Choanozoa Temporal range: [1] | |
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Codonosiga | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Amorphea |
Clade: | Obazoa |
(unranked): | Opisthokonta |
(unranked): | Holozoa |
(unranked): | Filozoa |
Clade: | Choanozoa Brunet and King, 2017 |
Subdivisions | |
Synonyms | |
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Choanozoa is a clade of opisthokont eukaryotes consisting of the choanoflagellates (Choanoflagellatea) and the animals (Animalia, Metazoa). The sister-group relationship between the choanoflagellates and animals has important implications for the origin of the animals. [2] The clade was identified in 2015 by Graham Budd and Sören Jensen, who used the name Apoikozoa. [3] The 2018 revision of the classification first proposed by the International Society of Protistologists in 2012 recommends the use of the name Choanozoa. [4]
A close relationship between choanoflagellates and animals has long been recognised, dating back at least to the 1840s. A particularly striking and famous similarity between the single-celled choanoflagellates and multicellular animals is provided by the collar cells of sponges and the overall morphology of the choanoflagellate cell. The relationship has since been confirmed by multiple molecular analyses. This proposed homology was however thrown into some doubt in 2013 by the still controversial suggestion that ctenophores, and not sponges, are the sister group to all other animals. [5] [6] More recent genomic work has suggested that choanoflagellates possess some of the important genetic machinery necessary for the multicellularity found in animals.[ citation needed ]
A synonym for the Choanozoa, Apoikozoa, derives from the ancient Greek for "colony" and "animal", referring to the ability of both animals and (some) choanoflagellates to form multicellular units. [4] While animals are permanently multicellular, the colony-building choanoflagellates are only sometimes so, which raises the question of whether or not the colony-building ability in both groups was present at the base of the entire clade, or whether it was independently derived within the animals and choanoflagellates.
The name "Choanozoa" was first used by protozoologist Thomas Cavalier-Smith in 1991 to refer to a group of basal protists that later proved not to form a clade. This group had the rank of phylum and contained all opisthokont protists while excluding both fungi and animals, making the group paraphyletic. Its classification was the following: [7]
The International Society of Protistologists rejected the use of this name for the paraphyletic group. Instead, since 2017, the name Choanozoa is considered appropriate for the clade that unites choanoflagellates and animals, since the Greek choanē (χοάνη), meaning 'funnel', refers to the collar, which is a synapomorphy (i.e. a unique characteristic) of the clade. A synonym of this clade, Apoikozoa, was used in previous years; however, it was rejected as being neither formally defined nor appropriate, since it refers to the ability to form colonies, a characteristic not unique to this clade. [4]
Although the last common ancestor of the Choanozoa cannot be reconstructed with certainty, Budd and Jensen suggest that these organisms formed benthic colonies that competed for space amongst other mat-forming organisms known to have existed during the Ediacaran Period some 635–540 million years ago. As such they would form an important link between the unicellular ancestors of the animals and the enigmatic "Ediacaran" organisms known from this interval, thus allowing some sort of reconstruction of the earliest animals and their ecology. [3] In the following cladogram, an indication is given of approximately how many million years ago (Mya) the clades diverged into newer clades. [8] [9] [10] (Note that the later Budd and Jensen paper gives significantly younger dates. See also Kimberella .) The Holomycota tree follows Tedersoo et al. [11]
Opisthokonta |
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1300 mya |
The choanoflagellates are a group of free-living unicellular and colonial flagellate eukaryotes considered to be the closest living relatives of the animals. Choanoflagellates are collared flagellates, having a funnel shaped collar of interconnected microvilli at the base of a flagellum. Choanoflagellates are capable of both asexual and sexual reproduction. They have a distinctive cell morphology characterized by an ovoid or spherical cell body 3–10 μm in diameter with a single apical flagellum surrounded by a collar of 30–40 microvilli. Movement of the flagellum creates water currents that can propel free-swimming choanoflagellates through the water column and trap bacteria and detritus against the collar of microvilli, where these foodstuffs are engulfed. This feeding provides a critical link within the global carbon cycle, linking trophic levels. In addition to their critical ecological roles, choanoflagellates are of particular interest to evolutionary biologists studying the origins of multicellularity in animals. As the closest living relatives of animals, choanoflagellates serve as a useful model for reconstructions of the last unicellular ancestor of animals. According to a 2021 study, crown group craspedids appeared 422.78 million years ago, Although a previous study from 2017 recovered the divergence of the crown group choanoflagellates (craspedids) at 786.62 million years.
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.
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.
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.
Viridiplantae constitute a clade of eukaryotic organisms that comprises approximately 450,000–500,000 species that play important roles in both terrestrial and aquatic ecosystems. They include the green algae, which are primarily aquatic, and the land plants (embryophytes), which emerged from within them. Green algae traditionally excludes the land plants, rendering them a paraphyletic group. However it is accurate to think of land plants as a kind of alga. Since the realization that the embryophytes emerged from within the green algae, some authors are starting to include them. They have cells with cellulose in their cell walls, and primary chloroplasts derived from endosymbiosis with cyanobacteria that contain chlorophylls a and b and lack phycobilins. Corroborating this, a basal phagotroph archaeplastida group has been found in the Rhodelphydia.
A protist or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a polyphyletic grouping of several independent clades that evolved from the last eukaryotic common ancestor.
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 unraveling the nature of the unicellular ancestor of animals, which has been proved to be much more complex than previously thought.
Corallochytrium belongs to the class of Corallochytrea within Teretosporea and is a sister group to Ichthyosporea. Corallochytrium limacisporum is the only species of Corallochytrium known so far. It was first discovered and named in the Arabian Sea’s coral lagoons by Kaghu-Kumar in 1987. It was first thought to be a member of the fungi-like thraustochytrids, however, this was later disproven due to Corallochytriums lack of cilia and sagenogenetosome. Little research has been done on the life cycle or morphology. Most research concerning this genus has been done to uncover the evolution of animals and fungi, as Corallochytrium possess both animal and fungal enzymatic trademarks.
Planulozoa is a clade which includes the Placozoa, Cnidaria and the Bilateria. The designation Planulozoa may be considered a synonym to Parahoxozoa. Within Planulozoa, the Placozoa may be a sister of Cnidaria to the exclusion of Bilateria. The clade excludes basal animals such as the Ctenophora, and Porifera (sponges). Although this clade was sometimes used to specify a clade of Cnidaria and Bilateria to the exclusion of Placozoa, this is no longer favoured due to recent data indicating a sister group relationship between Cnidaria and Placozoa, another study still supports Placozoa as sister to Cnidaria+Bilateria.
SAR or Harosa is a highly diverse clade of eukaryotes, often considered a supergroup, that includes stramenopiles (heterokonts), alveolates, and rhizarians. It is a node-based taxon, including all descendants of the three groups' last common ancestor, and comprises most of the now-rejected Chromalveolata. Their sister group has been found to be telonemids, with which they make up the TSAR clade.
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 78 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.
The Filozoa are a monophyletic grouping within the Opisthokonta. They include animals and their nearest unicellular relatives.
Varisulca was a proposed basal Podiate taxon. It encompassed several lineages of heterotrophic protists, most notably the ancyromonads (planomonads), collodictyonids (diphylleids), rigifilids and mantamonadids. Recent evidence suggests that the latter three are closely related to each other, forming a clade called CRuMs, but that this is unlikely to be specifically related to ancyromonads.
The Scotokaryotes (Cavalier-Smith) is a proposed basal Neokaryote clade as sister of the Diaphoretickes. Basal Scotokaryote groupings are the Metamonads, the Malawimonas and the Podiata. In this phylogeny the Discoba are sometimes seen as paraphyletic and basal Eukaryotes.
Pluriformea is a proposed sibling clade of the Filozoa, and consists of Syssomonas multiformis and the Corallochytrea. Together with the Ichthyosporea they form the Holozoa.
Tunicaraptor is a genus of marine microbial protists containing the single species Tunicaraptor unikontum, discovered in 2020 from marine waters of Chile. It is a lineage of predatorial flagellates closely related to animals. It has a rare feeding structure not seen in other opisthokonts.
Syssomonas is a monotypic genus of unicellular flagellated protists containing the species Syssomonas multiformis. It is a member of Pluriformea inside the lineage of Holozoa, a clade containing animals and their closest protistan relatives. It lives in freshwater habitats. It has a complex life cycle that includes unicellular amoeboid and flagellated phases, as well as multicellular aggregates, depending on the growth medium and nutritional state.
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.