Sharks are a group of 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 Selachimorpha (or Selachii) and are the sister group to the rays. However, the term “shark” has also been used for extinct members of the subclass Elasmobranchii outside the Selachimorpha, such as Cladoselache and Xenacanthus, as well as other Chondrichthyes such as the holocephalid eugenedontidans. Under this broader definition, the earliest known sharks date back to more than 420 million years ago. Acanthodians are often referred to as “spiny sharks”; though they are not part of Chondrichthyes proper, they are a paraphyletic assemblage leading to cartilaginous fish as a whole.
Since then, sharks have diversified into over 500 species. They range in size from the small dwarf lanternshark (Etmopterus perryi), a deep sea species of only 17 centimetres (6.7 in) in length, to the whale shark (Rhincodon typus), the largest fish in the world, which reaches approximately 12 metres (40 ft) in length. Sharks are found in all seas and are common to depths of 2,000 metres (6,600 ft). They generally do not live in freshwater although there are a few known exceptions, such as the bull shark and the river shark, which can survive and be found in both seawater and freshwater. Sharks have a covering of dermal denticles that protects their skin from damage and parasites in addition to improving their fluid dynamics. They have several sets of replaceable teeth.
Until the 16th century, sharks were known to mariners as “sea dogs”. The etymology of the word “shark” is uncertain. One theory is that it derives from the Yucatec Maya word xok, pronounced ‘shok’. Evidence for this etymology comes from the Oxford English Dictionary, which notes shark first came into use after Sir John Hawkins’ sailors exhibited one in London in 1569 and posted “sharke” to refer to the large sharks of the Caribbean Sea. However, the Middle English Dictionary records an isolated occurrence of the word shark (referring to a sea fish) in a letter written by Thomas Beckington in 1442, which rules out a New World etymology. An alternate etymology states that the original sense of the word was that of “predator, one who preys on others” from the Dutch schurk, meaning “villain, scoundrel” (cf. card shark, loan shark, etc.), which was later applied to the fish due to its predatory behaviour.
Evidence for the existence of sharks dates from the Ordovician period, 450–420 million years ago, before land vertebrates existed and before many plants had colonized the continents. Only scales have been recovered from the first sharks and not all paleontologists agree that these are from true sharks, suspecting that these scales are actually those of thelodont agnathans. The oldest generally accepted shark scales are from about 420 million years ago, in the Silurian period. The first sharks looked very different from modern sharks. The majority of modern sharks can be traced back to around 100 million years ago. Most fossils are of teeth, often in large numbers. Partial skeletons and even complete fossilized remains have been discovered. Estimates suggest that sharks grow tens of thousands of teeth over a lifetime, which explains the abundant fossils. The teeth consist of easily fossilized calcium phosphate, an apatite. When a shark dies, the decomposing skeleton breaks up, scattering the apatite prisms. Preservation requires rapid burial in bottom sediments.
Shark teeth are embedded in the gums rather than directly affixed to the jaw, and are constantly replaced throughout life. Multiple rows of replacement teeth grow in a groove on the inside of the jaw and steadily move forward in comparison to a conveyor belt; some sharks lose 30,000 or more teeth in their lifetime. The rate of tooth replacement varies from once every 8 to 10 days to several months. In most species, teeth are replaced one at a time as opposed to the simultaneous replacement of an entire row, which is observed in the cookiecutter shark. Tooth shape depends on the shark’s diet: those that feed on mollusks and crustaceans have dense and flattened teeth used for crushing, those that feed on fish have needle-like teeth for gripping, and those that feed on larger prey such as mammals have pointed lower teeth for gripping and triangular upper teeth with serrated edges for cutting. The teeth of plankton-feeders such as the basking shark are small and non-functional.
Shark skeletons are very different from those of bony fish and terrestrial vertebrates. Sharks and other cartilaginous fish (skates and rays) have skeletons made of cartilage and connective tissue. Cartilage is flexible and durable, yet is about half the normal density of bone. This reduces the skeleton’s weight, saving energy. Because sharks do not have rib cages, they can easily be crushed under their own weight on land.
Jaws of sharks, like those of rays and skates, are not attached to the cranium. The jaw’s surface (in comparison to the shark’s vertebrae and gill arches) needs extra support due to its heavy exposure to physical stress and its need for strength. It has a layer of tiny hexagonal plates called “tesserae”, which are crystal blocks of calcium salts arranged as a mosaic. This gives these areas much of the same strength found in the bony tissue found in other animals. Generally sharks have only one layer of tesserae, but the jaws of large specimens, such as the bull shark, tiger shark, and the great white shark, have two to three layers or more, depending on body size. The jaws of a large great white shark may have up to five layers. In the rostrum (snout), the cartilage can be spongy and flexible to absorb the power of impacts.