Temporal range: Early Jurassic–Present [1]
Common stingray, Dasyatis pastinaca
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Clade: Neoselachii
Superorder: Batoidea
Compagno, 1973
Spotted eagle ray, Aetobatus narinari

Batoidea is a superorder of cartilaginous fishes, 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.


See also: Batoid locomotion

Batoids are flat-bodied, and, like sharks, are cartilaginous fish, meaning they have a boneless skeleton made of a tough, elastic cartilage. Most batoids have five ventral slot-like body openings called gill slits that lead from the gills, but the Hexatrygonidae have six.[2] Batoid gill slits lie under the pectoral fins on the underside, whereas a shark's are on the sides of the head. Most batoids have a flat, disk-like body, with the exception of the guitarfishes and sawfishes, while most sharks have a spindle-shaped body. Many species of batoid have developed their pectoral fins into broad flat wing-like appendages. The anal fin is absent. The eyes and spiracles are located on top of the head. Batoids have a ventrally located mouth and can considerably protrude their upper jaw (palatoquadrate cartilage) away from the cranium to capture prey.[3] The jaws have euhyostylic type suspension, which relies completely on the hyomandibular cartilages for support.[4] Bottom-dwelling batoids breathe by taking water in through the spiracles, rather than through the mouth as most fish do, and passing it outward through the gills.


Batoids reproduce in a number of ways. As is characteristic of elasmobranchs, batoids undergo internal fertilization. Internal fertilization is advantageous to batoids as it conserves sperm, does not expose eggs to consumption by predators, and ensures that all the energy involved in reproduction is retained and not lost to the environment.[5] All skates and some rays are oviparous (egg laying) while other rays are ovoviviparous, meaning that they give birth to young which develop in a womb but without involvement of a placenta.[6]

The eggs of oviparous skates are laid in leathery egg cases that are commonly known as mermaid's purses and which often wash up empty on beaches in areas where skates are common.

Capture-induced premature birth and abortion (collectively called capture-induced parturition) occurs frequently in sharks and rays when fished.[6] Capture-induced parturition is rarely considered in fisheries management despite being shown to occur in at least 12% of live bearing sharks and rays (88 species to date).[6]


Most species live on the sea floor, in a variety of geographical regions – mainly in coastal waters, although some live in deep waters to at least 3,000 metres (9,800 ft). Most batoids have a cosmopolitan distribution, preferring tropical and subtropical marine environments, although there are temperate and cold-water species. Only a few species, like manta rays, live in the open sea, and only a few live in freshwater, while some batoids can live in brackish bays and estuaries.


Most batoids have developed heavy, rounded teeth for crushing the shells of bottom-dwelling species such as snails, clams, oysters, crustaceans, and some fish, depending on the species. Manta rays feed on plankton.


Batoids belong to the ancient lineage of cartilaginous fishes. Fossil denticles (tooth-like scales in the skin) resembling those of today's chondrichthyans date at least as far back as the Ordovician, with the oldest unambiguous fossils of cartilaginous fish dating from the middle Devonian. A clade within this diverse family, the Neoselachii, emerged by the Triassic, with the best-understood neoselachian fossils dating from the Jurassic. The oldest confirmed ray is Antiquaobatis, from the Pliensbachian of Germany.[7] The clade is represented today by sharks, sawfish, rays and skates.[8]


Molecular evidence refutes the hypothesis that skates and rays are derived sharks.[9] The monophyly of the skates, the stingrays, and the electric rays has long been generally accepted. Along with Rhinopristiformes, these comprise the four traditionally accepted major batoid lineages, as in Nelson's 2006 Fishes of the World. However, the exact phylogeny of the major batoid lineages, internally and with respect to one another, has been subject to diverse treatments. The following cladogram is based on a comprehensive morphological assessment of batoid phylogeny published in 2004:[10]


However, a 2011 study significantly reevaluated the phylogeny of batoids, using nuclear and mitochondrial DNA from 37 taxa, representing almost all recognized families and all of the traditional four major lineages. This is a far more numerous and diverse set of sample taxa than in any previous study, producing findings reflected in the cladogram below.[1]


Holocephali (incl. Chimaera)


Selachimorpha (Sharks)


Rajiformes (Skates)

Platyrhinidae (Thornbacks)

Torpediniformes (Electric rays)

"Guitarfishes 1" (Trygonorrhinidae)

"Guitarfishes 2" (incl. Pristidae (Sawfishes))

Zanobatidae (Panrays)

Myliobatoidei (Stingrays)

This study strongly confirmed the traditionally accepted internal monophyly of skates, stingrays, and electric rays. It also recovered panrays as sister to the stingrays, as older morphological analyses had suggested. However, it found the Rhinopristiformes, including the sawfishes and various "guitarfishes", to be paraphyletic, comprising two distinct clades. Referred to as "Guitarfishes 1" and "Guitarfishes 2", the former contains only the Trygonorrhinidae, while the latter contains the remainder of Rhinopristiformes (the families Glaucostegidae, Pristidae, Rhinidae, and Rhinobatidae). In addition, while traditional phylogenies often find electric rays to be the basalmost batoids, followed by the Rhinopristiformes, this analysis finds a polytomy between skates, electric rays, and thornbacks at the base of Batoidea, with weak support for skates being the actual most basal lineage, followed by a clade uniting the electric rays and thornbacks.

The Mesozoic Sclerorhynchoidea are basal or incertae sedis; they show features of the Rajiformes but have snouts resembling those of sawfishes. However, evidence indicates they are probably the sister group to sawfishes.[11]

Order Image Common name Family Genera Species Comment
Myliobatiformes Stingrays and relatives 10 29 223 1 16 33 [a][12]
Rajiformes Skates and relatives 5 36 270 4 12 26 [b]
Torpediniformes Electric rays 4 12 69 2 9 [c][13]
Rhinopristiformes Shovelnose rays and relatives 1 2 5-7 3-5 2 [d][14]
Early Eocene fossil stingray Heliobatis radians
Giant devil ray, Mobula mobular

Order Torpediniformes

Order Rhinopristiformes

* the placement of these families is uncertain

Order Rajiformes

Order Myliobatiformes


See also: List of threatened rays

According to a 2021 study in Nature, the number of oceanic sharks and rays has declined globally by 71% over the preceding 50 years, jeopardising "the health of entire ocean ecosystems as well as food security for some of the world's poorest countries". Overfishing has increased the global extinction risk of these species to the point where three-quarters are now threatened with extinction.[15][16][17] This is notably the case in the Mediterranean Sea - most impacted by unregulated fishing - where a recent international survey of the Mediterranean Science Commission concluded that only 38 species of rays and skates still subsisted.[18]

Differences between sharks and rays

All sharks and rays are cartilaginous fish, contrasting with bony fishes. Many rays are adapted for feeding on the bottom. Guitarfishes are somewhat between sharks and rays, displaying characteristics of both (though they are classified as rays).

Comparison of Elasmobranchid fish
Characteristic Sharks Guitar fish Rays
Shape laterally compressed spindle dorsoventrally compressed (flattened) disc
Spiracles not always present always present
Habitat usually pelagic surface feeders, though carpet sharks are demersal bottom feeders demersal / pelagic mix usually demersal bottom feeders
Eyes usually at the side of the head usually on top of the head
Gill openings on the sides ventral (underneath)
Pectoral fins distinct not distinct
Tail large caudal fin whose primary function is to provide main forward propulsion varies from thick tail as extension of body to a whip that can sting to almost no tail.
Locomotion swim by moving their tail (caudal fin) from side to side Guitar fish and sawfish have a caudal fin like sharks swim by flapping their pectoral fins like wings

See also


  1. ^ Myliobatiformes include stingrays, butterfly rays, eagle rays, and manta rays. They were formerly included in the order Rajiformes, but more-recent phylogenetic studies have shown that they are a monophyletic group, and that its more-derived members evolved their highly flattened shapes independently of the skates.[12]
  2. ^ Rajiformes include skates, guitarfishes, and wedgefishes. They are distinguished by the presence of greatly enlarged pectoral fins, which reach as far forward as the sides of the head, with a generally flattened body. The undulatory pectoral fin motion diagnostic to this taxon is known as rajiform locomotion. The eyes and spiracles are located on the upper surface of the body, and the gill slits on the underside. They have flattened, crushing teeth, and are generally carnivorous. Most species give birth to live young, although some lay eggs inside a protective capsule or mermaid's purse.
  3. ^ The electric rays have electric organs in their pectoral fin discs that generate electric current. They are used to immobilize prey and for defense. The current is strong enough to stun humans, and the ancient Greeks and Romans used these fish to treat ailments such as headaches.[13]
  4. ^ The sawfishes are shark-like in form, having tails used for swimming and smaller pectoral fins than most batoids. The pectoral fins are attached above the gills as in all batoids, giving the fishes a broad-headed appearance. They have long, flat snouts with a row of tooth-like projections on either side. The snouts are up to 1.8 metres (6 ft) long, and 30 centimetres (1 ft) wide, and are used for slashing and impaling small fishes and to probe in the mud for embedded animals. Sawfishes can enter freshwater rivers and lakes. Some species reach a total length of 6 metres (20 ft). All species of sawfish are endangered or critically endangered.[14]


  1. ^ a b Aschliman, Neil C.; Nishida, Mutsumi; Miya, Masaki; Inoue, Jun G.; Rosana, Kerri M.; Naylor, Gavin J.P. (2012). "Body plan convergence in the evolution of skates and rays (Chondrichthyes: Batoidea)". Molecular Phylogenetics and Evolution. 63 (1). Elsevier BV: 28–42. doi:10.1016/j.ympev.2011.12.012.
  2. ^ Martin, R. Aidan (February 2010). "Batoids: Sawfishes, Guitarfishes, Electric Rays, Skates, and Sting Rays". Elasmo research. ReefQuest Centre for Shark Research.
  3. ^ Motta, P.J.; Wilga, C.D. (2001). "Advances in the study of feeding behaviors, mechanisms, and mechanics of sharks". Environmental Biology of Fishes. 60 (1–3): 131–56. doi:10.1023/A:1007649900712. S2CID 28305317.
  4. ^ Wilga, C.A.D. (2008). "Evolutionary divergence in the feeding mechanism of fishes". Acta Geologica Polonica. 58: 113–20.
  5. ^ "Reproduction overall". Risk Section, Bedford Institute of Oceanography & Marine Fish Species. Canadian Shark Research Lab. Skates and rays of Atlantic Canada. Northwest Atlantic Fisheries Center. Archived from the original on 16 January 2015. Retrieved 27 May 2012.
  6. ^ a b c Adams, Kye R.; Fetterplace, Lachlan C.; Davis, Andrew R.; Taylor, Matthew D.; Knott, Nathan A. (January 2018). "Sharks, rays and abortion: The prevalence of capture-induced parturition in elasmobranchs". Biological Conservation. 217: 11–27. doi:10.1016/j.biocon.2017.10.010. S2CID 90834034. Archived from the original on 2019-02-23. Retrieved 2018-12-09.
  7. ^ Stumpf, Sebastian; Kriwet, Jürgen (2019). "A new Pliensbachian elasmobranch (Vertebrata, Chondrichthyes) assemblage from Europe, and its contribution to the understanding of late Early Jurassic elasmobranch diversity and distributional patterns". PalZ. 93 (4): 637–658. doi:10.1007/s12542-019-00451-4.
  8. ^ "Chondrichthyes: Fossil Record". University of California Museum of Paleontology. U.C. Berkeley.
  9. ^ Douady, C.J.; Dosay, M.; Shivji, M.S.; Stanhope, M.J. (2003). "Molecular phylogenetic evidence refuting the hypothesis of Batoidea (rays and skates) as derived sharks". Molecular Phylogenetics and Evolution. 26 (2): 215–221. doi:10.1016/S1055-7903(02)00333-0. PMID 12565032.
  10. ^ McEachran, J.D.; Aschliman, N. (2004). "Phylogeny of batoidea". In Carrier, J.C.; Musick, J.A.; Heithaus, M.R. (eds.). Biology of sharks and their relatives. Boca Raton, Florida: CRC Press. pp. 79–114.
  11. ^ Kriwet, Jürgen. "The systematic position of the Cretaceous sclerorhynchid sawfishes (Elasmobranchii, Pristiorajea)" (PDF). Archived from the original (PDF) on 2016-04-16. Retrieved 2016-04-04.
  12. ^ a b Nelson, J.S. (2006). Fishes of the World (fourth ed.). John Wiley. pp. 69–82. ISBN 978-0-471-25031-9.
  13. ^ a b Bullock, Theodore Holmes; Hopkins, Carl D.; Popper, Arthur N.; Fay, Richard R. (2005). Electroreception. Springer. pp. 5–7. ISBN 978-0-387-23192-1.
  14. ^ a b Faria, Vicente V.; McDavitt, Matthew T.; Charvet, Patricia; Wiley, Tonya R.; Simpfendorfer, Colin A.; Naylor, Gavin J.P. (2013). "Species delineation and global population structure of critically endangered sawfishes (Pristidae)". Zoological Journal of the Linnean Society. 167: 136–164. doi:10.1111/j.1096-3642.2012.00872.x.
  15. ^ Pacoureau, Nathan; Rigby, Cassandra L.; Kyne, Peter M.; Sherley, Richard B.; Winker, Henning; Carlson, John K.; Fordham, Sonja V.; Barreto, Rodrigo; Fernando, Daniel; Francis, Malcolm P.; Jabado, Rima W.; Herman, Katelyn B.; Liu, Kwang-Ming; Marshall, Andrea D.; Pollom, Riley A.; Romanov, Evgeny V.; Simpfendorfer, Colin A.; Yin, Jamie S.; Kindsvater, Holly K.; Dulvy, Nicholas K. (2021). "Half a century of global decline in oceanic sharks and rays". Nature. 589 (7843): 567–571. Bibcode:2021Natur.589..567P. doi:10.1038/s41586-020-03173-9. hdl:10871/124531. PMID 33505035. S2CID 231723355.
  16. ^ Briggs, Helen (28 January 2021). "Extinction: 'Time is running out' to save sharks and rays". BBC News. Retrieved 29 January 2021.
  17. ^ Richardson, Holly (27 January 2021). "Shark, ray populations have declined by 'alarming' 70 per cent since 1970s, study finds". ABC News. Australian Broadcasting Corporation. Retrieved 29 January 2021.
  18. ^ Guide of Mediterranean Skates and Rays. Oct. 2022. Mendez L., Bacquet A. and F. Briand.[1]