RESUMEN
In this study, we investigated the morphological and genetic variability of selected species belonging to the genus Chloridium sensu lato, some also referred to as chloridium-like asexual morphs and other undescribed morphologically similar fungi. These species do not conform to the revised generic concept and thus necessitate a re-evaluation in terms of taxonomy and phylogeny. The family Chaetosphaeriaceae (Chaetosphaeriales) encompasses a wide range of asexual morphotypes, and among them, the simplest form is represented by Chloridium sect. Chloridium. The morphological simplicity of the Chloridium morphotype has historically led to the amalgamation of numerous unrelated species, thereby creating a heterogeneous genus. By conducting phylogenetic reconstruction of four DNA loci and examining a set of 71 strains, including all available ex-type and other non-type strains as well as holotypes and other herbarium material, we were able to gain new insights into the relationships between these taxa. Phylogenetic analyses revealed that the studied species are distantly related to Chloridium sensu stricto and can be grouped into two orders in the Sordariomycetes. Within the Chaetosphaeriales, they formed nine well-separated genera in four clades, such as Cacumisporium, Caliciastrum gen. nov., Caligospora gen. nov., Capillisphaeria gen. nov., Curvichaeta, Fusichloridium, Geniculoseta gen. nov., Papillospora gen. nov., and Spicatispora gen. nov. We also established Chloridiopsiella gen. nov. and Chloridiopsis gen. nov. in Vermiculariopsiellales. Four new species and eight new combinations are proposed in these genera. Our study provides a clearer understanding of the genus Chloridium, its relationship to other morphologically similar fungi, and a new taxonomic treatment and molecular phylogeny to facilitate their accurate identification and classification in future research. Taxonomic novelties: New genera: Caliciastrum Réblová, Caligospora Réblová, Capillisphaeria Réblová, Chloridiopsiella Réblová, Chloridiopsis Réblová, Geniculoseta Réblová, Papillospora Réblová, Spicatispora Réblová; New species: Caliciastrum bicolor Réblová, Caligospora pannosa Réblová, Chloridiopsis syzygii Réblová, Gongromerizella silvana Réblová; New combinations: Caligospora dilabens (Réblová & W. Gams) Réblová, Capillisphaeria crustacea (Sacc.) Réblová, Chloridiopsiella preussii (W. Gams & Hol.-Jech.) Réblová, Chloridiopsis constrictospora (Crous et al.) Réblová, Geniculoseta preussii (W. Gams & Hol.-Jech.) Réblová, Papillospora hebetiseta (Réblová & W. Gams) Réblová, Spicatispora carpatica (Hol.-Jech. & Révay) Réblová, Spicatispora fennica (P. Karst.) Réblová; Epitypifications (basionyms): Chaetosphaeria dilabens Réblová & W. Gams, Chloridium cylindrosporum W. Gams & Hol.-Jech. Citation: Réblová M, Nekvindová J (2023). New genera and species with chloridium-like morphotype in the Chaetosphaeriales and Vermiculariopsiellales. Studies in Mycology 106: 199-258. doi: 10.3114/sim.2023.106.04.
RESUMEN
BACKGROUND: Hirsutella Pat genus, the asexual morphs of the Ophiocordyceps Sung, is globally distributed entomopathogenic fungi, which infect a variety of arthropods, mites and nematodes. The fungal species also have shown potential application in the field of biological control, bio-medicine and food development. Although these fungi are synonymized under Ophiocordyceps, formal taxonomic assignments remain necessary for classification of species in Hirsutella. However, due to the heterogeneity and complexity of Hirsutella genus, more detailed taxonomic and phylogenetic analyses are required to address the following subjects: (1) the relationships between the phialide morphological characteristics and phylogenetic information of Hirsutella with asexual morphs, (2) the origin and evolution of the phialide structure, and (3) host specificity and fungal pathogenicity. RESULTS: Five typical phialide structures are summarized, in which the variation in phialide characteristics overlaps well with phylogenetic information. A new member of the special twisted neck clade in the Hirsutella-like group, Ophiocordyceps retorta, was reported based on these analyses. The molecular clock calibration analysis based on one fossil record revealed that Hirsutella (asexual morph) species originated from a common ancestor approximately 102 million years ago (Mya) (Early Cretaceous, Lower Albian) and then resolved into two major lineages. One lineage was typically phialidic, which was a larger shape, including H. guyana, H. nodulosa and H. sinensis clades (86.9 Mya, 95% highest posterior density (HPD): 69.1-101.4 Mya). Another main lineage of the phialides was more diversified and smaller than the former, which included H. citriformis and H. thompsonii clades (71.9 Mya, 95% HPD: 41.8-99.6 Mya). CONCLUSIONS: Our results showed that certain phialide characteristics of Hirsutella were phylogenetically informative for two groups of taxa. The differentiation of the phialides structures in the major clades demonstrated a clear evolutionary path of Hirsutella (asexual morph) species, which exhibited two trends depending on the host size. Fungi in one of the groups displayed elongated conidiogenous cells with increased complexity of auxiliary structures from the mycelia. The species in another group reduced the volume of phialides and spores, which might be due to an energy-efficient strategy. These results suggested that a common origin allowed for diversification of given clades into separate niches. The distinct parallel evolutionary path combined with the specific phialides structure might result in the host specificity of Hirsutella (asexual morphs). A direct relationship between Hirsutella (asexual morphs) and the Cretaceous-Tertiary extinction was not found, which suggested that the diversity of phialides is more likely to be caused by long-term environmental adaptation and evolution rather than dramatic extinction events. This evolutionary result might correspond to the background of important biological and geological events in the late Cretaceous occurring near the divergence times of Hirsutella (asexual morphs).
Asunto(s)
Evolución Biológica , Hypocreales/fisiología , Animales , Fósiles , Hypocreales/clasificación , Filogenia , Especificidad de la Especie , Factores de TiempoRESUMEN
A spore assay was developed to measure the relative density of spores of the nematophagous fungus Hirsutella rhossiliensis in soil. Orchard soil containing H. rhossiliensis-parasitized Criconemella xenoplax was placed in vials and incubated for 0-120 days before the addition of probe nematodes, Heterorhabditis heliothidis juveniles. After 18 hours, H. heliothidis were extracted from the soil and examined for adhering spores of H. rhossiliensis. No spores were detected when H. heliothidis were added to freshly mixed soil, but the percentage of H. heliothidis with spores increased rapidly if soil was incubated undisturbed. Because mixing soil detaches spores from phialides, the results indicate that spores must be attached to phialides to adhere to nematodes. The spore assay was compared with a plate assay that measures the population density of H. rhossiliensis-parasitized C. xenoplax. Results from the two assays were highly correlated, suggesting that spores occur in three phases: reserves in nematodes that may be converted into spores; spores on phialides and therefore capable of adhering to nematodes; and spores detached from phialides and thus incapable of adhering to nematodes.