Ray (fish)
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Batoidea is a superorder of cartilaginous fishes, commonly known as rays. They and their close relatives, the
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 Selachimo ...
s, comprise the subclass
Elasmobranchii Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of ...
. 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 fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
s that are fused to the head, and
gill slit Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In con ...
s that are placed on their
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
surfaces.


Anatomy

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 Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
slot-like body openings called
gill slits Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of Chondrichthyes, cartilaginous fish such as sharks and Batoidea, rays, as well as deep-branching vertebrates ...
that lead from the
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
, but the
Hexatrygonidae The sixgill stingray (''Hexatrygon bickelli'') is a species of stingray and the only extant member of the family Hexatrygonidae. Although several species of sixgill stingrays have been described historically, they may represent variations in a ...
have six. Batoid gill slits lie under the
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
s 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
guitarfish The guitarfish, also referred to as shovelnose rays, 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 trop ...
es 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
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
s 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. The jaws have euhyostylic type suspension, which relies completely on the hyomandibular cartilages for support. 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.


Reproduction

Batoids reproduce in a number of ways. As is characteristic of elasmobranchs, batoids undergo
internal fertilization Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For inte ...
. 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. All skates and some rays are oviparous (egg laying) while other rays are
ovoviviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
, meaning that they give birth to young which develop in a womb but without involvement of a placenta. The eggs of oviparous skates are laid in leathery egg cases that are commonly known as
mermaid's purse An egg case or egg capsule, often colloquially called a mermaid's purse, is the casing that surrounds the eggs of oviparous sharks, skates and chimaeras. Egg cases typically contain one embryo, except for big skate and mottled skate egg c ...
s 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. Capture-induced parturition is rarely considered in
fisheries management The goal of fisheries management is to produce sustainable biological, environmental and socioeconomic benefits from renewable aquatic resources. Wild fisheries are classified as renewable when the organisms of interest (e.g., fish, shellfish, ...
despite being shown to occur in at least 12% of live bearing sharks and rays (88 species to date).


Habitat

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 . 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.


Feeding

Most batoids have developed heavy, rounded teeth for crushing the shells of bottom-dwelling species such as
snails A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gastro ...
,
clams Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two ...
, oysters,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s, and some
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
, depending on the species. Manta rays feed on
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
.


Evolution

Batoids belong to the ancient lineage of cartilaginous fishes. Fossil
denticle A denticle is any small tooth-like or bristle-like structure. "Denticle" may refer to: * Denticle (tooth feature), serrations on the teeth of dinosaurs, lizards, sharks, and mammals * Dermal denticles or placoid scales, in cartilaginous fishes * ...
s (tooth-like scales in the skin) resembling those of today's
chondrichthyan Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
s date at least as far back as the
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
, with the oldest unambiguous fossils of cartilaginous fish dating from the middle Devonian. A clade within this diverse family, the
Neoselachii Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gi ...
, emerged by the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
, with the best-understood neoselachian fossils dating from the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
. The oldest confirmed ray is ''
Antiquaobatis ''Antiquaobatis'' is an extinct genus of ray from the Early Jurassic (Late Pliensbachian) of Europe, containing the single species ''A. grimmenensis''. It is the oldest known described member of the Rajiformes, and is based on a single tooth fro ...
'', from the
Pliensbachian The Pliensbachian is an age (geology), age of the geologic timescale and stage (stratigraphy), stage in the stratigraphic column. It is part of the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series an ...
of
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. The clade is represented today by
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 Selachimo ...
s,
sawfish Sawfish, also known as carpenter sharks, are a family of rays characterized by a long, narrow, flattened rostrum, or nose extension, lined with sharp transverse teeth, arranged in a way that resembles a saw. They are among the largest fish ...
, rays and
skate Skate or Skates may refer to: Fish *Skate (fish), several genera of fish belonging to the family Rajidae * Pygmy skates, several genera of fish belonging to the family Gurgesiellidae * Smooth skates or leg skates, several genera of fish belongin ...
s.


Classification

The classification of batoids is currently undergoing revision; however, molecular evidence refutes the hypothesis that skates and rays are derived sharks. Nelson's 2006 '' Fishes of the World'' recognizes four orders. The
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
Sclerorhynchoidea are basal or ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
''; they show features of the Rajiformes but have snouts resembling those of sawfishes. However, evidence indicates they are probably the sister group to sawfishes. Phylogenetic tree of Batoidea: Order
Torpediniformes Fernando de Buen y Lozano was a Spanish ichthyologist and oceanographer Oceanography (), also known as oceanology and ocean science, is the scientific study of the oceans. It is an Earth science, which covers a wide range of topics, including ...
* Family
Hypnidae ''Hypnos monopterygius'', also known as the coffin ray or Australian numbfish, is a species of electric ray endemic to Australia, where it is common in inshore waters shallower than . It is the sole member of its genus ''Hypnos'', and family ...
(coffin rays) * Family
Narcinidae Narcinidae, or numbfishes, are a family of electric rays (order Torpediniformes). They are bottom-dwelling cartilaginous fishes with large, rounded pectoral fin discs and long tails. They can produce an electric discharge for defense, from whic ...
(numbfishes) * Family Narkidae (sleeper rays) * Family
Torpedinidae The family Torpedinidae contains 22 species of electric rays or torpedoes, flat cartilaginous fishes that produce electricity as a defense and feeding mechanism. They are slow-moving bottom-dwellers. The largest species is the Atlantic torped ...
(torpedo rays) Order
Rhinopristiformes Rhinopristiformes is an order of rays, cartilaginous fishes related to sharks, containing shovelnose rays and allied groups. Families * Family Glaucostegidae (giant guitarfishes) * Family Pristidae (sawfishes) * Family Rhinidae (wedgefishes) * ...
* Family Glaucostegidae (giant guitarfishes) * Family
Platyrhinidae The Platyrhinidae are a family of rays, commonly known as thornbacks due to their dorsal rows of large thorns. They resemble guitarfishes in shape. Though traditionally classified with stingrays, molecular evidence suggests they are more closely ...
* (fanrays) * Family
Pristidae Sawfish, also known as carpenter sharks, are a family of rays characterized by a long, narrow, flattened rostrum, or nose extension, lined with sharp transverse teeth, arranged in a way that resembles a saw. They are among the largest fish ...
(sawfishes) * Family Rhinidae (wedgefishes) * Family Rhinobatidae (guitarfishes) * Family
Trygonorrhinidae Trygonorrhinidae, the banjo rays, is a family of rays, comprising eight species in three genera. They were formerly classified in the family Rhinobatidae. Taxonomy * '' Aptychotrema'' Norman, 1926 ** '' Aptychotrema rostrata'' Shaw, 1794 (East ...
(banjo rays) * Family
Zanobatidae 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 ...
* (panrays) * the placement of these families is uncertain Order
Rajiformes 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 hea ...
* Family Anacanthobatidae (legskates) * Family Arhynchobatidae (softnose skates) * Family
Gurgesiellidae Pygmy skates are cartilaginous fish belonging to the family Gurgesiellidae in the superorder Batoidea of rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spin ...
(pygmy skates) * Family Rajidae (skates) Order
Myliobatiformes 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 grou ...
* Family Aetobatidae (pelagic eagle rays) * Family Dasyatidae (whiptail stingrays) * Family
Gymnuridae The butterfly rays are the rays forming the genus ''Gymnura'' and the family Gymnuridae. They are found in warm oceans worldwide, and occasionally in estuaries. The body of butterfly rays is flattened and surrounded by an extremely broad disc f ...
(butterfly rays) * Family
Hexatrygonidae The sixgill stingray (''Hexatrygon bickelli'') is a species of stingray and the only extant member of the family Hexatrygonidae. Although several species of sixgill stingrays have been described historically, they may represent variations in a ...
(sixgill stingrays) * Family Myliobatidae (devilrays) * Family
Plesiobatidae The deepwater stingray or giant stingaree (''Plesiobatis daviesi'') is a species of stingray and the sole member of the family Plesiobatidae. It is widely distributed in the Indo-Pacific, typically over fine sediments on the upper continental ...
(giant stingarees) * Family
Potamotrygonidae River stingrays or freshwater stingrays are Neotropical freshwater fishes of the family Potamotrygonidae in the order Myliobatiformes, one of the four orders of batoids, cartilaginous fishes related to sharks. They are found in rivers in tropic ...
(Neotropical stingrays) * Family Rhinopteridae (cownose rays) * Family
Urolophidae 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 formi ...
(stingarees) * Family
Urotrygonidae Urotrygonidae is a family of rays in the order Myliobatiformes, commonly referred to as the American round stingrays or round rays. They are native to the tropical and warm temperate marine waters of the Americas. The two genera in this family ...
(round stingrays)


Conservation

According to a 2021 study in ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...
'', 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. This is notably the case in the Mediterranean Sea - most impacted by unregulated fishing - where a recent international survey concluded that only 38 species of rays and skates still subsisted.Guide of Mediterranean Skates and Rays. Oct. 2022. Mendez L., Bacquet A. and F. Brian

/ref>


Differences between sharks and rays

All sharks and rays are cartilaginous fishes, Bony versus cartilaginous, contrasting with bony fishes. Many rays are adapted for feeding on the bottom.
Guitarfish The guitarfish, also referred to as shovelnose rays, 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 trop ...
es are somewhat between sharks and rays, displaying characteristics of both (though they are classified as rays).


See also

* List of cartilaginous fish#Superorder Batoidea *
Rhenanida Rhenanida ("Rhine (fish)") is an Order (biology), order of scaly placoderms. Unlike most other placoderms, the rhenanids' armor was made up of a mosaic of unfused scales and tubercles. The patterns and components of this "mosaic" correspond to t ...


Footnotes


References


Bibliography

* *


External links

* – database of bibliography of living/fossil sharks and rays (''see'' Chondrichtyes: Selachii) with more than 15 000 listed papers and many download links. * {{Authority control Extant Early Triassic first appearances Fish superorders