Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order of turbellarian flatworms.[2][3] About 400 species are known, but probably many more not yet described.[4]
| Acoela Temporal range: [1]
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| Many flatworm-like, orange individuals of the Waminoa acoel on a Plerogyra coral (whitish bubbles) | |
Scientific classification 
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| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Xenacoelomorpha |
| Subphylum: | Acoelomorpha |
| Order: | Acoela Uljanin, 1870 |
The etymology of "acoel" is from the Ancient Greek words ἀ (a), the alpha privative, expressing negation or absence, and κοιλία (koilía), meaning "cavity".[5][6] This refers to the fact that acoels have a structure lacking a fluid-filled body cavity.
Description
editAcoels are very small flattened worms, usually under 2 millimetres (0.079 in) in length, but some larger species, such as Symsagittifera roscoffensis, may reach up to 15 millimetres (0.59 in).[7] They are bilaterally symmetric and microscopic.[8]
They are found worldwide in marine and brackish waters, usually having a benthic lifestyle, although some species are epibionts.[9] Two species, Limonoposthia polonica and Oligochoerus limnophilus, live in freshwater.[10]
Species in the family Convolutidae often form endosymbiotic relationship with microalgae. In one of the genera, Waminoa, the algae are transmitted vertically from parents to offspring. In addition to Convolutidae, there appears to be a potential new and yet unnamed family of acoels that also live in relationships with microalgal endosymbionts.[11]
Members of the class Acoela lack a conventional gut, so that the mouth opens directly into the mesenchyme, i.e., the layer of tissue that fills the body.[12][13] Digestion is accomplished by means of a syncytium that forms a vacuole around ingested food. There are no epithelial cells lining the digestive vacuole, but in the families Diopisthoporidae, Hallangiidae, Hofsteniidae, and Solenofilomorphidae, and the genera Oligochoerus (Convolutidae) and Proporus (Proporidae), there is a short pharynx leading from the mouth to the vacuole.[14] All other bilateral animals (apart from tapeworms) have a gut lined with epithelial cells. As a result, the acoels appear to be solid-bodied.
As the basal lineage of bilateral animals, the Acoela provide interesting insights into early animal evolution and development.[15][16] The most thoroughly studied animal in this group is the species Isodiametra pulchra. Acoela used to be classified in the phylum Platyhelminthes. However, Acoela was separated from this phylum after molecular analyses showed that it had diverged before the three main bilaterian clades had formed, making flatworms as traditionally understood an evolutionary grade from which higher animals had evolved.
Taxonomy
editThe following sub-taxa are recognised in the order Acoela:[17]
- Family Actinoposthiidae Hooge, 2001
- Family Antigonariidae Dörjes, 1968
- Family Antroposthiidae Faubel, 1976
- Family Diopisthoporidae Westblad, 1940
- Family Nadinidae Dörjes, 1968
- Family Paratomellidae Dörjes, 1966
- Family Taurididae Kostenko, 1989
- Suborder Bursalia Jondelius et al., 2011
- Infraorder Crucimusculata Jondelius et al., 2011
- Family Dakuidae Hooge, 2003
- Family Isodiametridae Hooge & Tyler, 2005
- Family Otocelididae Westblad, 1948
- Family Proporidae Graff, 1882
- Superfamily Aberrantospermata Jondelius et al., 2011
- Family Convolutidae Graff, 1905
- Family Mecynostomidae Dörjes, 1968
- Infraorder Prosopharyngida Jondelius et al., 2011
- Family Hallangiidae Westblad, 1946
- Family Hofsteniidae Bock, 1923
- Family Solenofilomorphidae Dörjes, 1968
- Infraorder Crucimusculata Jondelius et al., 2011
References
edit- ^ Knaust, Dirk (2021-10-07). "A microbialite with its entombed benthic community from the Middle Triassic (Anisian-Ladinian) Muschelkalk Group of Germany". Palaeontographica Abteilung A. 320 (1–3): 1–63. doi:10.1127/pala/2021/0114. ISSN 0375-0442.
- ^ Cannon, Johanna Taylor; Vellutini, Bruno Cossermelli; Smith, Julian; Ronquist, Fredrik; Jondelius, Ulf; Hejnol, Andreas (2016). "Xenacoelomorpha is the sister group to Nephrozoa". Nature. 530 (7588): 89–93. Bibcode:2016Natur.530...89C. doi:10.1038/nature16520. ISSN 0028-0836. PMID 26842059. S2CID 205247296.
- ^ Jimenez-Guri E, Paps J, Garcia-Fernandez J, Salo E (2006), "Hox and ParaHox genes in Nemertodermatida, a basal bilaterian clade", Int. J. Dev. Biol., 50 (8): 675–9, doi:10.1387/ijdb.062167ej, PMID 17051477.
- ^ A New Species of Acoela Possessing a Middorsal Appendage with a Possible Sensory Function
- ^ Bailly, Anatole (1981-01-01). Abrégé du dictionnaire grec français. Paris: Hachette. ISBN 2010035283. OCLC 461974285.
- ^ Bailly, Anatole. "Greek-french dictionary online". www.tabularium.be. Retrieved 2020-03-02.
- ^ Hooge, M. D.; Tyler, S. (2005). "New tools for resolving phylogenies: a systematic revision of the Convolutidae (Acoelomorpha, Acoela)". Journal of Zoological Systematics and Evolutionary Research. 43 (2): 100–113. doi:10.1111/j.1439-0469.2005.00301.x. ISSN 0947-5745.
- ^ Achatz, Johannes G.; Chiodin, Marta; Salvenmoser, Willi; Tyler, Seth; Martinez, Pedro (2013). "The Acoela: on their kind and kinships, especially with nemertodermatids and xenoturbellids (Bilateria incertae sedis)". Organisms, Diversity & Evolution. 13 (2): 267–286. doi:10.1007/s13127-012-0112-4. ISSN 1439-6092. PMC 3789126. PMID 24098090.
- ^ Ogunlana, M. V.; Hooge, M. D.; Tekle, Y. I.; Benayahu, Y.; Barneah, O.; Tyler, Seth (2005). "Waminoa brickneri n. sp. (Acoela: Acoelomorpha) associated with corals in the Red Sea". Zootaxa. 1008: 1–11. doi:10.11646/zootaxa.1008.1.1.
- ^ Giribet, Gonzalo; Edgecombe, Gregory D. (3 March 2020). The Invertebrate Tree of Life. Princeton University Press. ISBN 978-0-691-17025-1.
- ^ Riewluang, Siratee; Wakeman, Kevin C. (2023). "Biodiversity of symbiotic microalgae associated with meiofaunal marine acoels in Southern Japan". PeerJ. 11: e16078. doi:10.7717/peerj.16078. PMC 10560497. PMID 37814628.
- ^ Boll, Piter Kehoma; Rossi, Ilana; Amaral, Silvana Vargas do; Oliveira, Simone Machado de; Müller, Eliara Solange; Lemos, Virginia Silva; Leal-Zanchet, Ana Maria (2013). "Platyhelminthes ou apenas semelhantes a Platyhelminthes? Relações filogenéticas dos principais grupos de turbelários". Neotropical Biology and Conservation (in Portuguese). 8 (1). doi:10.4013/nbc.2013.81.06. ISSN 2236-3777.
- ^ "The Platyhelminthes and the Acoela". Archived from the original on 2008-05-09. Retrieved 2009-03-21.
- ^ [The Acoela: on their kind and kinships, especially with nemertodermatids and xenoturbellids (Bilateria incertae sedis) The Acoela: on their kind and kinships, especially with nemertodermatids and xenoturbellids (Bilateria incertae sedis)]
- ^ Hejnol A, Martindale MQ (2008), "Acoel development supports a simple planula-like urbilaterian", Phil. Trans. R. Soc. B, 363 (1496): 1493–1501, doi:10.1098/rstb.2007.2239, PMC 2614228, PMID 18192185
- ^ Hejnol A, Martindale MQ (2008), "Acoel development indicates the independent evolution of the bilaterian mouth and anus", Nature, 456 (7220): 382–386, Bibcode:2008Natur.456..382H, doi:10.1038/nature07309, PMID 18806777, S2CID 4403355
- ^ "WoRMS - World Register of Marine Species - Acoela". www.marinespecies.org. Retrieved 2020-02-08.