Choanozoa

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Choanozoa
Temporal range: 665–0 Ma
M2-2cells.jpg
Codonosiga
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Amorphea
Clade: Obazoa
(unranked): Opisthokonta
(unranked): Holozoa
(unranked): Filozoa
Clade: Choanozoa
Brunet and King, 2017
Subdivisions
Synonyms
  • Apoikozoa Budd & Jensen, 2015
  • Choanimal Fairclough et al., 2013

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]

Contents

Introduction

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.

Nomenclature

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]

Evolutionary implications

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
1300 mya

Related Research Articles

<span class="mw-page-title-main">Choanoflagellate</span> Group of eukaryotes considered the closest living relatives of animals

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.

<span class="mw-page-title-main">Opisthokont</span> Group of eukaryotes which includes animals and fungi, among other groups

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.

<span class="mw-page-title-main">Amorphea</span> Group including fungi, animals and various protozoa

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.

<span class="mw-page-title-main">Apusozoa</span> Phylum of micro-organisms

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.

<span class="mw-page-title-main">Viridiplantae</span> Clade of archaeplastids including green algae and the land plants

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.

<span class="mw-page-title-main">Protist</span> Eukaryotes other than animals, plants or fungi

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.

<i>Capsaspora</i> Single-celled eukaryote genus

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.

<i>Corallochytrium</i> Genus of unicellular organisms

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.

<span class="mw-page-title-main">Planulozoa</span> Clade of animals

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.

<span class="mw-page-title-main">SAR supergroup</span> Eukaryotes superphylum

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.

<span class="mw-page-title-main">Filasterea</span> Basal Filozoan 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.

<span class="mw-page-title-main">Holozoa</span> Clade containing animals and some protists

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.

<span class="mw-page-title-main">Apusomonadidae</span> Group of microorganisms with two flagella

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.

<span class="mw-page-title-main">Filozoa</span> Monophyletic grouping within the Opisthokonta

The Filozoa are a monophyletic grouping within the Opisthokonta. They include animals and their nearest unicellular relatives.

<span class="mw-page-title-main">Varisulca</span> Proposed phylum of protists

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.

<span class="mw-page-title-main">Pluriformea</span> Clade of unicellular organisms

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.

<i>Syssomonas</i> Genus of protists

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.

<span class="mw-page-title-main">Amoeboflagellate</span> Cellular body type

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.

References

  1. Laura Wegener Parfrey; Daniel J G Lahr; Andrew H Knoll; Laura A Katz (16 August 2011). "Estimating the timing of early eukaryotic diversification with multigene molecular clocks" (PDF). Proceedings of the National Academy of Sciences of the United States of America . 108 (33): 13624–9. Bibcode:2011PNAS..10813624P. doi:10.1073/PNAS.1110633108. ISSN   0027-8424. PMC   3158185 . PMID   21810989. Wikidata   Q24614721.
  2. King, N.; Westbrook, M. J.; Young, S. L.; Kuo, A.; Abedin, M.; Chapman, J.; Fairclough, S.; Hellsten, U.; Isogai, Y.; Letunic, I. (February 14, 2008). "The genome of the choanoflagellate Monosiga brevicollis and the origin of metazoans". Nature. 451 (7180): 783–788. Bibcode:2008Natur.451..783K. doi:10.1038/nature06617. PMC   2562698 . PMID   18273011.
  3. 1 2 Budd, G. E.; Jensen, S. (2015). "The origin of the animals and a 'Savannah' hypothesis for early bilaterian evolution". Biological Reviews. 92 (1): 446–473. doi: 10.1111/brv.12239 . PMID   26588818.
  4. 1 2 3 Adl, Sina M.; Bass, David; Lane, Christopher E.; Lukeš, Julius; Schoch, Conrad L.; Smirnov, Alexey; Agatha, Sabine; Berney, Cedric; Brown, Matthew W. (2018-09-26). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/jeu.12691. PMC   6492006 . PMID   30257078.
  5. Ryan, J. F. (December 13, 2013). "The genome of the ctenophore Mnemiopsis leidyi and its implications for cell type evolution". Science. 342 (6164): 1242592. doi:10.1126/science.1242592. PMC   3920664 . PMID   24337300.
  6. Pisani, D.; Pett, W.; Dohrmann, M.; Feuda, R.; Rota-Stabelli, O.; Philippe, H.; Lartillot, N. & Wörheide, G. (December 15, 2015). "Genomic data do not support comb jellies as the sister group to all other animals". Proceedings of the National Academy of Sciences. 112 (50): 15402–7. Bibcode:2015PNAS..11215402P. doi: 10.1073/pnas.1518127112 . PMC   4687580 . PMID   26621703.
  7. Cavalier-Smith T (May 2022). "Ciliary transition zone evolution and the root of the eukaryote tree: implications for opisthokont origin and classification of kingdoms Protozoa, Plantae, and Fungi". Protoplasma. 259 (3): 487–593. doi:10.1007/s00709-021-01665-7. PMC   9010356 . PMID   34940909.
  8. Peterson, Kevin J.; Cotton, James A.; Gehling, James G.; Pisani, Davide (2008-04-27). "The Ediacaran emergence of bilaterians: congruence between the genetic and the geological fossil records". Philosophical Transactions of the Royal Society of London B: Biological Sciences. 363 (1496): 1435–1443. doi:10.1098/rstb.2007.2233. PMC   2614224 . PMID   18192191.
  9. Parfrey, Laura Wegener; Lahr, Daniel J. G.; Knoll, Andrew H.; Katz, Laura A. (2011-08-16). "Estimating the timing of early eukaryotic diversification with multigene molecular clocks". Proceedings of the National Academy of Sciences. 108 (33): 13624–13629. Bibcode:2011PNAS..10813624P. doi: 10.1073/pnas.1110633108 . PMC   3158185 . PMID   21810989.
  10. Hehenberger, Elisabeth; Tikhonenkov, Denis V.; Kolisko, Martin; Campo, Javier del; Esaulov, Anton S.; Mylnikov, Alexander P.; Keeling, Patrick J. (2017). "Novel Predators Reshape Holozoan Phylogeny and Reveal the Presence of a Two-Component Signaling System in the Ancestor of Animals". Current Biology. 27 (13): 2043–2050.e6. doi: 10.1016/j.cub.2017.06.006 . PMID   28648822.
  11. Tedersoo, Leho; Sánchez-Ramírez, Santiago; Kõljalg, Urmas; Bahram, Mohammad; Döring, Markus; Schigel, Dmitry; May, Tom; Ryberg, Martin; Abarenkov, Kessy (2018). "High-level classification of the Fungi and a tool for evolutionary ecological analyses". Fungal Diversity. 90 (1): 135–159. doi: 10.1007/s13225-018-0401-0 . ISSN   1560-2745.