New mycoparasitic species in the genera Niveomyces and Pseudoniveomycesgen. nov. (Hypocreales: Cordycipitaceae), with sporothrix-like asexual morphs, from Thailand.

Four new species of the genus Niveomyces are described from Thailand. They were found as mycoparasites on: Ophiocordyceps infecting flies (Diptera) for Niveomyces albus; ants (Hymenoptera) for N. formicidarum; and leafhoppers (Hemiptera) for N. hirsutellae and N. multisynnematus. A new genus, Pseudoniveomyces with two species: Pseudoniveo. blattae (type species), parasitic on Ophiocordyceps infecting cockroaches, and Pseudoniveo. arachnovorum, found on a spider egg sac, are also described. These fungi share a common feature which is a sporothrix-like asexual morph. Based on our molecular data, Sporothrix insectorum is shown to be affiliated to the genus Niveomyces, and thus a new combination N. insectorum comb. nov. is proposed. Niveomyces coronatus, N. formicidarum and N. insectorum formed the N. coronatus species complex found on ant-pathogenic Ophiocordyceps from different continents. Pseudoniveomyces species are distinguished from Niveomyces spp. based on the presence of fusoid macroconidia in culture and a red pigment diffused in the medium, resembling to Gibellula and Hevansia. The molecular phylogenetic analyses also confirmed its generic status. The host/substrates associated with the genera within Cordycipitaceae were mapped onto the phylogeny to demonstrate that mycoparasitism also evolved independently multiple times in this family. Citation: Kobmoo N, Tasanathai K, Araújo JPM, Noisripoom W, Thanakitpipattana D, Mongkolsamrit S, Himaman W, Houbraken J, Luangsa-ard JJ (2023). New mycoparasitic species in the genera Niveomyces and Pseudoniveomyces gen. nov. (Hypocreales: Cordycipitaceae), with sporothrix-like asexual morphs, from Thailand. Fungal Systematics and Evolution 12: 91-110. doi: 10.3114/fuse.2023.12.07.

Multi-locus phylogeny unmasks hidden species within the specialised spider-parasitic fungus, Gibellula (Hypocreales, Cordycipitaceae) in Thailand.

Over 80 species of hypocrealean fungi are reported as pathogens of spiders and harvestmen. Among these fungi, the genus Gibellula is highly regarded as a specialised spider-killer that has never been reported to infect other arthropods. While more than 20 species of Gibellula are known, few attempts to identify the infected spiders have been made despite the fact that the host specificity can help identify the fungal species. Here, we morphologically describe and illustrate eight new species of Gibellula and three new records from Thailand of known species along with the multi-gene phylogeny that clearly showed the segregation among the proposed species. Examination of the Gibellula-infected spider hosts identified Oxyopidae, Uloboridae and, for the first time, the ant-mimicking genus Myrmarachne. Taxonomic novelties: New species: Gibellula brevistipitata Kuephadungphan, Tasanathai & Luangsa-ard, G. longicaudata Tasanathai, Kuephadungphan & Luangsa-ard, G. longispora Kuephadungphan & Luangsa-ard, G. nigelii Kuephadungphan, Tasanathai & Luangsa-ard, G. parvula Kuephadungphan, Tasanathai & Luangsa-ard, G. pilosa Kuephadungphan, Tasanathai & Luangsa-ard, G. solita Kuephadungphan, Tasanathai & Luangsa-ard, G. trimorpha Tasanathai, Khonsanit, Kuephadungphan & Luangsa-ard. Citation: Kuephadungphan W, Petcharad B, Tasanathai K, Thanakitpipattana D, Kobmoo N, Khonsanit A, Samson RA, Luangsa-ard JJ (2022). Multi-locus phylogeny unmasks hidden species within the specialised spider-parasitic fungus, Gibellula (Hypocreales, Cordycipitaceae) in Thailand. Studies in Mycology 101: 245-286. doi: 10.3114/sim.2022.101.04.

Fungal pathogens occurring on Orthopterida in Thailand.

Two new fungal genera and six species occurring on insects in the orders Orthoptera and Phasmatodea (superorder Orthopterida) were discovered that are distributed across three families in the Hypocreales. Sixty-seven sequences generated in this study were used in a multi-locus phylogenetic study comprising SSU, LSU, TEF, RPB1 and RPB2 together with the nuclear intergenic region (IGR). These new taxa are introduced as Metarhizium gryllidicola, M. phasmatodeae, Neotorrubiella chinghridicola, Ophiocordyceps kobayasii, O. krachonicola and Petchia siamensis. Petchia siamensis shows resemblance to Cordyceps mantidicola by infecting egg cases (ootheca) of praying mantis (Mantidae) and having obovoid perithecial heads but differs in the size of its perithecia and ascospore shape. Two new species in the Metarhizium cluster belonging to the M. anisopliae complex are described that differ from known species with respect to phialide size, conidia and host. Neotorrubiella chinghridicola resembles Torrubiella in the absence of a stipe and can be distinguished by the production of whole ascospores, which are not commonly found in Torrubiella (except in Torrubiella hemipterigena, which produces multiseptate, whole ascospores). Ophiocordyceps krachonicola is pathogenic to mole crickets and shows resemblance to O. nigrella, O. ravenelii and O. barnesii in having darkly pigmented stromata. Ophiocordyceps kobayasii occurs on small crickets, and is the phylogenetic sister species of taxa in the 'sphecocephala' clade.

Revisiting Metarhizium and the description of new species from Thailand.

Over the last two decades the molecular phylogeny and classification of Metarhizium has been widely studied. Despite these efforts to understand this enigmatic genus, the basal lineages in Metarhizium are still poorly resolved. In this study, a phylogenetic framework is reconstructed for the Clavicipitaceae focusing on Metarhizium through increased taxon-sampling using five genomic loci (SSU, LSU, tef, rpb1, rpb2) and the barcode marker ITS rDNA. Multi-gene phylogenetic analyses and morphological characterisation of green-spored entomopathogenic Metarhizium isolates from Thailand and soil isolates of M. carneum and M. marquandii reveal their ecological, genetic and species diversity. Nineteen new species are recognised in the Metarhizium clade with narrow host ranges: two new species are found in the M. anisopliae complex - M. clavatum on Coleoptera larvae and M. sulphureum on Lepidoptera larvae; four new species are found in the M. flavoviride complex - M. biotecense and M. fusoideum on brown plant hoppers (Hemiptera), M. culicidarum on mosquitoes, M. nornnoi on Lepidoptera larvae; three new species M. megapomponiae, M. cicadae, M. niveum occur on cicadas; five new species M. candelabrum, M. cercopidarum, M. ellipsoideum, M. huainamdangense M. ovoidosporum occur on planthoppers, leafhoppers and froghoppers (Hemiptera); one new species M. eburneum on Lepidoptera pupae; and four new species M. phuwiangense, M. purpureum, M. purpureonigrum, M. flavum on Coleoptera . Of these 19 new species, seven produce a sexual morph (M. clavatum, M. eburneum, M. flavum, M. phuwiangense, M. purpureonigrum, M. purpureum, and M. sulphureum) and asexual morphs are found in the remaining new species and also in M. sulphureum, M. purpureonigrum and M. purpureum. Metarhizium blattodeae, M. koreanum and M. viridulum are new records for Thailand. An alternative neotype for Metarhizium anisopliae is proposed based on multi-gene and 5'tef analyses showing that CBS 130.71 from Ukraine is more suitable, being from a much closer geographical location to Metchnikoff's Metarhizium anisopliae. This isolate is distinct from the neotype of Metarhizium anisopliae var. anisopliae proposed by M. Tulloch from Ethiopia (ARSEF 7487). Six new genera are established for monophyletic clades subtending the core Metarhizium clade, including Keithomyces, Marquandomyces, Papiliomyces, Purpureomyces, Sungia, and Yosiokobayasia. Metarhizium carneum, M. aciculare, and M. neogunnii are combined in Keithomyces and one new combination for M. marquandii in Marquandomyces is proposed. Purpureomyces is introduced for species producing purple stromata including a new combination for M. khaoyaiense and two new species P. maesotensis and P. pyriformis. Papiliomyces contains two new combinations for M. liangshanense and Metacordyceps shibinensis. The genus Sungia is proposed for the Korean species M. yongmunense on Lepidoptera pupa and Yosiokobayasia for the Japanese species M. kusanagiense also on Lepidoptera pupa. A synoptic and dichotomous key to the accepted taxa is provided together with tables listing distinguishing morphological characters between species, host preferences, and geography.

Fungal Planet description sheets: 951-1041.

Novel species of fungi described in this study include those from various countries as follows: Antarctica, Apenidiella antarctica from permafrost, Cladosporium fildesense from an unidentified marine sponge. Argentina, Geastrum wrightii on humus in mixed forest. Australia, Golovinomyces glandulariae on Glandularia aristigera, Neoanungitea eucalyptorum on leaves of Eucalyptus grandis, Teratosphaeria corymbiicola on leaves of Corymbia ficifolia, Xylaria eucalypti on leaves of Eucalyptus radiata. Brazil, Bovista psammophila on soil, Fusarium awaxy on rotten stalks of Zea mays, Geastrum lanuginosum on leaf litter covered soil, Hermetothecium mikaniae-micranthae (incl. Hermetothecium gen. nov.) on Mikania micrantha, Penicillium reconvexovelosoi in soil, Stagonosporopsis vannaccii from pod of Glycine max. British Virgin Isles, Lactifluus guanensis on soil. Canada, Sorocybe oblongispora on resin of Picea rubens. Chile, Colletotrichum roseum on leaves of Lapageria rosea. China, Setophoma caverna from carbonatite in Karst cave. Colombia, Lareunionomyces eucalypticola on leaves of Eucalyptus grandis. Costa Rica, Psathyrella pivae on wood. Cyprus, Clavulina iris on calcareous substrate. France, Chromosera ambigua and Clavulina iris var. occidentalis on soil. French West Indies, Helminthosphaeria hispidissima on dead wood. Guatemala, Talaromyces guatemalensis in soil. Malaysia, Neotracylla pini (incl. Tracyllales ord. nov. and Neotracylla gen. nov.) and Vermiculariopsiella pini on needles of Pinus tecunumanii. New Zealand, Neoconiothyrium viticola on stems of Vitis vinifera, Parafenestella pittospori on Pittosporum tenuifolium, Pilidium novae-zelandiae on Phoenix sp. Pakistan, Russula quercus-floribundae on forest floor. Portugal, Trichoderma aestuarinum from saline water. Russia, Pluteus liliputianus on fallen branch of deciduous tree, Pluteus spurius on decaying deciduous wood or soil. South Africa, Alloconiothyrium encephalarti, Phyllosticta encephalarticola and Neothyrostroma encephalarti (incl. Neothyrostroma gen. nov.) on leaves of Encephalartos sp., Chalara eucalypticola on leaf spots of Eucalyptus grandis × urophylla, Clypeosphaeria oleae on leaves of Olea capensis, Cylindrocladiella postalofficium on leaf litter of Sideroxylon inerme, Cylindromonium eugeniicola (incl. Cylindromonium gen. nov.) on leaf litter of Eugenia capensis, Cyphellophora goniomatis on leaves of Gonioma kamassi, Nothodactylaria nephrolepidis (incl. Nothodactylaria gen. nov. and Nothodactylariaceae fam. nov.) on leaves of Nephrolepis exaltata, Falcocladium eucalypti and Gyrothrix eucalypti on leaves of Eucalyptus sp., Gyrothrix oleae on leaves of Olea capensis subsp. macrocarpa, Harzia metrosideri on leaf litter of Metrosideros sp., Hippopotamyces phragmitis (incl. Hippopotamyces gen. nov.) on leaves of Phragmites australis, Lectera philenopterae on Philenoptera violacea, Leptosillia mayteni on leaves of Maytenus heterophylla, Lithohypha aloicola and Neoplatysporoides aloes on leaves of Aloe sp., Millesimomyces rhoicissi (incl. Millesimomyces gen. nov.) on leaves of Rhoicissus digitata, Neodevriesia strelitziicola on leaf litter of Strelitzia nicolai, Neokirramyces syzygii (incl. Neokirramyces gen. nov.) on leaf spots of Syzygium sp., Nothoramichloridium perseae (incl. Nothoramichloridium gen. nov. and Anungitiomycetaceae fam. nov.) on leaves of Persea americana, Paramycosphaerella watsoniae on leaf spots of Watsonia sp., Penicillium cuddlyae from dog food, Podocarpomyces knysnanus (incl. Podocarpomyces gen. nov.) on leaves of Podocarpus falcatus, Pseudocercospora heteropyxidicola on leaf spots of Heteropyxis natalensis, Pseudopenidiella podocarpi, Scolecobasidium podocarpi and Ceramothyrium podocarpicola on leaves of Podocarpus latifolius, Scolecobasidium blechni on leaves of Blechnum capense, Stomiopeltis syzygii on leaves of Syzygium chordatum, Strelitziomyces knysnanus (incl. Strelitziomyces gen. nov.) on leaves of Strelitzia alba, Talaromyces clemensii from rotting wood in goldmine, Verrucocladosporium visseri on Carpobrotus edulis. Spain, Boletopsis mediterraneensis on soil, Calycina cortegadensisi on a living twig of Castanea sativa, Emmonsiellopsis tuberculata in fluvial sediments, Mollisia cortegadensis on dead attached twig of Quercus robur, Psathyrella ovispora on soil, Pseudobeltrania lauri on leaf litter of Laurus azorica, Terfezia dunensis in soil, Tuber lucentum in soil, Venturia submersa on submerged plant debris. Thailand, Cordyceps jakajanicola on cicada nymph, Cordyceps kuiburiensis on spider, Distoseptispora caricis on leaves of Carex sp., Ophiocordyceps khonkaenensis on cicada nymph. USA, Cytosporella juncicola and Davidiellomyces juncicola on culms of Juncus effusus, Monochaetia massachusettsianum from air sample, Neohelicomyces melaleucae and Periconia neobrittanica on leaves of Melaleuca styphelioides × lanceolata, Pseudocamarosporium eucalypti on leaves of Eucalyptus sp., Pseudogymnoascus lindneri from sediment in a mine, Pseudogymnoascus turneri from sediment in a railroad tunnel, Pulchroboletus sclerotiorum on soil, Zygosporium pseudomasonii on leaf of Serenoa repens. Vietnam, Boletus candidissimus and Veloporphyrellus vulpinus on soil. Morphological and culture characteristics are supported by DNA barcodes.

Novel and interesting Ophiocordyceps spp. (Ophiocordycipitaceae, Hypocreales) with superficial perithecia from Thailand.

Ophiocordyceps is a heterogeneous, species-rich genus in the order Hypocreales (Sordariomycetes, Ascomycota) that includes invertebrate-pathogenic taxa. In this study, seven new species in Ophiocordyceps producing superficial perithecia infecting various insect hosts (Lepidoptera, Hemiptera) are described from Thailand - Ophiocordyceps brunneinigra, O. brunneiperitheciata, O. geometridicola, O. multiperitheciata, O. pauciovoperitheciata, O. pseudoacicularis and O. spataforae. Phylogenetic analyses based on multigene loci comprising the large subunit of the ribosomal DNA (LSU), partial sequences of elongation factor 1-alpha (TEF) and the largest and second largest subunit of the RNA polymerase (RPB1, PRB2) strongly support these new species of Ophiocordyceps in the Ophiocordycipitaceae. They differ from species previously described species Ophiocordyceps acicularis, O. atewensis, O. cochlidiicola, and O. crinalis, in the shape and sizes of distinguishing characters such as perithecia, ascospores and conidia. We also report a new record of O. macroacicularis in Thailand.

Life cycle, host range and temporal variation of Ophiocordyceps unilateralis/Hirsutella formicarum on Formicine ants.

Ophiocordyceps unilateralis sensu lato infects ants, modifies their behavior, and is found in many countries around the world. One unifying concept of all such parasitic associations is that both the parasite and the host adapt to maximize their fitness and reproductive output. However, little is known about the reproductive life span of this pathogen or about its alternation between asexual and sexual states, even though these two states affect host population dynamics differently. To address these issues, a permanent plot in a tropical rainforest was surveyed over the course of two years to examine the development of O. unilateralis s.l. and the incidence of infection of Polyrhachis and Camponotus ants, which were found to be specifically attacked by this fungus in Thailand. We document here for the first time the life cycle of this pathogen over the long term, which provides fundamental knowledge for the understanding of this fascinating host-parasite interaction.

Molecular phylogenies reveal host-specific divergence of Ophiocordyceps unilateralis sensu lato following its host ants.

Ophiocordyceps unilateralis (Hypocreales, Ascomycetes) is an entomopathogenic fungus specific to formicine ants (Formicinae, Hymenoptera). Previous works have shown that the carpenter ant Camponotus leonardi acts as the principal host with occasional infections of ants from the genus Polyrhachis (sister genus of Camponotus). Observations were made on the permanent plots of Mo Singto, Khao Yai National Park of Thailand according to which O. unilateralis was found to occur predominantly on three host species: C. leonardi, C. saundersi and P. furcata. Molecular phylogenies of the elongation factor 1-α and ß-Tubulin genes indicate a separation of O. unilateralis samples into three clades, reflecting specificity to each of the three different ant species. Samples collected from P. furcata and from C. leonardi were found to form sister groups with samples from C. saundersi forming an outgroup to the latter. Additional samples collected from unidentified ant species of Camponotus and Polyrhachis were positioned as outgroups to those samples on identified species. These results demonstrate that O. unilateralis is clearly not a single phylogenetic species and comprises at least three species that are specific to different host ant species. These cryptic species may arise through recent events of speciation driven by their specificity to host ant species.