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Gnathifera (Spiralia)

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Gnathifera (from the Greek gnáthos, “jaw”, and the Latin -fera, “bearing”) is a clade of generally small spiralians characterized by complex jaws made of chitin. It comprises the phyla Gnathostomulida, Rotifera and Micrognathozoa.[1] Chaetognatha has recently been recognised as closely related to the group, with it either being included within Gnathifera[1] or the broader group Chaetognathifera.[2]

Gnathifera
Temporal range: Fortunian – Recent
Pseudosagitta maxima (Chaetognatha)
Brachionus quadridentatus (Rotifera)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Clade: Bilateria
Clade: Nephrozoa
(unranked): Protostomia
(unranked): Spiralia
Clade: Gnathifera
Ahlrichs, 1995
Phyla

Gnathiferans include some of the most abundant phyla. Rotifers are among the most diverse and abundant freshwater animals and chaetognaths are among the most abundant marine plankton.[3][4]

Description

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Comparison of the head region of Gnathifera

The most distinctive characteristic of gnathiferans is the presence of complex sclerotized mouthparts made of chitin.[1]

In most gnathiferans, the anus opens on the dorsal surface of the animal.[5][6][7] In micrognathozoans and gnathostomulids, the anus is transient and only forms during defecation.[6][7] Unlike other gnathiferans, in chaetognaths and the extinct Amiskwia the anus is located on the ventral surface in a subterminal position.[8][9]

Both Gnathostomulida and Micrognathozoa are acoelomates, rotifers are pseudocoelomates, and chaetognathans have a true coelom. Gnathostomulidans and chaetognathans are hermaphrodites, and Micrognathozoa appears to be parthenogenetic as no males have ever been observed, but it has been suggested undiscovered dwarf males could be responsible for fertilization. In rotifers both males and females occurs, except from the class Bdelloidea.[10][11][12][13][14][15][16]

Development

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All known gnathiferans are direct developers.[17] Though gnathiferans are included in Spiralia, rotifers and chaetognaths do not exhibit spiral cleavage.[17][1] Little is known of the development of micrognathozoans.[17] The development of gnathostomulids is poorly known, but they appear to exhibit spiral cleavage.[17][1]

Classification

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Spiralia
Summary of relationships of gnathiferans in recent studies, with disputed relationships represented as polytomies[1][5][18][19][20]

Gnathifera is a member of Spiralia. It is the sister taxon of a clade comprising all other spiralians.[1][19] An alternative phylogeny place Gnathifera into a main spiralian clade Platyzoa s.l. as sister clade to Mesozoa and Platyhelminthes.[21] 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.[5][18] However, based on molecular data, micrognathozoans may be more closely related to rotifers than chaetognaths.[1]

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.[19] Numerous hypotheses of rotifer interrelationships exist.[19][20]

The enigmatic phylum Cycliophora may belong to Gnathifera, but other studies suggest that it is more closely related to the Entoprocta.[22]

Fossil record

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Amiskwia sagittiformis a basal gnathiferan from Cambrian of Canada, which is much larger than modern gnathiferans

The fossil record of gnathiferans is poor. There are no known fossil gnathostomulids.[23] 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.[24][25] By contrast, the chaetognath fossil record, while still patchy, includes numerous Paleozoic specimens.[26] Protoconodonts are stem-group chaetognaths.[27] The earliest protoconodonts date to the Fortunian age of the Cambrian, and are among the oldest known bilaterians.[28] The enigmatic Cambrian taxon Amiskwia is a gnathiferan and may also be a stem-group chaetognath.[5][8] The Cambrian ectoparasite Inquicus appears to be a gnathiferan.[5]

List of Genera

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History

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Gnathifera was named in 1995 to unite gnathostomulids and rotifers. Micrognathozoans were soon added to this grouping. Chaetognaths, long considered a distinct lineage with no close relatives, were identified as gnathiferans in 2019.[1]

A similar grouping, Acanthognatha, was suggested in 1998 to unite gastrotrichs with gnathostomulids and rotifers.[29] However, gastrotrichs are more closely related to lophotrochozoans than gnathiferans.[19][1]

References

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  1. ^ a b c d e f g h i j 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. Bibcode:2019CBio...29E.312M. doi:10.1016/j.cub.2018.11.042. PMID 30639106.
  2. ^ Park, Tae-Yoon S.; Nielsen, Morten Lunde; Parry, Luke A.; Sørensen, Martin Vinther; Lee, Mirinae; Kihm, Ji-Hoon; Ahn, Inhye; Park, Changkun; de Vivo, Giacinto; Smith, M. Paul; Harper, David A. T.; Nielsen, Arne T.; Vinther, Jakob (2024-01-05). "A giant stem-group chaetognath". Science Advances. 10 (1): eadi6678. Bibcode:2024SciA...10I6678P. doi:10.1126/sciadv.adi6678. ISSN 2375-2548. PMC 10796117. PMID 38170772.
  3. ^ Suga K, Mark Welch D, Tanaka Y, Sakakura Y, Hagiwara A (2007). "Analysis of expressed sequence tags of the cyclically parthenogenetic rotifer Brachionus plicatilis". PLOS ONE. 2 (7): e671. Bibcode:2007PLoSO...2..671S. doi:10.1371/journal.pone.0000671. PMC 1925144. PMID 17668053.
  4. ^ Longhurst, Alan R. (1985). "The structure and evolution of plankton communities". Progress in Oceanography. 15 (1): 1–35. Bibcode:1985PrOce..15....1L. doi:10.1016/0079-6611(85)90036-9.
  5. ^ a b c d e 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. Bibcode:2019CBio...29E.881V. doi:10.1016/j.cub.2019.01.052. hdl:1983/51b1b6c1-0220-4469-977f-480e847a9101. PMID 30799238.
  6. ^ a b Knauss, Elizabeth B. (1979). "Indication of an anal pore in Gnathostomulida". Zoologica Scripta. 8 (1–4): 181–186. doi:10.1111/j.1463-6409.1979.tb00630.x. S2CID 84391082.
  7. ^ a b Kristensen, Reinhardt Møbjerg; Funch, Peter (2000). "Micrognathozoa: a new class with complicated jaws like those of Rotifera and Gnathostomulida". Journal of Morphology. 246 (1): 1–49. doi:10.1002/1097-4687(200010)246:1<1::AID-JMOR1>3.0.CO;2-D. PMID 11015715. S2CID 13294045.
  8. ^ a b Caron, Jean-Bernard; Cheung, Brittany (2019). "Amiskwia is a large Cambrian gnathiferan with complex gnathostomulid-like jaws". Communications Biology. 2: 164. doi:10.1038/s42003-019-0388-4. PMC 6499802. PMID 31069273.
  9. ^ Arnaud, Jean; Brunet, Michel; Casanova, Jean-Paul; Mazza, Jacques; Pasqualini, Vanina (1996). "Morphology and ultrastructure of the gut in Spadella cephaloptera (Chaetognatha)". Journal of Morphology. 228 (1): 27–44. doi:10.1002/(SICI)1097-4687(199604)228:1<27::AID-JMOR3>3.0.CO;2-M. PMID 29852579. S2CID 46918465.
  10. ^ The Gnathifera
  11. ^ Rotifers as live feed
  12. ^ Struck, Torsten H.; Wey-Fabrizius, Alexandra R.; Golombek, Anja; Hering, Lars; Weigert, Anne; Bleidorn, Christoph; Klebow, Sabrina; Iakovenko, Nataliia; Hausdorf, Bernhard; Petersen, Malte; Kück, Patrick; Herlyn, Holger; Hankeln, Thomas (July 2014). "Platyzoan Paraphyly Based on Phylogenomic Data Supports a Noncoelomate Ancestry of Spiralia". Molecular Biology and Evolution. 31 (7): 1833–1849. doi:10.1093/molbev/msu143. PMID 24748651.
  13. ^ Phylum Chaetognatha - Australian Faunal Directory
  14. ^ Morphology of the nervous system of monogonont rotifer Epiphanes senta with a focus on sexual dimorphism between feeding females and dwarf males
  15. ^ Yin, Xu-Wang; Tan, Bing-Bing; Zhou, Yan-Chun; Li, Xiao-Chun; Liu, Wei (2016). "Development time of male and female rotifers with sexual size dimorphism". Hydrobiologia. 767 (1): 27–35. doi:10.1007/s10750-015-2472-1.
  16. ^ Birky, C. William (2004). "Bdelloid rotifers revisited". Proceedings of the National Academy of Sciences of the United States of America. 101 (9): 2651–2652. Bibcode:2004PNAS..101.2651B. doi:10.1073/pnas.0308453101. ISSN 0027-8424. PMC 365675. PMID 14981265.
  17. ^ a b c d Hejnol, Andreas (2015). "Gnathifera". In Wanninger, A. (ed.). Evolutionary Developmental Biology of Invertebrates 2: Lophotrochozoa (Spiralia). Springer. pp. 1–12. doi:10.1007/978-3-7091-1871-9_1. ISBN 978-3-7091-1870-2.
  18. ^ 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. Bibcode:2017NatCo...8....9F. doi:10.1038/s41467-017-00020-w. PMC 5431905. PMID 28377584.
  19. ^ a b c d e 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 (15): 2000–2006. Bibcode:2015CBio...25.2000L. doi:10.1016/j.cub.2015.06.068. PMID 26212884.
  20. ^ a b Sielaff, Malte; Schmidt, Hanno; Struck, Torsten H.; Rosenkranz, David; Mark Welch, David B.; Hankeln, Thomas; Herlyn, Holger (2016). "Phylogeny of Syndermata (syn. Rotifera): Mitochondrial gene order verifies epizoic Seisonidea as sister to endoparasitic Acanthocephala within monophyletic Hemirotifera". Molecular Phylogenetics and Evolution. 96: 79–92. Bibcode:2016MolPE..96...79S. doi:10.1016/j.ympev.2015.11.017. PMID 26702959.
  21. ^ Drábková, Marie; Kocot, Kevin M.; Halanych, Kenneth M.; Oakley, Todd H.; Moroz, Leonid L.; Cannon, Johanna T.; Kuris, Armand; Garcia-Vedrenne, Ana Elisa; Pankey, M. Sabrina; Ellis, Emily A.; Varney, Rebecca; Štefka, Jan; Zrzavý, Jan (6 July 2022). "Different phylogenomic methods support monophyly of enigmatic 'Mesozoa' (Dicyemida Orthonectida, Lophotrochozoa)". Proceedings of the Royal Society B: Biological Sciences. 289 (1978). doi:10.1098/rspb.2022.0683. ISSN 1471-2954. PMC 9257288. PMID 35858055.
  22. ^ 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. PMID 21708760.
  23. ^ Piper, Ross (2013). Animal Earth: The Amazing Diversity of Living Creatures. Thames & Hudson.
  24. ^ Poinar, G. O.; Ricci, C. (1992). "Bdelloid rotifers in Dominican amber: evidence for parthenogenetic continuity". Experientia. 48 (4): 408–410. doi:10.1007/bf01923444. S2CID 13098228.
  25. ^ Waggoner, B. M.; Poinar, G. O. (1993). "Fossil habrotrochid rotifers in Dominican amber". Experientia. 49 (4): 354–357. doi:10.1007/bf01923421. S2CID 28087284.
  26. ^ 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 (1610): 627–633. doi:10.1098/rspb.2006.3761. PMC 2197202. PMID 17254986.
  27. ^ Szaniawski, Hubert (2002). "New evidence for the protoconodont origin of chaetognaths". Acta Palaeontologica Polonica. 47 (3): 405–419.
  28. ^ Kouchinsky, Artem; Bengtson, Stefan; Runnegar, Bruce; Skovsted, Christian; Steiner, Michael; Vendrasco, Michael (2011). "Chronology of early Cambrian biomineralization". Geological Magazine. 149 (2): 221–251. doi:10.1017/S0016756811000720.
  29. ^ Cavalier-Smith, T. (1998). "A revised six-kingdom system of life". Biological Reviews. 73 (3): 203–266. doi:10.1111/j.1469-185X.1998.tb00030.x. PMID 9809012. S2CID 6557779.
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