# Wallemiomycetes

**Wallemiomycetes** is a class of basidiomycete fungi erected in 2005 for the xerophilic mold genus *Wallemia* and its relatives, recognized on the basis of a distinctive mode of conidiogenesis (how a fungus forms and releases its asexual spores) and extreme tolerance of low water activity<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup>. The class contains the order Wallemiales and the family Wallemiaceae, and sits within the subphylum Wallemiomycotina of the phylum [Basidiomycota](https://www.edgepedia.org/basidiomycota)<sup>[2](https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=148960)</sup>. Its species are among the most salt- and sugar-tolerant eukaryotes known, and one member, *Wallemia ichthyophaga*, is described as the most halophilic fungus known to date<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup>.

| Key fact | Detail |
|---|---|
| Rank and authorship | Class Wallemiomycetes Zalar, de Hoog & Schroers, 2005, with the order Wallemiales, in the phylum Basidiomycota<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup> |
| Higher placement | Subphylum Wallemiomycotina; lineage: Basidiomycota, Wallemiomycotina, Wallemiomycetes, Wallemiales, Wallemiaceae, *Wallemia*<sup>[2](https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=148960)</sup> |
| Defining traits | Unique conidiogenesis with basauxic fertile hyphae and conidial units disarticulating into mostly four arthrospore-like conidia; extreme xerotolerance<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup> |
| Species at establishment | Three species segregated in 2005: *W. ichthyophaga*, *W. sebi*, and *W. muriae*<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup> |
| Divergence times | Subphylum Wallemiomycotina estimated to have split away approximately 487 million years ago; whole-proteome analysis dates Wallemiomycetes as a 250-million-year-old sister group of Agaricomycotina<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC6027281/)</sup><sup> • </sup><sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup> |
| Record physiology | *W. ichthyophaga* requires at least 10% NaCl and thrives in saturated salt solution<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup> |
| Compact genome | *W. ichthyophaga* genome 9.6 Mb with 1.67% repetitive sequences and 4,884 predicted protein-coding genes, among the smallest observed for fungi<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup> |

## Taxonomic history

The genus *Wallemia* was introduced by Johan-Olsen for *W. ichthyophaga*, discovered in 1885 on dried salted fish sent for examination by a fishery inspector named Mr. Wallem; *W. ichthyophaga* remained the only recognized species until von Arx recognized *Sporendonema sebi* as congeneric in 1970<sup>[5](https://pmc.ncbi.nlm.nih.gov/articles/PMC4446336/)</sup>. The taxonomic position of the genus remained uncertain for over a century. Terracina showed dolipore-like septal structures in *W. sebi*, and Moore interpreted these as a special kind of parenthesome, describing the family Wallemiaceae, first classified in the Filobasidiales within the Basidiomycota<sup>[6](https://pmc.ncbi.nlm.nih.gov/articles/PMC4370657/)</sup>.

**Resolution by sequencing.** Zalar, de Hoog, Schroers, Frank, and Gunde-Cimerman applied nuclear small subunit ribosomal DNA (SSU rDNA) analysis and found that *Wallemia* belonged to a highly supported clade with [Ustilaginomycetes](https://www.edgepedia.org/ustilaginomycetes) and Hymenomycetes, in an isolated position distantly related to the Filobasidiales and marked by numerous nucleotide substitutions not shared by any other fungus<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup>. To acknowledge its unique morphology, evolution, and xerotolerance, they proposed the new class Wallemiomycetes covering the new order Wallemiales<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup>. The same study segregated three species on differences in conidial size, xerotolerance, and ITS rDNA sequence, proposing the combination *Wallemia muriae* for *Torula epizoa* var. *muriae* and neotypifying all three<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup>.

The class then remained monogeneric until Nguyen and colleagues added two genera, *Geminibasidium* and *Basidioascus*, in a new order Geminibasidiales tentatively classified in Wallemiomycetes, although relatively distant from *Wallemia*<sup>[5](https://pmc.ncbi.nlm.nih.gov/articles/PMC4446336/)</sup>. Later treatments have variously ranked these three genera as distinct orders or classes, with *Wallemia* as sister to *Basidioascus* + *Geminibasidium*<sup>[7](https://pmc.ncbi.nlm.nih.gov/articles/PMC6899921/)</sup>.

## Phylogenetic position

Published studies have placed Wallemiomycetes at the base of the Basidiomycota, as a sister group to Agaricomycotina or Ustilaginomycotina or both, or simply as incertae sedis within the phylum<sup>[8](https://pmc.ncbi.nlm.nih.gov/articles/PMC3849046/)</sup>. A phylogenomic analysis aligning the whole proteomes of 14 fungi resolved the position of *Wallemia* as a sister group to, or the earliest diverging lineage of, Agaricomycotina, and supported Pucciniomycotina as the earliest diverging lineage among the Basidiomycota analyzed<sup>[8](https://pmc.ncbi.nlm.nih.gov/articles/PMC3849046/)</sup>. A 71-protein dataset reached the same placement, with Ustilaginomycotina as sister to the branch uniting Wallemiomycetes and the remaining Agaricomycotina<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC6027281/)</sup>. Genome-wide datasets preferentially support the sister relationship with Agaricomycotina, while ribosomal markers have given other topologies<sup>[7](https://pmc.ncbi.nlm.nih.gov/articles/PMC6899921/)</sup>.

The distinctiveness of the lineage was emphasized by its allocation to the newly distinguished subphylum Wallemiomycotina, estimated to have split away approximately 487 million years ago<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC6027281/)</sup>. The same whole-proteome chronogram dated the Wallemiomycetes stem as a 250-million-year-old sister group of Agaricomycotina<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup>. A 2024 phylogenomic outline of Basidiomycota treated Wallemiomycetes as one of the classes of the phylum within Wallemiomycotina<sup>[9](https://gredos.usal.es/bitstream/handle/10366/161602/He%282024%29_Phylogenomics%2C%20divergence%20times%20and%20notes%20of%20orders%20in%20Basidiomycota.pdf?isAllowed=y&sequence=1)</sup>.

## Ecology and physiology

On media with low water activity, *Wallemia* forms brown, powdery colonies, and the fungus is regarded as xerophilic or, at minimum, xerotolerant.<sup>[6](https://pmc.ncbi.nlm.nih.gov/articles/PMC4370657/)</sup> The genus is known from air, soil, dried food, and salt, and can be isolated from the hypersaline water of man-made salterns on different continents<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup>. Tests on xerotolerance indicated that *Wallemia* presents one of the most xerophilic fungal taxa, and xerotolerance is otherwise rare in the Basidiomycota<sup>[1](https://pubmed.ncbi.nlm.nih.gov/15928984/)</sup>.

The three species differ in their tolerance profiles. *W. sebi* can grow in a wide range of water activities, *W. muriae* grows only on media with slightly reduced water activities, and *W. ichthyophaga* is obligately halophilic, requiring water-activity-lowering solutes for in vitro growth<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. *W. ichthyophaga* requires at least 10% NaCl and thrives in saturated salt solution, and counterbalances the osmotic pressure of the surrounding medium by intracellular accumulation of a mixture of polyols, with glycerol as the major osmotically regulated solute<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup><sup> • </sup><sup>[8](https://pmc.ncbi.nlm.nih.gov/articles/PMC3849046/)</sup>. Morphological responses to high sugar concentrations differ from adaptation to high salt: *W. ichthyophaga* grew slowly in all sugar-based media, while *W. muriae* and *W. sebi* grew better<sup>[11](https://pubmed.ncbi.nlm.nih.gov/23931114/)</sup>.

## The genus Wallemia and its species

Beyond the three species recognized in 2005, a multi-locus phylogenetic approach resolved four additional species: *W. mellicola*, *W. canadensis*, and *W. tropicalis*, closely related to *W. sebi*, and *W. hederae*, phylogenetic sister of *W. ichthyophaga*<sup>[4](https://pmc.ncbi.nlm.nih.gov/articles/PMC6027281/)</sup>.

**The W. sebi species complex.** Previous ITS analyses had suggested that *W. sebi* was genetically heterogeneous, with at least two subclades<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. Multi-locus analyses using ITS and the protein-encoding genes MCM7, TSR1, RPB1, RPB2, and the new marker HAL2 confirmed that *W. sebi* presents a complex of at least four species, resolved as four strongly supported clades<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. The revision recognized *W. sebi* sensu stricto and described three new species, *W. mellicola*, *W. canadensis*, and *W. tropicalis*; the species differ in conidial size, xerotolerance, halotolerance, chaotolerance (tolerance to MgCl<sub>2</sub>), growth temperature regimes, extracellular enzyme activity profiles, and secondary metabolite patterns<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. Strains of the clade containing *W. sebi* sensu stricto typically originate from sea salt, hypersaline water, and air and dust in indoor environments worldwide<sup>[5](https://pmc.ncbi.nlm.nih.gov/articles/PMC4446336/)</sup>.

## Genomics and health relevance

The genome of the *W. ichthyophaga* type strain is unusually compact: 9.6 Mb with only 1.67% repetitive sequences, and 4,884 predicted protein-coding genes covering almost three quarters of the sequence, figures among the smallest observed for fungi<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup>. Of 639 differentially expressed genes across salinity limits, two thirds were more expressed at lower salinity<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup>. Contrary to the closely related *W. sebi*, *W. ichthyophaga* appears to have lost the ability for sexual reproduction<sup>[3](https://pubmed.ncbi.nlm.nih.gov/24034603/)</sup>. The species also has an unusually thick cell wall, reported as a possible mechanism of salt adaptation<sup>[8](https://pmc.ncbi.nlm.nih.gov/articles/PMC3849046/)</sup>.

**Health relevance.** *W. sebi* is the most frequently isolated and best-studied species, commonly involved in the spoilage of food with low water activity; its air-disseminated spores have been associated with allergies, bronchial asthma, and farmer's lung disease<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. [Human antibodies](https://www.edgepedia.org/human-antibodies) react to compounds produced by *W. sebi* spores, and metabolites present in spore and mycelial fragments caused inflammation and affected lung biology in an in vivo mouse model<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. The species produces the metabolites walleminol, walleminone, the azasteroid UCA1064-B, and the highly toxic wallimidione; walleminol is found in food contaminated by *W. sebi* and has an LD50 of 40 μg/ml for brine shrimp and a minimum inhibitory dose of 50 μg/ml for rat liver cells<sup>[10](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)</sup>. Species of *Wallemia* are among the few osmophilic fungi that can contaminate salted and dried cod<sup>[5](https://pmc.ncbi.nlm.nih.gov/articles/PMC4446336/)</sup>.

## References

1. [Zalar P, de Hoog GS, Schroers H-J, Frank JM, Gunde-Cimerman N (2005). Taxonomy and phylogeny of the xerophilic genus Wallemia (Wallemiomycetes and Wallemiales, cl. et ord. nov.). Antonie van Leeuwenhoek 87: 311–328.](https://pubmed.ncbi.nlm.nih.gov/15928984/)
2. [NCBI Taxonomy Browser: Wallemiomycetes](https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=148960)
3. [Zajc J and colleagues (2013). Genome and transcriptome sequencing of the halophilic fungus Wallemia ichthyophaga.](https://pubmed.ncbi.nlm.nih.gov/24034603/)
4. [The Genus Wallemia—From Contamination of Food to Health Threat, Microorganisms (2018)](https://pmc.ncbi.nlm.nih.gov/articles/PMC6027281/)
5. [Jančič S and colleagues (2015). A Taxonomic Revision of the Wallemia sebi Species Complex (PMC version)](https://pmc.ncbi.nlm.nih.gov/articles/PMC4446336/)
6. [Application of the Phylogenetic Species Concept to Wallemia sebi from House Dust and Indoor Air (PMC)](https://pmc.ncbi.nlm.nih.gov/articles/PMC4370657/)
7. [Fungal evolution: diversity, taxonomy and phylogeny of the Fungi, Biological Reviews (2019)](https://pmc.ncbi.nlm.nih.gov/articles/PMC6899921/)
8. [Genome and transcriptome sequencing of the halophilic fungus Wallemia ichthyophaga (PMC full text)](https://pmc.ncbi.nlm.nih.gov/articles/PMC3849046/)
9. [He and colleagues (2024). Phylogenomics, divergence times and notes of orders in Basidiomycota. Fungal Diversity.](https://gredos.usal.es/bitstream/handle/10366/161602/He%282024%29_Phylogenomics%2C%20divergence%20times%20and%20notes%20of%20orders%20in%20Basidiomycota.pdf?isAllowed=y&sequence=1)
10. [Jančič S and colleagues (2015). A Taxonomic Revision of the Wallemia sebi Species Complex. PLoS ONE 10(5): e0125933.](https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125933)
11. [Morphological responses to high sugar concentrations differ from adaptation to high salt concentrations in the xerophilic fungi Wallemia spp. Fungal Biology (2013).](https://pubmed.ncbi.nlm.nih.gov/23931114/)

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*Topic: Encyclopedia › Life and health › Microorganisms and fungi › Fungi and mycology › Basidiomycete taxa › Other basidiomycete classes › Wallemiomycetes*

*Initially written Sep 17, 2026 · Reviewed: Sep 19, 2026; Oct 21, 2026 · Edited: Sep 19, 2026; Oct 11, 2026; Oct 21, 2026 · Last review: Oct 21, 2026*

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