Vetulicola cuneata

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Vetulicola cuneata
Temporal range: Cambrian Stage 3
Vetilicola cuneata reconstruction.png
Reconstruction of V. cuneata
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade?:† Vetulicolia
Class:† Vetulicolida
Order:† Vetulicolata
Family:† Vetulicolidae
Genus:† Vetulicola
Species:
†V. cuneata
Binomial name
†Vetulicola cuneata
Hou, 1987

Vetulicola cuneata ("wedged-shaped ancient dweller") is a species of extinct marine animal from the Early Cambrian Chengjiang biota of China. It was described by Hou Xian-guang in 1987 from the Lower Cambrian Chiungchussu Formation, [1] [2] and became the first animal (type species) under an eponymous phylum Vetulicolia. [3]

Contents

Remarked as "enigmatic creatures" [4] and originally classified as an arthropod, it is recognised as a deuterostome along with other related species, for which a separate class (Vetulicolida), order (Vetulicolata), family (Vetulicolidae) and ultimately a new clade (Vetulicolia) were created based on its name. [3] [5]

Discovery and taxonomy

In 1984, Hou Xian-guang of the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, explored the Chiungchussu Formation in Chengjiang, China. At the Maotianshan Hill, from which the sediments called Maotianshan Shales were already established to be of Cambrian in origin, [6] he found a variety of animal fossils. The first species he formally reported was that of an arthropod Naraoia , described as an "unusual trilobite". [7] The rich fossils were eventually established as the Chengjiang biota. [8]

Among Hou's earliest collections were a group of bivalved arthropods, which he described in the journal Acta Palaeontologica Sinica in 1987. [2] One group which he categorised "larger bivalved arthropods" included five distinct species, one of which he named Vetulicola cuneata, [1] for its obvious ancient nature (Latin vetuli means "old" and cola, "inhabitant") and wedge-shaped beak-like body (cuneata for wedge-shaped). [4] Although Hou classified V. cuneata as a species of the phylum Arthopoda for its prominent body covering called carapace, a common feature among arthropods, he immediately noticed some issues. The Vetulicola body was divided into two separate parts, dorsal and ventral carapace, by clear longitudinal groove, which he realised was an odd feature for an arthropod that should normally lack such demarcating groove. He also mistakenly identified a short tail-like projection at the posterior region as consisting of two segments and attached to the ventral side of the body. [9]

In 1991, with the help of Jan Bergström (of the Swedish Museum of Natural History), Hou re-analysed the species from more than 100 specimens and resolved that the posterior projection typically consists of seven segments and arose (after turning upside-down the specimens) from the dorsal surface of the body. [10] Palaeontologists continued to consider the species as an arthropod although they understood the anomaly and missing features of Arthropoda. [11] [12] [13]

In 1997, Hou's colleagues Jun-yuan Chen and Guiqing Zhou extensively reviewed and re-eaxmined the Chengjian fossils. Among the new species they identified was Banffia confusa . [14] The genus Banffia was created by American palaeontologist Charles Doolittle Walcott in 1911 for a new species B. constricta from the Burgess Shale of British Columbia, Canada. [15] B. constricta was considered as member of the phylum Annelida for its apparent segmented worm-like body. [16] When Chen and Zhou compared B. confusa with V. cuneata, they realised the close relationship between the two specimens, having shared most defining features, so that they created a new class Vetulicolida for the two genus under Arthropoda. [9] [17]

In 1999, Huilin Luo and his colleagues at the Yunnan Institute of Geological Sciences described a new species Pomatrum ventralis as a closely related arthropod to V. cuneata. [9] [18] The same year, Degan Shu and his research team at the Northwest University in Xi'an, along with Simon Conway Morris at the University of Cambridge, described a new species Xidazoon stephanus which was recognised as related to primitive jawless fish. [19] In 2001, Shu and Conway Morris realised the need to reassess all the Chengjian species related to V. cuneata. With a report of a new species Didazoon haoae, they revised the entire classification adopting the class Vetulicolida and introduced the phylum Vetulicolia for all species of Vetulicola, Banffia, Pomatrum,Xidazoon and Didazoon. The new phylum was classified as member of the "primitive deuterostomes," a group that consists of animals ancestral to chordates. [5] [20]

Description

Fossil of Vetulicola cuneata Vetulicola cuneata uetsurikora kunieta.jpg
Fossil of Vetulicola cuneata

Vetulicola cuneata has a body composed of two distinct parts of approximately equal length. [1] The anterior part is rectangular in shape, and enclosed by a carapace-like structure of cuticular plates fused together, with a large, V-shaped mouth at the front end: there is a keel-like extension of the body wall on the top and belly. [5] It was earlier described as having four plates, [5] but closer examination showed that the apparent lateral groove that appeared to create two plates was not cutting through the entire body. The lateral plates are in fact joined at the posterior region, thus, only two plates are present. [9] The tail-like posterior section is slender, strongly cuticularised and placed dorsally. [1] It is made up of seven segments. [9] Paired openings connecting the pharynx to the outside run down the sides. These features are interpreted as possible primitive gill slits. [21] The entire body could be up to 7 cm long. [22]

Lifestyle

It is assumed that V. cuneata spent most or all of its time swimming in the water column. Sediment found within the gut suggest that it was a deposit-feeder, possibly swimming to and from favorable feeding sites. Some specimens that had individuals of the putative entoproct, Cotyledion tyloides attached to the terminal tail segment indicate that V. cuneata lay buried in the sediment, with only its terminal segment exposed, but, nothing about its anatomy, or the taphonomy of its fossils, suggest that it was a burrower. Rather, it was more likely that larval C. tyloides would occasionally settle on the terminal segment, then take advantage of serendipity to feed whenever their host discharged nutrient-rich fecal matter. [23]

V. cuneata and V. rectangulata have symbiotic association with a worm called Vermilituus gregarius. The worms are attached on the internal surface of the plates as a kind of endosymbionts. The nature of the symbiosis is not clear, as it could be either useful (commensal) or harmful (parasitic). [24]

Related Research Articles

<span class="mw-page-title-main">Vetulicolia</span> Extinct Cambrian group of animals

Vetulicolia is a group of bilaterian marine animals encompassing several extinct species from the Cambrian, and possibly Ediacaran, periods. As of 2023, the majority of workers favor placing Vetulicolians in the stem group of the Chordata, but some continue to favor a more crownward placement as a sister group to the Tunicata. It was initially erected as a monophyletic clade with the rank of phylum in 2001, with subsequent work supporting its monophyly. However, more recent research suggests that vetulicolians may be paraphyletic and form a basal evolutionary grade of stem chordates.

<span class="mw-page-title-main">Maotianshan Shales</span> Series of Early Cambrian deposits in the Chiungchussu Formation in China

The Maotianshan Shales (帽天山页岩) are a series of Early Cambrian sedimentary deposits in the Chiungchussu Formation, famous for their Konservat Lagerstätten, deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales form one of some forty Cambrian fossil locations worldwide exhibiting exquisite preservation of rarely preserved, non-mineralized soft tissue, comparable to the fossils of the Burgess Shale of British Columbia, Canada. They take their name from Maotianshan Hill in Chengjiang County, Yunnan Province, China.

<i>Yunnanozoon</i> Cambrian fossil chordate

Yunnanozoon lividum is an extinct species of bilaterian animal from the Lower Cambrian Chengjiang biota of Yunnan province, China. Its affinities have been long the subject of controversy.

<i>Vetulicola</i> Fossil genus of marine animal

Vetulicola is an extinct genus of marine animal discovered from the Cambrian of China. It is the eponymous member of the enigmatic taxon Vetulicolia, which is of uncertain affinities but may belong to the deuterostomes. 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.

<i>Pomatrum</i> Cambrian age animal

Pomatrum is an extinct vetulicolian, the senior synonym of Xidazoon; the latter taxon was described by Shu, et al. (1999) based on fossils found in the Qiongzhusi (Chiungchussu) Formation, Yu'anshan Member, Lower Cambrian, Haikou, (Kunming), about 50 km west of Chengjiang, China.

<span class="mw-page-title-main">Vetulicolidae</span> Extinct Cambrian family of vetulicolian animals

Vetulicolidae is a vetulicolian family from the Cambrian Stage 3 Maotianshan Shale and Sirius Passet Lagerstätte that consists of Vetulicola, Beidazoon, and Ooedigera. It is distinguished from the Didazoonidae by a harder body wall and the lack of an oral disc.

<span class="mw-page-title-main">Didazoonidae</span> Extinct Cambrian family of vetulicolid animals

Didazoonidae is a vetulicolian family within the order Vetulicolata. It is characterized by a relatively thin-walled, non-biomineralized body and a large, round anterior opening surrounded by an oral disc. It may be paraphyletic, even if the phylum Vetulicolia is monophyletic.

<span class="mw-page-title-main">Banffozoa</span> Extinct Cambrian group of marine animals

Banffozoa is an extinct class of bilaterians. Most workers place it in the Vetulicolia, but the protostome-like features of some members have motivated ongoing debate. Banffozoa consists of the order Banffiata as well as a dwarf "Form A" that has not been formally described or named. Skeemella has been placed incertae sedis in this class, but has more recently been placed with the Banffiidae. Banffozoa may be paraphyletic even if Vetulicolia is monophyletic.

<i>Chuandianella</i> Extinct genus of Cambrian animals

Chuandianella ovata is an extinct bivalved arthropod that lived during Cambrian Stage 3 of the Early Cambrian. It is the only species classified under the genus Chuandianella. Its fossils were recovered from the Chengjiang Biota in Yunnan, China.

<i>Miraluolishania</i> Extinct genus of lobopodians

Miraluolishania is an extinct lobopodian known from Chengjiang County in China. It is remarkable for the possession of lensed pit-eyes. The only species, Miraluolishania haikouensis, was described from the Maotianshan Shales at Haikou by Jianni Liu and Degan Shu in 2004. In 2009, a team of palaeontologists at the Yunnan University, led by Xiaoya Ma reported the discovery of 42 other specimens from Haikou. With the help of Swiss palaeontologist Jan Bergström, Ma and Hou came to the conclusion that all the specimens were the same species as Luolishania; another lobopod discovered from the Chengjian in 1989. Chengjian is 40 kms from Haikou and the fossil fauna are different. A reassessment by Liu and Shu's team at the Northwest University in 2008 established that Luolishania and Miraluolishania are distinct animals.

<i>Phlogites</i> Genus of ambulacrarians

Phlogites is a member of the extinct ambulacrarian stem group Cambroernida, occupying an intermediate position between the basal Herpetogaster and the more derived Eldonioidea. It is known from the Lower Cambrian Haikou Chengjiang deposits of China.

<i>Vetulicola rectangulata</i> Extinct animal from Cambrian of the Chengjiang biota of China

Vetulicola rectangulata is a species of extinct animal from the Early Cambrian of the Chengjiang biota of China. Regarded as a deuterostome, it has characteristic rectangular anterior body on which the posterior tail region is attached. It was described by Luo Huilin and Hu Shi-xue in 1999.

<i>Beidazoon</i> Extinct species of Cambrian organism

Beidazoon venustum is a marine deuterostome from the group Vetulicolia. It originates from the lower Cambrian Chengjiang biota of Yunnan Province, China, and was discovered in 2005. It is known as the smallest described vetulicolian, and for its surface being covered in many small nodes.

<span class="mw-page-title-main">Banffiidae</span> Extinct Cambrian family of animals

Banfiidae is a family of extinct banffozoan animals from North America and China. The family name is sometimes spelt Banffidae. It includes Banffia, Heteromorphus, and possibly Skeemella. The family may be paraphyletic. The family may be paraphyletic. A Heteromorphus-like dwarf "Form A" is allied with this group at the class level, but has not been formally described or assigned to this family.

<i>Vetulocystis</i> Genus of vetulocystids

Vetulocystis is an extinct genus type genus of the family Vetulocystidae. It was found in the Maotianshan Shales, which date to roughly 518 mya.

Luolishania is an extinct genus of lobopodian panarthropod and known from the Lower Cambrian Chiungchussu Formation of the Chengjiang County, Yunnan Province, China. A monotypic genus, it contains one species Luolishania longicruris. It was discovered and described by Hou Xian-Guang and Chen Jun-Yuan in 1989. It is one of the superarmoured Cambrian lobopodians suspected to be either an intermediate form in the origin of velvet worms (Onychophora) or basal to at least Tardigrada and Arthropoda. It is the basis of the family name Luolishaniidae, which also include other related lobopods such as Acinocricus, Collinsium, Facivermis, and Ovatiovermis. Along with Microdictyon, it is the first lobopodian fossil discovered from China.

<i>Lenisambulatrix</i> Extinct genus of Lobopodian

Lenisambulatrix is a genus of extinct worm belonging to the group Lobopodia and known from the Lower Cambrian Maotianshan shale of China. It is represented by a single species L. humboldti. The incomplete fossil was discovered and described by Qiang Ou and Georg Mayer in 2018. Due to its missing parts, its relationship with other lobopodians is not clear. It shares many structural features with another Cambrian lobopodian Diania cactiformis, a fossil of which was found alongside it.

Hou Xian-guang is a Chinese paleontologist at Yunnan University who made key discoveries in the Cambrian life of China around 518 myr. His first discovery of animal fossils from the Cambrian sediments at Chengjiang County, Yunnan Province, led to the establishment of the Chengjiang biota, an assemblage of various life forms during the Cambrian Period. The discovery of the Chengjiang biota, remarked as "among the most spectacular in this [20th] century", added to the better understanding of how animal forms originated and evolved during the so-called Cambrian explosion.

Cheungkongella is a fossil organism from the lower Cambrian Chengjiang lagerstatte, the affinity of which has been the subject of debate. It was announced as a "probable" tunicate while noting the lack of definitive Cambrian fossils from that group. However, this affinity was later disputed in a paper announcing the discovery of Shankouclava, also from Chengjiang, as the oldest known tunicate. Cheungkongella has been accepted as a distinct taxon and possible tunicate by multiple workers not involved in its discovery, but the dispute remains unresolved.

<i>Heteromorphus</i> Extinct genus of Cambrian organisms

Heteromorphus is an extinct genus of banffiid from the lower Cambrian Chengjiang lagerstatte. It contains one broadly accepted species, Heteromorphus confusus, as well as a proposed junior synonym, Heteromorphus longicaudatus that may prove to be a separate species as additional specimens are examined. A much smaller species labeled "Form A" is allied with Heteromorphus at the class level but has not been formally described or assigned to Heteromorphus itself.

References

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  2. 1 2 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.
  3. 1 2 Shu, Degan (2005). "On the Phylum Vetulicolia". Chinese Science Bulletin. 50 (20): 2342–2354. doi:10.1007/BF03183746. ISSN   1001-6538. S2CID   86827605.
  4. 1 2 McMenamin, Mark A. S. (2019). "Cambrian Chordates and Vetulicolians". Geosciences. 9 (8): 354. Bibcode:2019Geosc...9..354M. doi: 10.3390/geosciences9080354 . ISSN   2076-3263.
  5. 1 2 3 4 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.
  6. ↑ Hou, Xianguang; Bergström, Jan (2003-03-31). "The Chengjiang fauna — the oldest preserved animal community". Paleontological Research. 7 (1): 55–70. doi: 10.2517/prpsj.7.55 . ISSN   1342-8144.
  7. ↑ Zhang, T.W.; Hou, X.-G. (1985). "Preliminary notes on the occurrence of the unusual trilobite Naraoia in Asia". Acta Palaeontologica Sinica. 24 (6): 591–595.
  8. ↑ Briggs, Derek E. G. (2015). "Extraordinary fossils reveal the nature of Cambrian life: a commentary on Whittington (1975) 'The enigmatic animal Opabinia regalis , Middle Cambrian, Burgess Shale, British Columbia'". Philosophical Transactions of the Royal Society B: Biological Sciences. 370 (1666): 20140313. doi:10.1098/rstb.2014.0313. PMC   4360120 . PMID   25750235.
  9. 1 2 3 4 5 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.
  10. ↑ Hou, Xian-Guang; Bergström, Jan (2009). "The arthropods of the Lower Cambrian Chengjiang fauna, with relationships and evolutionary significance". In Simonetta, Alberto M.; Conway Morris, Simon (eds.). The Early Evolution of Metazoa and the Significance of Problematic Taxa. Cambridge University Press. p. 181. ISBN   978-0-521-11158-4.
  11. ↑ XianGuang, Hou; Ramsköld, Lars; Bergström, Jan (1991). "Composition and preservation of the Chengjiang fauna -a Lower Cambrian soft-bodied biota". Zoologica Scripta. 20 (4): 395–411. doi:10.1111/j.1463-6409.1991.tb00303.x. ISSN   0300-3256. S2CID   85077111.
  12. ↑ Briggs, Derek E. G.; Fortey, Richard A. (1992), Lipps, Jere H.; Signor, Philip W. (eds.), "The Early Cambrian Radiation of Arthropods", Origin and Early Evolution of the Metazoa, vol. 10, Boston, MA: Springer US, pp. 335–373, doi:10.1007/978-1-4899-2427-8_10, ISBN   978-1-4899-2429-2 , retrieved 2023-03-18
  13. ↑ Zhi-Wen, Jiang (1992), Lipps, Jere H.; Signor, Philip W. (eds.), "The Lower Cambrian Fossil Record of China", Origin and Early Evolution of the Metazoa, Topics in Geobiology, vol. 10, Boston, MA: Springer US, pp. 311–333, doi:10.1007/978-1-4899-2427-8_9, ISBN   978-1-4899-2429-2 , retrieved 2023-03-18
  14. ↑ Chen, Junyuan; Zhou, Guiqing (1997). "Biology of the Chengjiang Fauna". Collection and Research. 10: 11–105. ISSN   1726-2038.
  15. ↑ Walcott, Charles D. (1911). "Cambrian Geology and Paleontology II: No.5--Middle Cambrian Annelids" (PDF). Smithsonian Miscellaneous Collections. 57 (5): 109–144.
  16. ↑ Conway Morris, S. (1989-10-20). "Burgess Shale Faunas and the Cambrian Explosion". Science. 246 (4928): 339–346. doi:10.1126/science.246.4928.339. ISSN   0036-8075. PMID   17747916. S2CID   10491968.
  17. ↑ Chen, Junyuan; Zhou, Guiqing (1997). "Biology of the Chengjiang Fauna". In Iten, H.V. (ed.). The Cambrian Explosion and the Fossil Record. National Museum of Natural Science. pp. 11, 95.
  18. ↑ Luo, H.; Hu, Shi-xue; Chen, Liangzhong; Zhang, Shishan; Tao, Yong-shan (1999). Early Cambrian Chengjiang fauna from Kunming region, China. Yunnan Science and Technology Press. pp. 16, 129. S2CID   133363305.
  19. ↑ Shu, D.; Conway Morris, S.; Zhang, X-L.; Chen, L.; Li, Y.; Han, J. (1999). "A pipiscid-like fossil from the Lower Cambrian of south China". Nature. 400 (6746): 746–749. doi:10.1038/23445. ISSN   0028-0836. S2CID   204995356.
  20. ↑ 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.
  21. ↑ Ou, Qiang; Conway Morris, Simon; Han, Jian; Zhang, Zhifei; Liu, Jianni; Chen, Ailin; Zhang, Xingliang; Shu, Degan (2012). "Evidence for gill slits and a pharynx in Cambrian vetulicolians: implications for the early evolution of deuterostomes". BMC Biology. 10 (1): 81. doi: 10.1186/1741-7007-10-81 . PMC   3517509 . PMID   23031545.
  22. ↑ Li, Yu-Jing; Cong, Pei-Yun; Zhao, Jun; Hou, Xian-Guang (2015). "New observations on morphological variation of genus Vetulicola with quadrate carapace from the Cambrian Chengjiang and Guanshan biotas, South China". Palaeoworld. Geologic and biotic events and their relationships during the Early to Middle Paleozoic. 24 (1): 36–45. doi:10.1016/j.palwor.2014.12.006. ISSN   1871-174X.
  23. ↑ Zhang, Zhifei; et al. (January 2013). "A sclerite-bearing stem group entoproct from the early Cambrian and its implications". Scientific Reports. 3: 1066. Bibcode:2013NatSR...3E1066Z. doi:10.1038/srep01066. PMC   3548229 . PMID   23336066.
  24. ↑ Li, Yujing; Williams, Mark; Harvey, Thomas H. P.; Wei, Fan; Zhao, Yang; Guo, Jin; Gabbott, Sarah; Fletcher, Tom; Hou, Xianguang; Cong, Peiyun (2020). "Symbiotic fouling of Vetulicola, an early Cambrian nektonic animal". Communications Biology. 3 (1): 517. doi:10.1038/s42003-020-01244-1. PMC   7501249 . PMID   32948820.
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