Sharks

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 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 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 numerous sets of replaceable teeth.

Well-known species such as the tiger shark, blue shark, great white shark, mako shark, thresher shark, and hammerhead shark are apex predators—organisms at the top of their underwater food chain. Many shark populations are threatened by human activities. Since 1970, shark populations have been reduced by 71%, mostly from overfishing.

Contents

 * 1Etymology
 * 2Evolutionary history
 * 2.1Taxonomy
 * 3Anatomy
 * 3.1Teeth
 * 3.2Skeleton
 * 3.3Jaw
 * 3.4Fins
 * 3.5Dermal denticles
 * 3.6Tails
 * 4Physiology
 * 4.1Buoyancy
 * 4.2Respiration
 * 4.3Thermoregulation
 * 4.4Osmoregulation
 * 4.5Digestion
 * 4.6Fluoresence
 * 5Senses
 * 5.1Smell
 * 5.2Sight
 * 5.3Hearing
 * 5.4Electroreception
 * 5.5Lateral line
 * 6Life history
 * 6.1Reproduction
 * 6.1.1Sexual
 * 6.1.2Asexual
 * 6.2Brooding
 * 6.2.1Ovoviviparity
 * 6.2.2Oviparity
 * 6.2.3Viviparity
 * 7Behavior
 * 7.1Speed
 * 7.2Intelligence
 * 7.3Sleep
 * 8Ecology
 * 8.1Feeding
 * 8.2Range and habitat
 * 9Relationship with humans
 * 9.1Attacks
 * 9.2In captivity
 * 9.3In culture
 * 9.3.1In Hawaii
 * 9.3.2In American Samoa
 * 9.3.3In popular culture
 * 9.3.4Popular misconceptions
 * 10Threats to sharks
 * 10.1Fishery
 * 10.2Shark culling
 * 10.3Other threats
 * 11Conservation
 * 12See also
 * 13References
 * 14Further reading

Etymology
Until the 16th century, sharks were known to mariners as "sea dogs". This is still evidential in several species termed "dogfish," or the porbeagle.

The etymology of the word "shark" is uncertain, the most likely 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.

A now disproven[original research?] theory is that it derives from the Yucatec Maya word xok (pronounced 'shok'), meaning "fish". Evidence for this etymology came 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.[original research?]

Evolutionary history
A collection of Cretaceous shark teeth See also: Evolution of fish

Evidence for the existence of sharks dates from the Ordovician period, 450–420 million years ago, before land vertebrates existed and before a variety of 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. At this time the most common shark tooth is the cladodont, a style of thin tooth with three tines like a trident, apparently to help catch fish. 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.

Among the most ancient and primitive sharks is Cladoselache, from about 370 million years ago, which has been found within Paleozoic strata in Ohio, Kentucky, and Tennessee. At that point in Earth's history these rocks made up the soft bottom sediments of a large, shallow ocean, which stretched across much of North America. Cladoselache was only about 1 metre (3.3 ft) long with stiff triangular fins and slender jaws. Its teeth had several pointed cusps, which wore down from use. From the small number of teeth found together, it is most likely that Cladoselache did not replace its teeth as regularly as modern sharks. Its caudal fins had a similar shape to the great white sharks and the pelagic shortfin and longfin makos. The presence of whole fish arranged tail-first in their stomachs suggest that they were fast swimmers with great agility.

Most fossil sharks from about 300 to 150 million years ago can be assigned to one of two groups. The Xenacanthida was almost exclusive to freshwater environments. By the time this group became extinct about 220 million years ago, they had spread worldwide. The other group, the hybodonts, appeared about 320 million years ago and lived mostly in the oceans, but also in freshwater.[citation needed] The results of a 2014 study of the gill structure of an unusually well preserved 325-million-year-old fossil suggested that sharks are not "living fossils", but rather have evolved more extensively than previously thought over the hundreds of millions of years they have been around. Megalodon (top two, estimated maximum and conservative sizes) with the whale shark, great white shark, and a human for scale Modern sharks began to appear about 100 million years ago. Fossil mackerel shark teeth date to the Early Cretaceous. One of the most recently evolved families is the hammerhead shark (family Sphyrnidae), which emerged in the Eocene. The oldest white shark teeth date from 60 to 66 million years ago, around the time of the extinction of the dinosaurs. In early white shark evolution there are at least two lineages: one lineage is of white sharks with coarsely serrated teeth and it probably gave rise to the modern great white shark, and another lineage is of white sharks with finely serrated teeth. These sharks attained gigantic proportions and include the extinct megatoothed shark, C. megalodon. Like most extinct sharks, C. megalodon is also primarily known from its fossil teeth and vertebrae. This giant shark reached a total length (TL) of more than 16 metres (52 ft). C. megalodon may have approached a maxima of 20.3 metres (67 ft) in total length and 103 metric tons (114 short tons) in mass. Paleontological evidence suggests that this shark was an active predator of large cetaceans.

Taxonomy
Sharks belong to the superorder Selachimorpha in the subclass Elasmobranchii in the class Chondrichthyes. The Elasmobranchii also include rays and skates; the Chondrichthyes also include Chimaeras. It was thought that the sharks form a polyphyletic group: some sharks are more closely related to rays than they are to some other sharks, but current molecular studies support monophyly of both groups of sharks and batoids.

The superorder Selachimorpha is divided into Galea (or Galeomorphii), and Squalea (or Squalomorphii). The Galeans are the Heterodontiformes, Orectolobiformes, Lamniformes, and Carcharhiniformes. Lamnoids and Carcharhinoids are usually placed in one clade, but recent studies show the Lamnoids and Orectoloboids are a clade. Some scientists now think that Heterodontoids may be Squalean. The Squaleans are divided into Hexanchiformes and Squalomorpha. The former includes cow shark and frilled shark, though some authors propose both families to be moved to separate orders. The Squalomorpha contains the Squaliformes and the Hypnosqualea. The Hypnosqualea may be invalid. It includes the Squatiniformes, and the Pristorajea, which may also be invalid, but includes the Pristiophoriformes and the Batoidea.

There are more than 470 species of sharks split across twelve orders, including four orders of sharks that have gone extinct:


 * Carcharhiniformes: Commonly known as ground sharks, the order includes the blue, tiger, bull, grey reef, blacktip reef, Caribbean reef, blacktail reef, whitetip reef, and oceanic whitetip sharks (collectively called the requiem sharks) along with the houndsharks, catsharks, and hammerhead sharks. They are distinguished by an elongated snout and a nictitating membrane which protects the eyes during an attack.
 * Heterodontiformes: They are generally referred to as the bullhead or horn sharks.
 * Hexanchiformes: Examples from this group include the cow sharks and frilled sharks, which somewhat resembles a marine snake.
 * Lamniformes: They are commonly known as the mackerel sharks. They include the goblin shark, basking shark, megamouth shark, the thresher sharks, shortfin and longfin mako sharks, and great white shark. They are distinguished by their large jaws and ovoviviparous reproduction. The Lamniformes also include the extinct megalodon, Carcharodon megalodon.
 * Orectolobiformes: They are commonly referred to as the carpet sharks, including zebra sharks, nurse sharks, wobbegongs, and the whale shark.
 * Pristiophoriformes: These are the sawsharks, with an elongated, toothed snout that they use for slashing their prey.
 * Squaliformes: This group includes the dogfish sharks and roughsharks.
 * Squatiniformes: Also known as angel sharks, they are flattened sharks with a strong resemblance to stingrays and skates.
 * † Cladoselachiformes
 * † Hybodontiformes
 * † Symmoriida
 * † Xenacanthida (Xenacantiformes)

Anatomy
General anatomical features of sharks Main article: Shark anatomy

Teeth
Main article: Shark tooth The teeth of tiger sharks are oblique and serrated to saw through flesh 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.

Skeleton
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.

Jaw
The 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.

Fins
Fin skeletons are elongated and supported with soft and unsegmented rays named ceratotrichia, filaments of elastic protein resembling the horny keratin in hair and feathers. Most sharks have eight fins. Sharks can only drift away from objects directly in front of them because their fins do not allow them to move in the tail-first direction.

Dermal denticles
Further information: Fish scale § Placoid scales The dermal denticles of a lemon shark, viewed through a scanning electron microscope Unlike bony fish, sharks have a complex dermal corset made of flexible collagenous fibers and arranged as a helical network surrounding their body. This works as an outer skeleton, providing attachment for their swimming muscles and thus saving energy. Their dermal teeth give them hydrodynamic advantages as they reduce turbulence when swimming.

Tails
Tails provide thrust, making speed and acceleration dependent on tail shape. Caudal fin shapes vary considerably between shark species, due to their evolution in separate environments. Sharks possess a heterocercal caudal fin in which the dorsal portion is usually noticeably larger than the ventral portion. This is because the shark's vertebral column extends into that dorsal portion, providing a greater surface area for muscle attachment. This allows more efficient locomotion among these negatively buoyant cartilaginous fish. By contrast, most bony fish possess a homocercal caudal fin.

Tiger sharks have a large upper lobe, which allows for slow cruising and sudden bursts of speed. The tiger shark must be able to twist and turn in the water easily when hunting to support its varied diet, whereas the porbeagle shark, which hunts schooling fish such as mackerel and herring, has a large lower lobe to help it keep pace with its fast-swimming prey. Other tail adaptations help sharks catch prey more directly, such as the thresher shark's usage of its powerful, elongated upper lobe to stun fish and squid.

Buoyancy
Unlike bony fish, sharks do not have gas-filled swim bladders for buoyancy. Instead, sharks rely on a large liver filled with oil that contains squalene, and their cartilage, which is about half the normal density of bone. Their liver constitutes up to 30% of their total body mass. The liver's effectiveness is limited, so sharks employ dynamic lift to maintain depth while swimming. Sand tiger sharks store air in their stomachs, using it as a form of swim bladder. Bottom-dwelling sharks, like the nurse shark, have negative buoyancy, allowing them to rest on the ocean floor.

Some sharks, if inverted or stroked on the nose, enter a natural state of tonic immobility. Researchers use this condition to handle sharks safely.