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Holomycota

Holomycota or Nucletmyceta is the taxonomic group that includes fungi and its protists relatives: opisthosporidians and nucleariids [1],[2]. Holomycota, as well as Holozoa (the taxonomic group that includes animals and their unicellular relatives) compose the eukaryotic supergroup Opisthokonta [3],[4],[5]. The position of nucleariids, unicellular free-living phagotrophic amoebae [6], as the earliest lineage of Holomycota suggests that animals and fungi independently acquired complex multicellularity from a common unicellular ancestor and that the osmotrophic lifestyle (one of the fungal hallmarks) was originated later in the divergence of this eukaryotic lineage. Opisthosporidians is a recently proposed taxonomic group that includes aphelids [7], Microsporidia and Cryptomycota, three groups of endoparasites [8].

Interestingly, Rozella (Cryptomycota) is the earliest fungal genus in which chitin has been observed at least in some stages of their life cycle [8], although the chitinus cell wall (another fungal hallmark) and osmotrophy originated in a common ancestor of Blastocladiomycota and Chytridiomycota, which still contain some ancestral characteristics such as the flagellum in zoosporic stage [9]. The groups of fungi with the characteristic hyphal growth, Zoopagomycota, Mucuromycotina and Dikarya, originated from a common ancestor ~700 Mya [9]. Zoopagomycota are mostly pathogens of animals or other fungi, Mucuromycotina is a more diverse group including parasites, saprotrophs or ectomycorrhizal [8]. Dikarya is the group embracing Ascomycota and Basidiomycota, which comprise ~98% of the described fungal species [9]. Because of this rich diversity, Dikarya includes highly morphologically distinct groups, from hyphae or unicellular yeasts (such as the model organism Saccharomyces cerevisiae) to the complex multicellular fungi popularly known as mushrooms [9]. Interestingly and in opposite to animals and land plants, where complex multicellularity once, the inferred phylogenetic relationships indicate that fungi acquired and lost multicellularity multiple times along Ascomycota and Basidiomycota evolution [10].

References:

1-Liu, Y. et al. (2009). “Phylogenomic analyses predict sistergroup relationship of nucleariids and fungi and paraphyly of zygomycetes with significant support”. BMC Evol. Biol. 9: 272., doi:10.1186/1471-2148-9-272

2-Brown, M.W. et al. (2009). “Phylogeny of the “forgotten” cellular slime mold, Fonticula alba, reveals a key evolutionary branch within Opisthokonta”. Mol. Biol. Evol. 26 (12): 2699–709. PMID 19692665. doi:10.1093/molbev/msp185

3-Adl, S.M. et al. 2012. “The revised classification of eukaryotes”. Journal of Eukaryotic Microbiology, 59(5), 429-514. 10.1111/j.1550-7408.2012.00644.x

4-Cavalier-Smith, T. and E. E. Chao. 2003. “Phylogeny of Choanozoa, Apusozoa, and other protozoa and early eukaryote megaevolution”. J. Mol. Evol. 56:540-563. https://doi.org/10.1007/s00239-002-2424-z

5-Torruella, G. et al. (2015). “Phylogenomics reveals convergent evolution of lifestyles in close relatives of animals and fungi”. Current Biology 25 (18), 2404-2410. https://doi.org/10.1016/j.cub.2015.07.053

6-López-Escardó, D. et al. (2017), “Parvularia atlantis gen. et sp. nov., a Nucleariid Filose Amoeba (Holomycota, Opisthokonta)”. J. Eukaryot. Microbiol., doi:10.1111/jeu.12450

7- Karpov, S.A. et al (2014). “Morphology, phylogeny, and ecology of the aphelids (Aphelidea, Opisthokonta) and proposal for the new superphylum Opisthosporidia”. Front Microbiol. 5: 112. https://doi.org/10.3389/fmicb.2014.00112

8-Mary L. Berbee. et al. (2017). “Early Diverging Fungi: Diversity and Impact at the Dawn of Terrestrial Life”. Annual Review of Microbiology 71:1, 41-60. https://doi.org/10.1146/annurev-micro-030117-020324

9-Stajich, Jason E. et al. (2009). “The Fungi”. Current Biology 19 (18), R840 – R845. http://dx.doi.org/10.1016/j.cub.2009.07.004

10-Tu Anh Nguyen (2017). “Innovation and constraint leading to complex multicellularity in the Ascomycota”. Nature communications 8, doi:10.1038/ncomms14444

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