acoelomorpha

The term "acoelomorph" derives from the Ancient Greek words {{wikt-lang|grc|ἀ}} ({{grc-transl|ἀ}}), the alpha privative, expressing negation or absence, {{wikt-lang|grc|κοιλία}} ({{grc-transl|κοιλία}}), meaning "cavity", and {{wikt-lang|grc|μορφή}} ({{grc-transl|μορφή}}), meaning "form".{{Cite book | title=Abrégé du dictionnaire grec français | last=Bailly | first=Anatole | date=1981-01-01 | publisher=Hachette | isbn=2010035283 | location=Paris | oclc=461974285}}{{Cite web | url=http://www.tabularium.be/bailly/ | title=Greek-french dictionary online | last=Bailly | first=Anatole | website=www.tabularium.be | access-date=2020-02-27}} This refers to the fact that acoelomorphs have a structure lacking a fluid-filled body cavity.

File:Acoela from Indian Ocean.gif

File:Nemertodermatida species.png

The subphylum Acoelomorpha is divided into two classes. There are at least 408 described species, with a majority of these falling within the Crucimusculata infraorder in Acoela.

• Acoela comprise small flattened worms, classified into a dozen families.{{Cite web |editor1=Tyler, S. |editor2=Artois, T. |editor3=Schilling, S. |editor4=Hooge, M. |editor5=Bush, L.F. | date=2006–2020 | url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=2847 | title=World List of turbellarian worms: Acoelomorpha, Catenulida, Rhabditophora. Acoela | publisher=WoRMS - World Register of Marine Species | website=www.marinespecies.org | access-date=2020-02-12}}
• Nemertodermatida comprise millimetre-sized, mostly interstitial worms, distributed into two families.{{Cite web |editor1=Tyler, S. |editor2=Artois, T. |editor3=Schilling, S. |editor4=Hooge, M. |editor5=Bush, L.F. | date=2006–2020 | url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=2848 | title=World List of turbellarian worms: Acoelomorpha, Catenulida, Rhabditophora. Nemertodermatida | publisher=WoRMS - World Register of Marine Species | website=www.marinespecies.org | access-date=2020-02-12}}

The soft bodies of acoelomorphs and the lack of some of the key bilaterian traits make them difficult to classify.{{Cite journal | last1 = Petrov | first1 = A. | last2 = Hooge | first2 = M. | last3 = Tyler | first3 = S. | doi = 10.1002/jmor.10428 | title = Comparative morphology of the bursal nozzles in acoels (Acoela, Acoelomorpha) | journal = Journal of Morphology | volume = 267 | issue = 5 | pages = 634–648 | year = 2006 | pmid = 16485278 | s2cid = 32595353 }}{{Cite web | url=https://www.smithsonianmag.com/science-nature/why-you-should-care-about-acoelomorph-flatworms-17782785/ | title=Why You Should Care About Acoelomorph Flatworms | last=Laden | first=Greg | date=2009-10-01 | website=Smithsonian Magazine | language=en | access-date=2020-03-02}} Traditionally, based on phenotypic features, acoelomorphs were considered to belong to the phylum Platyhelminthes, which was long seen as the sister group to all other bilaterian phyla.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|date=June 2018}} 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,{{Cite journal | last1=Katayama | first1=Tomoe | last2=Yamamoto | first2=Masamichi | last3=Wada | first3=Hiroshi | last4=Satoh | first4=Noriyuki | year=1993 | title=Phylogenetic Position of Acoel Turbellarians Inferred from Partial 18S rDNA Sequences | url=https://repository.kulib.kyoto-u.ac.jp/dspace/bitstream/2433/57238/3/i0289-0003-10-3-529.pdf | journal=Zoological Science | volume=10 | issue=3 | pages=529–536 | issn=0289-0003| pmid=7764139 }}{{Cite journal | last1=Katayama | first1=T. | last2=Wada | first2=H. | last3=Furuya | first3=H. | last4=Satoh | first4=N. | last5=Yamamoto | first5=M. | date=1995-10-01 | title=Phylogenetic Position of the Dicyemid Mesozoa Inferred from 18S rDNA Sequences | journal=The Biological Bulletin | volume=189 | issue=2 | pages=81–90 | doi=10.2307/1542458 | pmid=8541419 | issn=0006-3185| jstor=1542458 | url=https://www.biodiversitylibrary.org/part/19914 }}{{Cite journal | last1=Carranza | first1=S. | last2=Baguñà | first2=J. | last3=Riutort | first3=M. | date=1997-05-01 | title=Are the Platyhelminthes a monophyletic primitive group? An assessment using 18S rDNA sequences. | url=https://academic.oup.com/mbe/article/14/5/485/994967 | journal=Molecular Biology and Evolution | language=en | volume=14 | issue=5 | pages=485–497 | doi=10.1093/oxfordjournals.molbev.a025785 | pmid=9159926 | issn=0737-4038| doi-access= | url-access=subscription }}{{cite journal | last1=Ruiz-Trillo | first1=Iñaki | last2=Riutort | first2=Marta | last3=Littlewood | first3=D. Timothy J. | last4=Herniou | first4=Elisabeth A. | last5=Baguñà | first5=Jaume | date=March 1999 | title=Acoel Flatworms: Earliest Extant Bilaterian Metazoans, Not Members of Platyhelminthes | journal=Science | volume=283 | issue=5409 | pages=1919–1923 | doi=10.1126/science.283.5409.1919 | pmid=10082465 | issn=0036-8075 | bibcode=1999Sci...283.1919R }} therefore involving the polyphyly of flatworms.{{Cite journal | last1 = Baguñà | first1 = J. | last2 = Riutort | first2 = M. | doi = 10.1139/z03-214 | title = Molecular phylogeny of the Platyhelminthes | journal = Canadian Journal of Zoology | volume = 82 | issue = 2 | pages = 168–193 | year = 2004 }}{{cite journal | last1=Baguñà | first1=Jaume | last2=Riutort | first2=Marta | title=The dawn of bilaterian animals: the case of acoelomorph flatworms | journal=BioEssays | volume=26 | issue=10 | year=2004 | pages=1046–57 | doi=10.1002/bies.20113 | pmid=15382134| s2cid=40453683 }}{{cite journal | last1=Ruiz-Trillo | first1=Iñaki | last2=Riutort | first2=Marta | last3=Fourcade | first3=H. Matthew | last4=Baguñà | first4=Jaume | last5=Boore | first5=Jeffrey L. | title=Mitochondrial genome data support the basal position of Acoelomorpha and the polyphyly of the Platyhelminthes | journal=Molecular Phylogenetics and Evolution | volume=33 | issue=2 | year=2004 | pages=321–32 | doi=10.1016/j.ympev.2004.06.002 | pmid=15336667| bibcode=2004MolPE..33..321R }}{{Cite journal | last1=Philippe | first1=Hervé | last2=Brinkmann | first2=Henner | last3=Martinez | first3=Pedro | last4=Riutort | first4=Marta | last5=Baguñà | first5=Jaume | date=2007-08-08 | editor-last=Volff | editor-first=Jean-Nicolas | title=Acoel Flatworms Are Not Platyhelminthes: Evidence from Phylogenomics | journal=PLOS ONE | language=en | volume=2 | issue=8 | pages=e717 | doi=10.1371/journal.pone.0000717 | issn=1932-6203 | pmc=1933604 | pmid=17684563| doi-access=free | bibcode=2007PLoSO...2..717P }}

Actually, Acoelomorpha appeared to constitute a separate, deep-branching phylum, kingpin of bilaterian evolution.{{Cite journal | last=Haszprunar | first=G. | date=1996 | title=Plathelminthes and Plathelminthomorpha — paraphyletic taxa | journal=Journal of Zoological Systematics and Evolutionary Research | language=en | volume=34 | issue=1 | pages=41–48 | doi=10.1111/j.1439-0469.1996.tb00808.x | issn=1439-0469| doi-access=free }} Yet their evolutionary affinities remain enigmatic as they might be the sister-group either to all other bilateral animals or to all deuterostomes. Resolving this debate would indicate whether acoelomorphs are simple or simplified. If they are the sister group to Bilateria, it would point to a simple body plan for the first bilaterian. Alternatively, if acoelomorphs are related to deuterostomes, this would imply that their organisation is the result of secondary simplification.{{Cite journal | last1=Ruiz-Trillo | first1=Iñaki | last2=Paps | first2=Jordi | date=2016-06-01 | title=Acoelomorpha: earliest branching bilaterians or deuterostomes? | journal=Organisms Diversity & Evolution | language=en | volume=16 | issue=2 | pages=391–399 | doi=10.1007/s13127-015-0239-1 | issn=1618-1077| doi-access=free | bibcode=2016ODivE..16..391R | hdl=1983/2686c832-dffe-4486-b3b2-17d4966fd64e | hdl-access=free }}

File:Xenoturbella japonica.jpg, the sister group to acoelomorphs]]

In addition, comparative analyses of morphological, developmental, and molecular characters raised two points.

• Xenoturbellida is the sister group to acoelomorphs, constituting the so-called Xenacoelomorpha clade.{{Cite journal | last1 = Philippe | first1 = H. | last2 = Brinkmann | first2 = H. | last3 = Copley | first3 = R. R. | last4 = Moroz | first4 = L. L. | last5 = Nakano | first5 = H. | last6 = Poustka | first6 = A. J. | last7 = Wallberg | first7 = A. | last8 = Peterson | first8 = K. J. | last9 = Telford | first9 = M. J. | title = Acoelomorph flatworms are deuterostomes related to Xenoturbella | journal = Nature | volume = 470 | pages = 255–258 | year = 2011 | pmid = 21307940 | doi = 10.1038/nature09676 | bibcode = 2011Natur.470..255P | issue=7333 | pmc=4025995}} The close evolutionary relationship between Acoelomorpha and Xenoturbella is supported by the morphology (structure of epidermal cilia{{cite journal | last1 = Lundin | first1 = K | year = 1998 | title = The epidermal ciliary rootlets of Xenoturbella bocki (Xenoturbellida) revisited: new support for a possible kinship with the Acoelomorpha (Platyhelminthes) | journal = Zoologica Scripta | volume = 27 | issue = 3 | pages = 263–270 | doi=10.1111/j.1463-6409.1998.tb00440.x| s2cid = 85324766 }}), the embryology (direct development without a feeding larval stage{{Cite journal | last2=Lundin | first2=K. | last3=Bourlat | first3=S. J. | last4=Telford | first4=M. J. | last5=Funch | first5=P. | last6=Nyengaard | first6=J. R. | last7=Obst | first7=M. | last8=Thorndyke | first8=M. C. | year=2013 | title=Xenoturbella bocki exhibits direct development with similarities to Acoelomorpha | journal=Nature Communications | volume=4 | pages=1537– | doi=10.1038/ncomms2556 | pmc=3586728 | pmid=23443565 | last1=Nakano | first1=H. | bibcode=2013NatCo...4.1537N}}), and the concatenation of hundreds of proteins.{{Cite journal | last1=Hejnol | first1=Andreas | last2=Obst | first2=Matthias | last3=Stamatakis | first3=Alexandros | last4=Ott | first4=Michael | last5=Rouse | first5=Greg W. | last6=Edgecombe | first6=Gregory D. | last7=Martinez | first7=Pedro | last8=Baguñà | first8=Jaume | last9=Bailly | first9=Xavier | last10=Jondelius | first10=Ulf | last11=Wiens | first11=Matthias | date=2009-12-22 | title=Assessing the root of bilaterian animals with scalable phylogenomic methods | journal=Proceedings of the Royal Society B: Biological Sciences | volume=276 | issue=1677 | pages=4261–4270 | doi=10.1098/rspb.2009.0896 | pmc=2817096 | pmid=19759036}}{{Cite journal | last1=Edgecombe | first1=G. D. | last2=Giribet | first2=G. | last3=Dunn | first3=C. W. | last4=Hejnol | first4=A. | last5=Kristensen | first5=R. M. | last6=Neves | first6=R. C. | last7=Rouse | first7=G. W. | last8=Worsaae | first8=K. | last9=Sørensen | first9=M. V. | year=2011 | title=Higher-level metazoan relationships: Recent progress and remaining questions | journal=Organisms Diversity & Evolution | volume=11 | issue=2 | pages=151–172 | doi=10.1007/s13127-011-0044-4 | bibcode=2011ODivE..11..151E | s2cid=32169826 | url=http://www.brown.edu/Faculty/Dunn_Lab/assets/Edgecombe_etal_2011.pdf}}
• 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 sophisticated models of sequence evolution in phylogenomic studies. There is a conflict between two evolutionary hypotheses, with Xenacoelomorpha being the sister group to Ambulacraria within deuterostomes (i.e., the Xenambulacraria hypothesis) on the one hand,{{cite journal | last1=Philippe | first1=Hervé | last2=Poustka | first2=Albert J. | last3=Chiodin | first3=Marta | last4=Hoff | first4=Katharina J. | last5=Dessimoz | first5=Christophe | last6=Tomiczek | first6=Bartlomiej | last7=Schiffer | first7=Philipp H. | last8=Müller | first8=Steven | last9=Domman | first9=Daryl | last10=Horn | first10=Matthias | last11=Kuhl | first11=Heiner | last12=Timmermann | first12=Bernd | last13=Satoh | first13=Noriyuki | last14=Hikosaka-Katayama | first14=Tomoe | last15=Nakano | first15=Hiroaki | last16=Rowe | first16=Matthew L. | last17=Elphick | first17=Maurice R. | last18=Thomas-Chollier | first18=Morgane | last19=Hankeln | first19=Thomas | last20=Mertes | first20=Florian | last21=Wallberg | first21=Andreas | last22=Rast | first22=Jonathan P. | last23=Copley | first23=Richard R. | last24=Martinez | first24=Pedro | last25=Telford | first25=Maximilian J. | title=Mitigating Anticipated Effects of Systematic Errors Supports Sister-Group Relationship between Xenacoelomorpha and Ambulacraria | journal=Current Biology | volume=29 | issue=11 | year=2019 | pages=1818–1826.e6 | issn=0960-9822 | doi=10.1016/j.cub.2019.04.009 | pmid=31104936 | bibcode=2019CBio...29E1818P | hdl=21.11116/0000-0004-DC4B-1| s2cid=155104811 | hdl-access=free }} and Xenacoelomorpha as sister group to all other bilaterians (i.e., the Nephrozoa hypothesis) on the other.{{Cite journal | doi = 10.1016/j.cub.2014.03.042 | title = Whole-Body Acoel Regeneration is Controlled by Wnt and Bmp-Admp Signaling | journal = Current Biology | volume = 24 | issue = 10 | pages = 1107–13 | year = 2014 | last1 = Srivastava | first1 = M. | last2 = Mazza-Curll | first2 = K. L. | last3 = Van Wolfswinkel | first3 = J. C. | last4 = Reddien | first4 = P. W. | pmid=24768051 | doi-access = free| bibcode = 2014CBio...24.1107S }}{{cite journal | last1=Cannon | first1=Johanna Taylor | last2=Vellutini | first2=Bruno Cossermelli | last3=Smith | first3=Julian | last4=Ronquist | first4=Fredrik | last5=Jondelius | first5=Ulf | last6=Hejnol | first6=Andreas | title=Xenacoelomorpha is the sister group to Nephrozoa | journal=Nature | volume=530 | issue=7588 | year=2016 | pages=89–93 | doi=10.1038/nature16520 | pmid=26842059 | url=http://urn.kb.se/resolve?urn=urn:nbn:se:nrm:diva-1844 | bibcode=2016Natur.530...89C| s2cid=205247296 }} However, the Nephrozoa hypothesis might reflect methodological errors resulting from model violations in the phylogenomic inference.

File:Symsagittifera roscoffensis(Jersey).jpg]]

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.

The epidermal cells of acoelomorphs are unable to proliferate, a feature that is only shared with rhabditophoran 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.{{cite journal|last1=Michalak|first1=Pawel|last2=Egger|first2=Bernhard|last3=Steinke|first3=Dirk|last4=Tarui|first4=Hiroshi|last5=De Mulder|first5=Katrien|last6=Arendt|first6=Detlev|last7=Borgonie|first7=Gaëtan|last8=Funayama|first8=Noriko|last9=Gschwentner|first9=Robert|last10=Hartenstein|first10=Volker|last11=Hobmayer|first11=Bert|last12=Hooge|first12=Matthew|last13=Hrouda|first13=Martina|last14=Ishida|first14=Sachiko|last15=Kobayashi|first15=Chiyoko|last16=Kuales|first16=Georg|last17=Nishimura|first17=Osamu|last18=Pfister|first18=Daniela|last19=Rieger|first19=Reinhard|last20=Salvenmoser|first20=Willi|last21=Smith|first21=Julian|last22=Technau|first22=Ulrich|last23=Tyler|first23=Seth|last24=Agata|first24=Kiyokazu|last25=Salzburger|first25=Walter|last26=Ladurner|first26=Peter|title=To Be or Not to Be a Flatworm: The Acoel Controversy|journal=PLOS ONE|volume=4|issue=5|pages=e5502|year=2009 |doi=10.1371/journal.pone.0005502|pmid=19430533|pmc=2676513|doi-access=free|bibcode=2009PLoSO...4.5502E }}

The nervous 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 the cephalization of the nerve system in more derived bilaterians.{{cite journal|last1=Perea-Atienza|first1=E.|last2=Gavilan|first2=B.|last3=Chiodin|first3=M.|last4=Abril|first4=J. F.|last5=Hoff|first5=K. J.|last6=Poustka|first6=A. J.|last7=Martinez|first7=P.|title=The nervous system of Xenacoelomorpha: a genomic perspective|journal=Journal of Experimental Biology|volume=218|issue=4|year=2015|pages=618–28 |doi=10.1242/jeb.110379|pmid=25696825|doi-access=free|hdl=2445/192702|hdl-access=free}} After decapitation, such a "brain" (rather, a cerebroid ganglion) regenerates in a few weeks.{{cite journal|last1= Sprecher|first1=Simon G.|last2= & al. |title=Functional brain regeneration in the acoel worm Symsagittifera roscoffensis|journal=Biology Open|volume=4|issue=12|year=2015|pages=1688–1695|doi=10.1242/bio.014266 |pmid=26581588|pmc=4736034}}

The sensory organs include a statocyst – which presumably helps them orient to gravity –, and, in some cases, ancestral pigment-spot ocelli capable of detecting light.

Acoelomorphs are simultaneous hermaphrodites, but have no gonads and no ducts associated with the female reproductive system. Instead, gametes are produced from the mesenchymal cells that fill the body between the epidermis and the digestive vacuole.{{cite book |author= Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|page= 229|isbn= 978-0-03-056747-6}}

{{Reflist}}

• [https://web.archive.org/web/20060912001722/http://devbio.umesci.maine.edu/styler/globalworming/pintro.html Phylogeny of Lower Worms of the Meiofauna (Acoelomorpha)]
• [https://web.archive.org/web/20060713063532/http://devbio.umesci.maine.edu/styler/turbellaria/turb2.php?action=1&code=12275 Acoelomorpha at the Turbellarian taxonomic database]
• [https://earthlingnature.wordpress.com/2013/05/30/acoelomorpha-a-phylogenetic-headache/ Acoelomorpha: A Phylogenetic Headache]. [https://earthlingnature.wordpress.com/ Earthling Nature]

{{Animalia}}

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Category:Animal subphyla