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Vetulicolia

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Vetulicolia
Temporal range: about 535–501 Ma Fortunian—Drumian
Fossil of Vetulicola cuneata
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Superphylum: Deuterostomia
Phylum: †Vetulicolia
Shu et al. 2001
Type species
†Vetulicola cuneata
Hou, 1987
Classes

Vetulicolia[note 1] is a phylum of bilaterian animals encompassing several extinct species belonging to the Cambrian period.[1][2] The phylum was created by Degan Shu and his research team in 2001,[3] and named after Vetulicola cuneata, the first species of the phylum described in 1987.[4]

The vetulicolian body plan comprises two parts: a voluminous rostral (anterior) forebody, tipped with an anteriorly positioned mouth and lined with a lateral row of five round to oval-shaped openings on each side, which have been interpreted as gills (or at least orifices in the vicinity of the pharynx); and a caudal (posterior) section that primitively comprises seven body segments and functions as a tail. All vetulicolians lack preserved appendages of any kind, having no legs, feelers or even eye spots.[5] The area where the anterior and posterior parts join is constricted.[6]

Their taxonomic affinity has been uncertain; they have been considered to represent stem- and crown-group arthropods, stem-group vertebrates,[7] and early deuterostomes (a group which as a whole includes the vertebrates, other invertebrate chordates, echinoderms and hemichordates). The general scientific consensus before 2001 considered them early limbless arthropods but now considers them early deuterostomes.[8] Vetulicolian fossils examined in 2014 show the presence of notochord-like structures, and it was concluded that vetulicolians are crown-group chordates and probably related to modern tunicates.[9] Research from 2017 rather indicates vetulicolians are related to Saccorhytus, another basal deuterostome group,[10] although another study shows possibility that Saccorhytus is ecdysozoan instead of deuterostome.[11]

Taxonomy and evolution

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See also: List of bilaterial animal orders

Vetulicolia as a phylum was created by Chinese palaeontologist Degan Shu and his research team at the Northwest University in Xi'an, China, along with Simon Conway Morris at the University of Cambridge in 2001.[3] The name was derived from Vetulicola cuneata, the first species described by Hou Xian-guang in 1987 from the Lower Cambrian Chiungchussu Formation in Chengjiang, China.[2][4] The purpose was to include all related Cambrian animals known under the families Didazoonidae (Didazoon, Pomatrum, and Xidazoon) and Vetulicolidae (Vetulicola, and Banffia) as a separate group of animals.[1][3] Other animals which may be related include the yunnanozoans.[3]

The taxonomic placement of the vetulicolians remains controversial. The original researcher Shu is of the opinion that the vetulicolians probably represent an early type of deuterostomes, and that this implies that segmentation in cephalochordates and vertebrates may be derived from the common ancestor of protostomes and deuterostomes.[7] However, Derek Briggs of Yale University and his team who described Skeemella from the Middle Cambrian of Utah regard it as having affinity to protostomes with important arthropod features, thus confounding assignment of Vetulicolia to Deuterostomia.[12] Thurston C. Lacalli of the University of Saskatchewan, Saskatoon, Canada, agreed that the group are deuterostomes, but suggested that the animals were more likely related to tunicates.[13]

Dominguez and Jefferies have argued, based on morphological analysis, that Vetulicola (and by implication, other vetulicolians) is a urochordate, and probably a stem-group larvacean. Some question the relation to tunicates and larvaceans, as there is no evidence of segmentation in tunicates, larval or adult, that is comparable to segmentation in vetulicolians, that the anus of urochordates is within the atrium, while that of vetulicolians is positioned at the terminal end of the tail, and, perhaps most importantly, there is no exhalant siphon, or analogous structure, seen in vetulicolians.[14] However, a discovery of a new vetulicolian, Nesonektris aldridgei, from Australia in 2014 supported a position close to urochordates for vetulicolians.[9]

A 2024 phylogenetic analysis by Mussini and colleagues found vetulicolians to be a paraphyletic group of stem-chordates, lying outside a clade formed by Yunnanozoon, Cathaymyrus, Pikaia and crown-chordates.[15]

Chordata
Vetulicolia

The monophyly of Deuterostomia is not strongly evidenced.[16][17] If deuterostomes are paraphyletic, pharyngeal slits were probably present in the common ancestor of Bilateria and vetulicolians may be stem-group protostomes that retained the ancestral pharyngeal slits.[17] If this is the case, banffids, which appear to lack pharyngeal slits, may be more closely related to crown-group protostomes than other vetulicolians.[17]

Ecology and lifestyle

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Fossil of Skeemella clavula

From their superficially tadpole-like forms, leaf or paddle-shaped tails, and various degrees of streamlining, it is assumed that all vetulicolians discovered to date were swimming animals that spent much, if not all, of their time living in water.[14] Some groups, like the genus Vetulicola, were more streamlined (complete with ventral keels) than other groups, such as the tadpole-like Didazoonidae.[14]

Because all vetulicolians had mouths which had no features for chewing or grasping, it is assumed that they were not predators.[14] Since vetulicolians possessed gill slits, many researchers regard these organisms as planktivores. The sediment infills in the guts of their fossils have caused some to suggest that they were deposit feeders. This idea has been contested, as deposit feeders tend to have straight guts, whereas the hindguts of vetulicolians were spiral-shaped. Some researchers propose that the vetulicolians were "selective deposit-feeders" which actively swam from one region of the seafloor to another, while supplementing their nutrition with filter-feeding.[14]

Classification

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Notes

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  1. ^ The taxon name, Vetulocolia, is derived from the type genus, Vetulicola, which is a compound Latin word composed of vetuli "old" and cola "inhabitant".

References

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  1. ^ a b Shu, Degan (2005). "On the Phylum Vetulicolia". Chinese Science Bulletin. 50 (20): 2342–2354. Bibcode:2005ChSBu..50.2342S. doi:10.1007/BF03183746. ISSN 1001-6538. S2CID 86827605.
  2. ^ a b McMenamin, Mark A. S. (2019). "Cambrian Chordates and Vetulicolians". Geosciences. 9 (8): 354. Bibcode:2019Geosc...9..354M. doi:10.3390/geosciences9080354. ISSN 2076-3263.
  3. ^ a b c d Shu, D. G.; Conway Morris, S.; Han, J.; Chen, L.; Zhang, X. L.; Zhang, Z. F.; Liu, H. Q.; Li, Y.; Liu, J. N. (2001-11-22). "Primitive deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China)". Nature. 414 (6862): 419–424. Bibcode:2001Natur.414..419S. doi:10.1038/35106514. ISSN 0028-0836. PMID 11719797. S2CID 4345484.
  4. ^ a b Briggs, Derek E. G.; Fortey, Richard A. (2005). "Wonderful strife: systematics, stem groups, and the phylogenetic signal of the Cambrian radiation". Paleobiology. 31 (2_Suppl): 94–112. doi:10.1666/0094-8373(2005)031[0094:WSSSGA]2.0.CO;2. ISSN 0094-8373. S2CID 44066226.
  5. ^ Ailin, C.; Hongzhen, F.; Maoyan, Z.H.U.; Dongsheng, M.A.; Ming, L.I. (2003), "A New Vetulicolian from the Early Cambrian Chengjiang Fauna in Yunnan of China", Acta Geologica Sinica, 77 (3): 281–287, doi:10.1111/j.1755-6724.2003.tb00742.x, S2CID 128895860[dead link]
  6. ^ a b McMenamin, Mark A. S. (11 August 2019). "Cambrian Chordates and Vetulicolians". Geosciences. 9 (8): 354. Bibcode:2019Geosc...9..354M. doi:10.3390/geosciences9080354.
  7. ^ a b Shu, Degan (2003). "A paleontological perspective of vertebrate origin". Chinese Science Bulletin. 48 (8): 725–735. Bibcode:2003ChSBu..48..725S. doi:10.1007/BF03187041. S2CID 85163902.
  8. ^ Dominguez, Patricio and Jeffries, Richard. (2003). Fossil evidence on the origin of appendicularians. Paper read at International Urochordate Meeting 2003. Abstract at [2] – URL retrieved June 22, 2006.
  9. ^ a b c García-Bellido, Diego C; Paterson, John R (2014). "A new vetulicolian from Australia and its bearing on the chordate affinities of an enigmatic Cambrian group". BMC Evolutionary Biology. 14: 214. doi:10.1186/s12862-014-0214-z. PMC 4203957. PMID 25273382.
  10. ^ Han, Jian; Conway Morris, Simon; Ou, Qiang; Shu, Degan; Huang, Hai (2017). "Meiofaunal deuterostomes from the basal Cambrian of Shaanxi (China)". Nature. 542 (7640): 228–231. Bibcode:2017Natur.542..228H. doi:10.1038/nature21072. PMID 28135722. S2CID 353780.
  11. ^ Liu, Yunhuan; Carlisle, Emily; Zhang, Huaqiao; Yang, Ben; Steiner, Michael; Shao, Tiequan; Duan, Baichuan; Marone, Federica; Xiao, Shuhai; Donoghue, Philip C. J. (2022-08-17). "Saccorhytus is an early ecdysozoan and not the earliest deuterostome". Nature. 609 (7927): 541–546. Bibcode:2022Natur.609..541L. doi:10.1038/s41586-022-05107-z. hdl:1983/454e7bec-4cd4-4121-933e-abeab69e96c1. ISSN 1476-4687. PMID 35978194. S2CID 251646316.
  12. ^ Briggs; et al. (2005). "A new metazoan from the Middle Cambrian of Utah and the nature of the Vetulicolia". Palaeontology. 48 (4): 681–686. doi:10.1111/j.1475-4983.2005.00489.x.
  13. ^ Lacalli, Thurston C. (2002). "Vetulicolians—are they deuterostomes? chordates?". BioEssays. 24 (3): 208–211. doi:10.1002/bies.10064. PMID 11891757.
  14. ^ a b c d e f Aldridge, Richard J.; Hou, Xian-guang; Siveter, David J.; Siveter, Derek J.; Sarah E., Gabbott (2007). "The systematics and phylogenetic relationships of vetulicolians". Palaeontology. 50: 131–168. doi:10.1111/j.1475-4983.2006.00606.x. S2CID 85722738.
  15. ^ Mussini, G.; Smith, M. P.; Vinther, J.; Rahman, I. A.; Murdock, D. J. E.; Harper, D. A. T.; Dunn, F. S. (2024). "A new interpretation of Pikaia reveals the origins of the chordate body plan". Current Biology. doi:10.1016/j.cub.2024.05.026.
  16. ^ Philippe, Hervé; Poustka, Albert J.; Chiodin, Marta; Hoff, Katharina J.; Dessimoz, Christophe; Tomiczek, Bartlomiej; Schiffer, Philipp H.; Müller, Steven; Domman, Daryl; Horn, Matthias; Kuhl, Heiner; Timmermann, Bernd; Satoh, Noriyuki; Hikosaka-Katayama, Tomoe; Nakano, Hiroaki; Rowe, Matthew L.; Elphick, Maurice R.; Thomas-Chollier, Morgane; Hankeln, Thomas; Mertes, Florian; Wallberg, Andreas; Rast, Jonathan P.; Copley, Richard R.; Martinez, Pedro; Telford, Maximilian J. (2019). "Mitigating Anticipated Effects of Systematic Errors Supports Sister-Group Relationship between Xenacoelomorpha and Ambulacraria". Current Biology. 29 (11): 1818–1826.e6. doi:10.1016/j.cub.2019.04.009. hdl:21.11116/0000-0004-DC4B-1. ISSN 0960-9822. PMID 31104936.
  17. ^ a b c Kapli, Paschalia; Natsidis, Paschalis; Leite, Daniel J.; Fursman, Maximilian; Jeffrie, Nadia; Rahman, Imran A.; Philippe, Hervé; Copley, Richard R.; Telford, Maximilian J. (2020-07-02). "Lack of support for Deuterostomia prompts reinterpretation of the first Bilateria". bioRxiv 10.1101/2020.07.01.182915.
  18. ^ Vinther, J.; Smith, M. P.; Harper, D. A. T. (2011). "Vetulicolians from the Lower Cambrian Sirius Passet Lagerstätte, North Greenland, and the polarity of morphological characters in basal deuterostomes". Palaeontology. 54 (3): 711–719. doi:10.1111/j.1475-4983.2011.01034.x. S2CID 85602402.
  19. ^ Chen, Feng, Ma, Li (2003). "A New Vetulicolian from the Early Cambrian Chengjiang Fauna in Yunnan of China". Acta Geologica Sinica. 77 (3): 281–287. doi:10.1111/j.1755-6724.2003.tb00742.x. S2CID 128895860.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  20. ^ Caron, Jean-Bernard (June 2005). "Banffia constricta, a putative vetulicolid from the Middle Cambrian Burgess Shale". Transactions of the Royal Society of Edinburgh: Earth Sciences. 96 (2): 95–111. doi:10.1017/S0263593300001255. ISSN 0263-5933. S2CID 128851764.
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