Shark

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From Wikipedia, the free encyclopedia

For other uses, see Shark (disambiguation).

Sharks

Temporal range: Early Jurassic–

Present

PreꞒ

Pg

Possible records extend back to

Early Permian
 A grey reef shark
(Carcharhinus amblyrhynchos)

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Chordata

Class: Chondrichthyes

Subclass: Elasmobranchii

Infraclass: Euselachii

Clade: Neoselachii

Subdivision: Selachimorpha
Nelson, 1984

Orders

 Galeomorphii
 Carcha
rhinifo
rmes
 Hetero
dontifo
 rmes
 Orecto
lobifor
mes
 Lamni
formes
 Squalomorphii
 Hexan
chifor
mes
 Pristio
phorif
ormes
 Squalif
ormes
 Squati
niform
es
 ?
†Synechodontifor
mes

Synonyms
Pleurotremata
Selachii

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[1] (or Selachii) and
are the sister group to the Batoidea (rays and kin). Some sources extend the term
"shark" as an informal category including extinct members
of Chondrichthyes (cartilaginous fish) with a shark-like morphology, such as hybodonts.
Shark-like chondrichthyans such as Cladoselache and Doliodus first appeared in
the Devonian Period (419–359 million years), though some fossilized chondrichthyan-
like scales are as old as the Late Ordovician (458–444 million years ago).[2] The earliest
confirmed modern sharks (selachimorphs) are known from the Early
Jurassic around 200 million years ago, with the oldest known member being Agaleus,
though records of true sharks may extend back as far as the Permian.[3]
Sharks range in size from the small dwarf lanternshark (Etmopterus perryi), a deep sea
species that is 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.[4] They are found in all seas and are common to depths up to 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 sharks, which can be found in both
seawater and freshwater, and the Ganges shark, which lives only in freshwater.
[5]
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.[6]

Several species are apex predators, which are organisms that are at the top of
their food chain. Select examples include the bull shark, tiger shark, great white
shark, mako sharks, thresher sharks, and hammerhead sharks.

Sharks are caught by humans for shark meat or shark fin soup. Many shark populations
are threatened by human activities. Since 1970, shark populations have been reduced
by 71%, mostly from overfishing.[7]

Etymology
Until the 16th century,[8] sharks were known to mariners as "sea dogs".[9] 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.[10]

A now disproven[original research?] theory is that it derives from the Yucatec

Maya word xook (pronounced [ʃoːk]), meaning 'shark'.[11] Evidence for this etymology came
from the Oxford English Dictionary, which notes that 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.[12]
[original research?]

Evolutionary history
See also: Evolution of fish
Fossil record
 Fossil shark tooth (size over 9 cm or 3.5 inches) with

crown, shoulder, root and root lobe A collection

of Cretaceous shark teeth
The oldest total-group chondrichthyans, known as acanthodians or "spiny sharks",
appeared during the Early Silurian, around 439 million years ago.[13] The oldest
confirmed members of Elasmobranchii sensu lato (the group containing all cartilaginous
fish more closely related to modern sharks and rays than to chimaeras) appeared
during the Devonian.[14] Anachronistidae, the oldest probable representatives of
Neoselachii, the group containing modern sharks (Selachimorpha) and rays (Batoidea)
to the exclusion of most extinct elasmobranch groups, date to the Carboniferous.
[15]
Selachiimorpha and Batoidea are suggested by some to have diverged during
the Triassic.[16] Fossils of the earliest true sharks may have appeared during
the Permian, based on remains of "synechodontiforms" found in the Early Permian of
Russia,[17] but if remains of "synechodontiformes" from the Permian and Triassic are true
sharks, they only had low diversity. Modern shark orders first appeared during the Early
Jurassic, and during the Jurassic true sharks underwent great diversification.
[18]
Selachimorphs largely replaced the hybodonts, which had previously been a
dominant group of shark-like fish during the Triassic and Early Jurassic.[19]

Taxonomy
Elasm
obran
chii Batoidea

Selachimorp Galeomorp
ha hii
 Phylogeny of living shark orders based
on mitochondrial DNA[20]

Shark fossil, Lebachacanthus senckenbergianus,

at Permian period
Sharks belong to the clade 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,[21] but current molecular studies support monophyly of both groups of
sharks and batoids.[22][23]

The clade Selachimorpha is divided into the superorders 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
that 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 that both families 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.[21][24]

There are more than 500 species of sharks split across thirteen orders, including
several orders of sharks that have gone extinct:[24][25]

 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, Otodus 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.
 Echinorhiniformes: This group includes the prickly shark and bramble shark.
Phylogenetic placement of this group has been ambiguous in scientific
studies.[26] They are sometimes given their own order, Echinorhiniformes.[25]
Anatomy
Main article: Shark anatomy
 General anatomical features of sharks
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.[27]

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.[28]

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.[29] Because
sharks do not have rib cages, they can easily be crushed under their own weight on
land.[30]

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.[31] 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.[29] In the rostrum (snout), the cartilage can be spongy and flexible to absorb
the power of impacts.[citation needed]

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.[32] 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. [30]

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.
[33]
Their dermal teeth give them hydrodynamic advantages as they reduce turbulence
when swimming.[34] Some species of shark have pigmented denticles that form complex
patterns like spots (e.g. Zebra shark) and stripes (e.g. Tiger shark). These markings are
important for camouflage and help sharks blend in with their environment, as well as
making them difficult for prey to detect.[35] For some species, dermal patterning returns to
healed denticles even after they have been removed by injury.[36]

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.[37]

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.[38] 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.[33] Their liver constitutes up to 30% of
their total body mass.[39] 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.[40]

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 spiracle lies 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 pelagic sharks.[28] 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 ventilators and would
presumably asphyxiate if unable to move. Obligate ram ventilation is also true of some
pelagic bony fish species.[41][42]

The respiratory and circulatory process begins when deoxygenated venous

blood travels to the shark's two-chambered heart. Here, the shark pumps blood to its
gills via the ventral aorta where it branches into afferent branchial arteries. Gas
exchange takes place in the gills and the reoxygenated blood flows into
the efferent branchial 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 venous blood re-enters the heart ventricle and the cycle repeats.[43]

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"). The common
thresher and bigeye thresher sharks have a similar mechanism for maintaining an
elevated body temperature.[44]

Larger species, like the whale shark, are able to conserve their body heat through sheer
size when they dive to colder depths, and the scalloped hammerhead close its mouth
and gills when they dives to depths of around 800 metres, holding its breath till it reach
warmer waters again.[45]

Osmoregulation
In contrast to bony fish, with the exception of the coelacanth,[46] the blood and other
tissue of sharks and Chondrichthyes is generally isotonic to their marine environments
because of the high concentration of urea (up to 2.5%[47]) 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.
[39]
When a shark dies, the urea is broken down to ammonia by bacteria, causing the
dead body to gradually smell strongly of ammonia.[48][49]

Research in 1930 by Homer W. Smith showed that sharks' urine does not contain
sufficient sodium to avoid hypernatremia, and it was postulated that there must be an
additional mechanism for salt secretion. In 1960 it was discovered at the Mount Desert
Island Biological Laboratory in Salsbury Cove, Maine that sharks have a type of salt
gland located at the end of the intestine, known as the "rectal gland", whose function is
the secretion of chlorides.[50]

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.[51] 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.[52]

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.[51]

Fluorescence
A few sharks appear fluorescent under blue light, such as the swell shark and the chain
catshark, where the fluorophore derives from a metabolite of kynurenic acid.[53]

Senses
Smell

The shape of the hammerhead shark's head may

enhance olfaction by spacing the nostrils further apart.
Sharks have keen olfactory senses, located in the short duct (which is not fused, unlike
bony fish) between the anterior and posterior nasal openings, with some species able to
detect as little as one part per million of blood in seawater.[54] The size of the olfactory
bulb varies across different shark species, with size dependent on how much a given
species relies on smell or vision to find their prey.[55] In environments with low visibility,
shark species generally have larger olfactory bulbs.[55] In reefs, where visibility is high,
species of sharks from the family Carcharhinidae have smaller olfactory bulbs.[55] Sharks
found in deeper waters also have larger olfactory bulbs.[56]

Sharks have the ability to determine the direction of a given scent based on the timing
of scent detection in each nostril.[57] This is similar to the method mammals use to
determine direction of sound.

They are more attracted to the chemicals found in the intestines of many species, and
as a result often linger near or in sewage outfalls. Some species, such as nurse sharks,
have external barbels that greatly increase their ability to sense prey.

Sight
 Eye of a bigeyed sixgill shark (Hexanchus nakamurai)
Shark eyes are similar to the eyes of other vertebrates, including
similar lenses, corneas and retinas, though their eyesight is well adapted to
the marine environment with the help of a tissue called tapetum lucidum. This tissue is
behind the retina and reflects light back to it, thereby increasing visibility in the dark
waters. The effectiveness of the tissue varies, with some sharks having
stronger nocturnal adaptations. Many sharks can contract and dilate their pupils, like
humans, something no teleost fish can do. Sharks have eyelids, but they do not blink
because the surrounding water cleans their eyes. To protect their eyes some species
have nictitating membranes. This membrane covers the eyes while hunting and when
the shark is being attacked. However, some species, including the great white
shark (Carcharodon carcharias), do not have this membrane, but instead roll their eyes
backwards to protect them when striking prey. The importance of sight in shark hunting
behavior is debated. Some believe that electro- and chemoreception are more
significant, while others point to the nictating membrane as evidence that sight is
important, since presumably the shark would not protect its eyes were they unimportant.
The use of sight probably varies with species and water conditions. The shark's field of
vision can swap between monocular and stereoscopic at any time.[58] A micro-
spectrophotometry study of 17 species of sharks found 10 had only rod
photoreceptors and no cone cells in their retinas giving them good night vision while
making them colorblind. The remaining seven species had in addition to rods a single
type of cone photoreceptor sensitive to green and, seeing only in shades of grey and
green, are believed to be effectively colorblind. The study indicates that an object's
contrast against the background, rather than colour, may be more important for object
detection.[59] [60][61]

Hearing
Although it is hard to test the hearing of sharks, they may have a sharp sense of
hearing and can possibly hear prey from many miles away.[62] The hearing sensitivity for
most shark species lies between 20 and 1000 Hz.[63] A small opening on each side of
their heads (not the spiracle) leads directly into the inner ear through a thin channel.
The lateral line shows a similar arrangement, and is open to the environment via a
series of openings called lateral line pores. This is a reminder of the common origin of
these two vibration- and sound-detecting organs that are grouped together as the
acoustico-lateralis system. In bony fish and tetrapods the external opening into the inner
ear has been lost.
Electroreception
Main article: Electroreception

Electromagnetic field receptors (ampullae of

Lorenzini) and motion detecting canals in the head of a shark
The ampullae of Lorenzini are the electroreceptor organs. They number in the hundreds
to thousands. Sharks use the ampullae of Lorenzini to detect the electromagnetic
fields that all living things produce.[64] This helps sharks (particularly the hammerhead
shark) find prey. The shark has the greatest electrical sensitivity of any animal. Sharks
find prey hidden in sand by detecting the electric fields they produce. Ocean
currents moving in the magnetic field of the Earth also generate electric fields that
sharks can use for orientation and possibly navigation.[65]

Lateral line
Main article: Lateral line
This system is found in most fish, including sharks. It is a tactile sensory system which
allows the organism to detect water speed and pressure changes near by.[66] The main
component of the system is the neuromast, a cell similar to hair cells present in the
vertebrate ear that interact with the surrounding aquatic environment. This helps sharks
distinguish between the currents around them, obstacles off on their periphery, and
struggling prey out of visual view. The shark can sense frequencies in the range of 25 to
50 Hz.[67]

Life history

The claspers of male spotted wobbegong

 Shark egg

The spiral egg case of a Port Jackson shark

Shark lifespans vary by species. Most live 20 to 30 years. The spiny dogfish has one of
the longest lifespans at more than 100 years.[68] Whale sharks (Rhincodon typus) may
also live over 100 years.[69] Earlier estimates suggested the Greenland
shark (Somniosus microcephalus) could reach about 200 years, but a recent study
found that a 5.02-metre-long (16.5 ft) specimen was 392 ± 120 years old (i.e., at least
272 years old), making it the longest-lived vertebrate known.[70][71]

Reproduction
Unlike most bony fish, sharks are K-selected reproducers, meaning that they produce a
small number of well-developed young as opposed to a large number of poorly
developed young. Fecundity in sharks ranges from 2 to over 100 young per
reproductive cycle.[72] Sharks mature slowly relative to many other fish. For
example, lemon sharks reach sexual maturity at around age 13–15.[73]

Sexual
Sharks practice internal fertilization.[74] The posterior part of a male shark's pelvic fins are
modified into a pair of intromittent organs called claspers, analogous to a mammalian
penis, of which one is used to deliver sperm into the female.[75]

Mating has rarely been observed in sharks.[76] The smaller catsharks often mate with the
male curling around the female. In less flexible species the two sharks swim parallel to
each other while the male inserts a clasper into the female's oviduct. Females in many
of the larger species have bite marks that appear to be a result of a male grasping them
to maintain position during mating. The bite marks may also come from courtship
behavior: the male may bite the female to show his interest. In some species, females
have evolved thicker skin to withstand these bites.[75]

Asexual
There have been a number of documented cases in which a female shark who has not
been in contact with a male has conceived a pup on her own through parthenogenesis.
[77][78]
The details of this process are not well understood, but genetic
fingerprinting showed that the pups had no paternal genetic contribution, ruling
out sperm storage. The extent of this behavior in the wild is unknown. Mammals are
now the only major vertebrate group in which asexual reproduction has not been
observed.

Scientists say that asexual reproduction in the wild is rare, and probably a last-ditch
effort to reproduce when a mate is not present. Asexual reproduction diminishes genetic
diversity, which helps build defenses against threats to the species. Species that rely
solely on it risk extinction. Asexual reproduction may have contributed to the blue
shark's decline off the Irish coast.[79]

Brooding
Sharks display three ways to bear their young, varying by
species, oviparity, viviparity and ovoviviparity.[80][81]

Ovoviviparity
Most sharks are ovoviviparous, meaning that the eggs hatch in the oviduct within the
mother's body and that the egg's yolk and fluids secreted by glands in the walls of the
oviduct nourishes the embryos. The young continue to be nourished by the remnants of
the yolk and the oviduct's fluids. As in viviparity, the young are born alive and fully
functional. Lamniforme sharks practice oophagy, where the first embryos to hatch eat
the remaining eggs. Taking this a step further, sand tiger shark pups cannibalistically
consume neighboring embryos. The survival strategy for ovoviviparous species is
to brood the young to a comparatively large size before birth. The whale shark is now
classified as ovoviviparous rather than oviparous, because extrauterine eggs are now
thought to have been aborted. Most ovoviviparous sharks give birth in sheltered areas,
including bays, river mouths and shallow reefs. They choose such areas for protection
from predators (mainly other sharks) and the abundance of food. Dogfish have the
longest known gestation period of any shark, at 18 to 24 months. Basking
sharks and frilled sharks appear to have even longer gestation periods, but accurate
data are lacking.[80]

Oviparity
Some species are oviparous, laying their fertilized eggs in the water. In most oviparous
shark species, an egg case with the consistency of leather protects the developing
embryo(s). These cases may be corkscrewed into crevices for protection. The egg case
is commonly called a mermaid's purse. Oviparous sharks include the horn
shark, catshark, Port Jackson shark, and swellshark.[80][82]

Viviparity
Viviparity is the gestation of young without the use of a traditional egg, and results in live
birth.[83] Viviparity in sharks can be placental or aplacental.[83] Young are born fully formed
and self-sufficient.[83] Hammerheads, the requiem sharks (such as the bull and blue
sharks), and smoothhounds are viviparous.[72][80]

Behavior
The classic view describes a solitary hunter, ranging the oceans in search of food.
However, this applies to only a few species. Most live far more social,
sedentary, benthic lives, and appear likely to have their own distinct personalities.
[84]
Even solitary sharks meet for breeding or at rich hunting grounds, which may lead
them to cover thousands of miles in a year.[85] Shark migration patterns may be even
more complex than in birds, with many sharks covering entire ocean basins.

Sharks can be highly social, remaining in large schools. Sometimes more than
100 scalloped hammerheads congregate around seamounts and islands, e.g., in
the Gulf of California.[39] Cross-species social hierarchies exist. For example, oceanic
whitetip sharks dominate silky sharks of comparable size during feeding.[72]

When approached too closely some sharks perform a threat display. This usually
consists of exaggerated swimming movements, and can vary in intensity according to
the threat level.[86]

Speed
In general, sharks swim ("cruise") at an average speed of 8 kilometres per hour
(5.0 mph), but when feeding or attacking, the average shark can reach speeds upwards
of 19 kilometres per hour (12 mph). The shortfin mako shark, the fastest shark and one
of the fastest fish, can burst at speeds up to 50 kilometres per hour (31 mph).
[87]
The great white shark is also capable of speed bursts. These exceptions may be due
to the warm-blooded, or homeothermic, nature of these sharks' physiology. Sharks can
travel 70 to 80 km in a day.[88]

Intelligence
Sharks possess brain-to-body mass ratios that are similar to mammals and birds,[89] and
have exhibited apparent curiosity and behavior resembling play in the wild.[90][91]

There is evidence that juvenile lemon sharks can use observational learning in their
investigation of novel objects in their environment.[92]

Sleep
All sharks need to keep water flowing over their gills in order for them to breathe;
however, not all species need to be moving to do this. Those that are able to breathe
while not swimming do so by using their spiracles to force water over their gills, thereby
allowing them to extract oxygen from the water. It has been recorded that their eyes
remain open while in this state and actively follow the movements of divers swimming
around them[93] and as such they are not truly asleep.

Species that do need to swim continuously to breathe go through a process known as

sleep swimming, in which the shark is essentially unconscious. It is known from
experiments conducted on the spiny dogfish that its spinal cord, rather than its brain,
coordinates swimming, so spiny dogfish can continue to swim while sleeping, and this
also may be the case in larger shark species.[93] In 2016 a great white shark was
captured on video for the first time in a state researchers believed was sleep swimming.
[94]

Ecology
Feeding
This section is about shark feeding. For the sport of shark feeding, see Shark baiting.

A great white shark attacking bait at Isla Guadalupe,

Mexico
Most sharks are carnivorous.[95] Basking sharks, whale sharks, and megamouth
sharks have independently evolved different strategies for filter feeding plankton:
basking sharks practice ram feeding, whale sharks use suction to take in plankton and
small fishes, and megamouth sharks make suction feeding more efficient by using
the luminescent tissue inside of their mouths to attract prey in the deep ocean. This type
of feeding requires gill rakers—long, slender filaments that form a very efficient sieve—
analogous to the baleen plates of the great whales. The shark traps the plankton in
these filaments and swallows from time to time in huge mouthfuls. Teeth in these
species are comparatively small because they are not needed for feeding.[95]

Other highly specialized feeders include cookiecutter sharks, which feed on flesh sliced
out of other larger fish and marine mammals. Cookiecutter teeth are enormous
compared to the animal's size. The lower teeth are particularly sharp. Although they
have never been observed feeding, they are believed to latch onto their prey and use
their thick lips to make a seal, twisting their bodies to rip off flesh.[39]

Some seabed–dwelling species are highly effective ambush predators. Angel

sharks and wobbegongs use camouflage to lie in wait and suck prey into their mouths.
[96]
Many benthic sharks feed solely on crustaceans which they crush with their
flat molariform teeth.

Other sharks feed on squid or fish, which they swallow whole. The viper dogfish has
teeth it can point outwards to strike and capture prey that it then swallows intact.
The great white and other large predators either swallow small prey whole or take huge
bites out of large animals. Thresher sharks use their long tails to stun shoaling fishes,
and sawsharks either stir prey from the seabed or slash at swimming prey with their
tooth-studded rostra.

The bonnethead shark is the only known omnivorous species. Its main prey is
crustaceans and mollusks, but it also eats a large amount of seagrass, and is able to
digest and extract nutrients from about 50% of the seagrass it consume.[97]

Many sharks, including the whitetip reef shark are cooperative feeders and hunt in
packs to herd and capture elusive prey. These social sharks are often migratory,
traveling huge distances around ocean basins in large schools. These migrations may
be partly necessary to find new food sources.[98]

Range and habitat

Sharks are found in all seas. They generally do not live in fresh water, with a few
exceptions such as the bull shark and the river shark which can swim both in seawater
and freshwater.[99] Sharks are common down to depths of 2,000 metres (7,000 ft), and
some live even deeper, but they are almost entirely absent below 3,000 metres
(10,000 ft). The deepest confirmed report of a shark is a Portuguese dogfish at 3,700
metres (12,100 ft).[100]

Relationship with humans

Attacks
Main article: Shark attack
 A sign warning about the presence of sharks in Salt

Rock, South Africa Snorkeler swims near a blacktip

reef shark. In rare circumstances involving poor visibility, blacktips may bite a human,
mistaking it for prey. Under normal conditions they are harmless and shy.
In 2006 the International Shark Attack File (ISAF) undertook an investigation into 96
alleged shark attacks, confirming 62 of them as unprovoked attacks and 16 as provoked
attacks. The average number of fatalities worldwide per year between 2001 and 2006
from unprovoked shark attacks is 4.3.[101]

Contrary to popular belief, only a few sharks are dangerous to humans. Out of more
than 470 species, only four have been involved in a significant number of fatal,
unprovoked attacks on humans: the great white, oceanic whitetip, tiger, and bull sharks.
[102][103]
These sharks are large, powerful predators, and may sometimes attack and kill
people. Despite being responsible for attacks on humans they have all been filmed
without using a protective cage.[104]

The perception of sharks as dangerous animals has been popularized by publicity given
to a few isolated unprovoked attacks, such as the Jersey Shore shark attacks of 1916,
and through popular fictional works about shark attacks, such as the Jaws film
series. Jaws author Peter Benchley, as well as Jaws director Steven Spielberg, later
attempted to dispel the image of sharks as man-eating monsters.[105]

To help avoid an unprovoked attack, humans should not wear jewelry or metal that is
shiny and refrain from splashing around too much.[106]

In general, sharks show little pattern of attacking humans specifically, part of the reason
could be that sharks prefer the blood of fish and other common preys.[107] Research
indicates that when humans do become the object of a shark attack, it is possible that
the shark has mistaken the human for species that are its normal prey, such as seals.[108]
[109]
This was further proven in a recent study conducted by researchers at the California
State University's Shark Lab. According to footage caught by the Lab's drones, juveniles
swam right up to humans in the water without any bites incidents. The lab stated that
the results showed that humans and sharks can co-exist in the water.[110]

In captivity
Main article: Sharks in captivity

A whale shark in Georgia Aquarium

Until recently, only a few benthic species of shark, such as hornsharks, leopard
sharks and catsharks, had survived in aquarium conditions for a year or more. This
gave rise to the belief that sharks, as well as being difficult to capture and transport,
were difficult to care for. More knowledge has led to more species (including the
large pelagic sharks) living far longer in captivity, along with safer transportation
techniques that have enabled long-distance transportation.[111] The great white shark had
never been successfully held in captivity for long periods of time until September 2004,
when the Monterey Bay Aquarium successfully kept a young female for 198 days before
releasing her.

Most species are not suitable for home aquaria, and not every species sold by pet
stores are appropriate. Some species can flourish in home saltwater aquaria.
[112]
Uninformed or unscrupulous dealers sometimes sell juvenile sharks like the nurse
shark, which upon reaching adulthood is far too large for typical home aquaria.[112] Public
aquaria generally do not accept donated specimens that have outgrown their housing.
Some owners have been tempted to release them.[112] Species appropriate to home
aquaria represent considerable spatial and financial investments as they generally
approach adult lengths of 3 feet (90 cm) and can live up to 25 years.[112]

In culture

Shark-themed nose art, made popular by the Flying

Tigers (pictured), is commonly seen on military aircraft.
In Hawaii
Sharks figure prominently in Hawaiian mythology. Stories tell of men with shark jaws on
their back who could change between shark and human form. A common theme was
that a shark-man would warn beach-goers of sharks in the waters. The beach-goers
would laugh and ignore the warnings and get eaten by the shark-man who warned
them. Hawaiian mythology also includes many shark gods. Among a fishing people, the
most popular of all aumakua, or deified ancestor guardians, are shark
aumakua. Kamaku describes in detail how to offer a corpse to become a shark. The
body transforms gradually until the kahuna can point the awe-struck family to the
markings on the shark's body that correspond to the clothing in which the beloved's
body had been wrapped. Such a shark aumakua becomes the family pet, receiving
food, and driving fish into the family net and warding off danger. Like all aumakua it had
evil uses such as helping kill enemies. The ruling chiefs typically forbade such sorcery.
Many Native Hawaiian families claim such an aumakua, who is known by name to the
whole community.[113]

Kamohoali'i is the best known and revered of the shark gods, he was the older and
favored brother of Pele,[114] and helped and journeyed with her to Hawaii. He was able to
assume all human and fish forms. A summit cliff on the crater of Kilauea is one of his
most sacred spots. At one point he had a heiau (temple or shrine) dedicated to him on
every piece of land that jutted into the ocean on the island of Molokai. Kamohoali'i was
an ancestral god, not a human who became a shark and banned the eating of humans
after eating one herself.[115][116] In Fijian mythology, Dakuwaqa was a shark god who was
the eater of lost souls.

In American Samoa
On the island of Tutuila in American Samoa (a U.S. territory), there is a location
called Turtle and Shark (Laumei ma Malie) which is important in Samoan culture—the
location is the site of a legend called O Le Tala I Le Laumei Ma Le Malie, in which two
humans are said to have transformed into a turtle and a shark.[117][118][119] According to
the U.S. National Park Service, "Villagers from nearby Vaitogi continue to reenact an
important aspect of the legend at Turtle and Shark by performing a ritual song intended
to summon the legendary animals to the ocean surface, and visitors are frequently
amazed to see one or both of these creatures emerge from the sea in apparent
response to this call."[117]

In popular culture
Main article: Sharks in popular culture
In contrast to the complex portrayals by Hawaiians and other Pacific Islanders, the
European and Western view of sharks has historically been mostly of fear and
malevolence.[120] Sharks are used in popular culture commonly as eating machines,
notably in the Jaws novel and the film of the same name, along with its sequels.
[121]
Sharks are threats in other films such as Deep Blue Sea, The Reef, and others,
although they are sometimes used for comedic effect such as in Finding Nemo and
the Austin Powers series. Sharks tend to be seen quite often in cartoons whenever a
scene involves the ocean. Such examples include the Tom and
Jerry cartoons, Jabberjaw, and other shows produced by Hanna-Barbera. They also are
used commonly as a clichéd means of killing off a character that is held up by a rope or
some similar object as the sharks swim right below them, or the character may be
standing on a plank above shark infested waters.[citation needed]

Popular misconceptions
A popular myth is that sharks are immune to disease and cancer, but this is not
scientifically supported. Sharks have been known to get cancer.[122][123] Both diseases
and parasites affect sharks. The evidence that sharks are at least resistant to cancer
and disease is mostly anecdotal and there have been few, if any, scientific
or statistical studies that show sharks to have heightened immunity to disease.[124] Other
apparently false claims are that fins prevent cancer[125] and treat osteoarthritis.[126] No
scientific proof supports these claims; at least one study has shown shark cartilage of
no value in cancer treatment.[127]

Threats to sharks
Further information: List of threatened sharks and Shark sanctuary

The annual shark catch has increased rapidly over

the last 60 years. The value of shark fins for shark fin
soup has led to an increase in shark catches where usually only the fins are taken,
while the rest of the shark is discarded, typically into the sea; health concerns
about BMAA in the fins now exists regarding consumption of the soup
 A 4.3-metre (14 ft), 540-kilogram (1,200 lb) tiger shark caught
in Kāne'ohe Bay, Oahu in 1966
Fishery
In 2008, it was estimated that nearly 100 million sharks were being killed by people
every year, due to commercial and recreational fishing.[128][129] In 2021, it was estimated
that the population of oceanic sharks and rays had dropped by 71% over the previous
half-century.[7]

Shark finning yields are estimated at 1.44 million metric tons (1.59 million short tons) for
2000, and 1.41 million metric tons (1.55 million short tons) for 2010. Based on an
analysis of average shark weights, this translates into a total annual mortality estimate
of about 100 million sharks in 2000, and about 97 million sharks in 2010, with a total
range of possible values between 63 and 273 million sharks per year.[130][131] Sharks are a
common seafood in many places, including Japan and Australia. In southern Australia,
shark is commonly used in fish and chips,[132] in which fillets are battered and deep-
fried or crumbed and grilled. In fish and chip shops, shark is called flake. In India, small
sharks or baby sharks (called sora in Tamil language, Telugu language) are sold in local
markets. Since the flesh is not developed, cooking the flesh breaks it into powder, which
is then fried in oil and spices (called sora puttu/sora poratu). The soft bones can be
easily chewed, they are considered a delicacy in coastal Tamil
Nadu. Icelanders ferment Greenland sharks to produce a delicacy called hákarl.
[133]
During a four-year period from 1996 to 2000, an estimated 26 to 73 million sharks
were killed and traded annually in commercial markets.[134]

Sharks are often killed for shark fin soup. Fishermen capture live sharks, fin them, and
dump the finless animal back into the water. Shark finning involves removing the fin with
a hot metal blade.[129] The resulting immobile shark soon dies from suffocation or
predators.[135] Shark fin has become a major trade within black markets all over the
world. Fins sell for about $300/lb in 2009.[136] Poachers illegally fin millions each year.
Few governments enforce laws that protect them.[131] In 2010 Hawaii became the first
U.S. state to prohibit the possession, sale, trade or distribution of shark fins. [137] From
1996 to 2000, an estimated 38 million sharks had been killed per year for harvesting
shark fins.[134] It is estimated by TRAFFIC that over 14,000 tonnes of shark fins were
exported into Singapore between 2005–2007 and 2012–2014.[138]

Shark fin soup is a status symbol in Asian countries and is erroneously considered
healthy and full of nutrients. Scientific research has revealed, however, that high
concentrations of BMAA are present in shark fins.[139] Because BMAA is a neurotoxin,
consumption of shark fin soup and cartilage pills, therefore, may pose a health risk.
[140]
BMAA is under study for its pathological role in neurodegenerative diseases such
as ALS, Alzheimer's disease, and Parkinson's disease.

Sharks are also killed for meat. European diners

consume dogfishes, smoothhounds, catsharks, makos, porbeagle and also skates and
rays.[141] However, the U.S. FDA lists sharks as one of four fish (with swordfish, king
mackerel, and tilefish) whose high mercury content is hazardous to children and
pregnant women.

Sharks generally reach sexual maturity only after many years and produce few offspring
in comparison to other harvested fish. Harvesting sharks before they reproduce
severely impacts future populations. Capture induced premature birth and abortion
(collectively called capture-induced parturition) occurs frequently in sharks/rays when
fished.[74] 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).[74]

The majority of shark fisheries have little monitoring or management. The rise in
demand for shark products increases pressure on fisheries.[40] Major declines in shark
stocks have been recorded—some species have been depleted by over 90% over the
past 20–30 years with population declines of 70% not unusual.[142] A study by
the International Union for Conservation of Nature suggests that one quarter of all
known species of sharks and rays are threatened by extinction and 25 species were
classified as critically endangered.[143][144]

Shark culling
Main article: Shark culling
In 2014, a shark cull in Western Australia killed dozens of sharks (mostly tiger sharks)
using drum lines,[145] until it was cancelled after public protests and a decision by the
Western Australia EPA; from 2014 to 2017, there was an "imminent threat" policy in
Western Australia in which sharks that "threatened" humans in the ocean were shot and
killed.[146] This "imminent threat" policy was criticized by senator Rachel Siewart for killing
endangered sharks.[147] The "imminent threat" policy was cancelled in March 2017.[148] In
August 2018, the Western Australia government announced a plan to re-introduce drum
lines (though, this time the drum lines are "SMART" drum lines).[149]

From 1962 to the present,[150] the government of Queensland has targeted and killed
sharks in large numbers by using drum lines, under a "shark control" program—this
program has also inadvertently killed large numbers of other animals such as dolphins;
it has also killed endangered hammerhead sharks.[151][152][153][154] Queensland's drum line
program has been called "outdated, cruel and ineffective".[154] From 2001 to 2018, a total
of 10,480 sharks were killed on lethal drum lines in Queensland, including in the Great
Barrier Reef.[155] From 1962 to 2018, roughly 50,000 sharks were killed by Queensland
authorities.[156]

The government of New South Wales has a program that deliberately kills sharks
using nets.[153][157] The current net program in New South Wales has been described as
being "extremely destructive" to marine life, including sharks.[158] Between 1950 and
2008, 352 tiger sharks and 577 great white sharks were killed in the nets in New South
Wales—also during this period, a total of 15,135 marine animals were killed in the nets,
including dolphins, whales, turtles, dugongs, and critically endangered grey nurse
sharks.[159] There has been a very large decrease in the number of sharks in eastern
Australia, and the shark-killing programs in Queensland and New South Wales are
partly responsible for this decrease.[156]

Kwazulu-Natal, an area of South Africa, has a shark-killing program using nets and
drum lines—these nets and drum lines have killed turtles and dolphins, and have been
criticized for killing wildlife.[160] During a 30-year period, more than 33,000 sharks have
been killed in KwaZulu-Natal's shark-killing program—during the same 30-year period,
2,211 turtles, 8,448 rays, and 2,310 dolphins were killed in KwaZulu-Natal.[160] Authorities
on the French island of Réunion kill about 100 sharks per year.[161]

Killing sharks negatively affects the marine ecosystem.[162][163] Jessica Morris of Humane
Society International calls shark culling a "knee-jerk reaction" and says, "sharks are top
order predators that play an important role in the functioning of marine ecosystems. We
need them for healthy oceans."[164]

George H. Burgess, the former[165] director of the International Shark Attack File,
"describes [shark] culling as a form of revenge, satisfying a public demand for blood and
little else";[166] he also said shark culling is a "retro-type move reminiscent of what people
would have done in the 1940s and 50s, back when we didn't have an ecological
conscience and before we knew the consequences of our actions."[166] Jane Williamson,
an associate professor in marine ecology at Macquarie University, says "There is no
scientific support for the concept that culling sharks in a particular area will lead to a
decrease in shark attacks and increase ocean safety."[167]

Other threats
Other threats include habitat alteration, damage and loss from coastal development,
pollution and the impact of fisheries on the seabed and prey species.[168] The 2007
documentary Sharkwater exposed how sharks are being hunted to extinction.[169]

Conservation
Further information: List of threatened sharks and Shark sanctuary
In 1991, South Africa was the first country in the world to declare Great White sharks a
legally protected species[170] (however, the KwaZulu-Natal Sharks Board is allowed to kill
great white sharks in its "shark control" program in eastern South Africa).[160]

Intending to ban the practice of shark finning while at sea, the United States Congress
passed the Shark Finning Prohibition Act in 2000.[171] Two years later the Act saw its first
legal challenge in United States v. Approximately 64,695 Pounds of Shark Fins. In 2008
a Federal Appeals Court ruled that a loophole in the law allowed non-fishing vessels
to purchase shark fins from fishing vessels while on the high seas.[172] Seeking to close
the loophole, the Shark Conservation Act was passed by Congress in December 2010,
and it was signed into law in January 2011.[173][174]

In 2003, the European Union introduced a general shark finning ban for all vessels of all
nationalities in Union waters and for all vessels flying a flag of one of its member states.
[175]
This prohibition was amended in June 2013 to close remaining loopholes.[176]

In 2009, the International Union for Conservation of Nature's IUCN Red List of
Endangered Species named 64 species, one-third of all oceanic shark species, as
being at risk of extinction due to fishing and shark finning.[177][178]

In 2010, the Convention on International Trade in Endangered Species (CITES)

rejected proposals from the United States and Palau that would have required countries
to strictly regulate trade in several species of scalloped hammerhead, oceanic
whitetip and spiny dogfish sharks. The majority, but not the required two-thirds of voting
delegates, approved the proposal. China, by far the world's largest shark market,
and Japan, which battles all attempts to extend the convention to marine species, led
the opposition.[179][180] In March 2013, three endangered commercially valuable sharks,
the hammerheads, the oceanic whitetip and porbeagle were added to Appendix 2
of CITES, bringing shark fishing and commerce of these species under licensing and
regulation.[181]

In 2010, Greenpeace International added the school shark, shortfin mako

shark, mackerel shark, tiger shark and spiny dogfish to its seafood red list, a list of
common supermarket fish that are often sourced from unsustainable fisheries.
[182]
Advocacy group Shark Trust campaigns to limit shark fishing. Advocacy
group Seafood Watch directs American consumers to not eat sharks.[183]

Under the auspices of the Convention on the Conservation of Migratory Species of Wild
Animals (CMS), also known as the Bonn Convention, the Memorandum of
Understanding on the Conservation of Migratory Sharks was concluded and came into
effect in March 2010. It was the first global instrument concluded under CMS and aims
at facilitating international coordination for the protection, conservation and
management of migratory sharks, through multilateral, intergovernmental discussion
and scientific research.
In July 2013, New York state, a major market and entry point for shark fins, banned the
shark fin trade joining seven other states of the United States and the three Pacific U.S.
territories in providing legal protection to sharks.[184]

In the United States, and as of January 16, 2019, 12 states including

(Massachusetts, Maryland, Delaware, California, Illinois, Hawaii, Oregon, Nevada, Rhod
e Island, Washington, New York and Texas) along with 3 U.S. territories (American
Samoa, Guam and the Northern Mariana Islands) have passed laws against the sale or
possession of shark fins.[185][186]

Several regions now have shark sanctuaries or have banned shark fishing—these
regions include American Samoa, the Bahamas, the Cook Islands, French
Polynesia, Guam, the Maldives, the Marshall Islands, Micronesia, the Northern Mariana
Islands, and Palau.[187][188][189]

In April 2020 researchers reported to have traced the origins of shark fins of
endangered hammerhead sharks from a retail market in Hong Kong back to their source
populations and therefore the approximate locations where the sharks were first caught
using DNA analysis.[190][191]

In July 2020 scientists reported results of a survey of 371 reefs in 58 nations estimating
the conservation status of reef sharks globally. No sharks have been observed on
almost 20% of the surveyed reefs and shark depletion was strongly associated with
both socio-economic conditions and conservation measures.[192][193] Sharks are
considered to be a vital part of the ocean ecosystem.

According to a 2021 study in Nature,[194] overfishing has resulted in a 71% global decline
in the number of oceanic sharks and rays over the preceding 50 years. The oceanic
whitetip, and both the scalloped hammerhead and great hammerheads are now
classified as critically endangered.[195] Sharks in tropical waters have declined more
rapidly than those in temperate zones during the period studied.[196] A 2021 study
published in Current Biology found that overfishing is currently driving over one-third of
sharks and rays to extinction.[197]