Temporal range: Ediacaran  - Recent
There are four phyla, which are very divergent in form, but are now known to be close relatives based on various ultrastructural and genetic similarities:
- Apicomplexa - parasitic protozoa that lack axonemal locomotive structures except in gametes
- Chromerida - a marine phylum of photosynthetic protozoa
- Ciliates - very common protozoa with many short cilia arranged in rows
- Dinoflagellates - mostly marine flagellates many of which have chloroplasts
The most notable shared characteristic is the presence of cortical alveoli, flattened vesicles packed into a continuous layer supporting the membrane, typically forming a flexible pellicle. In dinoflagellates they often form armor plates. Alveolates have mitochondria with tubular cristae and their flagella or cilia have a distinct structure.
The ancestors of this group appear to have been photosynthetic.
All sequenced mitochondrial genomes of ciliates and apicomplexia are linear.
The Apicomplexa and dinoflagellates may be more closely related to each other than to the ciliates. Both have plastids, and most share a bundle or cone of microtubules at the top of the cell. In apicomplexans this forms part of a complex used to enter host cells, while in some colorless dinoflagellates it forms a peduncle used to ingest prey. Various other genera are closely related to these two groups, mostly flagellates with a similar apical structure. These include free-living members in Oxyrrhis and Colponema, and parasites in Perkinsus, Parvilucifera, Rastrimonas and the ellobiopsids. In 2001, direct amplification of the rRNA gene in marine picoplankton samples revealed the presence of two novel alveolate linages, called group I and II. Group I has no cultivated relatives, while group II is related to the dinoflagellate parasite Amoebophrya, which was classified until now in the Syndiniales dinoflagellate order.
Relationships between some of these the major groups were suggested during the 1980s, and a specific relationship between all three was confirmed in the early 1990s by genetic studies, most notably by Gajadhar et al. Cavalier-Smith, introduced the formal name Alveolata in 1991, although at the time he actually considered the grouping to be a paraphyletic assemblage, rather than a monophyletic group.
The development of plastids among the alveolates is uncertain. Cavalier-Smith proposed the alveolates developed from a chloroplast-containing ancestor, which also gave rise to the Chromista (the chromalveolate hypothesis). However, as plastids only appear in relatively derived (as opposed to ancestral) groups, others argue the alveolates originally lacked them and possibly the dinoflagellates and Apicomplexa acquired them separately.
It seems likely that the common ancestor of this group was a myzocytotic predator with two heterodynamic flagella, micropores, trichocysts, rhoptries, micronemes, a polar ring and a coiled open sided conoid. This ancestor also probably possesed a plastid but it is presently not clear whether it was photosynthetic. Furthermore it is not clear whether extant perkinsids or colpodellids have retained this organelle.
Li, C.-W.; et al. (2007). "Ciliated protozoans from the Precambrian Doushantuo Formation, Wengan, South China". Geological Society, London, Special Publications. 286: 151–156. doi:10.1144/SP286.11. Cite uses deprecated parameter
- ^ "alveolate". Memidex (WordNet) Dictionary/Thesaurus. Retrieved 2011-01-26.
- ^ Zhang H, Campbell DA, Sturm NR, Dungan CF, Lin S (2011) Spliced leader RNAs, mitochondrial gene frameshifts and multi-protein phylogeny expand support for the genus Perkinsus as a unique group of Alveolates. PLoS One. 2011;6(5):e19933
- ^ Reyes-Prieto A, Moustafa A, Bhattacharya D. (2008) Multiple genes of apparent algal origin suggest ciliates may once have been photosynthetic. 18 (13):956-962
- ^ Barth D, Berendonk TU (2011) The mitochondrial genome sequence of the ciliate Paramecium caudatum reveals a shift in nucleotide composition and codon usage within the genus Paramecium. BMC Genomics. 12:272
- ^ López-García, P. et al. (2001). Unexpected diversity of small eukaryotes in deep-sea Antarctic plankton. Nature 409: 603-7.
- ^ Moon-van der Staay, S. Y. et al. (2001). Oceanic 18S rDNA sequences from picoplankton reveal unsuspected eukaryotic diversity. Nature 409: 607-10.
- ^ Gajadhar, A. A.; et al. (1991). "Ribosomal RNA sequences of Sarcocystis muris, Theilera annulata, and Crypthecodinium cohnii reveal evolutionary relationships among apicomplexans, dinoflagellates, and ciliates". Molecular and Biochemical Parasitology. 45: 147–153. doi:10.1016/0166-6851(91)90036-6.
- ^ Cavalier-Smith, T. (1991). Cell diversification in heterotrophic flagellates. In The Biology of Free-living Heterotrophic Flagellates, ed. D.J. Patterson & J. Larsen. pp. 113-131. Oxford University Press.
- ^ a b Janouskovec J, Horák A, Oborník M, Lukes J, Keeling PJ (2010) A common red algal origin of the apicomplexan, dinoflagellate, and heterokont plastids. Proc Natl Acad Sci USA 107(24):10949-10954
- ^ Kuvardina ON, Leander BS, Aleshin VV, Myl'nikov AP, Keeling PJ, Simdyanov TG (2002) The phylogeny of colpodellids (Alveolata) using small subunit rRNA gene sequences suggests they are the free living sister group to apicomplexans. J Eukaryot Microbiol 49(6):498-504
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