Elasmobranchii
Temporal range:
Great white shark
(Carcharodon carcharias)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Bonaparte, 1838
Superorders

Elasmobranchs lack swim bladders, and maintain buoyancy with oil that they store in their livers. Some deep sea sharks are targeted by fisheries for this liver oil, including the school, gulper and basking sharks (pictured).[1] All three of these species have been assessed by the IUCN as vulnerable due to overfishing.[2][3][4]

From a practical point of view the life-history pattern of elasmobranchs makes this group of animals extremely susceptible to over fishing. It is no coincidence that the commercially exploited marine turtles and baleen whales, which have life-history patterns similar to the sharks, are also in trouble.[5]

Elasmobranchii (/ɪˌlæzməˈbræŋki/[6]) is a subclass of Chondrichthyes or cartilaginous fish, including modern sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gill clefts opening individually to the exterior, rigid dorsal fins and small placoid scales on the skin. The teeth are in several series; the upper jaw is not fused to the cranium, and the lower jaw is articulated with the upper. The details of this jaw anatomy vary between species, and help distinguish the different elasmobranch clades. The pelvic fins in males are modified to create claspers for the transfer of sperm. There is no swim bladder; instead, these fish maintain buoyancy with large livers rich in oil.

The definition of the clade is unclear with respect to fossil chondrichthyans. It has been used by different authors as equivalent to Neoselachii (the clade including modern sharks and rays and their last common ancestor) or for all chondrichthyans more closely related to modern sharks and rays than to Holocephali (the clade containing chimaeras and their extinct relatives).[7] Important groups of elasmobranchs sensu lato include the hybodonts (Order Hybodontiformes), xenacanths (order Xenacanthformes) and Ctenacanthiformes, which are also often referred to as "sharks".

The name Elasmobranchii comes from the Ancient Greek words elasmo- ("plate") and bránchia ("gill"), referring to the broad, flattened gills which are characteristic of these fishes.

Description

Elasmobranchii is one of the two subclasses of cartilaginous fish in the class Chondrichthyes, the other being Holocephali (chimaeras).

Members of the elasmobranchii subclass have no swim bladders, five to seven pairs of gill clefts opening individually to the exterior, rigid dorsal fins, and small placoid scales. The teeth are in several series; the upper jaw is not fused to the cranium, and the lower jaw is articulated with the upper.

Extant elasmobranchs exhibit several archetypal jaw suspensions: amphistyly, orbitostyly, hyostyly, and euhyostyly. In amphistyly, the palatoquadrate has a postorbital articulation with the chondrocranium from which ligaments primarily suspend it anteriorly. The hyoid articulates with the mandibular arch posteriorly, but it appears to provide little support to the upper and lower jaws. In orbitostyly, the orbital process hinges with the orbital wall and the hyoid provides the majority of suspensory support.

In contrast, hyostyly involves an ethmoid articulation between the upper jaw and the cranium, while the hyoid most likely provides vastly more jaw support compared to the anterior ligaments. Finally, in euhyostyly, also known as true hyostyly, the mandibular cartilages lack a ligamentous connection to the cranium. Instead, the hyomandibular cartilages provide the only means of jaw support, while the ceratohyal and basihyal elements articulate with the lower jaw, but are disconnected from the rest of the hyoid.[8][9][10] The eyes have a tapetum lucidum. The inner margin of each pelvic fin in the male fish is grooved to constitute a clasper for the transmission of sperm. These fish are widely distributed in tropical and temperate waters.[11]

Many fish maintain buoyancy with swim bladders. However elasmobranchs lack swim bladders, and maintain buoyancy instead with large livers that are full of oil.[12] This stored oil may also function as a nutrient when food is scarce.[5][13] Deep sea sharks are usually targeted for their oil, because the livers of these species can weigh up to 20% of their total weight.[1]

Evolution

The oldest unambigous elasmobranch sensu lato, Phoebodus, has its earliest records in the Middle Devonian (Givetian), around 387.7 to 382.7 million years ago.[14] Several important groups of elasmobranchs, including Ctenacanthiformes and Hybodontiformes, had already emerged by the latest Devonian (Famennian).[15] During the Carboniferous, some ctenacanths would grow to sizes rivalling the modern great white shark.[16] During the Carboniferous and Permian, the xenacanths were abundant in both freshwater and marine environments, and would continue to exist into the Triassic with reduced diversity.[17] The hybodonts had achieved a high diversity by the Permian,[18] and would end up becoming the dominant group of elasmobranchs during the Triassic and Early Jurassic. Hybodonts were extensively present in both marine and freshwater environments.[19] While Neoselachii (the group of modern sharks and rays) had already appeared by the Triassic, they only had low diversity during this period would and only begin to extensively diversify from the Early Jurassic onwards, when modern orders of sharks and rays appeared.[20] This co-incided with the decline of the hybodonts, which had become minor components of marine environments by the Late Jurassic, but would remain common in freshwater environments into the Cretaceous.[21] The youngest remains of hybodonts date to the very end of the Cretaceous.[22]

Taxonomy

Recent molecular studies suggest the Batoidea are not derived selachians as previously thought. Instead, skates and rays are a monophyletic superorder within Elasmobranchii that shares a common ancestor with the selachians.[25][26]

See also

References

  1. 1 2 Vannuccini, Stefania (2002) Shark liver oil products Archived 2013-06-26 at the Wayback Machine In: Shark Utilization, Marketing and Trade, Fisheries Technical paper 389, FAO, Rome. ISBN 92-5-104361-2.
  2. Fowler, S.L. (2005). "Cetorhinus maximus". IUCN Red List of Threatened Species. 2005: e.T4292A10763893. doi:10.2305/IUCN.UK.2005.RLTS.T4292A10763893.en.
  3. Walker, T.I.; Rigby, C.L.; Pacoureau, N.; Ellis, J.; Kulka, D.W.; Chiaramonte, G.E.; Herman, K. (2020). "Galeorhinus galeus". IUCN Red List of Threatened Species. 2020: e.T39352A2907336. doi:10.2305/IUCN.UK.2020-2.RLTS.T39352A2907336.en. Retrieved 11 November 2021.
  4. Finucci, B.; Bineesh, K.K.; Cheok, J.; Cotton, C.F.; Dharmadi, Kulka, D.W.; Neat, F.C.; Pacoureau, N.; Rigby, C.L.; Tanaka, S.; Walker, T.I. (2020). "Centrophorus granulosus". IUCN Red List of Threatened Species. 2020: e.T162293947A2897883. doi:10.2305/IUCN.UK.2020-3.RLTS.T162293947A2897883.en. Retrieved 11 November 2021.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  5. 1 2 Hoenig, J.M. and Gruber, S.H. (1990) "Life-history patterns in the elasmobranchs: implications for fisheries management" Archived 2013-02-18 at the Wayback Machine In: Elasmobranchs as living resources: advances in the biology, ecology, systematics and the status of the fisheries, eds. J. H. L. Pratt, S. H. Gruber and T. Taniuchi, US Department of Commerce, NOAA technical report NMFS 90, pp.1–16.
  6. "Elasmobranch". Merriam-Webster.com Dictionary.
  7. Maisey, J. G. (April 2012). "What is an 'elasmobranch'? The impact of palaeontology in understanding elasmobranch phylogeny and evolution". Journal of Fish Biology. 80 (5): 918–951. doi:10.1111/j.1095-8649.2012.03245.x. PMID 22497368.
  8. Wilga, C.D. (2005). "Morphology and evolution of the jaw suspension in lamniform sharks". Journal of Morphology. 265 (1): 102–19. doi:10.1002/jmor.10342. PMID 15880740. S2CID 45227734.
  9. Wilga, C. D.; Motta, P. J.; Sanford, C. P. (2007). "Evolution and ecology of feeding in elasmobranchs". Integrative and Comparative Biology. 47 (1): 55–69. doi:10.1093/icb/icm029. PMID 21672820.
  10. Wilga, Cheryl A.D. (2008). "Evolutionary divergence in the feeding mechanism of fishes". Acta Geologica Polonica. 58 (2): 113–20. Archived from the original on 2018-08-19. Retrieved 2017-05-24.
  11. Bigelow, Henry B.; Schroeder, William C. (1948). Fishes of the Western North Atlantic. Sears Foundation for Marine Research, Yale University. pp. 64–65. ASIN B000J0D9X6.
  12. Oguri, M (1990) "A review of selected physiological characteristics unique to elasmobranchs" Archived 2013-02-18 at the Wayback Machine In: Elasmobranchs as living resources: advances in the biology, ecology, systematics and the status of the fisheries, eds. J. H. L. Pratt, S. H. Gruber and T. Taniuchi, US Department of Commerce, NOAA technical report NMFS 90, pp.49–54.
  13. Bone, Q.; Roberts, B. L. (2009). "The density of elasmobranchs". Journal of the Marine Biological Association of the United Kingdom. 49 (4): 913. doi:10.1017/S0025315400038017. S2CID 85871565.
  14. Frey, Linda; Coates, Michael; Ginter, Michał; Hairapetian, Vachik; Rücklin, Martin; Jerjen, Iwan; Klug, Christian (2019-10-09). "The early elasmobranch Phoebodus : phylogenetic relationships, ecomorphology and a new time-scale for shark evolution". Proceedings of the Royal Society B: Biological Sciences. 286 (1912): 20191336. doi:10.1098/rspb.2019.1336. ISSN 0962-8452. PMC 6790773. PMID 31575362.
  15. Schultze, H.-P., Bullecks, J., Soar, L. K., & Hagadorn, J. (2021). Devonian fish from Colorado’s Dyer Formation and the appearance of Carboniferous faunas in the Famennian. In A. Pradel, J. S. S. Denton, & P. Janvier (Eds.), Ancient Fishes and their Living Relatives: a Tribute to John G. Maisey (pp. 247–256.). Verlag Dr. Friedrich Pfeil.
  16. Maisey, John G.; Bronson, Allison W.; Williams, Robert R.; McKinzie, Mark (2017-05-04). "A Pennsylvanian 'supershark' from Texas". Journal of Vertebrate Paleontology. 37 (3): e1325369. doi:10.1080/02724634.2017.1325369. ISSN 0272-4634.
  17. Pauliv, Victor E.; Martinelli, Agustín G.; Francischini, Heitor; Dentzien-Dias, Paula; Soares, Marina B.; Schultz, Cesar L.; Ribeiro, Ana M. (December 2017). "The first Western Gondwanan species of Triodus Jordan 1849: A new Xenacanthiformes (Chondrichthyes) from the late Paleozoic of Southern Brazil". Journal of South American Earth Sciences. 80: 482–493. doi:10.1016/j.jsames.2017.09.007.
  18. Koot, Martha B.; Cuny, Gilles; Tintori, Andrea; Twitchett, Richard J. (March 2013). "A new diverse shark fauna from the Wordian (Middle Permian) Khuff Formation in the interior Haushi‐Huqf area, Sultanate of Oman". Palaeontology. 56 (2): 303–343. doi:10.1111/j.1475-4983.2012.01199.x. ISSN 0031-0239.
  19. Rees, J. A. N., and Underwood, C. J., 2008, Hybodont sharks of the English Bathonian and Callovian (Middle Jurassic): Palaeontology, v. 51, no. 1, p. 117-147.
  20. Underwood, Charlie J. (March 2006). "Diversification of the Neoselachii (Chondrichthyes) during the Jurassic and Cretaceous". Paleobiology. 32 (2): 215–235. Bibcode:2006Pbio...32..215U. doi:10.1666/04069.1. ISSN 0094-8373. S2CID 86232401.
  21. Rees, Jan; Underwood, Charlie J. (January 2008). "HYBODONT SHARKS OF THE ENGLISH BATHONIAN AND CALLOVIAN (MIDDLE JURASSIC)". Palaeontology. 51 (1): 117–147. doi:10.1111/j.1475-4983.2007.00737.x. ISSN 0031-0239.
  22. Carrillo-Briceño, Jorge D.; Cadena, Edwin A.; Dececchi, Alex T.; Larson, Hans C. E.; Du, Trina Y. (2016-01-01). "First record of a hybodont shark (Chondrichthyes: Hybodontiformes) from the Lower Cretaceous of Colombia". Neotropical Biodiversity. 2 (1): 81–86. doi:10.1080/23766808.2016.1191749. ISSN 2376-6808.
  23. Ebert, David A.; Fowler, Sarah; Dando, Marc (2021). Sharks of the world: a complete guide. Princeton: Princeton University Press. ISBN 978-0-691-20599-1.
  24. "WoRMS - World Register of Marine Species - Echinorhiniformes". Retrieved 2022-01-04.
  25. Winchell, Christopher J; Martin, Andrew P; Mallatt, Jon (2004). "Phylogeny of elasmobranchs based on LSU and SSU ribosomal RNA genes". Molecular Phylogenetics and Evolution. 31 (1): 214–24. doi:10.1016/j.ympev.2003.07.010. PMID 15019621.
  26. Douady, Christophe J.; Dosay, Miné; Shivji, Mahmood S.; Stanhope, Michael J. (2003). "Molecular phylogenetic evidence refuting the hypothesis of Batoidea (rays and skates) as derived sharks". Molecular Phylogenetics and Evolution. 26 (2): 215–21. doi:10.1016/S1055-7903(02)00333-0. PMID 12565032.
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