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.

Temporal range: Early Triassic-Recent[1]
Atlantic mobula lisbon
Manta ray, Mobula mobular
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Superorder: Batoidea

Batoidimorpha Myliobatoidea

Spotted Eagle Ray (Aetobatus narinari)2
Spotted eagle ray, Aetobatus narinari


Batoids are flat-bodied, and, like sharks, are cartilaginous marine 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 fishes 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/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 that 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 clade is represented today by sharks, sawfish, rays and skates.[7]













Phylogenetic tree of Batoidea[8]

The classification of batoids is currently undergoing revision; however, molecular evidence refutes the hypothesis that skates and rays are derived sharks.[9] Nelson's 2006 Fishes of the World recognizes four orders. 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[10]

Order Image Common name Family Genera Species Comment
Total CR IUCN 3 1.svg EN IUCN 3 1.svg VU IUCN 3 1.svg
Myliobatiformes Myliobatis aquila sasrája Stingrays and relatives 10 29 223 1 16 33 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.[11]
Rajiformes Amblyraja hyperborea1 Skates and relatives 5 36 270 4 12 26 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.
Torpediniformes Torpedo torpedo corsica2 Electric rays 4 12 69 2 9 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.[12]
Rhinopristiformes Sawfish genova Shovelnose rays and relatives 1 2 5-7 3-5 2 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.[13]

Order Torpediniformes

Order Rhinopristiformes

* the placement of these families is uncertain

Order Rajiformes

Order Myliobatiformes

Differences between sharks and rays

All sharks and rays are cartilaginous fishes, 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 sharks, guitar fishes and rays
Characteristic Shark Guitar fish Ray
Shark fish chondrichthyes Rhina anyclostoma australia Dasyatis brevicaudata 4x3
Shape laterally compressed spindle dorsoventrally compressed (flattened) disc 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 usually on top of the head
Gill openings on the sides ventral (underneath)
Pectoral fins distinct not distinct not distinct
Tail large caudal fin used for propulsion caudal fin that can be used for propulsion varies from thick tail as extension of body to a whip that can sting to almost no tail.
Locomotion swim by moving their caudal (tail) fin from side to side Guitar fish and sawfish have a caudal fin like the shark. swim by flapping their pectoral fins like wings.

See also



  1. ^ Stevens, J. & Last, P.R. (1998), Paxton, J.R. & Eschmeyer, W.N. (eds.), Encyclopedia of Fishes, San Diego: Academic Press, p. 60, ISBN 978-0-12-547665-2
  2. ^ Martin 2010.
  3. ^ Motta, PJ; Wilga, CD (2001), "Advances in the study of feeding behaviors, mechanisms, and mechanics of sharks", Environmental Biology of Fishes, 60: 131–56, doi:10.1023/A:1007649900712.
  4. ^ Wilga, CAD (2008), "Evolutionary divergence in the feeding mechanism of fishes", Acta Geologica Polonica, 58: 113–20.
  5. ^ "Reproduction overall". Skates and rays of Atlantic Canada. Canadian Shark Research Lab, Bedford Institute of Oceanography & Marine Fish Species, Risk Section, Northwest Atlantic Fisheries Center. Archived from the original on 2015-01-16. 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.
  7. ^ UCMP Berkeley "Chondrichthyes: Fossil Record"
  8. ^ McEachran & Aschliman 2004.
  9. ^ Douady, CJ; Dosay, M; Shivji, MS; Stanhope, MJ (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.
  10. ^ Kriwet, Jürgen. "The systematic position of the Cretaceous sclerorhynchid sawfishes (Elasmobranchii, Pristiorajea)" (PDF).
  11. ^ Nelson, J.S. (2006), Fishes of the World (fourth ed.), John Wiley, pp. 69–82, ISBN 978-0-471-25031-9
  12. ^ Bullock, Theodore Holmes; Hopkins, Carl D; Popper, Arthur N; Fay, Richard R (2005), Electroreception, Springer, pp. 5–7, ISBN 978-0-387-23192-1
  13. ^ 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.


  • Shark references. Database of bibliography of living/fossil sharks and rays (Chondrichtyes: Selachii) with more than 15.000 listed papers and many download links.
  • Martin, R Aidan (February 2010), "Batoids: Sawfishes, Guitarfishes, Electric Rays, Skates, and Sting Rays", Elasmo research, ReefQuest Centre for Shark Research.
  • McEachran, JD; Dunn, KA; Miyake, T (1996), "Interrelationships of the batoid fishes (Chondrichthyes: Batoidea)", Interrelationships of Fishes, Academic Press.
  • McEachran, JD; Aschliman, N (2004), "Phylogeny of batoidea", in Carrier, JC; Musick, JA; Heithaus, MR (eds.), Biology of sharks and their relatives, Boca Raton: CRC Press, pp. 79–114.
  • Nelson, Joseph S (2006) Fishes of the World Edition 4, illustrated. John Wiley & Sons. ISBN 9780471756446.

External links


Arhynchobatidae is a family of skates and is commonly known as the softnose skates. It belongs to the order Rajiformes in the superorder Batoidea of rays. At least 104 species have been described, in 13 genera. Softnose skates have at times been placed in the same family as hardnose skates, but most recent authors recognize them as a distinct family. Members of the Arhynchobatidae can be distinguished from hardnose skates in having a soft and flexible snout, as well as a more or less reduced rostrum.


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.


Dalpiazia is a prehistoric genus of sclerorhynchid sawfish whose fossils are found in rocks dating from the Maastrichtian stage in Morocco. It was named in honor of Ernst Stromer.

The validity of Dalpiazia has been questioned on the basis of its similarities to Ischyrhiza, a subgenus of Onchosaurus.


Elasmobranchii () 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.


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


The guitarfish are a family, Rhinobatidae, of rays. The guitarfish are known for an elongated body with a flattened head and trunk and small, ray-like wings. The combined range of the various species is tropical, subtropical, and warm temperate waters worldwide.


Pygmy skates are cartilaginous fish belonging to the family Gurgesiellidae in the superorder Batoidea of rays. Nineteen species in three genera are known.


Ischyrhiza is an extinct genus of sawfish from the Late Cretaceous and Early Paleogene, belonging to the primitive Batoidea family Sclerorhynchidae. Fossils of the genus have been found in Canada, the United States, the Aguja Formation of Mexico, the Tamayama Formation of Japan, the Dukamaje Formation of Niger, the El Molino Formation of Bolivia, the Quiriquina Formation of Chile, and the Chota Formation of Peru.

Knifetooth sawfish

The knifetooth sawfish (Anoxypristis cuspidata), also known as the pointed sawfish or narrow sawfish, is a species of sawfish in the family Pristidae, part of the Batoidea, a superorder of cartilaginous fish that includes the rays and skates. Sawfish display a circumglobal distribution in warm marine and freshwater habitats. Their extant biodiversity is limited to five species belonging to two genera (Pristis and Anoxypristis). The sawfishes are characterised by the long, narrow, flattened rostrum or extension on their snout. This is lined with sharp transverse teeth, arranged in a way that resembles the teeth of a saw. It is found in the shallow coastal waters and estuaries of the Indo-West Pacific, ranging from the Red Sea and the Persian Gulf to southern Japan, Papua New Guinea and northern Australia. It is the only living member of the genus Anoxypristis, but was previously included in the genus Pristis. Compared to that genus, Anoxypristis has a narrower rostral saw with numerous teeth on the distal part and no teeth on the basal one-quarter (toothless section about one-sixth in juveniles). This endangered species reaches a length of up to 3.5 m (11.5 ft).In addition to the living Anoxypristis cuspidata, this genus includes a few extinct species that are only known from fossil remains.


Myliobatiformes is one of the four orders of batoids, cartilaginous fishes related to sharks. They were formerly included in the order Rajiformes, but more recent phylogenetic studies have shown the myliobatiforms to be a monophyletic group, and its more derived members evolved their highly flattened shapes independently of the skates.


The panrays are a genus, Zanobatus, of rays found in coastal parts of the warm East Atlantic Ocean, ranging from Morocco to Angola. It is the only genus in the family Zanobatidae, which traditionally has been included in the Myliobatiformes order, but based on genetic evidence it is now in Rhinopristiformes or a sister taxon to Rhinopristiformes.The two species of panrays are generally poorly known and one of the species was only scientifically described in 2016. They are up to about 60 cm (2 ft) long, and brownish above with a heavily mottled, blotched or barred dark pattern. They are ovoviviparous and feed on benthic invertebrates.


Rajiformes is one of the four orders in the superorder Batoidea, flattened cartilaginous fishes related to sharks. Rajiforms 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 head and the gill slits are on the underside of the body. Most species give birth to live young, although some lay eggs with a horny capsule ("mermaid's purse").


Rhinobatos is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to Acroteriobatus, Glaucostegus, and Pseudobatos.


Rhinopristiformes is an order of rays, cartilaginous fishes related to sharks, containing shovelnose rays and allied groups.


Sclerorhynchidae is an extinct family of cartilaginous fish from the Cretaceous and Paleogene belonging to the primitive Batoidea suborder Sclerorhynchiformes.

Skate (fish)

Skates are cartilaginous fish belonging to the family Rajidae in the superorder Batoidea of rays. More than 150 species have been described, in 17 genera. Softnose skates and pygmy skates were previously treated as subfamilies of Rajidae (Arhynchobatinae and Gurgesiellinae), but are now considered as distinct families. Alternatively, the name "skate" is used to refer to the entire order of Rajiformes (families Anacanthobatidae, Arhynchobatidae, Gurgesiellidae and Rajidae).Members of Rajidae are distinguished by their stiff snout and a rostrum that is not reduced.

Transtillaspis batoidea

Transtillaspis batoidea is a species of moth of the family Tortricidae. It is found in Peru.


The Urolophidae are a family of rays in the order Myliobatiformes, commonly known as stingarees or round stingrays. This family formerly included the genera Urobatis and Urotrygon of the Americas, which are presently recognized as forming their own family Urotrygonidae. Stingarees are found in the Indo-Pacific region, with the greatest diversity off Australia. They are sluggish, bottom-dwelling fish that have been recorded from shallow waters close to shore to deep waters over the upper continental slope. Measuring between 15 and 80 cm (5.9 and 31.5 in) long, these rays have oval to diamond-shaped pectoral fin discs and relatively short tails that terminate in leaf-shaped caudal fins, and may also have small dorsal fins and lateral skin folds. Most are smooth-skinned, and some have ornate dorsal color patterns.

Stingarees feed on or near the sea floor, consuming small invertebrates and occasionally bony fishes. They are aplacental viviparous, meaning their embryos emerge from eggs inside the uterus, and are sustained to term first by yolk and later by maternally produced histotroph ("uterine milk"). As far is known, the gestation period lasts around a year and litter sizes tend to be small. Stingarees have one or two relatively large, venomous stinging spines on their tail for defense, with which they can inflict a painful wound on humans. Generally, stingarees have no economic value. Some species form a substantial component of the bycatch of commercial trawl fisheries.

Extant cartilaginous fish orders
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