The subphylum Saccharomycotina is a lineage in the fungal phylum Ascomycota that exhibits levels of genomic diversity similar to those of plants and animals. The Saccharomycotina consist of more than 1 200 known species currently divided into 16 families, one order, and one class. Species in this subphylum are ecologically and metabolically diverse and include important opportunistic human pathogens, as well as species important in biotechnological applications. Many traits of biotechnological interest are found in closely related species and often restricted to single phylogenetic clades. However, the biotechnological potential of most yeast species remains unexplored. Although the subphylum Saccharomycotina has much higher rates of genome sequence evolution than its sister subphylum, Pezizomycotina , it contains only one class compared to the 16 classes in Pezizomycotina . The third subphylum of Ascomycota , the Taphrinomycotina , consists of six classes and has approximately 10 times fewer species than the Saccharomycotina . These data indicate that the current classification of all these yeasts into a single class and a single order is an underappreciation of their diversity. Our previous genome-scale phylogenetic analyses showed that the Saccharomycotina contains 12 major and robustly supported phylogenetic clades; seven of these are current families ( Lipomycetaceae , Trigonopsidaceae , Alloascoideaceae , Pichiaceae , Phaffomycetaceae , Saccharomycodaceae , and Saccharomycetaceae ), one comprises two current families ( Dipodascaceae and Trichomonascaceae ), one represents the genus Sporopachydermia , and three represent lineages that differ in their translation of the CUG codon (CUG-Ala, CUG-Ser1, and CUG-Ser2). Using these analyses in combination with relative evolutionary divergence and genome content analyses, we propose an updated classification for the Saccharomycotina , including seven classes and 12 orders that can be diagnosed by genome content. This updated classification is consistent with the high levels of genomic diversity within this subphylum and is necessary to make the higher rank classification of the Saccharomycotina more comparable to that of other fungi, as well as to communicate efficiently on lineages that are not yet formally named.
more »
« less
This content will become publicly available on May 18, 2026
Orders of Ascomycota
Ascomycota, the most speciose phylum of fungi, is a complex entity, comprising three diversesubphyla: Pezizomycotina, Saccharomycotina, and Taphrinomycotina. The largest and most diversesubphylum, Pezizomycotina, is a rich tapestry of 16 classes and 171 orders. Saccharomycotina, thesecond largest subphylum, is a diverse collection of seven classes and 12 orders, whileTaphrinomycotina, the smallest, is a unique assembly of six classes and six orders. Over the pastdecade, numerous taxonomic studies have focused on the generic, family, and class classifications ofAscomycota. These efforts, well-documented across various databases, are crucial for acomprehensive understanding of the classification. However, the study of taxonomy at the ordinallevel, a crucial tier in the taxonomic hierarchy, has been largely overlooked. In a global collaborationwith mycologists and lichenologists, this study presents the first comprehensive information on theorders within Pezizomycotina and Taphrinomycotina. The recent taxonomic classification ofSaccharomycotina has led to the exclusion of this subphylum from the present study, as an immediaterevision is not necessary. Each order is thoroughly discussed, highlighting its historical significance,current status, key identification characteristics, evolutionary relationships, ecological and economicroles, future recommendations, and updated family-level classification. Teaching diagrams for thelife cycles of several orders, viz. Asterinales, Helotiales, Hypocreales, Laboulbeniales, Meliolales,Mycosphaerellales, Ophiostomatales, Pezizales, Pleosporales, Phyllachorales, Rhytismatales,Sordariales, Venturiales, Xylariales (Pezizomycotina) and Pneumocystidales,Schizosaccharomycetales and Taphrinales (Taphrinomycotina) are provided. Each diagram is explained with a representative genus/genera of their sexual and asexual cycles of each order. WithinPezizomycotina, Dothideomycetes contains the highest number of orders, with 57, followed bySordariomycetes (52 orders), Lecanoromycetes (21 orders), Eurotiomycetes and Leotiomycetes (12orders each), Laboulbeniomycetes (3 orders), and Arthoniomycetes and Xylonomycetes (2 orderseach). Candelariomycetes, Coniocybomycetes, Geoglossomycetes, Lichinomycetes, Orbiliomycetes,Pezizomycetes, Sareomycetes, and Xylobotryomycetes each contain a single order, whileThelocarpales and Vezdaeales are treated as incertae sedis within Pezizomycotina. Notably, theclasses Candelariomycetes, Coniocybomycetes, Geoglossomycetes, Sareomycetes, andXylonomycetes, all recently grouped under Lichinomycetes, are treated as separate classes based onphylogenetic analysis and current literature. Within Lecanoromycetes, the synonymization ofSporastatiales with Rhizocarpales and Sarrameanales with Schaereriales is not supported in thephylogenetic analysis. These orders are retained separately, and the justifications are provided undereach section as well as in the discussion. Within Leotiomycetes, the order Medeolariales, which wasonce considered part of Helotiales, is treated as a distinct order based on phylogenetic evidence. Theclassification of Medeolariales may change as more data becomes available from different generegions. Lahmiales (Leotiomycetes) is not included in the phylogenetic analysis due to a lack ofmolecular data. Sareomycetes and Xylonomycetes are treated as separate classes. Spathulosporamixed with Lulworthiales and the inclusion of Spathulosporales within Lulworthiomycetidae issupported and extant molecular sampling is important to resolve the phylogenetic boundaries ofmembers of this subclass. The majority of the classes of Pezizomycotina and Taphrinomycotinaformed monophyletic clades in the phylogenetic analysis conducted based on SSU, LSU, 5.8S, TEFand RPB2 sequence data. However, Arthoniomycetes nested with the basal lineage ofDothideomycetes and formed a monophyletic clade also known as the superclass, Dothideomyceta.In Taphrinomycotina, a single order is accepted within each class.
more »
« less
- Award ID(s):
- 2018215
- PAR ID:
- 10618822
- Author(s) / Creator(s):
- ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; more »
- Publisher / Repository:
- Mycosphere
- Date Published:
- Journal Name:
- mycosphere
- Volume:
- 16
- Issue:
- 1
- ISSN:
- 2077-7019
- Page Range / eLocation ID:
- 536 to 1411
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
-
Abstract Botrytis cinereaPers. Fr. (teleomorph:Botryotinia fuckeliana) is a necrotrophic fungal pathogen that attacks a wide range of plants. This updated pathogen profile explores the extensive genetic diversity ofB. cinerea, highlights the progress in genome sequencing, and provides current knowledge of genetic and molecular mechanisms employed by the fungus to attack its hosts. In addition, we also discuss recent innovative strategies to combatB. cinerea. TaxonomyKingdom: Fungi, phylum: Ascomycota, subphylum: Pezizomycotina, class: Leotiomycetes, order: Helotiales, family: Sclerotiniaceae, genus:Botrytis, species:cinerea. Host rangeB. cinereainfects almost all of the plant groups (angiosperms, gymnosperms, pteridophytes, and bryophytes). To date, 1606 plant species have been identified as hosts ofB. cinerea. Genetic diversityThis polyphagous necrotroph has extensive genetic diversity at all population levels shaped by climate, geography, and plant host variation. PathogenicityGenetic architecture of virulence and host specificity is polygenic using multiple weapons to target hosts, including secretory proteins, complex signal transduction pathways, metabolites, and mobile small RNA. Disease control strategiesEfforts to controlB. cinerea, being a high‐diversity generalist pathogen, are complicated. However, integrated disease management strategies that combine cultural practices, chemical and biological controls, and the use of appropriate crop varieties will lessen yield losses. Recently, studies conducted worldwide have explored the potential of small RNA as an efficient and environmentally friendly approach for combating grey mould. However, additional research is necessary, especially on risk assessment and regulatory frameworks, to fully harness the potential of this technology.more » « less
-
ABSTRACT Yeasts in the subphylum Saccharomycotina are found across the globe in disparate ecosystems. A major aim of yeast research is to understand the diversity and evolution of ecological traits, such as carbon metabolic breadth, insect association, and cactophily. This includes studying aspects of ecological traits like genetic architecture or association with other phenotypic traits. Genomic resources in the Saccharomycotina have grown rapidly. Ecological data, however, are still limited for many species, especially those only known from species descriptions where usually only a limited number of strains are studied. Moreover, ecological information is recorded in natural language format limiting high throughput computational analysis. To address these limitations, we developed an ontological framework for the analysis of yeast ecology. A total of 1,088 yeast strains were added to the Ontology of Yeast Environments (OYE) and analyzed in a machine‐learning framework to connect genotype to ecology. This framework is flexible and can be extended to additional isolates, species, or environmental sequencing data. Widespread adoption of OYE would greatly aid the study of macroecology in the Saccharomycotina subphylum.more » « less
-
Mycoviruses are widely distributed across fungi, including the yeasts of the Saccharomycotina subphylum. This manuscript reports the first double-stranded RNA (dsRNA) virus isolated from Pichia membranifaciens. This novel virus has been named Pichia membranifaciens virus L-A (PmV-L-A) and is a member of the Totiviridae. PmV-L-A is 4579 bp in length, with RNA secondary structures similar to the packaging, replication, and frameshift signals of totiviruses that infect Saccharomycotina yeasts. PmV-L-A was found to be part of a monophyletic group within the I-A totiviruses, implying a shared ancestry between mycoviruses isolated from the Pichiaceae and Saccharomycetaceae yeasts. Energy-minimized AlphaFold2 molecular models of the PmV-L-A Gag protein revealed structural conservation with the Gag protein of Saccharomyces cerevisiae virus L-A (ScV-L-A). The predicted tertiary structure of the PmV-L-A Pol and other homologs provided a possible mechanism for totivirus RNA replication due to structural similarities with the RNA-dependent RNA polymerases of mammalian dsRNA viruses. Insights into the structure, function, and evolution of totiviruses gained from yeasts are essential because of their emerging role in animal disease and their parallels with mammalian viruses.more » « less
-
IntroductionEukaryotic life depends on the functional elements encoded by both the nuclear genome and organellar genomes, such as those contained within the mitochondria. The content, size, and structure of the mitochondrial genome varies across organisms with potentially large implications for phenotypic variance and resulting evolutionary trajectories. Among yeasts in the subphylum Saccharomycotina, extensive differences have been observed in various species relative to the model yeastSaccharomyces cerevisiae, but mitochondrial genome sampling across many groups has been scarce, even as hundreds of nuclear genomes have become available. MethodsBy extracting mitochondrial assemblies from existing short-read genome sequence datasets, we have greatly expanded both the number of available genomes and the coverage across sparsely sampled clades. ResultsComparison of 353 yeast mitochondrial genomes revealed that, while size and GC content were fairly consistent across species, those in the generaMetschnikowiaandSaccharomycestrended larger, while several species in the order Saccharomycetales, which includesS. cerevisiae, exhibited lower GC content. Extreme examples for both size and GC content were scattered throughout the subphylum. All mitochondrial genomes shared a core set of protein-coding genes for Complexes III, IV, and V, but they varied in the presence or absence of mitochondrially-encoded canonical Complex I genes. We traced the loss of Complex I genes to a major event in the ancestor of the orders Saccharomycetales and Saccharomycodales, but we also observed several independent losses in the orders Phaffomycetales, Pichiales, and Dipodascales. In contrast to prior hypotheses based on smaller-scale datasets, comparison of evolutionary rates in protein-coding genes showed no bias towards elevated rates among aerobically fermenting (Crabtree/Warburg-positive) yeasts. Mitochondrial introns were widely distributed, but they were highly enriched in some groups. The majority of mitochondrial introns were poorly conserved within groups, but several were shared within groups, between groups, and even across taxonomic orders, which is consistent with horizontal gene transfer, likely involving homing endonucleases acting as selfish elements. DiscussionAs the number of available fungal nuclear genomes continues to expand, the methods described here to retrieve mitochondrial genome sequences from these datasets will prove invaluable to ensuring that studies of fungal mitochondrial genomes keep pace with their nuclear counterparts.more » « less
An official website of the United States government
