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 andXenacanthus. Under this broader definition, the earliest known sharks date from more than 420 million years ago.[1]
Since then, sharks have diversified into over 470 species. They range in size from the small dwarf lanternshark (Etmopterus perryi), a deep seaspecies 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 (39 ft). Sharks are found in all seas and are common to depths of 2,000 metres (6,600 ft). They generally do not live in freshwateralthough there are a few known exceptions, such as the bull shark and the river shark, which can survive in both seawater and freshwater.[2] They breathe through five to seven gill slits. 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.[3]
Well-known species such as the great white shark, tiger shark, blue shark, mako shark, and the hammerhead shark are apex predators—organisms at the top of their underwater food chain.
Many shark populations are threatened by human activities.
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.[21] Because sharks do not have rib cages, they can easily be crushed under their own weight on land.[22]
Jaw
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.[23] 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.[21] 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.[24] 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.[22]
Dermal denticles
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.[25] Their dermal teeth give them hydrodynamic advantages as they reduce turbulence when swimming.[19]
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 ahomocercal caudal fin.[26]
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.[27] 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.
Physiology
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.[25] Their liver constitutes up to 30% of their total body mass.[28] The liver's effectiveness is limited, so sharks employ dynamic lift to maintain depth when not swimming.Sand tiger sharks store air in their stomachs, using it as a form of swim bladder. Most sharks need to constantly swim in order to breathe and cannot sleep very long without sinking (if at all). However, certain species, like the nurse shark, are capable of pumping water across their gills, allowing them to rest on the ocean bottom.[29]
Some sharks, if inverted or stroked on the nose, enter a natural state of tonic immobility. Researchers use this condition to handle sharks safely.[30]
Respiration
Like other fish, sharks extract oxygen from seawater as it passes over their gills. Unlike other fish, shark gill slits are not covered, but lie in a row behind the head. A modified slit called a spiraclelies just behind the eye, which assists the shark with taking in water during respiration and plays a major role in bottom–dwelling sharks. Spiracles are reduced or missing in active pelagicsharks.[20] While the shark is moving, water passes through the mouth and over the gills in a process known as "ram ventilation". While at rest, most sharks pump water over their gills to ensure a constant supply of oxygenated water. A small number of species have lost the ability to pump water through their gills and must swim without rest. These species are obligate ram ventilatorsand would presumably asphyxiate if unable to move. Obligate ram ventilation is also true of some pelagic bony fish species.[31]
The respiration and circulation process begins when deoxygenated blood travels to the shark's two-chambered heart. Here the shark pumps blood to its gills via the ventral aorta artery where it branches into afferent brachial arteries. Reoxygenation takes place in the gills and the reoxygenated blood flows into the efferent brachial arteries, which come together to form the dorsal aorta. The blood flows from the dorsal aorta throughout the body. The deoxygenated blood from the body then flows through the posterior cardinal veins and enters the posterior cardinal sinuses. From there blood enters the heart ventricle and the cycle repeats.[32]
Thermoregulation
Most sharks are "cold-blooded" or, more precisely, poikilothermic, meaning that their internal body temperature matches that of their ambient environment. Members of the family Lamnidae (such as the shortfin mako shark and the great white shark) are homeothermic and maintain a higher body temperature than the surrounding water. In these sharks, a strip of aerobic red muscle located near the center of the body generates the heat, which the body retains via a countercurrent exchange mechanism by a system of blood vessels called the rete mirabile ("miraculous net"). Thecommon thresher shark has a similar mechanism for maintaining an elevated body temperature, which is thought to have evolved independently[not in citation given].[33]
Osmoregulation
In contrast to bony fish, with the exception of the coelacanth,[34] the blood and other tissue of sharks and Chondrichthyes is generally isotonic to their marine environments because of the high concentration of urea and trimethylamine N-oxide (TMAO), allowing them to be in osmotic balance with the seawater. This adaptation prevents most sharks from surviving in freshwater, and they are therefore confined to marine environments. A few exceptions exist, such as the bull shark, which has developed a way to change its kidney function to excrete large amounts of urea.[28] When a shark dies, the urea is broken down to ammonia by bacteria, causing the dead body to gradually smell strongly of ammonia.[35][36]
Digestion
Digestion can take a long time. The food moves from the mouth to a J-shaped stomach, where it is stored and initial digestion occurs.[37] Unwanted items may never get past the stomach, and instead the shark either vomits or turns its stomachs inside out and ejects unwanted items from its mouth.
One of the biggest differences between the digestive systems of sharks and mammals is that sharks have much shorter intestines. This short length is achieved by the spiral valve with multiple turns within a single short section instead of a long tube-like intestine. The valve provides a long surface area, requiring food to circulate inside the short gut until fully digested, when remaining waste products pass into the cloaca.[37]

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