The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads as well. These four groups are all anaerobic (many being aerotolerant anaerobes), occurring mostly as symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.
A number of parabasalids and oxymonads are found in termite guts, and play an important role in breaking down the cellulose found in wood. Some other metamonads are parasites.
These flagellates are unusual in lacking mitochondria. Originally they were considered among the most primitive eukaryotes, diverging from the others before mitochondria appeared. However, they are now known to have lost mitochondria secondarily, and retain both organelles and nuclear genes derived from them. Mitochondrial relics include hydrogenosomes, which produce hydrogen, and small structures called mitosomes.
It now appears the Metamonada are, together with Malawimonas, sister clades of the Podiata.
All of these groups are united by having flagella or basal bodies in characteristic groups of four, which are often associated with the nucleus, forming a structure called a karyomastigont. In addition, the genera Carpediemonas and Trimastix are now known to be close relatives of the retortamonad-diplomonad line and the oxymonads, respectively. Both are free-living and amitochondriate.
The metamonads were thought to make up part of the Excavata, a paraphyletic eukaryotic supergroup including flagellates with feeding grooves and their close relatives. Their relationships are uncertain, and they do not always appear together on molecular trees. It is possible that the metamonads as defined here do not form a monophyletic subgroup.
The following higher level treatment from 2013 is based on works of Cavalier-Smith with amendments within Fornicata from Yubukia, Simpson & Leander.
Metamonada were once again proposed to be basal eukaryotes in 2018.
- Phylum Metamonada (Grassé 1952) Cavalier-Smith 1987 emend. Cavalier-Smith 2003
- Subphylum Anaeromonada Cavalier-Smith 1997 emend. 2003
- Subphylum Trichozoa Cavalier-Smith 1996 emend. Cavalier-Smith 2003 stat. n. 2013
- Superclass Fornicata Simpson 2003 stat. n. Cavalier-Smith 2013
- Superclass Parabasalia Honigberg 1973 stat. n. Cavalier-Smith 2003
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^ Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (2016-06-01). "187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution". Molecular Phylogenetics and Evolution. 99: 275–296. doi:10.1016/j.ympev.2016.03.023. PMID 27001604.
^ Cavalier-Smith T (November 2003). "The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa". Int. J. Syst. Evol. Microbiol. 53 (Pt 6): 1741–58. doi:10.1099/ijs.0.02548-0. PMID 14657102.
^ Cavalier-Smith T (2013). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". Eur. J. Protistol. 49 (2): 115–178. doi:10.1016/j.ejop.2012.06.001. PMID 23085100.
^ Yubukia; Simpson; Leander (2013). "Comprehensive Ultrastructure of Kipferlia bialata Provides Evidence for Character Evolution within the Fornicata (Excavata)". Protist. 164 (3): 423–439. doi:10.1016/j.protis.2013.02.002. PMID 23517666.
^ Krishnan, Arunkumar; Burroughs, A. Max; Iyer, Lakshminarayan; Aravind, L. (2018-07-04). "The unexpected provenance of components in eukaryotic nucleotide-excision-repair and kinetoplast DNA-dynamics from bacterial mobile elements". bioRxiv: 361121. doi:10.1101/361121.