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Chrompodellids are a of single-celled belonging to the supergroup. It comprises two different groups of : the colpodellids, , and the chromerids, that live as symbionts of . These groups were independently discovered and described, but molecular analyses demonstrated that they are intermingled in a that is the closest relative to , and they became collectively known as chrompodellids. Due to the history of their research, they are variously known in biological classification as Chromerida or Colpodellida (ICZN)/ Colpodellales (ICN).


Description and life cycle
Chrompodellids are a of unicellular containing two functionally different groups: the "chromerids" and the predatory "colpodellids". Like other , they present tubular and highly flattened cortical alveoli with underneath. They exhibit a -like structure similar to that of , with an , a micropore and a rostrum. They live as with two anisokont (i.e. differently sized) that are heterodynamic (i.e. move in different patterns). Some species exhibit thin in their anterior flagellum, while others bear bulbs. Some species are capable of forming .


Colpodellids
Colpodellids, represented by the genera , , and , are free-living . They live as biflagellated single cells, 5–20 μm in length, with an open and used to hunt. They present genetic sequences of non-photosynthetic , evidence of their ancestry.

Some species, considered , do not prey cells, but rather fully or partially "suck" their contents, a process known as , common among . They feed on and other , such as , , , and . After feeding, they internalize their flagella, become and divide into tetrads, similarly to the development of zoospores in . The cells conjugate after leaving the cyst, which could imply a sexual stage.


Chromerids
Chromerids, represented by the genera and , are , and are thus considered . They exist in with . For most of their life cycle, they live as round () brownish immobile called autospores, surrounded by a thick resistant . They contain one in each cell, with , , and .

The two genera are markedly different from each other, both in and life cycles. autospores are 5–7 μm in diameter. They asexually reproduce through binary division to develop autosporangia, which in turn harbor 2–4 autospores under an additional membrane. They also form , up to 15 μm in diameter, capable of generating 2–10 flagellated that strongly resemble colpodellids. This dispersal process is similar to the of apicomplexans. Sexual reproduction has not been observed. Under adverse environmental conditions, they form resistant that remain viable for years. Similarly to apicomplexans, they undergo closed , without dissolving the . In addition, Chromera produces high amounts of an exclusive type of .

autospores, by contrast, start measuring 3 μm and grow up to 40 μm before transforming into sporangia that generate dozens of autospores or zoospores. There are two types of Vitrella zoospores: one is generated by budding from the mother cell and exhibits flagella outside the , the other develops axonemes and flagella within their cytoplasm and are ejected from the mother cell after maturing, though both types lack a pseudo-. Some zoospores fuse, possibly representing a sexual stage in the life cycle. In addition, Vitrella produces .


Evolution
Chrompodellids are the closest living relatives of the parasites, which evolved from a ancestor, making chromerids the last remaining photosynthetic members of an otherwise parasitic clade within . The apicomplexans, chrompodellids, perkinsids and dinoflagellates constitute the clade , characterized by the and derived from an event of secondary endosymbiosis with a . The photosynthetic ability of these plastids was eventually lost in apicomplexans, colpodellids, perkinsids and other groups that transitioned into a predatory or parasitic lifestyle. The following cladogram summarizes alveolate relationships and the internal relationships among most genera within the chrompodellid clade (chromerids marked with asterisks):


Systematics

Taxonomic history
In 1993, protozoologist Thomas Cavalier-Smith described the order Colpodellida (under the , later regularized as Colpodellales in accordance to the ) to contain what he considered one of the "most primitive flagellate apicomplexans", the genus . This order was introduced in the class Apicomonadea along with the . Cavalier-Smith treats this class as a member of the phylum , while "true" apicomplexans are united under the name Sporozoa. Although the inclusion of colpodellids within apicomplexans was not supported by other authors, studies demonstrated that they were .

The first chromerid alga, , was discovered and isolated from in 2001. It was described in 2008 as the first member of a new phylum Chromerida, followed by in 2012. They showed morphological resemblance to colpodellids and other . In the following years, studies reported the evolutionary proximity between colpodellids and chromerid algae. This was supported by the discovery of retained vestigial in some colpodellid species. In 2015 there was strong support for a containing the two groups, phylogenetically mixed with each other, which rendered both as . The clade was given the provisional name " chrompodellids", later referred to as Chrompodellida by posterior studies.

Between 2004 and 2017, Cavalier-Smith retained the classification scheme of Apicomonadea, from which he excluded Perkinsida, leaving only colpodellids and chromerids across multiple orders. In addition, several genera of were added on the basis of morphological data: Algovora, Microvorax and Dinomonas. Due to lacking , these genera have been excluded from later classifications. Two genera, and , were eventually rejected in his classification, but later revisions by other authors maintain them as independent genera supported by molecular data.

The treatment of chrompodellids as a subgroup of , under the name of Apicomonadea, was rejected by the International Society of Protistologists. In a 2019 revision of classification, emended the previous name Colpodellida to contain all chrompodellids, and treated it as a direct subgroup of , independent from Apicomplexa. Later, advocated for this treatment as a separate , and regularized it under the name of Chromerida or Chromeridophyta, composed of a single class Colpodellophyceae and a single order Colpodellales, in accordance to the nomenclatural rules of the ICN. However, other authors consider them a subgroup of the phylum , together with apicomplexans, and .


Classification
As of 2023, chrompodellids are divided into four families and seven genera:

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