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"Acanthognatha" is a similar grouping, with the addition of the [[gastrotrich]]s.<ref name="pmid9809012">{{cite journal |author=Cavalier-Smith T |title=A revised six-kingdom system of life |journal=Biol Rev Camb Philos Soc |volume=73 |issue=3 |pages=203–66 |date=August 1998 |pmid=9809012 |doi= 10.1111/j.1469-185X.1998.tb00030.x|url=http://www3.interscience.wiley.com/resolve/openurl?genre=article&sid=nlm:pubmed&issn=1464-7931&date=1998&volume=73&issue=3&spage=203|archive-url=https://archive.today/20121205040851/http://www3.interscience.wiley.com/resolve/openurl?genre=article&sid=nlm:pubmed&issn=1464-7931&date=1998&volume=73&issue=3&spage=203|dead-url=yes|archive-date=2012-12-05}}</ref>
"Acanthognatha" is a similar grouping, with the addition of the [[gastrotrich]]s.<ref name="pmid9809012">{{cite journal |author=Cavalier-Smith T |title=A revised six-kingdom system of life |journal=Biol Rev Camb Philos Soc |volume=73 |issue=3 |pages=203–66 |date=August 1998 |pmid=9809012 |doi= 10.1111/j.1469-185X.1998.tb00030.x|url=http://www3.interscience.wiley.com/resolve/openurl?genre=article&sid=nlm:pubmed&issn=1464-7931&date=1998&volume=73&issue=3&spage=203|archive-url=https://archive.today/20121205040851/http://www3.interscience.wiley.com/resolve/openurl?genre=article&sid=nlm:pubmed&issn=1464-7931&date=1998&volume=73&issue=3&spage=203|dead-url=yes|archive-date=2012-12-05}}</ref>


==Description==
Possible relationships are as in the cladogram:<ref>{{Cite web|url=http://www.zmuc.dk/InverWeb/Dyr/Limnognathia/phylogeny/phylogeny_UK.htm|title=www.zmuc.dk|last=|first=|date=|website=|access-date=}}</ref><ref>{{Cite journal|last=Fröbius|first=Andreas C.|last2=Funch|first2=Peter|date=2017-04-04|title=Rotiferan Hox genes give new insights into the evolution of metazoan bodyplans|url=http://www.nature.com/articles/s41467-017-00020-w|journal=Nature Communications|language=En|volume=8|issue=1|pages=9|doi=10.1038/s41467-017-00020-w|pmid=28377584|pmc=5431905|issn=2041-1723}}</ref><ref name=marletaz19/><ref name=vinther19/>


The most distinctive characteristic of gnathiferans is the presence of complex sclerotized mouthparts made of chitin.

===Development===

All known gnathiferans are direct developers.<ref name=hejnol15/> Though gnathiferans are included in Spiralia, rotifers and chaetognaths do not exhibit spiral cleavage. Little is known of the development of micrognathozoans.

==Classification==
{{cladogram|caption=Summary of relationships of gnathiferans in recent studies<ref name=marletaz19/><ref name=vinther19/><ref name=frobius17/><ref name=laumer15/><ref name=sielaff16/>
|clades=
{{Clade|{{clade
{{Clade|{{clade
|label1='''Gnathifera'''
|label1='''Gnathifera'''
|1={{clade
|1={{clade
|1=[[Gnathostomulida]]
|1=[[Gnathostomulida]]
|2={{Clade
|2={{Clade
|1=[[Micrognathozoa]]
|1=[[Micrognathozoa]]
|label2=[[Syndermata]]
|2=[[Chaetognatha]]
|label3=[[Rotifera]]
|2={{clade
|1=[[Seisonida]]
|3={{clade
|2={{clade
|1=[[Seisonida]]
|2=[[Acanthocephala]]
|1={{Clade
|2=[[Chaetognatha]]
|3=[[Bdelloidea]]
|label1=[[Rotifera]]
|4=[[Monogononta]]
}}
|1={{clade
}}
|1=[[Bdelloidea]]
|2=[[Monogononta]]
}}}}
|2=[[Acanthocephala]]
}}
}}
}}
}}
}}
|2=[[Platytrochozoa]]
|2=[[Platytrochozoa]]
}}|style=font-size:100%; line-height:100%|label1=[[Spiralia]]}}
}}|style=font-size:100%; line-height:100%|label1=[[Spiralia]]}}
}}
Gnathifera is a member of Spiralia. It is the sister taxon of a clade comprising all other spiralians.<ref name=marletaz19/><ref name=laumer15/> Before the cladistic era, most gnathiferans were regarded as [[aschelminth]]s, a grouping now recognized as [[polyphyly|polyphyletic]].

Chaetognaths exhibit numerous morphological similarities to rotifers, suggesting that they may be sister taxa.<ref name=vinther19/><ref name=frobius17/> However, based on molecular data, micrognathozoans may be more closely related to rotifers than chaetognaths.<ref name=marletaz19/>

The internal relationships of Rotifera are uncertain. Rotifera comprises four subclades: [[Seisonida]], [[Acanthocephala]], [[Bdelloidea]], and [[Monogononta]]. Acanthocephalans were traditionally excluded from Rotifera, but it is now known that rotifers are paraphyletic without including acanthocephalans. Some taxonomists call the clade of rotifers including acanthocephalans Syndermata, but others continue to use Rotifera and regard acanthocephalans as rotifers.<ref name=laumer15/>

The engimatic phylum [[Cycliophora]] may belong to Gnathifera, but other studies suggest that it is more closely related to the [[Entoprocta]].


==Fossil record==
==Fossil record==
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{{Reflist|refs=
{{Reflist|refs=
<ref name=caron19>{{cite journal | first1 = Jean-Bernard | last1 = Caron | first2 = Brittany | last2 = Cheung | year = 2019 | title = Amiskwia is a large Cambrian gnathiferan with complex gnathostomulid-like jaws | url = https://www.nature.com/articles/s42003-019-0388-4 | journal = Communications Biology | volume = 2 | doi = 10.1038/s42003-019-0388-4 | doi-access = free }}</ref>
<ref name=caron19>{{cite journal | first1 = Jean-Bernard | last1 = Caron | first2 = Brittany | last2 = Cheung | year = 2019 | title = Amiskwia is a large Cambrian gnathiferan with complex gnathostomulid-like jaws | url = https://www.nature.com/articles/s42003-019-0388-4 | journal = Communications Biology | volume = 2 | doi = 10.1038/s42003-019-0388-4 | doi-access = free }}</ref>
<ref name=frobius17>{{cite journal | last1 = Fröbius | first1 = Andreas C. | last2 = Funch | first2 = Peter | year = 2017 | title = Rotiferan Hox genes give new insights into the evolution of metazoan bodyplans | url = http://www.nature.com/articles/s41467-017-00020-w | journal = Nature Communications | volume = 8 | issue = 1 | pages = 9 | doi = 10.1038/s41467-017-00020-w | pmid = 28377584 | pmc = 5431905 }}</ref>
<ref name=hejnol15>{{cite journal | doi = 10.1007/978-3-7091-1871-9_1 }}</ref>
<ref name=kristensen02>{{cite journal | last1 = Kristensen | first1 = Reinhardt Møbjerg | title = An Introduction to Loricifera, Cycliophora, and Micrognathozoa | journal = Integrative and Comparative Biology | volume=42 | issue = 3| year = 2002 | pages = 641–651 | doi=10.1093/icb/42.3.641 }}</ref>
<ref name=kristensen02>{{cite journal | last1 = Kristensen | first1 = Reinhardt Møbjerg | title = An Introduction to Loricifera, Cycliophora, and Micrognathozoa | journal = Integrative and Comparative Biology | volume=42 | issue = 3| year = 2002 | pages = 641–651 | doi=10.1093/icb/42.3.641 }}</ref>
<ref name=laumer15>{{cite journal | first1 = Christopher E. | last1 = Laumer | first2 = Nicolas | last2 = Bekkouche | first3 = Alexandra | last3 = Kerbl | first4 = Freya | last4 = Goetz | first5 = Ricardo C. | last5 = Neves | first6 = Martin V. | last6 = Sørensen | first7 = Reinhardt M. | last7 = Kristensen | first8 = Andreas | last8 = Hejnol | first9 = Casey W. | last9 = Dunn | first10 = Gonzalo | last10 = Giribet | first11 = Katrine | last11 = Worsaae | year = 2015 | title = Spiralian phylogeny informs the evolution of microscopic lineages | journal = Current Biology | volume = 25 | pages = 2000–2006 | doi = 10.1016/j.cub.2015.06.068 }}</ref>
<ref name=marletaz19>{{Cite journal | last5 = Rokhsar | first5 = Daniel S.| last4 = Satoh | first4 = Noriyuki | last3 = Goto | first3 = Taichiro | last2 = Peijnenburg | first2 = Katja T. C. A. | last1 = Marlétaz | first1 = Ferdinand | year = 2019 | title = A new spiralian phylogeny places the enigmatic arrow worms among gnathiferans | url = https://www.cell.com/current-biology/abstract/S0960-9822(18)31541-0 | url-access = subscription | journal = Current Biology | volume = 29 | issue = 2 | pages = 312–318.e3 | doi=10.1016/j.cub.2018.11.042 | pmid = 30639106}}</ref>
<ref name=marletaz19>{{Cite journal | last5 = Rokhsar | first5 = Daniel S.| last4 = Satoh | first4 = Noriyuki | last3 = Goto | first3 = Taichiro | last2 = Peijnenburg | first2 = Katja T. C. A. | last1 = Marlétaz | first1 = Ferdinand | year = 2019 | title = A new spiralian phylogeny places the enigmatic arrow worms among gnathiferans | url = https://www.cell.com/current-biology/abstract/S0960-9822(18)31541-0 | url-access = subscription | journal = Current Biology | volume = 29 | issue = 2 | pages = 312–318.e3 | doi=10.1016/j.cub.2018.11.042 | pmid = 30639106}}</ref>
<ref name=piper13>{{cite book | first1 = Ross | last1 = Piper | title = Animal Earth: The Amazing Diversity of Living Creatures | publisher = Thames & Hudson | year = 2013 }}</ref>
<ref name=piper13>{{cite book | first1 = Ross | last1 = Piper | title = Animal Earth: The Amazing Diversity of Living Creatures | publisher = Thames & Hudson | year = 2013 }}</ref>
<ref name=poinar92>{{cite journal | first1 = G. O. | last1 = Poinar | first2 = C. | last2 = Ricci | year = 1992 | title = Bdelloid rotifers in Dominican amber: evidence for parthenogenetic continuity | journal = Experientia | volume = 48 | pages = 408–410 }}</ref>
<ref name=poinar92>{{cite journal | first1 = G. O. | last1 = Poinar | first2 = C. | last2 = Ricci | year = 1992 | title = Bdelloid rotifers in Dominican amber: evidence for parthenogenetic continuity | journal = Experientia | volume = 48 | pages = 408–410 }}</ref>
<ref name=sielaff16>{{cite journal | doi = 10.1016/j.ympev.2015.11.017 }}</ref>
<ref name=szaniawski02>{{cite journal | first1 = Hubert | last1 = Szaniawski | year = 2002 | title = New evidence for the protoconodont origin of chaetognaths | journal = Acta Palaeontologica Polonica | volume = 47 | issue = 3 | pages = 405–419 | url = https://www.app.pan.pl/article/item/app47-405.html }}</ref>
<ref name=szaniawski02>{{cite journal | first1 = Hubert | last1 = Szaniawski | year = 2002 | title = New evidence for the protoconodont origin of chaetognaths | journal = Acta Palaeontologica Polonica | volume = 47 | issue = 3 | pages = 405–419 | url = https://www.app.pan.pl/article/item/app47-405.html }}</ref>
<ref name=vannier07>{{cite journal | first1 = J. | last1 = Vannier | first2 = M. | last2 = Steiner | first3 = E. | last3 = Renvoisé | first4 = S.-X. | last4 = Hu | first5 = J.-P. | last5 = Casanova | year = 2007 | title = Early Cambrian origin of modern food webs: evidence from predator arrow worms | journal = Proceedings of the Royal Society B | volume = 274 | pages = 627–633 | doi = 10.1098/rspb.2006.3761 | pmc = 2197202 }}</ref>
<ref name=vannier07>{{cite journal | first1 = J. | last1 = Vannier | first2 = M. | last2 = Steiner | first3 = E. | last3 = Renvoisé | first4 = S.-X. | last4 = Hu | first5 = J.-P. | last5 = Casanova | year = 2007 | title = Early Cambrian origin of modern food webs: evidence from predator arrow worms | journal = Proceedings of the Royal Society B | volume = 274 | pages = 627–633 | doi = 10.1098/rspb.2006.3761 | pmc = 2197202 }}</ref>

Revision as of 22:49, 21 May 2019

Gnathifera
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
(unranked): Spiralia
Clade: Gnathifera
Ahlrichs, 1995
Phyla

Gnathifera, from the Greek gnathos, “jaw”, and the Latin fera, “to bear,” is a clade of spiralians comprising the phyla Gnathostomulida, Rotifera, Micrognathozoa, and Chaetognatha.[1][2] It may also include the Cycliophora.[3]

"Acanthognatha" is a similar grouping, with the addition of the gastrotrichs.[4]

Description

The most distinctive characteristic of gnathiferans is the presence of complex sclerotized mouthparts made of chitin.

Development

All known gnathiferans are direct developers.[5] Though gnathiferans are included in Spiralia, rotifers and chaetognaths do not exhibit spiral cleavage. Little is known of the development of micrognathozoans.

Classification

Spiralia
Summary of relationships of gnathiferans in recent studies[1][6][7][8][9]

Gnathifera is a member of Spiralia. It is the sister taxon of a clade comprising all other spiralians.[1][8] Before the cladistic era, most gnathiferans were regarded as aschelminths, a grouping now recognized as polyphyletic.

Chaetognaths exhibit numerous morphological similarities to rotifers, suggesting that they may be sister taxa.[6][7] However, based on molecular data, micrognathozoans may be more closely related to rotifers than chaetognaths.[1]

The internal relationships of Rotifera are uncertain. Rotifera comprises four subclades: Seisonida, Acanthocephala, Bdelloidea, and Monogononta. Acanthocephalans were traditionally excluded from Rotifera, but it is now known that rotifers are paraphyletic without including acanthocephalans. Some taxonomists call the clade of rotifers including acanthocephalans Syndermata, but others continue to use Rotifera and regard acanthocephalans as rotifers.[8]

The engimatic phylum Cycliophora may belong to Gnathifera, but other studies suggest that it is more closely related to the Entoprocta.

Fossil record

The fossil record of gnathiferans is poor. There are no known fossil gnathostomulids.[10] Fossils of the extant rotifer genus Habrotrocha are known from Dominican amber dating to the late Eocene, but rotifers are otherwise only known from the Holocene.[11][12] By contrast, the chaetognath fossil record, while still patchy, includes numerous Paleozoic specimens.[13] Protoconodonts are stem-group chaetognaths.[14] The enigmatic Cambrian taxon Amiskwia is a gnathiferan and may also be a stem-group chaetognath.[6][15] The Cambrian ectoparasite Inquicus appears to be a gnathiferan.[6]

References

  1. ^ a b c d Marlétaz, Ferdinand; Peijnenburg, Katja T. C. A.; Goto, Taichiro; Satoh, Noriyuki; Rokhsar, Daniel S. (2019). "A new spiralian phylogeny places the enigmatic arrow worms among gnathiferans". Current Biology. 29 (2): 312–318.e3. doi:10.1016/j.cub.2018.11.042. PMID 30639106.
  2. ^ Suga K, Mark Welch D, Tanaka Y, Sakakura Y, Hagiwara A (1 August 2007). Ellegren H (ed.). "Analysis of expressed sequence tags of the cyclically parthenogenetic rotifer Brachionus plicatilis". PLoS ONE. 2 (7): e671. doi:10.1371/journal.pone.0000671. PMC 1925144. PMID 17668053.{{cite journal}}: CS1 maint: unflagged free DOI (link) Open access icon
  3. ^ Kristensen, Reinhardt Møbjerg (2002). "An Introduction to Loricifera, Cycliophora, and Micrognathozoa". Integrative and Comparative Biology. 42 (3): 641–651. doi:10.1093/icb/42.3.641.
  4. ^ Cavalier-Smith T (August 1998). "A revised six-kingdom system of life". Biol Rev Camb Philos Soc. 73 (3): 203–66. doi:10.1111/j.1469-185X.1998.tb00030.x. PMID 9809012. Archived from the original on 2012-12-05. {{cite journal}}: Unknown parameter |dead-url= ignored (|url-status= suggested) (help)
  5. ^ . doi:10.1007/978-3-7091-1871-9_1. {{cite journal}}: Cite journal requires |journal= (help); Missing or empty |title= (help)
  6. ^ a b c d Vinther, Jakob; Parry, Luke A. (2019). "Bilateral jaw elements in Amiskwia sagittiformis bridge the morphological gap between gnathiferans and chaetognaths". Current Biology. 29 (5): 881–888.e1. doi:10.1016/j.cub.2019.01.052. PMID 30799238.
  7. ^ a b Fröbius, Andreas C.; Funch, Peter (2017). "Rotiferan Hox genes give new insights into the evolution of metazoan bodyplans". Nature Communications. 8 (1): 9. doi:10.1038/s41467-017-00020-w. PMC 5431905. PMID 28377584.
  8. ^ a b c Laumer, Christopher E.; Bekkouche, Nicolas; Kerbl, Alexandra; Goetz, Freya; Neves, Ricardo C.; Sørensen, Martin V.; Kristensen, Reinhardt M.; Hejnol, Andreas; Dunn, Casey W.; Giribet, Gonzalo; Worsaae, Katrine (2015). "Spiralian phylogeny informs the evolution of microscopic lineages". Current Biology. 25: 2000–2006. doi:10.1016/j.cub.2015.06.068.
  9. ^ . doi:10.1016/j.ympev.2015.11.017. {{cite journal}}: Cite journal requires |journal= (help); Missing or empty |title= (help)
  10. ^ Piper, Ross (2013). Animal Earth: The Amazing Diversity of Living Creatures. Thames & Hudson.
  11. ^ Poinar, G. O.; Ricci, C. (1992). "Bdelloid rotifers in Dominican amber: evidence for parthenogenetic continuity". Experientia. 48: 408–410.
  12. ^ Waggoner, B. M.; Poinar, G. O. (1993). "Fossil habrotrochid rotifers in Dominican amber". Experientia. 49: 354–357.
  13. ^ Vannier, J.; Steiner, M.; Renvoisé, E.; Hu, S.-X.; Casanova, J.-P. (2007). "Early Cambrian origin of modern food webs: evidence from predator arrow worms". Proceedings of the Royal Society B. 274: 627–633. doi:10.1098/rspb.2006.3761. PMC 2197202.
  14. ^ Szaniawski, Hubert (2002). "New evidence for the protoconodont origin of chaetognaths". Acta Palaeontologica Polonica. 47 (3): 405–419.
  15. ^ Caron, Jean-Bernard; Cheung, Brittany (2019). "Amiskwia is a large Cambrian gnathiferan with complex gnathostomulid-like jaws". Communications Biology. 2. doi:10.1038/s42003-019-0388-4.