Chondrichthyes (; ) is a

class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...

jawed fish Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant taxon, extant vertebrates, including all living bony fishes (bot ...

that contains the cartilaginous fish or chondrichthyans, which all have

skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...

s primarily composed of

cartilage Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...

. They can be contrasted with the

Osteichthyes Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...

or ''bony fish'', which have skeletons primarily composed of

bone tissue A bone is a Stiffness, rigid Organ (biology), organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red blood cell, red and white blood cells, store minerals, provi ...

. Chondrichthyes are aquatic

vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

s with

paired fins Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only b ...

, paired

nare Nare may refer to: People * Daniel Nare, Botswanan football manager * Tadu Teshome Nare (born 2001), Ethiopian runner * Théophile Naré (born 1966), Catholic bishop of Kaya, Burkina Faso Other meanings * Nareh (given name) (or Nare, Nara), Armeni ...

placoid scale A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as w ...

conus arteriosus The infundibulum (also known as ''conus arteriosus'') is a conical pouch formed from the upper and left angle of the right ventricle in the chordate heart, from which the pulmonary trunk arises. It develops from the bulbus cordis. Typically, the ...

in the

heart The heart is a muscular Organ (biology), organ found in humans and other animals. This organ pumps blood through the blood vessels. The heart and blood vessels together make the circulatory system. The pumped blood carries oxygen and nutrie ...

, and a lack of opercula and

swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ (anatomy), organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift ...

s. Within the infraphylum

Gnathostomata Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant vertebrates, including all living bony fishes (both ray-finned ...

, cartilaginous fishes are distinct from all other jawed vertebrates. The class is divided into two subclasses:

Elasmobranchii Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including modern sharks ( division Selachii), and batomorphs (division Batomorphi, including rays, skates, and sawfish). Members of this subclass are characterised by h ...

(

shark Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...

s, rays, skates and

sawfish Sawfish, also known as carpenter sharks, are a family of very large 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 lar ...

) and

Holocephali Holocephali (Sometimes spelled Holocephala; Romanization of Greek, Greek for "complete head" in reference to the fusion of Palatoquadrate, upper jaw with the rest of the skull) is a Subclass (biology), subclass of Chondrichthyes, cartilaginous fi ...

(

chimaera Chimaeras are Chondrichthyes, cartilaginous fish in the order (biology), order Chimaeriformes (), known informally as ghost sharks, rat fish (not to be confused with rattails), spookfish, or rabbit fish; the last two names are also applied to B ...

s, sometimes called ghost sharks, which are sometimes separated into their own class). Extant chondrichthyans range in size from the finless sleeper ray to the over

whale shark The whale shark (''Rhincodon typus'') is a slow-moving, filter feeder, filter-feeding carpet shark and the largest known Extant taxon, extant fish species. The largest confirmed individual had a length of . The whale shark holds many records for ...

Anatomy

Skeleton

The skeleton is cartilaginous. The

notochord The notochord is an elastic, rod-like structure found in chordates. In vertebrates the notochord is an embryonic structure that disintegrates, as the vertebrae develop, to become the nucleus pulposus in the intervertebral discs of the verteb ...

is gradually replaced by a vertebral column during development, except in

, where the notochord stays intact. In some deepwater sharks, the column is reduced. As they do not have

bone marrow Bone marrow is a semi-solid biological tissue, tissue found within the Spongy bone, spongy (also known as cancellous) portions of bones. In birds and mammals, bone marrow is the primary site of new blood cell production (or haematopoiesis). It i ...

red blood cell Red blood cells (RBCs), referred to as erythrocytes (, with -''cyte'' translated as 'cell' in modern usage) in academia and medical publishing, also known as red cells, erythroid cells, and rarely haematids, are the most common type of blood cel ...

s are produced in the

spleen The spleen (, from Ancient Greek '' σπλήν'', splḗn) is an organ (biology), organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The spleen plays important roles in reg ...

and the epigonal organ (special tissue around the

gonad A gonad, sex gland, or reproductive gland is a Heterocrine gland, mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gon ...

s, which is also thought to play a role in the immune system). They are also produced in the Leydig's organ, which is only found in certain cartilaginous fishes. The subclass

, which is a very specialized group, lacks both the Leydig's and epigonal organs.

Appendages

Apart from

electric ray The electric rays are a group of rays, flattened cartilaginous fish with enlarged pectoral fins, composing the order Torpediniformes . They are known for being capable of producing an electric discharge, ranging from 8 to 220 volts, depending ...

s, which have a thick and flabby body, with soft, loose skin, chondrichthyans have tough skin covered with dermal teeth (again, Holocephali is an exception, as the teeth are lost in adults, only kept on the clasping organ seen on the caudal ventral surface of the male), also called

s (or ''dermal denticles''), making it feel like sandpaper. In most species, all dermal denticles are oriented in one direction, making the skin feel very smooth if rubbed in one direction and very rough if rubbed in the other. Originally, the pectoral and pelvic girdles, which do not contain any dermal elements, did not connect. In later forms, each pair of fins became ventrally connected in the middle when scapulocoracoid and puboischiadic bars evolved. In rays, the pectoral fins are connected to the head and are very flexible. One of the primary characteristics present in most sharks is the heterocercal tail, which aids in locomotion.

Body covering

Chondrichthyans have tooth-like scales called

dermal denticle A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scale (zoology), scales, which can also provide effective Underwater camouflage, camouflage through the us ...

s or placoid scales. Denticles usually provide protection, and in most cases, streamlining. Mucous glands exist in some species, as well. It is assumed that their oral teeth evolved from dermal denticles that migrated into the mouth, but it could be the other way around, as the

teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...

bony fish '' Denticeps clupeoides'' has most of its head covered by dermal teeth (as does, probably, '' Atherion elymus'', another bony fish). This is most likely a secondary evolved characteristic, which means there is not necessarily a connection between the teeth and the original dermal scales. The old

placoderms Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos plate' and δέρμα 'dermaskin') are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Dev ...

did not have teeth at all, but had sharp bony plates in their mouth. Thus, it is unknown whether the dermal or oral teeth evolved first. It has even been suggested that the original bony plates of ''all'' vertebrates are now gone and that the present scales are just modified teeth, even if both the teeth and body armor had a common origin a long time ago. However, there is currently no evidence of this.

Respiratory system

All chondrichthyans breathe through five to seven pairs of

gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...

s, depending on the species. In general, pelagic species must keep swimming to keep oxygenated water moving through their gills, whilst demersal species can actively pump water in through their spiracles and out through their gills. However, this is only a general rule and many species differ. A spiracle is a small hole found behind each eye. These can be tiny and circular, such as found on the nurse shark (''Ginglymostoma cirratum''), to extended and slit-like, such as found on the wobbegongs (Orectolobidae). Many larger, pelagic species, such as the mackerel sharks (Lamnidae) and the thresher sharks (Alopiidae), no longer possess them.

Nervous system

In chondrichthyans, the nervous system is composed of a small brain, 8–10 pairs of cranial nerves, and a spinal cord with spinal nerves. They have several sensory organs which provide information to be processed.

Ampullae of Lorenzini Ampullae of Lorenzini (: ''ampulla'') are electroreceptors, sense organs able to detect electric fields. They form a network of mucus-filled pores in the skin of Chondrichthyes, cartilaginous fish (sharks, Ray (fish), rays, and chimaeras) and of ...

are a network of small jelly filled pores called electroreceptors which help the fish sense electric fields in water. This aids in finding prey, navigation, and sensing temperature. The

lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelia ...

system has modified epithelial cells located externally which sense motion, vibration, and pressure in the water around them. Most species have large well-developed eyes. Also, they have very powerful nostrils and

olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...

organs. Their inner ears consist of 3 large

semicircular canals The semicircular canals are three semicircular interconnected tubes located in the innermost part of each ear, the inner ear. The three canals are the lateral, anterior and posterior semicircular canals. They are the part of the bony labyrinth, ...

which aid in balance and orientation. Their sound detecting apparatus has limited range and is typically more powerful at lower frequencies. Some species have

electric organs Electricity is the set of physical phenomena associated with the presence and motion of matter possessing an electric charge. Electricity is related to magnetism, both being part of the phenomenon of electromagnetism, as described by Maxwel ...

which can be used for defense and predation. They have relatively simple brains with the forebrain not greatly enlarged. The structure and formation of myelin in their nervous systems are nearly identical to that of tetrapods, which has led evolutionary biologists to believe that Chondrichthyes were a cornerstone group in the evolutionary timeline of myelin development.

Immune system

Like all other jawed vertebrates, members of Chondrichthyes have an

adaptive immune system The adaptive immune system (AIS), also known as the acquired immune system, or specific immune system is a subsystem of the immune system that is composed of specialized cells, organs, and processes that eliminate pathogens specifically. The ac ...

Reproduction

Fertilization is internal. Development is usually live birth (

ovoviviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a "bridging" form of reproduction between egg-laying oviparity, oviparous and live-bearing viviparity, viviparous reproduction. Ovoviviparous animals possess embryos that develo ...

species) but can be through eggs (

oviparous Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...

). Some rare species are

viviparous In animals, viviparity is development of the embryo inside the body of the mother, with the maternal circulation providing for the metabolic needs of the embryo's development, until the mother gives birth to a fully or partially developed juve ...

. There is no parental care after birth; however, some chondrichthyans do guard their eggs. Capture-induced premature birth and abortion (collectively called capture-induced parturition) occurs frequently in sharks/rays when fished. Capture-induced parturition is often mistaken for natural birth by recreational fishers and is rarely considered in commercial fisheries management despite being shown to occur in at least 12% of live bearing sharks and rays (88 species to date).

Classification

The class Chondrichthyes has two subclasses: the subclass

(

s, rays, skates, and sawfish) and the subclass

(

s). To see the full list of the species, click

here Here may refer to: Music * ''Here'' (Adrian Belew album), 1994 * ''Here'' (Alicia Keys album), 2016 * ''Here'' (Cal Tjader album), 1979 * ''Here'' (Edward Sharpe album), 2012 * ''Here'' (Idina Menzel album), 2004 * ''Here'' (Merzbow album), ...

Evolution

Cartilaginous fish are considered to have evolved from acanthodians. The discovery of ''

Entelognathus ''Entelognathus primordialis'' (“primordial complete jaw”) is an early placoderm from the late Silurian (Ludlow epoch) of Qujing, Yunnan, 419 million years ago. A team led by Min Zhu of the Academy of Sciences' Institute of Vertebrate Paleon ...

'' and several examinations of acanthodian characteristics indicate that bony fish evolved directly from placoderm like ancestors, while acanthodians represent a paraphyletic assemblage leading to Chondrichthyes. Some characteristics previously thought to be exclusive to acanthodians are also present in basal cartilaginous fish. In particular, new phylogenetic studies find cartilaginous fish to be well nested among acanthodians, with ''

Doliodus ''Doliodus'' is a genus of cartilaginous fish known from the Early Devonian period of what is now New Brunswick, Canada. It includes a single named species, either ''D. problematicus'' or ''D. latispinosus'', which was originally described based o ...

'' and '' Tamiobatis'' being the closest relatives to Chondrichthyes. Recent studies vindicate this, as ''

'' had a mosaic of chondrichthyan and acanthodian traits. Dating back to the Middle and Late

Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...

Period, many isolated scales, made of

dentine Dentin ( ) (American English) or dentine ( or ) (British English) () is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by enamel on the crown ...

and bone, have a structure and growth form that is chondrichthyan-like. They may be the remains of

stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...

-chondrichthyans, but their classification remains uncertain. The earliest unequivocal fossils of acanthodian-grade cartilaginous fishes are '' Qianodus'' and '' Fanjingshania'' from the early Silurian (

Aeronian In the geologic timescale, the Aeronian is an geologic age, age of the Llandovery Epoch of the Silurian geologic period, Period of the Paleozoic geologic era, Era of the Phanerozoic geologic eon, Eon that began 440.8 ± 1.2 annum, Ma and ended 43 ...

) of

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, China around 439 million years ago, which are also the oldest unambiguous remains of any jawed vertebrates. ''Shenacanthus vermiformis'', which lived 436 million years ago, had thoracic armour plates resembling those of placoderms. By the start of the Early Devonian, 419 million years ago,

es had divided into three distinct groups: the now extinct

placoderm Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...

s (a paraphyletic assemblage of ancient armoured fishes), the

bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...

es, and the clade that includes spiny sharks and early

cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...

. The modern bony fishes, class

, appeared in the late

Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...

or early Devonian, about 416 million years ago. The first abundant genus of shark, ''

Cladoselache ''Cladoselache'' ("branch shark") is an Extinction, extinct genus of shark-like chondrichthyan (cartilaginous fish) from the Late Devonian (Famennian) of North America. It was similar in body shape to modern Lamnidae, lamnid sharks (such as Isuru ...

'', appeared in the oceans during the Devonian Period. The first cartilaginous fishes evolved from ''

''-like spiny shark ancestors.

Taxonomy

Subphylum

Vertebrata Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Verte ...

└─Infraphylum Gnathostomata ├─

Placodermi Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos plate' and δέρμα 'dermaskin') are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the De ...

— ''extinct'' (armored gnathostomes) └

Eugnathostomata Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant vertebrates, including all living bony fishes (both ray-finned ...

(true jawed vertebrates) ├─

Acanthodii Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes). They are currently considered to represent a paraphyletic grade of various fish lineages basal to extant Chondrichthyes, which includes living sharks, rays, and ...

(stem cartilaginous fish) └─Chondrichthyes (true cartilaginous fish) ├─

(chimaeras + several extinct clades) └

(shark and rays) ├─ Selachii (true sharks) └─

Batoidea Batomorphi is a Division (taxonomy), division of Chondrichthyes, cartilaginous fishes, commonly known as rays, this taxon is also known as the Order (biology), superorder Batoidea, but the 5th edition of ''Fishes of the World'' classifies it as ...

(rays and relatives) *Note: Lines show evolutionary relationships.