Shark

Sharks are a group of elasmobranch fish characterized by a

cartilaginous skeleton, five to seven gill slits on the sides of the Sharks
head, and pectoral fins that are not fused to the head. Modern sharks Temporal range: Ludfordian-
are classified within the clade Selachimorpha (or Selachii) and are Present, 425–0 Ma
the sister group to the rays. However, the term "shark" has also
been used for extinct members of the subclass Elasmobranchii PreꞒ Ꞓ O S D C P T J K PgN
[1]
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.[2] 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.[3] Sharks are found in all seas and are common to depths of Clockwise from top left: spiny dogfish,
2,000 metres (6,600 ft). They generally do not live in freshwater
Japanese sawshark, whale shark,
although there are a few known exceptions, such as the bull shark
and the river shark, which can be found in both seawater and great white shark, horn shark, frilled
freshwater.[4] Sharks have a covering of dermal denticles that shark, scalloped hammerhead and
protects their skin from damage and parasites in addition to Australian angelshark representing
improving their fluid dynamics. They have numerous sets of
the orders Squaliformes,
replaceable teeth.[5]
Pristiophoriformes, Orectolobiformes,
Well-known species such as the tiger shark, blue shark, great white Lamniformes, Heterodontiformes,
shark, mako shark, thresher shark, and hammerhead shark are apex Hexanchiformes, Carcharhiniformes
predators—organisms at the top of their underwater food chain.
and Squatiniformes respectively.
Many shark populations are threatened by human activities.
Scientific classification
Kingdom: Animalia
Contents Phylum: Chordata
Etymology Class: Chondrichthyes
Evolutionary history
Subclass: Elasmobranchii
Taxonomy
Infraclass: Euselachii
Anatomy
Teeth Superorder: Selachimorpha
Skeleton Orders
Jaw
Fins
 Dermal denticles Carcharhiniformes
Tails Heterodontiformes
Hexanchiformes
Physiology Lamniformes
Buoyancy Orectolobiformes
Respiration Pristiophoriformes
Thermoregulation Squaliformes
Osmoregulation Squatiniformes
Digestion † Cladoselachiformes
† Hybodontiformes
Biofluoresence
† Symmoriida
Senses † Xenacanthida
Smell (Xenacantiformes) †
Sight Elegestolepis
Hearing
† = extinct
Electroreception
Lateral line Synonyms

Life history
Reproduction Pleurotremata
Sexual
Asexual
Brooding
Ovoviviparity
Oviparity
Viviparity
Behavior
Speed
Intelligence
Sleep
Ecology
Feeding
Range and habitat
Relationship with humans
Attacks
In captivity
In culture
In Hawaii
In American Samoa
In popular culture
Popular misconceptions
Threats to sharks
Fishery
Shark culling
Other threats
Conservation
See also
References
Further reading

Etymology
Until the 16th century,[6] sharks were known to mariners as "sea dogs".[7] 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.[8]

A now disproven theory is that it derives from the Yucatec Maya word xok (pronounced 'shok'), meaning
"fish".[9] 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.[10]

Evolutionary history
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.[2] 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.[11] The oldest generally
accepted shark scales are from about 420 million years ago, in the
A collection of Cretaceous shark
Silurian period.[11] The first sharks looked very different from modern
teeth
sharks.[12] 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.[13] 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,[12] 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.[12] 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.[14][15] 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. 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.[16]

Modern sharks began to appear about 100 million years ago.[13]

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.[17] 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 Megalodon (top two, estimated
megatoothed shark, C. megalodon. Like most extinct sharks, C. maximum and conservative sizes)
megalodon is also primarily known from its fossil teeth and vertebrae. with the whale shark, great white
shark, and a human for scale
This giant shark reached a total length (TL) of more than 16 metres
(52 ft).[18][19] 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.[20] Paleontological evidence suggests that this shark was an active predator of large cetaceans.[20]

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,[21] but current molecular
studies support monophyly of both groups of
sharks and batoids.[22][23]

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.[21][24]
There are more than 470 species of sharks split across twelve orders, including four orders of sharks that have
gone extinct:[24]

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

Teeth

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 General anatomical features of sharks
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.[25]

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.[26]

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.
The teeth of tiger sharks are oblique
This reduces the skeleton's weight, saving energy.[27] Because sharks
and serrated to saw through flesh
do not have rib cages, they can easily be crushed under their own
weight on land.[28]

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.[29] 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.[27] 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.[30] 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.[28]

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.[31]
Their dermal teeth give them hydrodynamic advantages as they
reduce turbulence when swimming.[32]

Tails
The dermal denticles of a lemon
shark, viewed through a scanning
Tails provide thrust, making speed and acceleration dependent on tail
electron microscope
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.[33]

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.[34] 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.[31] Their liver constitutes up to 30% of their total body mass.[35] 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.[36]

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.[26] 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.[37][38]

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.[39]

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.[40]

Osmoregulation

In contrast to bony fish, with the exception of the coelacanth,[41] 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%[42]) 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.[35] When a shark dies, the urea is broken down to ammonia
by bacteria, causing the dead body to gradually smell strongly of ammonia.[43][44]

Research in 1930 by Homer W. Smith showed that sharks' urine doesn't 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.[45]

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.[46] 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.[47]

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.[46]

Biofluoresence

Biofluorescence is a characteristic of a few shark species, such as the swell shark and the chain catshark, the
mechanism of which is unique among marine animals and depends upon a small molecule metabolite.[48]

Senses

Smell

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.[49]

Sharks have the ability to determine the direction of a given scent based on the timing of scent detection in
each nostril.[50] 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

Shark eyes are similar to

the eyes of other The shape of the hammerhead shark's
vertebrates, including head may enhance olfaction by spacing
similar lenses, corneas and the nostrils further apart.
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
Eye of a Bigeyed sixgill shark
the tissue varies, with some sharks having stronger nocturnal
(Hexanchus nakamurai)
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. 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.[51] A micro-spectrophotometry study of 17 species of shark 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.[52]
[53][54]

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.[55] 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

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.[56] 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.[57]

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.[58] 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
Electromagnetic field receptors (ampullae of
helps sharks distinguish between the currents around
Lorenzini) and motion detecting canals in the head
them, obstacles off on their periphery, and struggling of a shark
prey out of visual view. The shark can sense frequencies
in the range of 25 to 50 Hz.[59]

Life history
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.[60]
Whale sharks (Rhincodon typus) may also live over 100 years.[61]
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.[62][63]

The claspers of male spotted

Reproduction wobbegong

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.[64]
Sharks mature slowly relative to many other fish. For example, lemon
sharks reach sexual maturity at around age 13–15.[65]

Sexual

Sharks practice internal fertilization.[66] 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.[67]

Mating has rarely been observed in sharks.[68] 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 Shark egg
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.[67]

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.[69][70] The details of this
process are not well understood, but genetic fingerprinting showed
that the pups had no paternal genetic contribution, ruling out sperm The spiral egg case of a Port
storage. The extent of this behavior in the wild is unknown. Mammals Jackson shark
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.[71]

Brooding

Sharks display three ways to bear their young, varying by species, oviparity, viviparity and
ovoviviparity.[72][73]

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.[72]

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.[72][74]

Viviparity
Viviparity is the gestation of young without the use of a traditional egg, and results in live birth.[75] Viviparity
in sharks can be placental or aplacental.[75] Young are born fully formed and self-sufficient.[75] Hammerheads,
the requiem sharks (such as the bull and blue sharks), and smoothhounds are viviparous.[64][72]

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.[76] Even solitary sharks meet for breeding or at rich hunting grounds, which may lead
them to cover thousands of miles in a year.[77] 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.[35] Cross-species social hierarchies
exist. For example, oceanic whitetip sharks dominate silky sharks of comparable size during feeding.[64]

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.[78]

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).[79] 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.[80]

Intelligence

Sharks possess brain-to-body mass ratios that are similar to mammals and birds,[81] and have exhibited
apparent curiosity and behavior resembling play in the wild.[82][83]

There is evidence that juvenile lemon sharks can use observational learning in their investigation of novel
objects in their environment.[84]

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[85] 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.[85] In 2016 a great white shark was captured on
video for the first time in a state researchers believed was sleep swimming.[86]
Ecology

Feeding

Most sharks are carnivorous.[87] 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.[87]

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.[35]

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.[88] 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
Unlike many other sharks, the great
viper dogfish has teeth it can point outwards to strike and capture prey white shark is not actually an apex
that it then swallows intact. The great white and other large predators predator in all of its natural
either swallow small prey whole or take huge bites out of large environments, as it is sometimes
animals. Thresher sharks use their long tails to stun shoaling fishes, hunted by orcas
and sawsharks either stir prey from the seabed or slash at swimming
prey with their tooth-studded rostra.

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.[89]

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.[90] 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).[91]

Relationship with humans

Attacks
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.[92]

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.[93][94] These
A sign warning about the presence of
sharks are large, powerful predators, and may sometimes attack and
sharks in Salt Rock, South Africa
kill people. Despite being responsible for attacks on humans they
have all been filmed without using a protective cage.[95]

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.[96]
Snorkeler swims near a blacktip reef
shark. In rare circumstances involving
To help avoid an unprovoked attack, humans should not wear
poor visibility, blacktips may bite a
jewelry or metal that is shiny and refrain from splashing around
human, mistaking it for prey. Under normal
too much.[97] conditions they are harmless and shy.

In general, sharks show little pattern of attacking humans

specifically. 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.[98][99]

In captivity

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.[100] 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.[101] Uninformed or unscrupulous dealers
sometimes sell juvenile sharks like the nurse shark, which upon
reaching adulthood is far too large for typical home aquaria.[101]
Public aquaria generally do not accept donated specimens that have
outgrown their housing. Some owners have been tempted to release
them.[101] Species appropriate to home aquaria represent considerable
spatial and financial investments as they generally approach adult A whale shark in Georgia Aquarium
lengths of 3 feet (90 cm) and can live up to 25 years.[101]
In culture

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.[102]

Kamohoali'i is the best known and revered of the shark gods, he was the older and favored brother of
Pele,[103] 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.[104][105] 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.[106][107][108] 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."[106]

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.[109] 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.[110] 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.

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.[111][112] 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.[113] Other apparently false claims
are that fins prevent cancer[114] and treat osteoarthritis.[115] No scientific proof supports these claims; at least
one study has shown shark cartilage of no value in cancer treatment.[116]

Threats to sharks

Fishery

It is estimated that 100 million sharks are killed by people every

year, due to commercial and recreational fishing.[117][118]

Shark finning yields are estimated at 1.44 million metric tons for
2000, and 1.41 million 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 The annual shark catch has increased
between 63 and 273 million sharks per year. [119][120] Sharks are rapidly over the last 60 years.
a common seafood in many places, including Japan and
Australia. In the Australian state of Victoria, shark is the most
commonly used fish in fish and chips, 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.[121] During a four-year period from 1996 to
2000, an estimated 26 to 73 million sharks were killed and traded The value of shark fins for shark fin
annually in commercial markets. [122] soup has led to an increase in shark
catches where usually only the fins
Sharks are often killed for shark fin soup. Fishermen capture live are taken, while the rest of the shark
sharks, fin them, and dump the finless animal back into the water. is discarded, typically into the sea;
health concerns about BMAA in the
Shark finning involves removing the fin with a hot metal blade.[118]
fins now exists regarding
The resulting immobile shark soon dies from suffocation or
consumption of the soup
predators.[123] Shark fin has become a major trade within black
markets all over the world. Fins sell for about $300/lb in 2009.[124]
Poachers illegally fin millions each year. Few governments enforce laws that protect them.[120] In 2010
Hawaii became the first U.S. state to prohibit the possession, sale, trade or distribution of shark fins.[125] From
1996 to 2000, an estimated 38 million sharks had been killed per year for harvesting shark fins.[122] It is
estimated by TRAFFIC that over 14,000 tonnes of shark fins were exported into Singapore between 2005–
2007 and 2012–2014.[126]

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.[127]
Because BMAA is a neurotoxin, consumption of shark fin soup and cartilage pills, therefore, may pose a
health risk.[128] BMAA is under study for its pathological role in neurodegegerative 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.[129]
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.[66] 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).[66]

The majority of shark fisheries have little monitoring or management. The rise A 14-foot (4.3 m), 1,200-
in demand for shark products increases pressure on fisheries.[36] Major pound (540 kg) tiger shark
declines in shark stocks have been recorded—some species have been caught in Kāne'ohe Bay,
depleted by over 90% over the past 20–30 years with population declines of Oahu in 1966
70% not unusual.[130] 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.[131][132]

Shark culling

In 2014, a shark cull in Western Australia killed dozens of sharks (mostly tiger sharks) using drum lines,[133]
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.[134] This "imminent threat" policy was criticized by senator Rachel Siewart for
killing endangered sharks.[135] The "imminent threat" policy was cancelled in March 2017.[136] 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).[137]

From 1962 to the present,[138] 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.[139][140][141][142] Queensland's drum line program has been called "outdated, cruel and
ineffective".[142] From 2001 to 2018, a total of 10,480 sharks were killed on lethal drum lines in Queensland,
including in the Great Barrier Reef.[143] From 1962 to 2018, roughly 50,000 sharks were killed by
Queensland authorities.[144]

The government of New South Wales has a program that deliberately kills sharks using nets.[141][145] The
current net program in New South Wales has been described as being "extremely destructive" to marine life,
including sharks.[146] 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.[147] 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.[144]

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.[148] 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.[148]
Authorities on the French island of Réunion kill about 100 sharks per year.[149]

Killing sharks negatively affects the marine ecosystem.[150][151] 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."[152]

George H. Burgess, the former[153] director of the International Shark Attack File, "describes [shark] culling
as a form of revenge, satisfying a public demand for blood and little else";[154] 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."[154] 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."[155]

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.[156] The 2007 documentary Sharkwater exposed how sharks are
being hunted to extinction.[157]

Conservation
In 1991, South Africa was the first country in the world to declare Great White sharks a legally protected
species[158] (however, the KwaZulu-Natal Sharks Board is allowed to kill great white sharks in its "shark
control" program in eastern South Africa).[148]

Intending to ban the practice of shark finning while at sea, the United States Congress passed the Shark
Finning Prohibition Act in 2000.[159] 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.[160] 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.[161][162]

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.[163] This prohibition was amended in June
2013 to close remaining loopholes.[164]

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.[165][166]

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.[167][168] 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.[169]
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.[170] Advocacy group Shark Trust campaigns to limit shark fishing. Advocacy group
Seafood Watch directs American consumers to not eat sharks.[171]

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.[172]

In the United States, and as of January 16, 2019, 12 states including (Massachusetts, Maryland, Delaware,
California, Illinois, Hawaii, Oregon, Nevada, Rhode 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.[173][174]

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.[175][176]

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.[177][178]

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.[179][180]
Sharks are considered to be a vital part of the ocean ecosystem.

See also
List of sharks
List of prehistoric cartilaginous fish genera
Osteichthyes
Marine vertebrate
Outline of sharks
Shark meat

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General references

Castro, Jose (1983). The Sharks of North American Waters (https://archive.org/details/sharksof

northame00jose). College Station: Texas A&M University Press. ISBN 978-0-89096-143-8.
OCLC 183037060 (https://www.worldcat.org/oclc/183037060).
Stevens, John D. (1987). Sharks (https://archive.org/details/sharks00stev). New York: NY Facts
on File Publications. ISBN 978-0-8160-1800-0. OCLC 15163749 (https://www.worldcat.org/ocl
c/15163749).
Pough, F. H.; Janis, C. M.; Heiser, J. B. (2005). Vertebrate Life (7th ed.). New Jersey: Pearson
Education Ltd. ISBN 978-0-13-127836-3. OCLC 54822028 (https://www.worldcat.org/oclc/5482
2028).
Clover, Charles (2004). The End of the Line: How overfishing is changing the world and what
we eat. London: Ebury Press. ISBN 978-0-09-189780-2.
Owen, David (2009). Shark: In Peril in the Sea. New South Wales: Allen and Unwin. ISBN 978-
1-74175-032-4.

Further reading
Musick, John A and Musick, Susanna (2011) "Sharks" (http://www.fao.org/docrep/015/i2389e/i2
389e.zip) In: Review of the state of world marine fishery resources, pages 245–254, FAO
Fisheries technical paper 569, FAO, Rome. ISBN 978-92-5-107023-9.
Sharks Falling Prey To Humans' Appetites (http://news.nationalgeographic.com/news/2002/06/
0603_020603_shark1_2.html) National Geographic, 28 October 2010.

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