
Batoidea is a
superorder of
cartilaginous fishes
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 ...
, 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 Selachi ...
s, 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 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 slits that are placed on their
ventral 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 slot-like body openings called
gill slits 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 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
guitarfishes and
sawfishes
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 wit ...
, 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. 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 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, 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 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.
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).
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,
clams,
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 gro ...
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% ...
, 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 cr ...
.
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
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. ...
, 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
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, with the best-understood neoselachian fossils dating from the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), 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 J ...
. The oldest confirmed ray is ''
Antiquaobatis'', from the
Pliensbachian
The Pliensbachian is an age of the geologic timescale and stage in the stratigraphic column. It is part of the Early or Lower Jurassic Epoch or Series and spans the time between 190.8 ± 1.5 Ma and 182.7 ± 1.5 Ma (million years ago). The ...
of
Germany
Germany, officially the Federal Republic of Germany (FRG),, is a country in Central Europe. It is the most populous member state of the European Union. Germany lies between the Baltic and North Sea to the north and the Alps to the sou ...
. 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 Selachi ...
s,
sawfish, rays and
skates.
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 Creta ...
Sclerorhynchoidea
Sclerorhynchoidei is an extinct suborder of rajiform rays that had long rostra with large denticles similar to sawfishes and sawsharks. This feature was convergently evolved and their closest living relatives are actually skates. While they ...
are
basal
Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''.
Science
* Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure
* Basal (medicine), a minimal level that is nec ...
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.
Phylogenetic tree of Batoidea:

Order
Torpediniformes
* Family
Hypnidae (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 wh ...
(numbfishes)
* Family
Narkidae
Members of the family Narkidae are commonly known as sleeper rays. They are restricted to the temperate and tropical Indo-West Pacific from South Africa to Japan to Indonesia, and are exclusively marine and are absent from freshwater habitats. Th ...
(sleeper rays)
* Family
Torpedinidae (torpedo rays)
Order
Rhinopristiformes
* Family
Glaucostegidae
''Glaucostegus'', also known as giant guitarfishes, is a genus of large Indo-Pacific rays, with a single species, '' Glaucostegus cemiculus'', in the East Atlantic, and Mediterranean. They were formerly classified in the family Rhinobatidae but ...
(giant guitarfishes)
* Family
Platyrhinidae* (fanrays)
* Family
Pristidae (sawfishes)
* Family
Rhinidae
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.
T ...
(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 (Eas ...
(banjo rays)
* Family
Zanobatidae* (panrays)
* the placement of these families is uncertain
Order
Rajiformes
* Family
Anacanthobatidae
The Anacanthobatidae, the smooth skates or leg skates, are a family of skates found at depths below in the Indian, Pacific and Atlantic Oceans.
They lack the dorsal denticles (sharp, tooth-like scales) of other rays, hence their name, from Gr ...
(legskates)
* Family
Arhynchobatidae (softnose skates)
* Family
Gurgesiellidae (pygmy skates)
* Family
Rajidae (skates)
Order
Myliobatiformes
* Family
Aetobatidae (pelagic eagle rays)
* Family
Dasyatidae (whiptail stingrays)
* Family
Gymnuridae (butterfly rays)
* Family
Hexatrygonidae (sixgill stingrays)
* Family
Myliobatidae (devilrays)
* Family
Plesiobatidae (giant stingarees)
* Family
Potamotrygonidae (Neotropical stingrays)
* Family
Rhinopteridae
''Rhinoptera'' is a genus of ray commonly known as the cownose rays. This genus is the only member of the family Rhinopteridae.
Species
There are currently 8 recognized extant (living) species in this genus:
* '' Rhinoptera adspersa'' J. P. M� ...
(cownose rays)
* Family
Urolophidae (stingarees)
* Family
Urotrygonidae (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 ar ...
'', 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
Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in the ...
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. Guitarfishes 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 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 the plates of ar ...
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