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Understanding the systematics and evolution of clavicipitoid fungi has been greatly aided by the application of molecular phylogenetics. They are now classified in three families, largely driven by reevaluation of the morphologically and ecologically diverse genus Cordyceps. Although reevaluation of morphological features of both sexual and asexual...
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The phylogeny and systematics of cordycipitoid fungi have been extensively studied in the last two decades. However, systematic positions of some taxa in the family Cordycipitaceae have not yet been thoroughly resolved. In this study, a new phylogenetic framework of Cordycipitaceae is reconstructed using multigene (nrSSU, nrLSU, tef-1α, rpb1 and rp...
Ophiocordycipitaceae is a diverse family comprising ecologically, economically, medicinally, and culturally important fungi. The family was recognized due to the polyphyly of the genus Cordyceps and the broad diversity of the mostly arthropod-pathogenic lineages of Hypocreales. The other two cordyceps-like families, Cordycipitaceae and Clavicipitac...
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... Based on 5~7 gene loci, Sung et al. reclassified a large number of Cordyceps-related species into three families: Cordycipitaceae, Ophiocordycipitaceae, and Clavicipitaceae [46]. Polycephalomyces and Ophiocordyceps share many of the same morphological characteristics, such as Hirsutella-like asexual forms and a consistent sexual spore structure [47,48]. Our study found many types of endophytic fungi that have never been previously reported. ...
Endosymbiotic fungi play an important role in the growth and development of insects. Understanding the endosymbiont communities hosted by the brown planthopper (BPH; Nilaparvata lugens Stål), the most destructive pest in rice, is a prerequisite for controlling BPH rice infestations. However, the endosymbiont diversity and dynamics of the BPH remain poorly studied. Here, we used circular consensus sequencing (CCS) to obtain 87,131 OTUs (operational taxonomic units), which annotated 730 species of endosymbiotic fungi in the various developmental stages and tissues. We found that three yeast-like symbionts (YLSs), Polycephalomyces prolificus, Ophiocordyceps heteropoda, and Hirsutella proturicola, were dominant in almost all samples, which was especially pronounced in instar nymphs 4–5, female adults, and the fat bodies of female and male adult BPH. Interestingly, honeydew as the only in vitro sample had a unique community structure. Various diversity indices might indicate the different activity of endosymbionts in these stages and tissues. The biomarkers analyzed using LEfSe suggested some special functions of samples at different developmental stages of growth and the active functions of specific tissues in different sexes. Finally, we found that the incidence of occurrence of three species of Malassezia and Fusarium sp. was higher in males than in females in all comparison groups. In summary, our study provides a comprehensive survey of symbiotic fungi in the BPH, which complements the previous research on YLSs. These results offer new theoretical insights and practical implications for novel pest management strategies to understand the BPH–microbe symbiosis and devise effective pest control strategies.
... Chaverri et al. (2005) initiated this molecular exploration by providing LSU, TEF, and RPB1 data for Polycephalomyces formosus and Polycephalomyces ramosopulvinatus (current name: Pleurocordyceps ramosopulvinata). Ban et al. (2009) used a 504-base-pair LSU fragment, but it fell short in resolving deep fungal nodes (Kepler et al., 2013). Different loci were selected for the analysis of novel species, with Wang et al. (2014) using a 4-loci (SSU, LSU, TEF, and RPB1), Wang et al. (2015b) using a 5-loci (SSU, LSU, TEF, RPB1, and RPB2), and Wang et al. (2015a) and Xiao et al. (2018) utilizing a 6-loci (ITS, SSU, LSU, TEF, and RPB1, and RPB2). ...
... Different loci were selected for the analysis of novel species, with Wang et al. (2014) using a 4-loci (SSU, LSU, TEF, and RPB1), Wang et al. (2015b) using a 5-loci (SSU, LSU, TEF, RPB1, and RPB2), and Wang et al. (2015a) and Xiao et al. (2018) utilizing a 6-loci (ITS, SSU, LSU, TEF, and RPB1, and RPB2). The phylogenetic placement of Polycephalomyces or the segregation of new genera from Polycephalomyces was analyzed using both 5-loci (SSU, LSU, TEF, RPB1, and RPB2) and 6-loci (ITS, SSU, LSU, TEF, RPB1, and RPB2) (Kepler et al., 2013;Matočec et al., 2014;Wang et al., 2021). Building on this molecular groundwork, Xiao et al. (2023) established a new family, Polycephalomycetaceae, accommodating three genera (Perennicordyceps, Pleurocordyceps, and Polycephalomyces) and comprising 28 species using 6 loci (ITS, SSU, LSU, TEF, RPB1, and RPB2). ...
... Most species in Polycephalomycetaceae are found in tropical and subtropical regions, with fewer taxa found in temperate regions (Van Vooren and Audibert, 2005;Wang et al., 2012Wang et al., , 2015aMatočec et al., 2014;Xiao et al., 2018Xiao et al., , 2023. A high diversity of polycephalomycetous fungi has been found in China and Japan (Kobayasi, 1939(Kobayasi, , 1941Kobayasi and Shimizu, 1982;Chen et al., 1984;Wang et al., 2012Wang et al., , 2014Wang et al., , 2015aWang et al., ,b, 2021Kepler et al., 2013;Quandt et al., 2014;Yang et al., 2020;Xiao et al., 2023). ...
Entomopathogenic fungi comprise an ecologically important group of specialized pathogens infecting other fungi, invertebrates, and plants. These fungi are species-rich with high diversity and broad distribution worldwide. The majority of entomopathogenic fungi belong to clavicipitoids, which consist of the hypocrealean families, Clavicipitaceae, Cordycipitaceae, Ophiocordycipitaceae, and Polycephalomycetaceae. The latter is a newly established entomopathogenic family that recently separated from the family Ophiocordycipitaceae to accommodate the genera, Perennicordyceps, Pleurocordyceps , and Polycephalomyces . In recent years, Polycephalomycetaceae has been enriched with parasitic and hyperparasitic fungi. With 16 species spread across China, Ecuador, Japan, and Thailand, Pleurocordyceps is the most speciose genus in the family. In this study, we expand the number of taxa in the genus by introducing four new Pleurocordyceps species from China, namely, P. clavisynnema, P. multisynnema, P. neoagarica , and P. sanduensis . We provide detailed descriptions and illustrations and infer genus-level phylogenies based on a combined 6-loci gene sequence dataset comprising the internal transcribed spacer gene region (ITS), small subunit ribosomal RNA gene region (SSU), large subunit rRNA gene region (LSU), translation elongation factor 1-alpha gene region (TEF-1α), RNA polymerase II largest subunit gene region (RPB1), and RNA polymerase II second largest subunit (RPB2). This study contributes to knowledge with regard to the diversity of Pleurocordyceps specifically and entomopathogenic Hypocreales more broadly.
... Cordyceps is a unique group of macrofungi that parasitizes insect larvae and pupae. More than 750 species have been reported, belonging to the order Ascomycota and classified into three families: Cordycipitaceae, Ophiocordycipitaceae, and Clavicipitaceae [13][14][15]. Ophiocordyceps (Ophiocordycipitaceae) is one of the most populous genera of entomopathogenic fungi, of which Ophiocordyceps gracilis (O. gracilis) is well-known traditional Chinese medicine [16]. ...
Utilizing mycoremediation is an important direction for managing heavy metal pollution. Zn2+ pollution has gradually become apparent, but there are few reports about its pollution remediation. Here, the Zn2+ remediation potential of Paraisaria dubia, an anamorph of the entomopathogenic fungus Ophiocordyceps gracilis, was explored. There was 60% Zn2+ removed by Paraisaria dubia mycelia from a Zn2+-contaminated medium. To reveal the Zn2+ tolerance mechanism of Paraisaria dubia, transcriptomic and metabolomic were executed. Results showed that Zn2+ caused a series of stress responses, such as energy metabolism inhibition, oxidative stress, antioxidant defense system disruption, autophagy obstruction, and DNA damage. Moreover, metabolomic analyses showed that the biosynthesis of some metabolites was affected against Zn2+ stress. In order to improve the tolerance to Zn2+ stress, the metabolic mechanism of metal ion transport, extracellular polysaccharides (EPS) synthesis, and microcycle conidiation were activated in P. dubia. Remarkably, the formation of microcycle conidiation may be triggered by reactive oxygen species (ROS) and mitogen-activated protein kinase (MAPK) signaling pathways. This study supplemented the gap of the Zn2+ resistance mechanism of Paraisaria dubia and provided a reference for the application of Paraisaria dubia in the bioremediation of heavy metals pollution.
... Screening for cellulolytic activity in this entomopathogenic species is not commonly done because it is mainly studied for its biological control abilities in multiple agricultural crops [69][70][71]. Beauveria has also been found as an endophyte, along with Purpureocillium [72]. An isolate of Purpureocillium lilacinum has been reported as a biodeteriogen of indoor materials able to grow in alkaline materials, producing damage in limestones and plasters of cultural heritage in Russia [73]. ...
We studied the physicochemical characteristics and mycobiota associated to five key historic documents from Costa Rica, including the Independence Act of Costa Rica from 1821. We used nondestructive techniques (i.e., ATR-FTIR and XRF) to determine paper and ink composition. Results show that some documents are composed of cotton-based paper, whereas others were made of wood cellulose with an increased lignin content. We also determined that the ink employed in some of the documents is ferrogallic. Cultivation and molecular techniques were used to characterize the fungi inhabiting the documents. In total, 22 fungal isolates were obtained: 15 from the wood-cellulose-based documents and seven from the other three cotton-based. We also tested the cellulolytic activity of the recovered fungi; 95% of the fungi presented cellulolytic activity correlated to their ability to cause deterioration of the paper. Results suggest that cotton-based paper is the most resistant to fungal colonization and that most of the isolates have cellulolytic activity. This work increases the knowledge of the fungal diversity that inhabits historic documents and its relationship with paper composition and provides valuable information to develop strategies to conserve and restore these invaluable documents.
... The order Hypocreales contains the widest diversity of animal parasites among the kingdom Fungi. Most hypocrealean fungi are parasites of plants and arthropods, especially insects, although some species are known to parasitize spiders, nematodes, rotifers and even immunocompromised humans, as well as other fungi (Samson et al. 1988, Kepler et al. 2013, Lombard et al. 2015, Araújo & Hughes 2016. The genus Ophio-cordyceps comprises approximately 300 species, strictly associated with insects belonging to 13 orders (Crous et al. 2004, Araújo & Hughes 2016. ...
... In order to investigate the relationships of Torrubiellomyces, we sampled Polycephalomyces s.lat. (Polycephalomyces insertae sedis, Pleurocordyceps and Perennicordyceps); including a range of species representing distinct ecologies, such as animal and fungal parasites (Kepler et al. 2013, Matočec et al. 2014, Xiao et al. 2018, Wang et al. 2021 (Fig. 3). Our phylogeny suggests the Polycephalomyces clade as the most basal lineage within Ophiocordycipitaceae (BS = 53) and strongly supports Torrubiellomyces as a distinct genus (BS = 98), closely related to a clade strictly associated with insects: Perennicordyceps cuboidea, Pe. ...
... Paracuboidea, Pe. ryogamiensis (all on Coleoptera), and Pe. prolificus (on Hemiptera) (Fig. 3) (Kepler et al. 2013). ...
During surveys in central Florida of the zombie-ant fungus Ophiocordyceps camponoti-floridani, which manipulates the behavior of the carpenter ant Camponotus floridanus, two distinct fungal morphotypes were discovered associated with and purportedly parasitic on O. camponoti-floridani. Based on a combination of unique morphology, ecology and phylogenetic placement, we discovered that these morphotypes comprise two novel lineages of fungi. Here, we propose two new genera, Niveomyces and Torrubiellomyces, each including a single species within the families Cordycipitaceae and Ophiocordycipitaceae, respectively. We generated de novo draft genomes for both new species and performed morphological and multi-loci phylogenetic analyses. The macromorphology and incidence of both new species, Niveomyces coronatus and Torrubiellomyces zombiae, suggest that these fungi are mycoparasites since their growth is observed exclusively on O. camponoti-floridani mycelium, stalks and ascomata, causing evident degradation of their fungal hosts. This work provides a starting point for more studies into fungal interactions between mycopathogens and entomopathogens, which have the potential to contribute towards efforts to battle the global rise of plant and animal mycoses.
Citation: Araújo JPM, Lebert BM, Vermeulen S, et al. 2022. Masters of the manipulator: two new hypocrealean general, Niveomyces (Cordycipitaceae) and Torrubiellomyces (Ophiocordycipitaceae), parasitic on the zombie ant fungus Ophiocordyceps camponoti-floridani. Persoonia 49: 171–194. https://doi.org/10.3767/persoonia.2022.49.05.
... Only three species, that is, P. paludosus, P. cylindrosporus, and P. tomentosus, were described in the last century (Mains, 1948;Samson et al., 1981;Seifert, 1986). Until recently, after the recombination of Paecilomyces sinensis (Wang et al., 2012) and several species of Cordyceps s. l. into the genus (Kepler et al., 2013), more and more new species have been discovered and described, especially from China and Southeast Asia (Crous et al., 2017;Wang et al., 2015aWang et al., , 2015bXiao et al., 2018;Yang et al., 2020). A total of 24 species names are currently recorded by the Index Fungorum (4 April 2022, http://www.index ...
... Ten species were included in the new genus including P. sinensis. In multigene phylogenetic analyses, species of Polycephalomyces s. l. usually formed a distinct clade sister to Ophiocordycipitaceae, although this sister relationship did not receive much statistical confidence (Kepler et al., 2013;Wang et al., 2021). In other words, the family-level taxonomic position of Polycephalomyces s. l. was not fully resolved; species in this group were tentatively placed in Ophiocordycipitaceae in most researches (e.g., Kepler et al., 2013;Xiao et al., 2018), which was accepted by the Index Fungorum. ...
... In multigene phylogenetic analyses, species of Polycephalomyces s. l. usually formed a distinct clade sister to Ophiocordycipitaceae, although this sister relationship did not receive much statistical confidence (Kepler et al., 2013;Wang et al., 2021). In other words, the family-level taxonomic position of Polycephalomyces s. l. was not fully resolved; species in this group were tentatively placed in Ophiocordycipitaceae in most researches (e.g., Kepler et al., 2013;Xiao et al., 2018), which was accepted by the Index Fungorum. Polycephalomyces s. l. may represent a new family that is different from the three existing families of clavicipitoid fungi , that is, Cordycipitaceae, Clavicipitaceae, and Ophiocordycipitaceae. ...
The complete mitochondrial (mt) genome of the ex-type strain of Pleurocordyceps sinensis, a fungus originally isolated from Ophiocordyceps sinensis, was sequenced, and assembled as a single circular DNA of 31,841 bp. The mt genome encoded 15 conserved proteins (rps3, cox1, cox2, cox3, cob, atp6, atp8, atp9, nad1, nad2, nad3, nad4, nad4L, nad5, and nad6), 2 rRNA (rnl and rns), and 25 tRNA, as well as 10 additional non-conserved open reading frames (ncORFs). Comparative analyses showed that mt genomes within the order Hypocreales encoded the same number and synteny of conserved protein coding genes despite an obvious size variation among this group of fungi. Phylogenetic analyses using 14 conserved protein sequences revealed that this fungus may not belong to the current designated family Ophiocordycipitaceae but is more closely related to the species of Clavicipitaceae. The mt genome presented herein would give valuable information on reconstructing the evolutionary history of clavicipitaceous fungi and also aid in resolving the family-level taxonomic assignment of Polycephalomyces s. l. species.
... A partition homogeneity test was conducted in PAUP* 4.0a166 [39], and the results indicated that there were no conflicts among the data partitions. PartitionFinder V1.1.1 identified eleven data partitions: nine corresponding to the three codon positions in each of the proteincoding genes (tef-1α, rpb1, and rpb2) and one each for nrLSU and nrSSU [42,43]. The results showed that the phylogenetic signals of the five genes were congruent (p = 0.02). ...
The taxonomy and phylogeny of the genus Tolypocladium are herein revised based on the most comprehensive dataset to date. Two species-level phylogenies of Tolypocladium were constructed: a single-gene phylogeny (ITS) of 35 accepted species and a multigene phylogeny (nrSSU, nrLSU, tef-1α, rpb1, and rpb2) of 27 accepted species. Three new species, Tolypocladium pseudoalbum sp. nov., Tolypocladium subparadoxum sp. nov., and Tolypocladium yunnanense sp. nov., are described in the present study. The genetic divergences of four markers (ITS, tef-1α, rpb1 and rpb2) among Tolypocladium species are also reported. The results indicated that species of Tolypocladium were best delimited by rpb1 sequence data, followed by the sequence data for the rpb2, tef-1α, and ITS provided regions. Finally, a key to the 48 accepted species of Tolypocladium worldwide is provided.
... Within Hypocreales, Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae have significant medicinal, economic and ecological value (Quandt et al. 2014;Kepler et al. 2017). These three families are well known as the clavicipitoid fungi (Humber and Richard 2008;Kepler et al. 2013). ...
... The taxonomy and systematics of clavicipitoids have been subject to frequent reevaluation (Sung et al. 2007;Kepler et al. 2013;Quandt et al. 2014;Spatafora et al. 2015;Maharachchikumbura et al. 2015Maharachchikumbura et al. , 2016Mongkolsamriit et al. 2019;Wijayawardene et al. 2018Wijayawardene et al. , 2020. Previously, clavicipitoid fungi were recognized as the single family Clavicipitaceae (Eriksson, 1982;Rogerson, 1970;Eriksson & Hawksworth, 1985, 1995White et al., 2000). ...
... Kobayasi (1941) established Polycephalomyces to accommodate Polycephalomyces formosus, an asexual morph species, which produces polycephalous synnemata and has been described as entomopathogenic fungus. The taxon has had a long history of being recognised as incertae sedis in Hypocreales (Kepler et al. 2013;Matočec et al. 2014). Its characters have been described as acremonium-like and hirsutella-like, while a sexual morph has yet to be identified (Kobayasi 1941;Kepler et al. 2013). ...
Clavicipitoid fungi comprise three families, namely Clavicipitaceae , Cordycipitaceae , and Ophiocordycipitaceae . They are found worldwide and are specialized pathogens of invertebrate, plant and fungal hosts. Over the last decade, morphology- and phylogeny-based studies on clavicipitoid fungi have increased. The latter have revealed that Polycephalomyces , Perennicordyceps and Pleurocordyceps consistently cluster together.These genera are currently considered as members of Ophiocordycipitaceae . Nonetheless, information with regard to their diversity and ecology remains sparse. To fill this gap, we collected 29 fresh specimens from insect and fungal substrates from tropical and subtropical evergreen forests in Thailand and southwestern China. We performed detailed morphological analyses and constructed photoplates for all isolated fungi. We used extensive taxon sampling and a dataset comprising internal transcribed spacer gene region (ITS), small subunit ribosomal RNA gene region (SSU), large subunit rRNA gene region (LSU), translation elongation factor 1-alpha gene region (TEF-1α), RNA polymerase II largest subunit gene region (RPB1) and RNA polymerase II second largest subunit (RPB2) to infer order-, family and genus-level phylogenetic trees. Based on these biphasic analyses, we segregate Polycephalomyces , Perennicordyceps , and Pleurocordyceps from Ophiocordycipitaceae and introduce the new family Polycephalomycetaceae to accomodate these three genera. The majority of species in this family have a vast range of insect and fungal hosts. The sexual morph of Polycephalomycetaceae has stromatic ascomata, long stipes, thick peridium, and cylindrical secondary spores. The asexual morph is characterized by colonies on the host surface or synnemata with stipes on the host, one or two types of phialides, and cylindrical to fusiform conidia. We expand the number of taxa in the new family by introducing seven new species ( Polycephalomyces albiramus , Perennicordyceps lutea , Pleurocordyceps parvicapitata , Pleurocordyceps lanceolatus , Pleurocordyceps nutansis , Pleurocordyceps heilongtanensis , Pleurocordyceps vitellina ), nine new hosts, and one new combination ( Perennicordyceps elaphomyceticola ). The results herein hint at a high level of diversity for Polycephalomycetaceae . Future investigations focusing on obtaining additional collections and specimens from different geographical areas would help to reveal not only the extent of the group’s diversity, but also resolve its deeper phylogenetic placement.
... Partition homogeneity test was conducted using PAUP* 4.0b10 (Swofford 2002), the result revealed that there was no significant conflict among different data partitions. Program Par-titionFinder V1.1.1 identified eleven data partitions, one each for LSU and SSU, and nine for each of the three codon positions for the protein coding genes TEF, RPB1, and RPB2 (Lanfear et al. 2012;Kepler et al. 2013). The results showed that the phylogenetic signals for the five genes were congruent (P = 0.02). ...
Two Tolypocladium species, Tolypocladium reniformisporum sp. nov., and T. cylindrosporum, were isolated from the sclerotium of a naturally occurring Ophiocordyceps sinensis specimen in the Baima Snow Mountains of Deqin County, Yunnan Province in southwestern China. They were described and illustrated by morphological characters and growth-temperature relationships. Phylogenetic analyses based on a 5-gene (SSU, LSU, TEF, RPB1, and RPB2) dataset and sequences of the internal transcribed spacer regions (ITS) showed that T. reniformisporum was phylogenetically closely related to T. cylindrosporum and other similar species. Tolypocladium reniformisporum was morphologically distinguishable from other Tolypocladium species because of its slow-growing colonies, short phialides, and smaller reniform conidia. Growth tests showed that the isolated T. reniformisporum and T. cylindrosporum strains were psychrotrophic, with optimal growth temperatures of 18–22 ℃, and minimum growth temperatures below 5 ℃.
... Only three species, i.e., P. paludosus, P. cylindrosporus and P. tomentosus, were described in the last century (Mains 1948; Samson et al. 1981;Seifert 1986). Until recently, after the recombination of Paecilomyces sinensis (Wang et al. 2012) and several species of Cordyceps s. l. into this genus (Kepler et al. 2013), more and more new species were discovered and described, especially from China and Southeast Asia (Wang et proposed a new genus Pleurocordyceps for one of the subclade within the 'Polycephalomyces clade' based on morphological and molecular analyses. Ten species were included in the new genus including P. sinensis. ...
... Ten species were included in the new genus including P. sinensis. The taxonomic position of species of Polycephalomyces based on the type P. formosus was not resolved (Kepler et al. 2013), though species of this group were tentatively placed in Ophiocordycipitaceae in Index Fungorum. ...
The complete mitochondrial (mt) genome of Pleurocordyceps sinensis , a fungus originally isolated from Ophiocordyceps sinensis was sequenced, and assembled as a single circular DNA of 31,841 bp. The mt genome encoded 15 conserved proteins ( rps3 , cox1 , cox2 , cox3 , cob , atp6 , atp8 , atp9 , nad1 , nad2 , nad3 , nad4 , nad4L , nad5 and nad6 ), 2 rRNA ( rnl and rns ) and 25 tRNA, as well as 10 additional non-conserved open reading frames. Comparative analyses showed that mt genomes within the order Hypocreales encoded same number and synteny of conserved protein coding genes despite an obvious size variation among this group of fungi. Phylogenetic analyses using 14 conserved protein sequences revealed that this fungus may not belong to the current designated family Ophiocordycipitaceae but is more closely related to the species of Clavicipitaceae . The mt genome presented herein would aid to clarify the phylogenetic position of species of Polycephalomyces s. l. and also gave valuable information on reconstructing the evolutionary history of clavicipitaceous fungi.
