Acoelomorpha is asubphylum of very simple and smallsoft-bodied animals withplanula-like features which live inmarine orbrackish waters. They usually live between grains ofsediment, swimming asplankton, or crawling on other organisms, such asalgae andcorals.[1] With the exception of two acoel freshwater species, all known acoelomorphs are marine.[2]
The term "acoelomorph" derives from theAncient Greek wordsἀ (a), thealpha privative, expressing negation or absence,κοιλία (koilía), meaning "cavity", andμορφή (morphḗ), meaning "form".[3][4] This refers to the fact that acoelomorphs have a structure lacking a fluid-filled body cavity.
Various members of the Acoela class.[5]Various members of the Nemertodermatida class.
The subphylum Acoelomorpha is divided into two classes. There are at least 408 described species, with a majority of these falling within theCrucimusculata infraorder in Acoela.
Acoela comprise small flattened worms, classified into a dozen families.[6]
Nemertodermatida comprise millimetre-sized, mostly interstitial worms, distributed into two families.[7]
The soft bodies of acoelomorphs and the lack of some of the key bilaterian traits make them difficult to classify.[8][9] Traditionally, based onphenotypic features, acoelomorphs were considered to belong to the phylumPlatyhelminthes, which was long seen as the sister group to all otherbilaterian phyla.[10] However, a series of molecular phylogenetics studies at the hinge between the 20th and 21st centuries demonstrated that they are fast-evolving organisms not closely related to platyhelminthes,[11][12][13][14] therefore involving thepolyphyly of flatworms.[15][16][17][18]
Actually, Acoelomorpha appeared to constitute a separate,deep-branching phylum, kingpin of bilaterian evolution.[19] Yet their evolutionary affinities remain enigmatic as they might be the sister-group either to all other bilateral animals[15][16][17] or to all deuterostomes.[18] Resolving this debate would indicate whether acoelomorphs are simple or simplified. If they are the sister group toBilateria, it would point to a simplebody plan for the first bilaterian. Alternatively, if acoelomorphs are related todeuterostomes, this would imply that their organisation is the result of secondary simplification.[20]
In addition, comparative analyses of morphological, developmental, and molecular characters raised two points.
Xenoturbellida is the sister group to acoelomorphs, constituting the so-calledXenacoelomorpha clade.[21] The close evolutionary relationship between Acoelomorpha andXenoturbella is supported by the morphology (structure of epidermalcilia[22]), the embryology (direct development without a feeding larval stage[23]), and the concatenation of hundreds of proteins.[24][21][25]
The phylogenetic placement of Xenacoelomorpha among bilaterian animals is not yet well defined, despite increased taxon and gene sampling, (re)-analyses of published data sets, and use of more sophisticatedmodels of sequence evolution inphylogenomic studies. There is a conflict between two evolutionary hypotheses, with Xenacoelomorpha being the sister group toAmbulacraria withindeuterostomes (i.e., theXenambulacraria hypothesis) on the one hand,[26] and Xenacoelomorpha as sister group to all other bilaterians (i.e., theNephrozoa hypothesis) on the other.[25][27][28] However, the Nephrozoa hypothesis might reflect methodological errors resulting from model violations in the phylogenomic inference.[26]
Acoelomorphs resemble flatworms in many respects, but have a simpler anatomy, not even having a gut. Like flatworms, they have no circulatory or respiratory systems, but they also lack an excretory system. They lack body cavities (acoelomate structure), a hindgut or an anus.[1]
Theepidermal cells of acoelomorphs are unable to proliferate, a feature that is only shared withrhabditophoran flatworms and was for some time considered a strong evidence for the position of Acoelomorpha within Platyhelminthes. In both groups, the epidermis is renewed from mesodermal stem cells.[29]
Thenervous system of acoelomorphs is formed by a set of longitudinal nerve bundles beneath the ciliated epidermis. Close to the anterior end, these bundles are united by a ring commissure, but do not form a true brain, although it is hypothesized that such organization was the precursor of thecephalization of the nerve system in more derived bilaterians.[30] After decapitation, such a "brain" (rather, acerebroid ganglion) regenerates in a few weeks.[31]
The sensory organs include astatocyst – which presumably helps them orient to gravity –, and, in some cases,ancestral pigment-spotocelli capable of detecting light.[32]
Acoelomorphs are simultaneoushermaphrodites, but have nogonads and no ducts associated with the female reproductive system. Instead,gametes are produced from themesenchymal cells that fill the body between the epidermis and the digestive vacuole.[32]
^abCannon, L. R. G. (1986)Turbellaria of the World. A guide to families and genera. Brisbane, Queensland Museum, 136 p.
^Harzsch, Steffen; Purschke, Günter (2016-01-01).Structure and evolution of invertebrate nervous systems. Oxford University Press. pp. 56–61.ISBN978-0-19-968220-1.OCLC951605913.
^Bailly, Anatole (1981-01-01).Abrégé du dictionnaire grec français. Paris: Hachette.ISBN2010035283.OCLC461974285.
^Petrov, A.; Hooge, M.; Tyler, S. (2006). "Comparative morphology of the bursal nozzles in acoels (Acoela, Acoelomorpha)".Journal of Morphology.267 (5):634–648.doi:10.1002/jmor.10428.PMID16485278.S2CID32595353.
^Conway-Morris, S.; George, J. D.; Gibson R.; Platt, H. M. (1985)The Origins and relationships of lower invertebrates. Oxford, Clarendon Press, 397 p.[page needed]
^abRuiz-Trillo, Iñaki; Riutort, Marta; Fourcade, H. Matthew; Baguñà, Jaume; Boore, Jeffrey L. (2004). "Mitochondrial genome data support the basal position of Acoelomorpha and the polyphyly of the Platyhelminthes".Molecular Phylogenetics and Evolution.33 (2):321–32.Bibcode:2004MolPE..33..321R.doi:10.1016/j.ympev.2004.06.002.PMID15336667.
^Lundin, K (1998). "The epidermal ciliary rootlets ofXenoturbella bocki (Xenoturbellida) revisited: new support for a possible kinship with the Acoelomorpha (Platyhelminthes)".Zoologica Scripta.27 (3):263–270.doi:10.1111/j.1463-6409.1998.tb00440.x.S2CID85324766.
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