Holomycota | |
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Nuclearia | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Amorphea |
Clade: | Obazoa |
(unranked): | Opisthokonta |
Clade: | Holomycota Liu et al., 2009 |
Groups [1] | |
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Synonyms | |
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Holomycota or Nucletmycea are a basal Opisthokont clade as sister of the Holozoa. It consists of the Cristidiscoidea and the kingdom Fungi. The position of nucleariids, unicellular free-living phagotrophic amoebae, [3] 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, [4] Microsporidia and Cryptomycota, three groups of endoparasites. [5]
Rozella (Cryptomycota) is the earliest diverging fungal genus in which chitin has been observed at least in some stages of their life cycle, [5] although the chitinous 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. [6] The groups of fungi with the characteristic hyphal growth, Zoopagomycota, Mucoromycotina and Dikarya, originated from a common ancestor ~700 Mya. [6] Zoopagomycota are mostly pathogens of animals or other fungi, Mucoromycotina is a more diverse group including parasites, saprotrophs or ectomycorrhizal. [5] Dikarya is the group embracing Ascomycota and Basidiomycota, which comprise ~98% of the described fungal species. [6] 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. [6] Contrary to animals and land plants with complex multicellularity, the inferred phylogenetic relationships indicate that fungi acquired and lost multicellularity multiple times along Ascomycota and Basidiomycota evolution. [7]
The phylogenetic tree depicts the fungi and their close relationship to other organisms, based on the work of Philippe Silar [8] and "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research". [9] [10] The holomycota tree is following Tedersoo et al. [11] [12] [13]
Opisthokonts |
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Nucleariida is a group of amoebae with filose pseudopods, known mostly from soils and freshwater. They are distinguished from the superficially similar vampyrellids mainly by having mitochondria with discoid cristae, in the absence of superficial granules, and in the way they consume food.
The opisthokonts are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and Breviata comprise the larger clade Obazoa.
Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or protists, but are now known to be fungi, or a sister group to fungi. These fungal microbes are obligate eukaryotic parasites that use a unique mechanism to infect host cells. They have recently been discovered in a 2017 Cornell study to infect Coleoptera on a large scale. So far, about 1500 of the probably more than one million species are named. Microsporidia are restricted to animal hosts, and all major groups of animals host microsporidia. Most infect insects, but they are also responsible for common diseases of crustaceans and fish. The named species of microsporidia usually infect one host species or a group of closely related taxa. Approximately 10 percent of the species are parasites of vertebrates —several species, most of which are opportunistic, can infect humans, in whom they can cause microsporidiosis.
Amorphea is a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the members of this clade were originally described and proposed by Thomas Cavalier-Smith in 2002.
Cristidiscoidea or Nucleariae is a proposed basal holomycota clade in which Fonticula and Nucleariida emerged, as sister of the fungi. Since it is close to the divergence between the main lineages of fungi and animals, the study of Cristidiscoidea can provide crucial information on the divergent lifestyles of these groups and the evolution of opisthokonts and slime mold multicellularity. The holomycota tree is following Tedersoo et al.
The Apusozoa are a paraphyletic phylum of flagellate eukaryotes. They are usually around 5–20 μm in size, and occur in soils and aquatic habitats, where they feed on bacteria. They are grouped together based on the presence of an organic shell or theca under the dorsal surface of the cell.
Capsaspora is a monotypic genus containing the single species Capsaspora owczarzaki. C. owczarzaki is a single-celled eukaryote that occupies a key phylogenetic position in our understanding of the origin of animal multicellularity, as one of the closest unicellular relatives to animals. It is, together with Ministeria vibrans, a member of the Filasterea clade. This amoeboid protist has been pivotal to unravel the nature of the unicellular ancestor of animals, which has been proved to be much more complex than previously thought.
Filasterea is a proposed basal Filozoan clade of single-celled ameboid eukaryotes that includes Ministeria and Capsaspora. It is a sister clade to the Choanozoa in which the Choanoflagellatea and Animals appeared, originally proposed by Shalchian-Tabrizi et al. in 2008, based on a phylogenomic analysis with dozens of genes. Filasterea was found to be the sister-group to the clade composed of Metazoa and Choanoflagellata within the Opisthokonta, a finding that has been further corroborated with additional, more taxon-rich, phylogenetic analyses.
Holozoa is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa and the protists Choanoflagellata, Filasterea, Pluriformea and Ichthyosporea. Along with fungi and some other groups, Holozoa is part of the Opisthokonta, a supergroup of eukaryotes. Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic.
The apusomonads are a group of protozoan zooflagellates that glide on surfaces, and mostly consume prokaryotes. They are of particular evolutionary interest because they appear to be the sister group to the Opisthokonts, the clade that includes both animals and fungi. Together with the Breviatea, these form the Obazoa clade.
Ancyromonadida or Planomonadida is a small group of biflagellated protists found in the soil and in aquatic habitats, where they feed on bacteria. Includes freshwater or marine organisms, benthic, dorsoventrally compressed and with two unequal flagellae, each emerging from a separate pocket. The apical anterior flagellum can be very thin or end in the cell membrane, while the posterior flagellum is long and is inserted ventrally or laterally. The cell membrane is supported by a thin single-layered theca and the mitochondrial crests are discoidal/flat.
The Filozoa are a monophyletic grouping within the Opisthokonta. They include animals and their nearest unicellular relatives.
Cryptomycota , Rozellida, or Rozellomycota are a clade of micro-organisms that are either fungi or a sister group to fungi. They differ from classical fungi in that they lack chitinous cell walls at any trophic stage in their lifecycle, as reported by Jones and colleagues in 2011. Despite their unconventional feeding habits, chitin has been observed in the inner layer of resting spores, and in immature resting spores for some species of Rozella, as indicated with calcofluor-white stain as well as the presence of a fungal-specific chitin synthase gene.
The following outline is provided as an overview of and topical guide to life forms:
Aphelida is a phylum of Fungi that appears to be the sister to true fungi.
Opisthosporidia is a superphylum of intracellular parasites with amoeboid vegetative stage, defined as a common group of eukaryotic groups Microsporidia, Cryptomycota and Aphelidea. They have been considered to represent a monophyletic lineage with shared ecological and structural features, being a sister clade of the Fungi. Together with the Fungi they represent a sister clade of the Cristidiscoidea, together forming the Holomycota.
Pirum gemmata is a unicellular eukaryote that belongs to the Ichthyosporea clade, a group of protists closely related to animals. P. gemmata was isolated from the gut contents of a marine invertebrate, specifically the detritivorous peanut worm Phascolosoma agassizii.
Parvularia atlantis is a filopodiated amoeba which was isolated from a lake in Atlanta and deposited in the American Type Culture Collection (ATCC) under the name Nuclearia sp. ATCC 50694 on 1997 by TK Sawyer. It was classified under the genus Nuclearia and morphologically resembles to Nuclearia species, although it is smaller. Later it was determined that it phylogenetically belongs to a new nucleariid lineage., distantly related to Nuclearia and Fonticula genera – the other two previously described nucleriid genera.
Aphelidium tribonemae is a species within the Aphelid group. Their classification in the kingdom Fungi is a subject of controversy. Some argue for the classification of aphelids as ‘fungal animals', and for a period of time in the 1950s, aphids were classified as protists due to their amoeboid stage. Recently, molecular phylogenetics placed the aphelids within Opisthosporidia, a super phylum within Opisthokonta. Aphelids have posterior uniflagellate zoospores which place them as Opisthokonts. They are an early diverging lineage in Kingdom Fungi. While the aphelid group only contains three genera, it spans many both freshwater and marine ecosystems.
An amoeboflagellate is any eukaryotic organism capable of behaving as an amoeba and as a flagellate at some point during their life cycle. Amoeboflagellates present both pseudopodia and at least one flagellum, often simultaneously.