SAR is a highly diverse

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

s, often considered a supergroup, that includes

stramenopile The stramenopiles, also called heterokonts, are protists 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, an ...

s (heterokonts), alveolates, and rhizarians. It is a node-based

taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...

(under the Sar name), 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. Harosa is sometimes used synonymously with TSAR.

Etymology

The name SAR is an

acronym An acronym is a type of abbreviation consisting of a phrase whose only pronounced elements are the initial letters or initial sounds of words inside that phrase. Acronyms are often spelled with the initial Letter (alphabet), letter of each wor ...

derived from the first letters of its three constituent clades; it has been alternatively spelled RAS. The term Harosa (at the subkingdom level) has also been used, with Stramenopiles replaced by its synonym Heterokonta in this variant of the acronym.

History of discovery

Before the discovery of the SAR supergroup,

stramenopiles The stramenopiles, also called heterokonts, are Protist, protists 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 sur ...

and alveolates were classified in the supergroup Chromalveolata alongside haptophytes and cryptomonads, being believed to have acquired plastids through secondary endosymbiosis of

red algae Red algae, or Rhodophyta (, ; ), make up one of the oldest groups of eukaryotic algae. The Rhodophyta comprises one of the largest Phylum, phyla of algae, containing over 7,000 recognized species within over 900 Genus, genera amidst ongoing taxon ...

through a common ancestor. Meanwhile,

Rhizaria The Rhizaria are a diverse and species-rich clade 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 ...

was traditionally considered to be a separate supergroup. More recent

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

studies confirmed that stramenopiles and alveolates diverged with rhizarians as part of the SAR lineage. This clade has been found by later phylogenomic studies to be robustly characterized compared to other supergroups. This group excludes haptophytes and cryptomonads, hypothesized to have acquired plastids in separate endosymbiosis events, leading Okamoto ''et al.'' (2009) to propose the clade Hacrobia to accommodate them.

Diversity

The SAR supergroup encompasses a variety of morphologies and ecological niches, from microscopic zoo- and

phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...

to massive kelp forests. The group includes both photosynthetic and non-photosynthetic forms. Photosynthesis arose independently across various

and alveolate lineages through secondary or higher-order

endosymbiosis An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...

events, acquiring plastids of red algal origin, while chlorarachniophyte

rhizaria The Rhizaria are a diverse and species-rich clade 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 ...

ns captured plastids from

green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...

, retaining vestigial

nucleomorph Nucleomorphs are small, vestigial eukaryotic nuclei found between the inner and outer pairs of membranes in certain plastids. They are thought to be vestiges of red and green algal nuclei that were engulfed by a larger eukaryote. Because the nucl ...

s. It has been estimated that SAR encompasses up to half of all eukaryotic diversity. Owing to the clade's discovery through phylogenomics, there are no known synapomorphies uniting its various members. 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. 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.

Internal phylogeny

A 2021 analysis places Alveolata and Stramenopiles in Halvaria, as sister to Rhizaria.

Notes

References