Elasmobranchii

Elasmobranchii (/ɪˌlæzməˈbræŋkiaɪ/[8]) is a subclass of Chondrichthyes or cartilaginous fish, including the sharks (superorder Selachii) and the 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 earliest elasmobranch fossils came from the Devonian and many surviving orders date back to the Cretaceous, or even earlier. Many species became extinct during the Permian and there was a burst of adaptive radiation during the Jurassic.

Elasmobranchii
Temporal range: Wenlock–Recent[1]
White shark
Great white shark
(Carcharodon carcharias)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Bonaparte, 1838
Superorders
Cetorhinus maximus by greg skomal
Evolution of cartilaginous fishes
Radiation of elasmobranchs, based on Michael Benton, 2005.[6]

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.[9][10][11] 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.[12]

Many fish maintain buoyancy with swim bladders. However elasmobranchs lack swim bladders, and maintain buoyancy instead with large livers that are full of oil.[13] This stored oil may also function as a nutrient when food is scarce.[7][14] Deep sea sharks are usually targeted for their oil, because the livers of these species can weigh up to 20% of their total weight.[2]

Evolution

Fossilised shark teeth are known from the early Devonian, around 400 million years ago. During the following Carboniferous period, the sharks underwent a period of diversification, with many new forms evolving. Many of these became extinct during the Permian, but the remaining sharks underwent a second burst of adaptive radiation during the Jurassic, around which time the skates and rays first appeared. Many surviving orders of elasmobranch date back to the Cretaceous, or earlier.[15]

Habitats

Elasmobranchs are mostly a marine taxon, but we know several species that live in freshwater environment (approximately 60 species which represent only 5% of the 1154 described species). They can be divided into two groups.

The euryhaline elasmobranchs, which are marine species that may survive and reproduce in freshwater environment, and the obligated freshwater elasmobranchs. The second group contains elasmobranchs that only lives in freshwater environment their entire life. This group contains only one clade: the subfamily Potamotrygoninae. This clade is endemic to one specific region (which means that they can only be seen in those regions): tropical, subtropical water and wetland of South America.

Recent researches in Paraná river[16] have shown that obligated freshwater elasmobranchs were more susceptible to anthropogenic threats as overfishing and destruction of habitats due to the very small place they live in compared to the marine species.

Taxonomy

Compagno's 2005 Sharks of the World arranges the class as follows:

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.[17][18]

See also

References

  1. ^ Märss, Tiiu; Gagnier, Pierre-Yves (2001). "A new chondrichthyan from the Wenlock, Lower Silurian, of Baillie-Hamilton Island, the Canadian Arctic". Journal of Vertebrate Paleontology. 21 (4): 693–701. doi:10.1671/0272-4634(2001)021[0693:ANCFTW]2.0.CO;2.
  2. ^ a b Vannuccini, Stefania (2002) Shark liver oil products In: Shark Utilization, Marketing and Trade, Fisheries Technical paper 389, FAO, Rome. ISBN 92-5-104361-2.
  3. ^ Fowler, S.L. (2005). "Cetorhinus maximus". The IUCN Red List of Threatened Species. 2005: e.T4292A10763893. doi:10.2305/IUCN.UK.2005.RLTS.T4292A10763893.en. Retrieved 24 December 2017.
  4. ^ "Galeorhinus galeus (School shark)". Iucnredlist.org. 2005-06-17. Retrieved 2013-03-26.
  5. ^ Guallart; et al. (2006). "Centrophorus granulosus". IUCN Red List of Threatened Species. Version 2006. International Union for Conservation of Nature. Retrieved 11 May 2006.
  6. ^ Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Fig 7.13 on page 185.
  7. ^ a b Hoenig, J.M. and Gruber, S.H. (1990) "Life-history patterns in the elasmobranchs: implications for fisheries management" 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.
  8. ^ "Elasmobranchii". Merriam-Webster Dictionary.
  9. ^ 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.
  10. ^ 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.
  11. ^ Wilga, Cheryl A.D. (2008). "Evolutionary divergence in the feeding mechanism of fishes". Acta Geologica Polonica. 58 (2): 113–20.
  12. ^ 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.
  13. ^ Oguri, M (1990) "A review of selected physiological characteristics unique to elasmobranchs" 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.
  14. ^ 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.
  15. ^ Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. p. 26. ISBN 978-1-84028-152-1.
  16. ^ Lucifora, Luis O.; Balboni, Leandro; Scarabotti, Pablo A.; Alonso, Francisco A.; Sabadin, David E.; Solari, Agustín; Vargas, Facundo; Barbini, Santiago A.; Mabragaña, Ezequiel; Díaz De Astarloa, Juan M. (2017). "Decline or stability of obligate freshwater elasmobranchs following high fishing pressure". Biological Conservation. 210: 293–298. doi:10.1016/j.biocon.2017.04.028.
  17. ^ 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.
  18. ^ 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.

External links

Asteracanthus

Asteracanthus is an extinct genus of Elasmobranchii (sharks belonging to the family Hybodontidae).

Australian ghostshark

The Australian ghostshark (Callorhinchus milii) is a cartilaginous fish (Chondrichthyes) belonging to the subclass Holocephali (chimaera). Sharks, rays and skates are the other members of the cartilaginous fish group and are grouped under the subclass Elasmobranchii. Alternative names include elephant shark, makorepe (in Māori), whitefish, plough-nose chimaera, or elephant fish. It is found off southern Australia, including Tasmania, and south of East Cape and Kaipara Harbour in New Zealand, at depths of 0–200 m (0–656 ft).

Batoidea

Batoidea is a superorder of cartilaginous fish commonly known as rays. They and their close relatives, the sharks, comprise the subclass Elasmobranchii. Rays are the largest group of cartilaginous fishes, with well over 600 species in 26 families. Rays are distinguished by their flattened bodies, enlarged pectoral fins that are fused to the head, and gill slits that are placed on their ventral surfaces.

Chondrichthyes

Chondrichthyes (; from Greek χονδρ- chondr- 'cartilage', ἰχθύς ichthys 'fish') is a class that contains the cartilaginous fishes: they are jawed vertebrates with paired fins, paired nares, scales, a heart with its chambers in series, and skeletons made of cartilage rather than bone. The class is divided into two subclasses: Elasmobranchii (sharks, rays, skates, and sawfish) and Holocephali (chimaeras, sometimes called ghost sharks, which are sometimes separated into their own class).

Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.

Chondrostei

Chondrostei are primarily cartilaginous fish showing some degree of ossification. It is thought that the cartilaginous condition is derived, and that the ancestors of this group were bony fish with fully ossified skeletons. Members of this group share with the Elasmobranchii certain features such as the possession of spiracles, a heterocercal tail and the absence of scales. Nevertheless, the fossil record suggests they have more in common with the teleosts. The Chondrostei is probably a paraphyletic grouping; the fifty-two living species are divided among two orders, the Acipenseriformes (sturgeons and paddlefishes), and the Polypteriformes (reedfishes and bichirs).

Cladodus

Cladodus is an extinct genus of cartilaginous fishes in the family Cladoselachidae. As the name implies, they are a type of cladodont, primitive sharks with teeth designed to snag fish and swallow them whole, instead of sawing off chunks to swallow.

Fossils of Cladodus have been found in Barkip, Scotland and in the Pitkin Formation (Carboniferous period) in Arkansas, United States.

Duffinselache

Duffinselache (Duffin's shark) is an extinct genus of basal selachimorph elasmobranchii cartilaginous fish known from the Late Triassic (Rhaetian stage) of England. It was first named by Duffin in 1998 as a species of Polyacrodus. Plamen S. Andreev and Gilles Cuny in 2012 reassigned to its own genus, and its type species is Duffinselache holwellensis.

Euselachii

Euselachii are an infraclass of a class of cartilaginous fish. This group includes sharks and rays.

Galeomorphii

Galeomorphii is a superorder of cartilaginous fishes which includes all modern sharks except the dogfish and its relatives. They are sometimes called galea or galean sharks. There are about 300 living species in 23 families. Galean sharks are divided into four orders: the Heterodontiformes, Orectolobiformes, Lamniformes, and Carcharhiniformes

Hopleacanthus

Hopleacanthus is an extinct genus of shark.

Hybodontiformes

Hybodontiformes, also called hybodonts, are an extinct subset of Elasmobranchii (sharks, skates and rays) which existed from the Devonian to the Miocene. They form the group of sharks closest to neoselachians, the clade of modern sharks and rays. Hybodonts were named and are distinguished based on their conical tooth shape. They comprised the main group of Jurassic sharks in Europe and North America. They survived into the Late Cretaceous before going extinct, possibly due to competition from other sharks, though forms like Miosynechodus endured as recently as the Miocene. Lonchidion was one of the last hybodonts — its distinctive serrated fine spines occur in freshwater deposits from Wyoming alongside the fossils of the last dinosaurs, including Tyrannosaurus rex and Triceratops. Hybodontiformes are identified in the fossil record predominantly based on distinct teeth and fin spines. They were known to live in both fresh and salt water environments.

Neoselachii

Neoselachii are a grouping of cartilaginous fishes in subclass Elasmobranchii of class Chondrichthyes. It contains all living (extant) sharks, skates, and rays. The first known neoselachian is the Devonian (possible) hexanchiforme Mcmurdodus.

Protospinax

Protospinax is an extinct genus of cartilaginous fish found in the Solnhofen limestones of southern Bavaria. It is a difficult taxon to accommodate in taxonomies. Formerly known from only two specimens, further museum specimens were discovered at the Museum of Comparative Zoology of Harvard University in the 1990s, having been misidentified as Squatina and Heterodontus.

Protospinax was about 1.7 m in length.

Ptychodus

Ptychodus is a genus of extinct hybodontiform sharks. As well as a genus of durophagous (shell-crushing) sharks from the Late Cretaceous. Fossils of Ptychodus teeth are found in plenty in many of the Late Cretaceous marine sediments. There are many species among the Ptychodus that have been uncovered on all the continents around the globe. Such species are Ptychodus mortoni, P. decurrens, P. marginalis, P. mammillaris, P. rugosus and P. latissimus to name a few. They died out approximately 85 million years ago in the Western Interior Sea, where a majority of them were found.

A recent publication found that Ptychodus are classified as neoselachian versus hybodont or batoid.

Shark

Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorpha (or Selachii) and are the sister group to the rays. However, the term "shark" has also been used for extinct members of the subclass Elasmobranchii outside the Selachimorpha, such as Cladoselache and Xenacanthus, as well as other Chondrichthyes such as the holocephalid eugenedontidans.

Under this broader definition, the earliest known sharks date back to more than 420 million years ago. Acanthodians are often referred to as "spiny sharks"; though they are not part of Chondrichthyes proper, they are a paraphyletic assemblage leading to cartilaginous fish as a whole. Since then, sharks have diversified into over 500 species. They range in size from the small dwarf lanternshark (Etmopterus perryi), a deep sea species of only 17 centimetres (6.7 in) in length, to the whale shark (Rhincodon typus), the largest fish in the world, which reaches approximately 12 metres (40 ft) in length. Sharks are found in all seas and are common to depths of 2,000 metres (6,600 ft). They generally do not live in freshwater although there are a few known exceptions, such as the bull shark and the river shark, which can be found in both seawater and freshwater. Sharks have a covering of dermal denticles that protects their skin from damage and parasites in addition to improving their fluid dynamics. They have numerous sets of replaceable teeth.Well-known species such as the great white shark, tiger shark, blue shark, mako shark, thresher shark, and hammerhead shark are apex predators—organisms at the top of their underwater food chain. Many shark populations are threatened by human activities.

Squalomorphii

Squalomorphii is a superorder of cartilaginous fishes, generally characterized by lacking traits such as an anal fin, nictitating membrane, or suborbital shelves in the cranium. Also called squalea, or squalean sharks. There are about 163 living species in 11 families. Squalean sharks are divided into four orders: the Hexanchiformes, Squaliformes, Squatiniformes, and Pristiophoriformes.

Symmoriida

Symmoriida is an extinct order of ratfish that contains three families. It was synonymized subjectively with Cladodontida by Lund (1986); it was corrected as Symmoriiformes by Maisey (2008). It was assigned to Cladoselachii by Goto et al. (1988); to Elasmobranchii by Williams (1998), and to Chondrichthyes by Sepkoski in 2002 and by Maisey in 2008. In the fossil record, they appear at the beginning of the Carboniferous. Most of them died out at the start of the Permian; however, a single tooth described by Guinot et al. (2013) from the Valanginian of France indicates that members of the family Falcatidae might have survived until the Early Cretaceous.

Tassiliodus

Tassiliodus is an extinct genus of euselachian chondrichthyan known from the Early Devonian of southern Algeria.

William Toby White

William Toby White is an Australian ichthyologist. He studies speciation and biodiversity of shark, ray, and skate species (subclass Elasmobranchii) through morphological and molecular systematics.

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