Batoidea

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Batoidea
Temporal range: Early Jurassic–Present
Atlantic mobula lisbon.jpg
Giant devil ray, Mobula mobular
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Clade: Neoselachii
Superorder: Batoidea
Compagno, 1973
Orders
Synonyms
Spotted eagle ray, Aetobatus narinari Spotted Eagle Ray (Aetobatus narinari)2.jpg
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.

Contents

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 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.

Reproduction

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]

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 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.

Feeding

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.

Evolution

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]

Classification

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]

Chondrichthyes

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]

Chondrichthyes

Holocephali (incl. Chimaera) Chimaera mon.JPG

Elasmobranchii

Selachimorpha (Sharks) Shark fish chondrichthyes.jpg

Batoidea

Rajiformes (Skates) Raja montagui2.jpg

Platyrhinidae (Thornbacks)

Torpediniformes (Electric rays) Torpedo nobiliana.jpg

"Guitarfishes 1" (Trygonorrhinidae) Zapteryx brevirostris.jpg

"Guitarfishes 2" (incl. Pristidae (Sawfishes)) Pristis microdon.jpg

Zanobatidae (Panrays)

Myliobatoidei (Stingrays) Mobula mobular.jpg

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]

OrderImageCommon nameFamilyGeneraSpeciesComment
Total CR IUCN 3 1.svg EN IUCN 3 1.svg VU IUCN 3 1.svg
Myliobatiformes Myliobatis aquila sasraja.jpg Stingrays and relatives102922311633 [lower-alpha 1] [12]
Rajiformes Amblyraja hyperborea1.jpg Skates and relatives53627041226 [lower-alpha 2]
Torpediniformes Torpedo torpedo corsica2.jpg Electric rays 4126929 [lower-alpha 3] [13]
Rhinopristiformes Sawfish genova.jpg Shovelnose rays and relatives125-73-52 [lower-alpha 4] [14]
Early Eocene fossil stingray Heliobatis radians Heliobatis radians Green River Formation (cropped).jpg
Early Eocene fossil stingray Heliobatis radians

Order Torpediniformes

Order Rhinopristiformes

* the placement of these families is uncertain

Order Rajiformes

Order Myliobatiformes

Conservation

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
Shark fish chondrichthyes.jpg Rhina anyclostoma australia.jpg Dasyatis brevicaudata 4x3.jpg
Shape laterally compressed spindle dorsoventrally compressed (flattened) disc
Spiracles not always presentalways present
Habitatusually pelagic surface feeders, though carpet sharks are demersal bottom feeders demersal / pelagic mixusually demersal bottom feeders
Eyesusually at the side of the headusually on top of the head
Gill openingson the sidesventral (underneath)
Pectoral fins distinctnot distinct
Taillarge caudal fin whose primary function is to provide main forward propulsionvaries 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 sideGuitar fish and sawfish have a caudal fin like sharksswim by flapping their pectoral fins like wings

See also

Footnotes

  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]

Related Research Articles

<span class="mw-page-title-main">Chondrichthyes</span> Class of jawed cartilaginous fishes

Chondrichthyes is a class of jawed fish that contains the cartilaginous fish or chondrichthyians, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or bony fish, which have skeletons primarily composed of bone tissue. Chondrichthyes are aquatic vertebrates with paired fins, paired nares, placoid scales, conus arteriosus in the heart, and a lack of opecula and swim bladders. Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.

<span class="mw-page-title-main">Stingray</span> Suborder of fishes

Stingrays are a group of sea rays, a type of cartilaginous fish. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae, Plesiobatidae, Urolophidae (stingarees), Urotrygonidae, Dasyatidae, Potamotrygonidae, Gymnuridae and Myliobatidae . There are about 220 known stingray species organized into 29 genera.

<span class="mw-page-title-main">Myliobatiformes</span> Order of cartilaginous fishes

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.

<span class="mw-page-title-main">Skate (fish)</span> Family of fishes

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, but are now considered as distinct families. Alternatively, the name "skate" is used to refer to the entire order of Rajiformes.

<span class="mw-page-title-main">Elasmobranchii</span> Subclass of fishes

Elasmobranchii is a subclass of Chondrichthyes or cartilaginous fish, including modern sharks, rays, skates, and sawfish. 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.

<span class="mw-page-title-main">Guitarfish</span> Family of cartilaginous fishes

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 tropical, subtropical, and warm temperate waters worldwide.

<span class="mw-page-title-main">Rajiformes</span> Order of fishes in the superorder Batoidea

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 enclosed in a horny capsule.

<span class="mw-page-title-main">Urolophidae</span> Family of cartilaginous fishes

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 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.

<span class="mw-page-title-main">Sixgill stingray</span> Species of cartilaginous fish

The sixgill stingray 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 single, widespread species. This flabby, heavy-bodied fish, described only in 1980, is unique among rays in having six pairs of gill slits rather than five. Growing up to 1.7 m (5.6 ft) long, it has a rounded pectoral fin disc and a long, triangular, and flexible snout filled with a gelatinous substance. It is brownish above and white below, and lacks dermal denticles.

<span class="mw-page-title-main">Smalltooth sawfish</span> Species of cartilaginous fish

The smalltooth sawfish is a species of sawfish in the family Pristidae. It is found in shallow tropical and subtropical waters in coastal and estuarine parts of the Atlantic. Reports from elsewhere are now believed to be misidentifications of other species of sawfish. It is a critically endangered species that has disappeared from much of its historical range.

<i>Rhina ancylostoma</i> Species of cartilaginous fish

Rhina ancylostoma, also known as the bowmouth guitarfish, shark ray or mud skate, is a species of ray and a member of the family Rhinidae. Its evolutionary affinities are not fully resolved, though it may be related to true guitarfishes and skates. This rare species occurs widely in the tropical coastal waters of the western Indo-Pacific, at depths of up to 90 m (300 ft). Highly distinctive in appearance, Rhina ancylostoma has a wide and thick body with a rounded snout and large shark-like dorsal and tail fins. Its mouth forms a W-shaped undulating line, and there are multiple thorny ridges over its head and back. It has a dorsal color pattern of many white spots over a bluish gray to brown background, with a pair of prominent black markings over the pectoral fins. This large species can reach a length of 2.7 m (8.9 ft) and weight of 135 kg (298 lb).

<span class="mw-page-title-main">Smooth butterfly ray</span> Species of fish

The smooth butterfly ray is a species of cartilaginous fish in the family Gymnuridae. It is a member of the order Myliobatiformes, which contains 10 total families. Its natural habitats are shallow seas, subtidal aquatic beds, estuarine waters, and coastal saline lagoons. Its common name is derived from its compressed body, pectoral fins that are wider than their length, and overall diamond shape.

<span class="mw-page-title-main">Thornback guitarfish</span> Species of cartilaginous fish

The thornback guitarfish is a species of ray in the family Platyrhinidae, and the only member of its genus. Despite its name and appearance, it is more closely related to electric rays than to true guitarfishes of the family Rhinobatidae. This species ranges from Tomales Bay to the Gulf of California, generally in inshore waters no deeper than 6 m (20 ft). It can be found on or buried in sand or mud, or in and near kelp beds. Reaching 91 cm (36 in) in length, the thornback guitarfish has a heart-shaped pectoral fin disc and a long, robust tail bearing two posteriorly positioned dorsal fins and a well-developed caudal fin. The most distinctive traits of this plain-colored ray are the three parallel rows of large, hooked thorns that start from the middle of the back and run onto the tail.

<span class="mw-page-title-main">Finless sleeper ray</span> Species of cartilaginous fish

The finless sleeper ray is a species of electric ray in the family Narkidae, and the sole member of its genus. It is found over the continental shelf of Southeast Asia from the eastern Andaman Sea to Vietnam and Borneo. Typically growing no more than 15 cm (5.9 in) long, it may be the smallest cartilaginous fish. The finless sleeper ray is the only electric ray that lacks dorsal fins. It has an oval pectoral fin disc that varies from longer than wide to wider than long, depending on age, and a short, robust tail that terminates in a short, deep caudal fin. The trailing margins of its pelvic fins are sexually dimorphic, being more concave in males.

<span class="mw-page-title-main">Arhynchobatidae</span> Family of fishes

Arhynchobatidae is a family of skates whose members are 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.

<span class="mw-page-title-main">Wedgefish</span> Family of cartilaginous fishes

Wedgefishes are rays of the family Rhinidae, comprising eleven species in three genera. Classified in the order Rhinopristiformes along with guitarfishes and sawfishes, they have also been known as giant guitarfishes or sharkfin guitarfishes.

<span class="mw-page-title-main">Clearnose skate</span> Species of cartilaginous fish

The clearnose skate is a species of cartilaginous fish in the family Rajidae. R. eglanteria is also known by other common names such as the brier skate and summer skate. Clearnose skates are easily identified by the translucent patches on either side of their snouts and their mottled dorsal surface. They are found along the Atlantic and Gulf coasts of the United States in shallow waters of the continental shelf.

<span class="mw-page-title-main">Rhinopristiformes</span> Order of cartilaginous fishes

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

Batoids are a superorder of cartilaginous fish consisting of skates, rays and other fish all characterized by dorsoventrally flattened bodies and large pectoral fins fused to the head. This distinctive morphology has resulted in several unique forms of locomotion. Most Batoids exhibit median paired fin swimming, utilizing their enlarged pectoral fins. Batoids that exhibit median paired fin swimming fall somewhere along a spectrum of swimming modes from mobuliform to rajiform based on the number of waves present on their fin at once. Of the four orders of Batoidae this holds truest for the Myliobatiformes (rays) and the Rajiformes (skates). The two other orders: Rhinopristiformes and Torpediniformes exhibit a greater degree of body caudal fin swimming.

References

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Bibliography