 2.5–4 μm. Terminal conidia globose to ellipsoid, sometimes hila evident, 2–4 × 2–3.5 μm. Microcyclic conidiogenesis absent. In vitro on SNA: Mycelium abundant, submerged, overgrowing whole culture dishes, hyphae straight to slightly sinuous, septate and branched, olive to brown, and thick-walled, 2–3.5 µm wide. Conidiophores erect, occasionally branched, brown, thick-walled, 33–118 × 2.5–4 µm. Ramoconidia olive to brown, narrowly ellipsoid to cylindrical-oblong, subcylindrical, septate or aseptate, smoothand thick-walled, 6–17.5 (–20) × 2–5 µm. Conidia in simple and branched acropetal chains, light olive, aseptate, smoothand thin-walled, ellipsoidovoid, obovoid, fusiform, 3–6.5 × 2–4 µm. Intercalary conidia oval to ellipsoid, with hila protuberant, 3.5–6.5 × 2.5–4 μm. Terminal conidia globose to ellipsoid, 2.5–4 × 2–4 μm. Culture characteristics. Conidia of all types can germinate within 12 h in sterile distilled water, with germination tubes forming from any part of the conidial body. Colonies on MEA attaining 40–50 mm diameter, after 1 week at 25 °C, greyish-green to greyish toward regular margins, velvety, sporulation profuse, radially furrowed, aerial mycelium abundant, reverse dark green. Colonies 19 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China on PDA attaining 35–40 mm diameter, after 1 week at 25 °C, grey-olivaceous to iron grey, pale greenish grey toward regular margins, reverse dark green. Colonies on SNA attaining 20–30 mm diameter, after 1 week at 25 °C, smoke-grey to olivaceous-grey, aerial mycelium loose, reverse olivaceous to brown. Host. Kuwanaspis howardi (Diaspididae). Habitat. Subtropical evergreen broad-leaved forests, particularly bamboo forests, are common habitats. Kuwanaspis howardi often causes damage in the lower canopy of bamboo forests. Occasionally, Cladosporium kuwanaspidis can be observed on bamboo culms. Scale insects are more easily found in shaded and sheltered areas. Distribution. China, Sichuan Province, Meishan City. Figure 4. Cladosporium kuwanaspidis. A–C. Symptoms and appearance of colonies observed on Kuwanaspis howardi. D–G. Conidiophores. H. Ramoconidia (white arrows), secondary ramoconidia (red arrows), and terminal conidia (yellow arrows). I–K. Colonies on MEA, PDA, and SNA for 7 days. Scale bars: 200 µm (B, C); 10 µm (D–H). 20 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Material examined. CHINA • Sichuan Province, Meishan City, Hongya County. Infected scale insects (Kuwanaspis howardi) were found on the culms of bamboo (Pleioblastus amarus), 29°41.88'N, 103°14.04'E, alt. 540 m, 9 Mar. 2021, C.L. Yang, YCL202103004 (living culture SICAUCC 25-0063); • ibid. YCL202103004-1 (SICAU 25-0083, living culture SICAUCC 25-0064). Notes. The ITS base-pair comparison between Cladosporium kuwanaspidis (SICAUCC 25-0063) and the phylogenetically affiliated ex-type culture of C. perangustum (CBS 125996) shows no nucleotide differences. However, nucleotide differences in the act and tef1-α regions are 1.86% (4 bp, 0 gap) and 4.01% (15 bp, 2 gaps), respectively. Morphologically, C. perangustum has larger ramoconidia compared to those observed in C. kuwanaspidis on both the host and SNA medium (25–45 µm vs. 6–13.5 µm and 6–20 µm, respectively). In addition, compared to the length of intercalary conidia (4–19 µm) observed in C. perangustum (CBS 125996), our isolates exhibit shorter intercalary conidia on both the host and PDA medium (3.5–7.5 µm and 3.5–6.5 µm, respectively) (Bensch et al. 2010). Cladosporium guizhouense S.Y. Wang, Yong Wang bis & Y. Li, MycoKeys 91: 151–168 (2022) MycoBank No: MB 842407 Fig. 5 Description. Parasitic on aphids from Telosma cordata (Burm. f.) Merr. (Apocynaceae). Sexual morph: Not observed. Asexual morph: Hyphomycetous. Mycelium superficial and immersed, with abundant sporulation on the surface of aphids. Conidiophores erect, fasciculate, macronematous, cylindrical, subnodulose or nodulose, geniculate, septate, branched, pale brown, slightly roughened to verruculose, thick-walled, and pronounced loci, 111–367 × 3.5–5 μm. Conidiogenous cells integrated, terminal or intercalary, cylindrical, cylindrical-oblong, sometimes geniculate, 20–65 × 3–6 μm, conidiogenous loci at the apex (2–5) or in lateral shoulders (0–4). Ramoconidia olive-green, 0–4-septate, ellipsoidal to subcylindrical, smoothand thick-walled, 10–26 × 3.5–7.5 μm. Secondary ramoconidia pale brown, oblong, oblong-ellipsoid, 0–1septate, 1–4 distal hila, 8.5–13.5 × 3.5–7.5 μm. Conidia numerous, catenate, forming short branched chains, aseptate, olive to light olive, smoothand thin-walled, variable in size and shape, ellipsoid-ovoid, obovoid, and fusiform, 4–10 × 3–6.5 μm. Intercalary conidia aseptate, olive to light olive, ellipsoid-ovoid, fusiform, 6–10 × 4–6.5 μm. Terminal conidia aseptate, olive to light olive, obovoid, 4–6.5 × 3–5.5 μm. Microcyclic conidiogenesis absent. In vitro on SNA: Mycelium abundant, submerged, overgrowing whole culture dishes, hyphae straight to slightly sinuous, septate and branched, light olive-green to olive-green, and thick-walled, 1.5–4.5 µm wide. Conidiophores erect, branched, light olive-green, thick-walled, 43–163 × 2.5–4 µm. Ramoconidia pale olivegreen, narrowly ellipsoid to cylindrical-oblong, subcylindrical, septate or aseptate, smoothand thick-walled, 6–27 × 3–5 µm. Conidia in simple and branched acropetal chains, light olive, aseptate, smoothand thin-walled, variable in size and shape, ellipsoid-ovoid, obovoid, fusiform, 3–8 × 2.5–4 µm. Intercalary conidia light olive, aseptate, smoothand thin-walled, mostly ellipsoid, 3.5–8 × 2.5–4 µm. Terminal conidia light olive, aseptate, smoothand thin-walled, mostly ovoid, 3–4.5 × 2.5–3.5 µm. 21 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Culture characteristics. All conidia can germinate within 12 h in sterile distilled water, with tubes forming from any part. Colonies on MEA attaining 35–45 mm diameter, after 1 week at 25 °C, dark green, white to greyish toward regular margins, velvety, sporulation profuse, radially furrowed, aerial mycelium abundant, reverse dark green. Colonies on PDA attaining 30–40 mm diameter, after 1 week at 25 °C, greyishgreen, greyish toward irregular margins, velvety, sporulation profuse, radially furrowed, aerial mycelium abundant, reverse dark green. Colonies on OA attaining 30–40 mm diameter, after 1 week at 25 °C, greyish olivaceous, white toward irregular margins, sporulation profuse, reverse olivaceous. Colonies on SNA attaining 25–30 mm diameter, after 1 week at 25 °C, olivaceous, flat, white toward regular and wide margins, aerial mycelium loose, reverse olivaceous. Host. Aphids (Aphididae), leaves of plants, and uredinia of Hemileia vastatrix (Pucciniaceae). Figure 5. Cladosporium guizhouense. A. Symptoms observed on host. B, C. Appearance of colonies on aphids. D–H. Conidiophores. I. Ramoconidia (white arrows), secondary ramoconidia (red arrows), and terminal conidia (yellow arrows). J–M. Colonies on MEA, PDA, OA, and SNA for 7 days. Scale bars: 1000 µm (B); 200 µm (C); 20 µm (D–I). 22 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Habitat. In tropical and subtropical regions, it can infect insects, act saprophytically on plant tissues, and function as a fungicolous fungus on the uredinia of rust. During the middle and late stages of aphid damage, numerous aphids die on the underside of plant leaves, becoming enveloped in mycelial tissue. Distribution. China, Sichuan Province, Guangan City. China, Guizhou Province, Guiyang City. Ethiopia, Oromia Region, Illubabor Gore. Brazil, Minas Gerais, Viçosa. Material examined. CHINA • Sichuan Province, Guangan City, Yuechi County. Infected aphids were found on the underside of the leaves of Telosma cordata, 30°44.18"N, 106°30.80"E, alt. 480 m, 31 Jan. 2020, X.L. Xu, XXL202001003 (SICAU 25-0076, living culture SICAUCC 25-0057); • ibid. XXL202001003-1 (SICAU 25-0077, living culture SICAUCC 25-0058). Notes. Phylogenetically, our collections grouped with isolates of Cladosporium guizhouense within the C. cladosporioides complex, showing strong statistical support (92% MLBS, 0.97 BIPP) (Fig. 1). A total of 6 bp nucleotide differences were observed between our isolate SICAUCC 25-0057 and the ex-type culture of C. guizhouense (GUCC 401.7), with 0 bp in ITS, 2 bp in act, and 4 bp in tef1-α (Wang et al. 2022b). Morphologically, our isolate is fully consistent with C. guizhouense, showing no noticeable differences (Wang et al. 2022b; Pereira et al. 2024). Moelleriella eucalypti X.L. Xu, Feng Liu & C.L. Yang, sp. nov. MycoBank No: MB 858370 Fig. 6 Etymology. In reference to the generic name of host plants. Diagnosis. Similar to Moelleriella sinensis in having somewhat similar stromata, M. eucalypti differs by having longer and wider paraphyses, shorter and wider conidiogenous cells, and wider conidia. Type. CHINA • Sichuan Province, Dazhou City, Heibaoshan National Forest Park. Infected scale insects were found on the underside of leaves of Eucalyptus sp. (Myrtaceae), 31°55.30'N, 107°47.75'E, alt. 690 m, 21 Jul. 2022, Feng Liu, LF202207001A (SICAU 25-0072 – holotype preserved in the Herbarium of Sichuan Agricultural University). Description. Parasitic on scale insects found on eucalyptus leaves. Sexual morph: Not observed. Asexual morph: Coelomycetous. Stromata yellow to dark orange in fresh specimens, pale yellow to white in old, entirely covering the insect hosts, thickened pulvinate, umbonate to hemispheric, cottony, sessile, globose to subglobose, tubercules on the surface, 1.5–2.5 mm in diameter, 1–2 mm in high. Conidiomata orifice scattered or circularly arranged, oval or elongate flask shaped, narrow orifices, 120–460 × 80–155 μm. Phialides hyaline, stick-shaped to cylindrical, 10–18 × 1–2.5 μm. Conidia hyaline, fusiform, yellow conidial masses, 8–12 × 1.5–3 μm. Paraphyses present, hyaline, filiform, 90–180 × 1.0–1.6 μm. Culture characteristics. No germination was observed due to the specimens being dried and the culture not being obtained. Host. Scale insects (Coccidae). Habitat. Subtropical monsoon evergreen broad-leaf forest. Scale insects were infected on the underside of eucalyptus leaves. Distribution. China, Sichuan Province, Dazhou City. 23 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China Material examined. CHINA • Sichuan Province, Dazhou City, Heibaoshan National Forest Park. Infected scale insects were found on the underside of leaves of Eucalyptus sp., 31°55.30'N, 107°47.75'E, alt. 690 m, 21 Jul. 2022, Feng Liu, LF202207001A (SICAU 25-0072); • ibid. LF202207001B (SICAU 25-0073); • ibid. LF202207001C (SICAU 25-0074); • ibid. LF202207001D (SICAU 25-0075). Notes. Phylogenetically, our collections are closely related to Moelleriella sinensis, showing few sequence differences with the type strain CGMCC3.18911, specifically 1 bp in LSU (0.12%, 0 gap) and 8 bp in rpb1 (1.10%, 0 gap), while the tef1-α sequence is invalid. Moelleriella sinensis was first described on whitefly nymphs (Hemiptera) and later found on scale insects (Coccidae) in Thailand (Chen et al. 2020; Khonsanit et al. 2021). The described specimens of M. sinensis are primarily characterized by flat to umbonate, pale yellow pulvinate stromata and fusiform conidia. However, our collections differ from M. sinensis in having pulvinate stromata that are yellow to dark orange when fresh, pale yellow to white when old, and thicker in appearance. Additionally, they have longer and wider paraphyses (90–180 × 1.0–1.6 μm vs. 43.2–68.9 × 0.6–0.8 μm), shorter and wider conidiogenous cells (10–18 × 1–2.5 μm vs. up Figure 6. Moelleriella eucalypti. A. Fungus-infected scale insect on the underside of leaves. B–D. Stromata containing conidiomata with conidial masses. E–I. Section of stroma showing conidioma. J–M. Phialides and conidia at the tips with paraphyses. N. Conidia. Scale bars: 500 mm (B–D); 200 μm (E, F); 50 μm (H, I); 10 μm (J, K). 24 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China to 30 μm × 0.8–1.3 μm), and wider conidia (1.5–3 μm vs. 1.3–1.8 μm). Hence, we introduce M. eucalypti as a new species, based on the distinct morphological differences observed. Moelleriella boehmeriae X.L. Xu & C.L. Yang, sp. nov. MycoBank No: MB 858372 Fig. 7 Etymology. In reference to the generic name of host plants. Diagnosis. Similar to Moelleriella jinuoana in having yellow to orange, globose stromata with a narrow hypothallus, M. boehmeriae differs by its slower growth on PDA, unique colony characteristics, and wider conidia. Type. CHINA • Sichuan Province, Leshan City, Muchuan County. Infected scale insects were found on the stems of Boehmeria spicata (Thunb.) Thunb., 28°47.91'N, 103°55.63'E, alt. 900 m, 12 Mar. 2021, C.L. Yang, YCL202103003 (SICAU 25-0080 – holotype preserved in the Herbarium of Sichuan Agricultural University; living culture SICAUCC 25-0061 – ex-holotype stored in the Culture Collection in Sichuan Agricultural University). Description. Parasitic on scale insect from Boehmeria spicata (Urticaceae). Sexual morph: Not observed. Asexual morph: Coelomycetous. Stromata yellow to orange in fresh specimens, entirely covering the insect hosts, thickened pulvinate, globose, tubercules on the surface, closely aggregated, some with narrow hypothallus, 0.8–2.5 mm diameter. Hyphae of stromata form compact textura epidermoidea. Conidiomata simple depressions of surface, round or irregular shape, no mature conidiomata observed. Culture characteristics. The conidia germinate within 12 h in sterile distilled water, with germ tubes extending from both ends of the conidia. Colonies on PDA slowgrowing, attaining a diameter of 5–7 mm in 28 days at 25 °C. Colonies compact pulvinate, surface velutinous, pale yellow to yellow. Conidial masses usually abundant, yellow. Reverse of colony pale orange. Conidiomata U-shaped, subcircular to circular. Conidia unicellular, hyaline, smooth, fusoid with rounded ends, 9.5–12.5 × 3–4 µm. No paraphyses were observed. Host. Scale insects (Coccidae). Habitat. Subtropical monsoon evergreen broad-leaf forests serve as the habitat for Moelleriella boehmeriae, which can infect scale insects found on the stems of Boehmeria spicata. Infections are typically observed at lower heights, close to the ground. Distribution. China, Sichuan Province, Leshan City. Material examined. CHINA • Sichuan Province, Leshan City, Muchuan County. Infected scale insects were found on the stems of Boehmeria spicata, 28°47.91'N, 103°55.63'E, alt. 900 m, 12 Mar. 2021, C.L. Yang, YCL202103003 (SICAU 25-0080, living culture SICAUCC 25-0061); • ibid. YCL202103003-1 (SICAU 25-0081, living culture SICAUCC 25-0062). Notes. Phylogenetic analysis revealed that Moelleriella boehmeriae clusters within the Globose clade and is closely related to M. jinuoana (Fig. 2), sharing characteristics such as yellow to orange, globose stromata with a narrow hypothallus (Wang et al. 2024). Although detailed morphological comparisons were limited due to the absence of mature conidiomata in our specimen, 25 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China M. boehmeriae differs by its slower growth on PDA and distinct colony characteristics. The colonies of M. boehmeriae are yellow on the front and light orange on the reverse, whereas those of M. jinuoana are pale orange to orange on the front and brownish on the reverse. Additionally, M. boehmeriae has wider conidia (3–4 µm) observed in culture compared to M. jinuoana, which has conidia measuring 2.1–2.9 µm on the substrate. In addition, M. boehmeriae differs from the type strain M. jinuoana (YHH MJBP2309031) by 16 bp (1.76%, 0 gap) in the tef1α region, 2 bp (0.28%, 0 gap) in the rpb1 region, and 4 bp (0.47%, 2 gaps) in the LSU region, respectively. Moelleriella cinnamomum X.L. Xu & C.L. Yang, sp. nov. MycoBank No: MB 858373 Fig. 8 Etymology. In reference to the generic name of host plants. Figure 7. Moelleriella boehmeriae. A, B. Immature stromata over scale insects. C. Conidioma formed in PDA. D. Phialides with conidia at the tips. E–G. Phialides with developing conidia. H. Conidia. I. Germinating conidia. J, K. Colonies obverse and reverse on PDA at 25 °C after 3 weeks. L. Conidial mass on PDA. Scale bars: 1000 µm (B); 50 µm (C); 20 µm (D); 10 µm (E–I). 32 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China and uredinia of Hemileia vastatrix on leaves of Coffea arabica from Africa (Pereira et al. 2024). This is the first report of C. guizhouense from aphids on Telosma cordata (Apocynaceae). The scale insect Kuwanaspis howardi is commonly distributed and harmful to bamboo. In our previous study, two Podonectria species and a Microcera species associated with native bamboo plants were discovered (Xu et al. 2021a). Cladosporium kuwanaspidis is the second entomopathogenic species found on bamboo (Pleioblastus amarus) in our investigations. As entomopathogenic C. cladosporioides is a potential candidate for biocontrol against whiteflies and aphids (Islam et al. 2019; Nicoletti et al. 2024), and as endophytic Cladosporium sp. has demonstrated insecticidal activity of α-glucosidase inhibitors against Spodoptera litura in vitro (Singh et al. 2015), more Cladosporium species need to be investigated as potential agents of biocontrol. The genus Moelleriella (Ascomycota, Hypocreales, Clavicipitaceae) was established to accommodate M. sulphurea (Bresadola 1896) and is characterized by globose to pyriform perithecia immersed in a brightly colored stroma with openings protruding from the stroma on scale insects (Coccidae, Homoptera) or whiteflies (Aleyrodidae, Homoptera), as well as multi-septate ascospores that disarticulate at the septa inside the ascus. In contrast, species of Hypocrella and the subsequently proposed genus Samuelsia, having Aschersonia-like anamorphs, have ascospores that remain whole (Petch 1921; Mains 1959; Kirk et al. 2001; Chaverri et al. 2008). The genus currently has about 52 records (https://www.speciesfungorum.org/Names/Names.asp, accessed on 4 November 2024), and species have been reported in Belize, Bolivia, Costa Rica, Côte d’Ivoire, China, Ecuador, Ghana, Guatemala, Guiana, Honduras, Jamaica, Mexico, Panama, the Philippines, Thailand, Trinidad, Venezuela, and Vietnam, frequently from Thailand, indicating it is more common in tropical regions (Liu et al. 2006; Chaverri et al. 2008; Qiu et al. 2009; Mongkolsamrit et al. 2011b, 2015; Li et al. 2016; Tibpromma et al. 2017; Chen et al. 2020; Yuan et al. 2020; Khonsanit et al. 2021). Moelleriella libera has been widely used due to its parasitism of large populations of whiteflies and scale insects in the wild (Zhang et al. 2018; Ingle et al. 2019; Qasim et al. 2020). In China, M. ochracea has previously been recorded from Fujian Province on homopteran cadavers (Qiu et al. 2009). In recent years, several species have been reported from the subtropical regions of Yunnan and Fujian provinces, viz. M. gracilispora on whitefly nymphs (Hemiptera) (Yuan et al. 2020), M. sinensis infecting whitefly nymphs (Hemiptera) (Chen et al. 2020), M. puerensis on whiteflies (Wang et al. 2022c), M. simaoensis on whiteflies (Yang et al. 2023a), and M. jinuoana and M. longzhuensis on scale insects and whiteflies (Wang et al. 2024). Based on our observation, four Moelleriella species were newly recorded on scale insects inhabiting Boehmeria spicata, Citrus sp., and Eucalyptus sp. in Sichuan Province. Moelleriella sinensis, M. cinnamomum, and M. citrus were accommodated in the Effuse clade (Chaverri et al. 2008; Wang et al. 2024), which is characterized by effuse to thin, pulvinate stromata with loose hyphal tissue, mostly having hypothalli. The new species M. boehmeriae was nested in the Globose clade (Chaverri et al. 2008), characterized by globose, darker stromata with compact tissue. Podonectriaceae was proposed as a family in the Pleosporales to accommodate the genus Podonectria, which was confirmed by phylogenetic analyses (Dao et al. 2016; Yang et al. 2019b). Podonectria was traditionally recorded with a Tetracrium-like conidial stage (Petch 1921; Rossman 1978). Previous studies confirmed the link between the sexual morphs in Podonectria and the asexual morphs 33 IMA Fungus 16: e170123 (2025), DOI: 10.3897/imafungus.16.170123 Chunlin Yang et al.: Novel entomopathogenic fungi infecting scale insects and aphids in China in Tetracrium with identical molecular sequences (Dao et al. 2015, 2016; Xu et al. 2021a). In addition, an asexual genus, Tetranacrium, typified by T. gramineum (Hudson and Sutton 1964), was recorded as an associated anamorph of Podonectria gahnia on scale insects. However, although the shape and development of conidia in the associated Tetranacrium are correlated with Tetracrium, the conidiomata in Tetranacrium are pycnidia, whereas in Tetracrium they are sporodochia. The relationship between Tetranacrium and Podonectria requires further phylogenetic and taxonomic studies with more samples. In this study, our collections had hyphomycetous anamorphs, which were consistent with Podonectria, identified based on the phylogenetic results of combined ITS, LSU, SSU, tef1-α, and rpb2 data. Most species of Podonectria are associated with armored scale insects on Citrus spp. Podonectria multiarmata is the third entomopathogenic species associated with Kuwanaspis howardi on bamboo (Pleioblastus amarus) in China. Conclusion Seven entomopathogenic fungi from Sichuan Province, China, are described, including six new species and one newly recorded species. These species are morphologically simple hyphomycetes or coelomycetes, with detailed identification based on morphological characteristics and phylogenetic analyses. These fungi warrant attention, particularly those associated with major agricultural and forestry pests. We predict that southwestern China harbors a rich diversity of entomopathogenic fungi, with many species yet to be discovered and evaluated. These fungi represent a crucial resource for future drug development and hold significant potential for pest management in agriculture and forestry. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of AI No use of AI was reported. Adherence to national and international regulations Not applicable. Funding This research was financed by the Sichuan Science and Technology Program of China (No. 2024NSFSC1192). Author contributions C-LY, X-LX, and FL collected samples. C-LY, X-LX, and FL isolated cultures and performed DNA isolation and PCR amplification. C-LY, X-LX, X-YL, and FL analyzed data. C-LY and X-LX wrote the original draft. Z-Z, QGX, and Y-GL reviewed and edited the draft. 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