SAR Temporal range: | |
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Scientific classification | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
Clade: | TSAR |
Clade: | SAR Burki et al., 2007 |
Subtaxa | |
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Synonyms | |
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SAR or Harosa is a highly diverse clade of eukaryotes, often considered a supergroup, [2] that includes stramenopiles (heterokonts), alveolates, and rhizarians. [3] [4] [5] It is a node-based taxon, including all descendants of the three groups' last common ancestor, [6] and comprises most of the now-rejected Chromalveolata. [2] Their sister group has been found to be telonemids, with which they make up the TSAR clade. [7] [nb 1]
The name SAR is an acronym derived from the first letters of its three constituent clades; [nb 2] it has been alternatively spelled "RAS". [6] [10] The term "Harosa" (at the subkingdom level) has also been used, with Stramenopiles replaced by its synonym Heterokonta in this variant of the acronym. [11]
Before the discovery of the SAR supergroup, stramenopiles and alveolates were classified in the supergroup Chromalveolata alongside haptophytes and cryptomonads, being believed to have acquired plastids through secondary endosymbiosis of red algae through a common ancestor. [2] Meanwhile, Rhizaria was traditionally considered to be a separate supergroup. More recent phylogenetic studies confirmed that stramenopiles and alveolates diverged with rhizarians as part of the SAR lineage. [12] This clade has been found by later phylogenomic studies to be robustly characterized compared to other supergroups. [7]
This groups excludes haptophytes and cryptomonads, hypothesized to have acquired plastids in separate endosymbiosis events, [13] leading Okamoto et al. (2009) to propose the clade Hacrobia to accommodate them. [14]
The SAR supergroup encompasses a variety of morphologies and ecological niches, from microscopic zoo- and phytoplankton to massive kelp forests. The group includes both photosynthetic and non-photosynthetic forms. Photosynthesis arose independently across various stramenopiles and alveolates lineages through secondary or higher-order endosymbiosis events, acquiring plastids of red algal origin, [15] [13] while chlorarachniophyte rhizarians captured plastids from green algae, retaining vestigial nucleomorphs. [16]
It has been estimated that SAR encompasses up to half of all eukaryotic diversity. [2]
Owing to the clade's discovery through phylogenomics, there are no known synapomorphies uniting its various members. [3] This was already the case for its subclade Rhizaria, established earlier through similar means. On the other hand, Stramenopiles is well-defined morphologically, characterized by an anterior flagellum with tripartite bristles (mastigonemes), while Alveolata is united by the presence of cortical alveoli. [17]
Nonetheless, studies of telonemids, believed to be the sister group to SAR, have revealed characteristics such as tripartite hair and peripheral vacuoles, potentially homologous to similar structures in stramenopiles and alveolates. This brings into light the possibility of these structures being ancestrally shared by the clade, with cortical alveoli originating from peripheral vacuoles under this hypothesis. [7]
A 2021 analysis places Alveolata and Stramenopiles in Halvaria, as sister to Rhizaria. [13]
TSAR |
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The Stramenopiles, also called Heterokonts, are a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have been secondarily lost. Stramenopiles represent one of the three major clades in the SAR supergroup, along with Alveolata and Rhizaria.
The alveolates are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with tubulocristate mitochondria into the SAR supergroup.
Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota. The group was first suggested by Simpson and Patterson in 1999 and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged. They are considered to be a basal flagellate lineage.
Chromista is a proposed but polyphyletic biological kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all eukaryotes whose plastids contain chlorophyll c and are surrounded by four membranes. If the ancestor already possessed chloroplasts derived by endosymbiosis from red algae, all non-photosynthetic Chromista have secondarily lost the ability to photosynthesise. Its members might have arisen independently as separate evolutionary groups from the last eukaryotic common ancestor.
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. They are the natural predators of many species of bacteria. They are closely related to the phylum Retaria, comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria.
The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthetic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon.
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.
A bikont is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled and multi-celled organisms are members of the group, and these, as well as the presumed ancestor, have two flagella.
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in 1981. Chromalveolata was proposed to represent the organisms descended from a single secondary endosymbiosis involving a red alga and a bikont. The plastids in these organisms are those that contain chlorophyll c.
The Archaeplastida are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It also includes the non-photosynthetic lineage Rhodelphidia, a predatorial (eukaryotrophic) flagellate that is sister to the Rhodophyta, and probably the microscopic picozoans. The Archaeplastida have chloroplasts that are surrounded by two membranes, suggesting that they were acquired directly through a single endosymbiosis event by phagocytosis of a cyanobacterium. All other groups which have chloroplasts, besides the amoeboid genus Paulinella, have chloroplasts surrounded by three or four membranes, suggesting they were acquired secondarily from red or green algae. Unlike red and green algae, glaucophytes have never been involved in secondary endosymbiosis events.
In the classification of eukaryotes, Cabozoa was a taxon proposed by Cavalier-Smith. It was a putative clade comprising the Rhizaria and Excavata. More recent research places the Rhizaria with the Alveolata and Stramenopiles instead of the Excavata, however, so "Cabozoa" is polyphyletic.
Telonemia is a phylum of microscopic eukaryotes commonly known as telonemids. They are unicellular free-living flagellates with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes.
Ochrophytes, also known as heterokontophytes or stramenochromes, are a group of algae. They are the photosynthetic stramenopiles, a group of eukaryotes, organisms with a cell nucleus, characterized by the presence of two unequal flagella, one of which has tripartite hairs called mastigonemes. In particular, they are characterized by photosynthetic organelles or plastids enclosed by four membranes, with membrane-bound compartments called thylakoids organized in piles of three, chlorophyll a and c as their photosynthetic pigments, and additional pigments such as β-carotene and xanthophylls. Ochrophytes are one of the most diverse lineages of eukaryotes, containing ecologically important algae such as brown algae and diatoms. They are classified either as phylum Ochrophyta or Heterokontophyta, or as subphylum Ochrophytina within phylum Gyrista. Their plastids are of red algal origin.
The cryptomonads-haptophytes assemblage is a proposed but disputed monophyletic grouping of unicellular eukaryotes that are not included in the SAR supergroup. Several alternative names have been used for the group, including Hacrobia ; CCTH ; and "Eukaryomonadae".
Diaphoretickes is a major group of eukaryotic organisms, with over 400,000 species. The majority of the earth's biomass that carries out photosynthesis belongs to Diaphoretickes.
Halvaria is a taxonomic grouping of protists that includes Alveolata and Stramenopiles (Heterokonta).
Cryptista is a clade of alga-like eukaryotes. It is most likely related to Archaeplastida which includes plants and many algae, within the larger group Diaphoretickes.
Haptista is a proposed group of protists made up of centrohelids and haptophytes. Phylogenomic studies indicate that Haptista, together with Ancoracysta twista, forms a sister clade to the SAR+Telonemia supergroup, but it may also be sister to the Cryptista (+Archaeplastida). It is thus one of the earliest diverging Diaphoretickes.
The cortical alveolum is a cellular organelle consisting of a vesicle located under the cytoplasmic membrane, to which they give support. The term "corticate" comes from an evolutionary hypothesis about the common origin of kingdoms Plantae and Chromista, because both kingdoms have cortical alveoli in at least one phylum. At least three protist lineages exhibit these structures: Telonemia, Alveolata and Glaucophyta.
A supergroup, in evolutionary biology, is a large group of organisms that share one common ancestor and have important defining characteristics. It is an informal, mostly arbitrary rank in biological taxonomy that is often greater than phylum or kingdom, although some supergroups are also treated as phyla.
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