Archezoa
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In biology, Archezoa is a term that has been introduced by several authors to refer to a group of organisms (a taxon). Authors include Josef Anton Maximilian Perty,
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; ; 16 February 1834 – 9 August 1919) was a German zoologist, natural history, naturalist, eugenics, eugenicist, Philosophy, philosopher, physician, professor, marine biology, marine biologist and artist ...
and in the 20th century by
Thomas Cavalier-Smith Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow (21 October 1942 – 19 March 2021), was a professor of evolutionary biology in the Department of Zoology, at the University of Oxford. His research has led to discov ...
in his classification system. Each author used the name to refer to different arrays of organisms. This reuse by later authors of the same taxon name for different groups of organisms is widely criticized in taxonomy because the inclusion of the name in a sentence (e.g. "Archezoa have no olfactory organs") does not make sense unless the particular usage is specified (e.g. "Archezoa ''sensu'' Cavalier-Smith (1987) have no olfactory organs"). Nonetheless, all uses of 'Archezoa' are now obsolete.


Archezoa ''sensu'' Cavalier-Smith (1987)

Cavalier-Smith proposed the term 'Archezoa' for a paraphyletic (see
Paraphyly Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
) territory of
eukaryotes The eukaryotes ( ) constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, seaweeds, and many unicellular organisms are eukaryotes. They constitute a major group of ...
that primitively lacked mitochondria. Like Margulis and others before (see Pelomyxa), Cavalier-Smith argued that the initial ancestor of eukaryotes emerged prior to the endosymbiotic acquisition (see
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 ...
) of mitochondria. The same paraphyletic territory was referred to as 'Hypochondria' by others. The argument for Archezoa sensu Cavalier-Smith was never universally accepted because of conflicting information, and was dropped when the contrary argument, that amitochondriates were descendants of eukaryotes with mitochondria, became dominant. Eukaryotes that eventually acquired a bacterial endosymbiont that became the mitochondria were placed in a taxonomic group which Cavalier-Smith called the Metakaryota, whereas the Archezoa represented an earlier
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
group to which Cavalier-Smith variously assigned the diplomonads, ''Entamoeba'',
Microsporidia Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore.Franzen, C. (2005). How do Microsporidia inva ...
, oxymonads, parabasalids ( Parabasalids), pelobionts (see '' Pelomyxa''), retortamonads, trichomonads, and ''Trimastix'' (see
Cavalier-Smith's system of classification The initial version of a classification system of life by British zoologist Thomas Cavalier-Smith appeared in 1978. This initial system continued to be modified in subsequent versions that were published until he died in 2021. As with classificati ...
). With the rejection of 'Archeozoa', the meaning of the term 'Metakaryota' became the same as 'Eukaryota' (see
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 ...
), and Metakaryota became superfluous.


Original mitochondria lost

Eukaryotic protists lacking mitochondria were discovered to have experienced secondary mitochondrial loss, meaning that their ancestors once possessed mitochondria but that these mitochondria had, over time, been transformed, reduced, or lost. In some of these organisms, mitochondria had degraded into simpler double-membrane bound organelles known as mitosomes and hydrogenosomes. Some of both types of organelles are known to have fully lost their genome. Initial discoveries found that amitochondriate organisms appeared to express mitochondrial Hsp60 and Hsp70 proteins from the nuclear DNA of the organism. This indicated that the ancestors of these organisms once possessed mitochondria which expressed these proteins, but that these genes had migrated to their nuclear DNA over time as a result of endosymbiotic gene transfer. As a result, the argument that some extant eukaryotes lacking mitochondria had emerged from the eukaryotic lineage before mitochondria were acquired was falsified.


Long branch attraction

An argument for the Archezoa group was that amitochondriate protists appeared to branch off early on from the eukaryotic lineage in phylogenetic analyses. This corroborated the supposition that Archezoa were more closely linked to primitive eukaryotes that evolved prior to the endosymbiotic process that generated the mitochondria. However, this early divergence later turned out to be a class of systematic errors in phylogenetic analysis called "
long branch attraction In phylogenetics, long branch attraction (LBA) is a form of systematic error whereby distantly related lineages are incorrectly inferred to be closely related. LBA arises when the amount of molecular or morphological change accumulated within a lin ...
".


References

Obsolete eukaryote taxa Biological hypotheses Mitochondria Kingdoms (biology) {{Eukaryote-stub