Biology:Balhuticaris

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Balhuticaris is a genus of extinct bivalved (referring to the carapace) hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. This extremely multisegmented (with over 100 segments) arthropod is the largest member of the group, and it was even one of the largest animals of the Cambrian, with individuals reaching lengths of 245 mm (9 in). Fossils of this animal suggests that gigantism occurred in more groups of Arthropoda than had been previously thought.[1] It also presents the possibility that bivalved arthropods were very diverse, and filled in a lot of ecological niches.[1][2][3]

The hymenocarines were an order of primitive mandibulates, the arthropod group that includes crustaceans, insects, myriapods and their relatives, that lived throughout the Cambrian period.[2] This group was extremely diverse and attained a wide variety of ecological niches and body plans.[2][3] Several dozen species are known from deposits of Cambrian, and ranged in size from smaller species like Fibulacaris nereidis reaching a length of 2 cm (0.79 in) long,[2] to larger ones like B. voltae.[1]

Discovery and Etymology

This arthropod was described in 2022 based on 11 specimens found in the Burgess Shale between 2014 and 2018, more specifically in the Marble canyon locality. By 2020, scientists realized that these fossils represented a new species. Because of how they were preserved the fossils were found two dimensional in several carbonaceous films. The holotype specimen and several others are nearly or fully complete with possible neural and other soft tissues having been preserved. Balhuticaris is named after Balhūt, a giant fish from Persian cosmography, as well as the Latin caris ("crab"). The specific epithet voltae is derived from the Catalan volta, meaning vault, referring to the shape of the carapace when seen from the front.[1]

Description

Balhuticaris was the largest bivalved arthropod in the fossil record, beating the previous holders of this title Nereocaris exilis and Tuzoia. This animal's body was very long, and had extreme segmentation compared to other Cambrian arthropods, with over 100 distinct segments. In total this creature had about 110 pairs of biramous limbs, the most of any Cambrian-aged arthropod. Covering the head of this creature was a large carapace that resembles an arch or other curved structure. This structure only covers the frontmost part of this arthropod but it does extend ventrally beyond its appendages.[1]

Classification

In several studies performed, Balhuticaris was found to be a member of the Hymenocarina. More specifically it was found to be most closely related to the genus Odaraia and its relatives. Although they are normally regarded as pancrustaceans, this study found the Hymenocarina to occupy a more basal branch of the mandibulates.[1] This cladogram shows the position of B. voltae in relation to other arthropods by López et al., 2022.[1]

Artiopoda

Eoredlichia

 
Bradoriids
 

Kunmingella

 
 

Kunyangella

 
 
Hymenocarina
 
 
 
 
 

Perspicaris recondita

 
 

Perspicaris dictynna

 
 
 

Canadaspis

 
 
 
 

Branchiocaris

 
 

Tokummia

 
 
 
 
 
 
 
 
 
 
 
 
 

Nereocaris exilis

 
 
 

Fibulacaris

 
 

Pakucaris

 
 
 
 

Nereocaris briggsi

 
 
 

Balhuticaris

 
 
 
 
 

Odaraia

 
 

Jugatacaris

 
 
Fuxianhuiids
 

Fuxianhuia

 
 

Chengjiangocaris

 
 
 
 
 

Erjiecaris

 
 
 

Loricicaris

 
 
 

Plenocaris

 
 
 
 

Clypecaris

 
 

Ercaicunia

 
 
 
 
 

Chuandianella

 
 

Waptia

 
 
 
 
 
 

Pectocaris

 
 

Pauloterminus

 
 
 

Cladogram of Hymenocarina, following Izquierdo-López and Caron, (2024), which recovered Balhuticaris within Odaraiidae:[4]

 
Hymenocarina
 
 
 

Tuzoia 70px

 
 

Perspicaris70px

 
 
 
 

Pectocaris 70px

 
Protocarididae
 

Loricicaris70px

 
 
 

Tokummia 70px

 
 

Branchiocaris 70px

 
 
 
 
 
 
 
 
 

Plenocaris 70px

 
 
 

Ercaicunia 70px

 
 

Clypecaris70px

 
 
 
 
 

Pauloterminus 70px

 
 
 

Canadaspis 70px

 
 
 

Waptia 70px

 
 

Chuandianella 70px

 
 
 
 
 
 
 

Vermontcaris

 
Odaraiidae
 

Odaraia 70px

 
 
 

Jugatacaris 70px

 
 
 
 

Fibulacaris 70px

 
 

Pakucaris 70px

 
 
 
 

Balhuticaris 70px

 
 

Nereocaris 70px

 
 
 
 
 
 
 
 
 

Lifestyle

This hymenocarine most likely engaged in a fast-paced nektonic (free swimming) lifestyle. Its large size means that it was safe from most of the other predatory fauna of its environment. Many features of the fossils evidence a free swimming, pelagic lifestyle. Examples being the presence of a tripartite caudal ramus, a feature only found in hymenocarines,[1][2] and that the carapace goes ventrally beyond the legs, which would have heavily impaired this arthropods ability to crawl on the ocean floor. Its eyes also have a similar shape seen in modern pelagic crustaceans. What this arthropod ate has been a difficult question to answer due to the lack of cephalic appendages in the fossils. Modern day arthropods of a similar size like lobsters, stomatopods, and giant isopods are mainly scavengers or predators. B. voltae however does not possess features that would suggest this, like chelate limbs and gnathobases.[1][5] Suspension and deposit feeding can also be readily ruled out due to lack of features needed for these lifestyles in B. voltae. Currently it is thought to have suctioned in prey in water currents through a ventral groove. The animal probably swam while it fed, similar to leptostracan and anostracan crustaceans.[1][6][7] It is possible that this animal swam in an upside down, or in an inverted position.[1] This is not unheard of, as many other free-swimming arthropods like anostracans,[7] pelagic trilobites like the Telephinids,[8] xiphosurans,[1] and other odaraiid hymenocarines, like Odaraia and Fibulacaris swam in inverted positions.[1][2]

Paleoecology

The Burgess Shale is a middle Cambrian aged Lagerstätte that lies in British Columbia in Canada.[9] This site was the first of its kind to have been discovered and provided great insights into the soft bodied fauna of the early Paleozoic.[10] Dozens of creatures have been preserved at this site including lobopodians, stem-group and total-group Arthropoda, worms, primitive chordates, echinoderms, sponges, as well as other animal groups.[11][12] This animal was one of the largest of its time, with only the giant radiodonts like Anomalocaris surpassing it in size.[1] More specifically, this animal was found in Marble Canyon. This Lagerstätte produced new taxa including other hymenocarines like Tokummia, Fibulacaris and Pakucaris, large hurdiid radiodonts like Cambroraster and Titanokorys, as well as other arthropods like the megacheiran genus Yawunik, the isoxyid genus Surusicaris, and the basal chelicerate Mollisonia plenovenatrix. Worms have also been found at this site, like the annelid Kootenayscolex. Well-preserved specimens of primitive chordate Metaspriggina are also known from there.[1][13][14][15][16][17][18][19][20][21]

See also

  • Paleobiota of the Burgess Shale

References

  1. ↑ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 1.10 1.11 1.12 1.13 1.14 Izquierdo-López, Alejandro (July 15, 2022). "Extreme multisegmentation in a giant bivalved arthropod from the Cambrian Burgess Shale". iScience 25 (7). doi:10.1016/j.isci.2022.104675. PMID 35845166. Bibcode: 2022iSci...25j4675I. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 Izquierdo-López, Alejandro; Caron, Jean-Bernard (2019). "A possible case of inverted lifestyle in a new bivalved arthropod from the Burgess Shale". Royal Society Open Science 6 (11). doi:10.1098/rsos.191350. PMID 31827867. Bibcode: 2019RSOS....691350I. 
  3. ↑ 3.0 3.1 O'Flynn, Robert; Williams, Mark; Yu, Mengxiao; Harvey, Thomas; Liu, Yu (2022-02-11). "A new euarthropod with large frontal appendages from the early Cambrian Chengjiang biota". Palaeontologia Electronica 25: a6. doi:10.26879/1167. https://www.researchgate.net/publication/358519106. 
  4. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (August 2024). "The Cambrian Odaraia alata and the colonization of nektonic suspension-feeding niches by early mandibulates" (in en). Proceedings of the Royal Society B: Biological Sciences 291 (2027). doi:10.1098/rspb.2024.0622. ISSN 1471-2954. PMID 39043240. 
  5. ↑ Briones-Fourzán, Patricia; Lozano-Alvarez, Enrique (1 July 1991). "Aspects of the biology of the giant isopod Bathynomus giganteus A. Milne Edwards, 1879 (Flabellifera: Cirolanidae), off the Yucatan Peninsula". Journal of Crustacean Biology 11 (3): 375–385. doi:10.2307/1548464. Bibcode: 1991JCBio..11..375B. 
  6. ↑ J. K. Lowry (October 2, 1999). "Leptostraca". Crustacea, the Higher Taxa: Description, Identification, and Information Retrieval. Australian Museum. http://www.crustacea.net/crustace/www/leptostr.htm. 
  7. ↑ 7.0 7.1 Denton Belk (2007). "Branchiopoda". The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon (4th ed.). University of California Press. pp. 414–417. ISBN 978-0-520-23939-5. https://books.google.com/books?id=64jgZ1CfmB8C&pg=PA416. 
  8. ↑ McCormick, T.; Fortey, R.A. (1998). "Independent testing of a paleobiological hypothesis: the optical design of two Ordovician pelagic trilobites reveals their relative paleobathymetry". Paleobiology 24 (2): 235–253. doi:10.1666/0094-8373(1998)024[0235:ITOAPH2.3.CO;2]. 
  9. ↑ Butterfield, N. J. (2003-02-01). "Exceptional Fossil Preservation and the Cambrian Explosion". Integrative and Comparative Biology 43 (1): 166–177. doi:10.1093/icb/43.1.166. ISSN 1540-7063. PMID 21680421. https://academic.oup.com/icb/article/43/1/166/604533. 
  10. ↑ Butterfield, N.J. (2006). "Hooking some stem-group" worms": fossil lophotrochozoans in the Burgess Shale". BioEssays 28 (12): 1161–6. doi:10.1002/bies.20507. PMID 17120226. 
  11. ↑ Conway Morris, S. (1986). "Community structure of the Middle Cambrian Phyllopod Bed (Burgess Shale)". Palaeontology 29 (3): 423–467. http://palaeontology.palass-pubs.org/pdf/Vol%2029/Pages%20423-467.pdf. Retrieved 28 April 2009. 
  12. ↑ "The largest Cambrian animal, Anomalocaris, Burgess Shale, British Columbia". Philosophical Transactions of the Royal Society B 309 (1141): 569–609. 1985. doi:10.1098/rstb.1985.0096. Bibcode: 1985RSPTB.309..569W. 
  13. ↑ Aria, Cédric; Caron, Jean-Bernard (2017). "Burgess Shale fossils illustrate the origin of the mandibulate body plan" (in en). Nature 545 (7652): 89–92. doi:10.1038/nature22080. ISSN 1476-4687. PMID 28445464. Bibcode: 2017Natur.545...89A. https://www.nature.com/articles/nature22080. 
  14. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (2019-11-13). "A possible case of inverted lifestyle in a new bivalved arthropod from the Burgess Shale". Royal Society Open Science 6 (11). doi:10.1098/rsos.191350. ISSN 2054-5703. PMID 31827867. Bibcode: 2019RSOS....691350I. 
  15. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (2021). Zhang, Xi-Guang. ed. "A Burgess Shale mandibulate arthropod with a pygidium: a case of convergent evolution" (in en). Papers in Palaeontology 7 (4): 1877–1894. doi:10.1002/spp2.1366. ISSN 2056-2799. Bibcode: 2021PPal....7.1877I. https://onlinelibrary.wiley.com/doi/10.1002/spp2.1366. 
  16. ↑ Caron, J.-B.; Moysiuk, J. (2021). "A giant nektobenthic radiodont from the Burgess Shale and the significance of hurdiid carapace diversity". Royal Society Open Science 8 (9). doi:10.1098/rsos.210664. ISSN 2054-5703. PMID 34527273. Bibcode: 2021RSOS....810664C. 
  17. ↑ Aria, Cédric; Caron, Jean-Bernard (2015-06-03). "Cephalic and Limb Anatomy of a New Isoxyid from the Burgess Shale and the Role of "Stem Bivalved Arthropods" in the Disparity of the Frontalmost Appendage". PLOS ONE 10 (6). doi:10.1371/journal.pone.0124979. ISSN 1932-6203. PMID 26038846. Bibcode: 2015PLoSO..1024979A. 
  18. ↑ Aria, Cédric; Caron, Jean-Bernard; Gaines, Robert (2015). Zhang, Xi-Guang. ed. "A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny" (in en). Palaeontology 58 (4): 629–660. doi:10.1111/pala.12161. Bibcode: 2015Palgy..58..629A. 
  19. ↑ Aria, Cédric; Caron, Jean-Bernard (2019). "A middle Cambrian arthropod with chelicerae and proto-book gills" (in en). Nature 573 (7775): 586–589. doi:10.1038/s41586-019-1525-4. ISSN 1476-4687. PMID 31511691. Bibcode: 2019Natur.573..586A. https://www.nature.com/articles/s41586-019-1525-4. 
  20. ↑ Nanglu, Karma; Caron, Jean-Bernard (2018-01-22). "A New Burgess Shale Polychaete and the Origin of the Annelid Head Revisited" (in English). Current Biology 28 (2): 319–326.e1. doi:10.1016/j.cub.2017.12.019. ISSN 0960-9822. PMID 29374441. Bibcode: 2018CBio...28E.319N. 
  21. ↑ Conway Morris, Simon; Caron, Jean-Bernard (2014-08-01). "A primitive fish from the Cambrian of North America". Nature 512 (7515): 419–422. doi:10.1038/nature13414. ISSN 0028-0836. PMID 24919146. Bibcode: 2014Natur.512..419M. https://ui.adsabs.harvard.edu/abs/2014Natur.512..419M. 

Wikidata ☰ Q113155693 entry Template:Hymenocarina



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