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ABOUT MEGALODONS:

Megalodon (Carcharocles megalodon), meaning "big tooth", is an extinct species of shark that lived approximately 23 to 2.6 million years ago (mya), during the Early Miocene to the end of the Pliocene. It was formerly thought to belong to the family Lamnidae, making it closely related to the great white shark (Carcharodon carcharias). However presently there is near unanimous consensus that it belongs to the extinct family Otodontidae, which diverged from the ancestry of the great white shark during the Early Cretaceous. Its genus placement is still debated, authors placing it in either Carcharocles, Megaselachus, Otodus, or Procarcharodon.

Scientists suggest that megalodon looked like a stockier version of the great white shark, though it may have looked similar to the basking shark (Cetorhinus maximus) or the sand tiger shark (Carcharias taurus). Regarded as one of the largest and most powerful predators to have ever lived, fossil remains of megalodon suggest that this giant shark reached a length of 18 meters (59 ft). Their large jaws could exert a bite force of up to 110,000 to 180,000 newtons (24,000 - 41,000 lbf). Their teeth were thick and robust, built for grabbing prey and breaking bone.

Megalodon probably had a major impact on the structure of marine communities. The fossil record indicates that it had a cosmopolitan distribution. It probably targeted large prey, such as whales, seals, and giant turtles. Juveniles inhabited warm coastal waters where they would feed on fish and small whales. Unlike the great white, which attacks prey from the soft underside, megalodon probably used its strong jaws to break through the chest cavity and puncture the heart and lungs of its prey.

The animal faced competition from whale-eating cetaceans, such as Livyatan and ancient killer whales (Orcinus citoniensis), which likely contributed to its extinction. As it preferred warmer waters, it is thought that oceanic cooling associated with the onset of the ice ages, coupled with the lowering of sea levels and resulting loss of suitable nursery areas, may have also contributed to its decline. A reduction in the diversity of baleen whales and a shift in their distribution toward polar regions may have reduced megalodon's primary food source. The extinction of the shark appeared to affect other animals; for example, the size of baleen whales increased significantly after the shark had disappeared.

References to the creature occur prominently in several pop culture media.

According to Renaissance accounts, gigantic, triangular fossil teeth often found embedded in rocky formations were once believed to be the petrified tongues, or glossopetrae, of dragons and snakes. This interpretation was corrected in 1667 by Danish naturalist Nicolas Steno, who recognized them as shark teeth, and famously produced a depiction of a shark's head bearing such teeth. He described his findings in the book The Head of a Shark Dissected, which also contained an illustration of a megalodon tooth.[6][7][8]

Megalodon has been portrayed in several works of fiction, including films and novels, and continues to be a popular subject for fiction involving sea monsters.[82] Three individual megalodon, two adults and one juvenile, were portrayed in BBC's 2003 TV series Sea Monsters, where it is defined as a "hazard" of the era.[83] The History Channel's Jurassic Fight Club portrays a megalodon attacking a Brygmophyseter sperm whale in Japan.[84] Several films depict megalodon, such as Shark Attack 3: Megalodon and the Mega Shark series (for instance Mega Shark Versus Giant Octopus and Mega Shark Versus Crocosaurus).[82] The shark appears in the 2017 videogame Ark: Survival Evolved.[85] Some stories, such as Jim Shepard's Tedford and the Megalodon, portray a rediscovery of the shark.[86] Steve Alten's Meg: A Novel of Deep Terror portrays the shark having preyed on dinosaurs with its prologue and cover artwork depicting megalodon killing a Tyrannosaurus in the sea.[87] The sequels to the book also star megalodon: The Trench, Meg: Primal Waters, Meg: Hell's Aquarium, and Meg: Origins,[82] and there is a film adaptation entitled The Meg released on August 10, 2018.[88]

Animal Planet's pseudo-documentary Mermaids: The Body Found included an encounter 1.6 mya between a pod of mermaids and a megalodon.[89] Later, in August 2013, the Discovery Channel opened its annual Shark Week series with another film for television, Megalodon: The Monster Shark Lives,[90] a controversial docufiction about the creature that presented alleged evidence in order to suggest that megalodon was still alive. This program received criticism for being completely fictional; for example, all of the supposed scientists depicted were paid actors. In 2014, Discovery re-aired The Monster Shark Lives, along with a new one-hour program, Megalodon: The New Evidence, and an additional fictionalized program entitled Shark of Darkness: Wrath of Submarine, resulting in further backlash from media sources and the scientific community.[39][91][92]

There have been a few alleged sightings of large sharks, purportedly megalodon, measuring anywhere from 10 to 90 meters (40 to 300 ft) throughout the 20th century; they all lack supporting evidence. One Polynesian myth regards a 30-meter (100 ft) shark called Lord of the Deep. Reports of supposedly fresh megalodon teeth, such as those made by HMS Challenger in 1873 which were erroneously dated to be around 11,000 to 24,000 years old, are probably teeth that were well-preserved by a thick mineral-crust precipitate of manganese dioxide, and so had a lower decomposition rate and retained a white color during fossilization. Fossil megalodon teeth can vary in color from off-white to dark browns and greys, and some fossil teeth may have been redeposited into a younger stratum. The claims that megalodon could remain elusive in the depths, similar to the megamouth shark which was discovered in 1976, are unlikely as the shark lived in warm coastal waters and probably could not survive in the cold and nutrient-poor deep sea environment.[93]


Carcharocles megalodon

Relationship between megalodon and other sharks, including the great white shark (Carcharodon carcharias)[15]
While the earliest megalodon remains were reported from the Late Oligocene dated to around 28 million years ago (mya),[16][17] competing figures still exist as to when it evolved, such as 16 mya and 23 mya.[18] It is believed that megalodon became extinct around the end of the Pliocene, probably about 2.6 mya;[18][19] reported Pleistocene megalodon teeth, younger than 2.6 million years old, are considered to be unreliable claims.[19]

Megalodon is now considered to be a member of the family Otodontidae, genus Carcharocles, as opposed to its previous classification into Lamnidae, genus Carcharodon.[18][19][20][21] Megalodon's classification into Carcharodon was due to dental similarity with the great white shark, but most authors currently believe that this is due to convergent evolution. In this model, the great white shark is more closely related to the shark Isurus hastalis than to megalodon, as evidenced by more similar dentition in those two sharks; megalodon teeth have much finer serrations than great white shark teeth. The great white shark is more closely related to the mako shark (Isurus spp.), with a common ancestor around 4 mya.[9][15] Proponents of the former model, wherein megalodon and the great white shark are more closely related, argue that the differences between their dentition are minute and obscure.[22]:23–25

A black megalodon tooth and two white great white shark teeth above a centimeter scale, the megalodon tooth extends between the zero and thirteen-and-a-half centimeter marks. One great white tooth extends between the eleven and thirteen centimeter marks, and the other extends between from the thirteen and sixteen centimeter marks.
Megalodon tooth with two great white shark teeth
The genus Carcharocles currently contains four species: C. auriculatus, C. angustidens, C. chubutensis, and C. megalodon.[14]:30–31 The genus was proposed by D. S. Jordan and H. Hannibal in 1923 to contain C. auriculatus. In the 1980s, megalodon was assigned to Carcharocles.[9][14]:30 Before this, in 1960, the genus Procarcharodon was erected by French ichthyologist Edgard Casier, which included those four sharks and was considered separate from the great white shark. It is now considered a junior synonym of Carcharocles.[14]:30 The genus Palaeocarcharodon was erected alongside Procarcharodon to represent the beginning of the lineage, and, in the model wherein megalodon and the great white shark are closely related, their last common ancestor. It is believed to be an evolutionary dead-end and unrelated to the Carcharocles sharks by authors who reject that model.[22]:70

A great white shark swimming a few meters below the surface, above a school of much smaller fish.
The great white shark (Carcharodon carcharias) and megalodon were previously thought to be close relatives.[9][15]
Another model of the evolution of this genus, also proposed by Casier in 1960, is that the direct ancestor of the Carcharocles is the shark Otodus obliquus, which lived from 60 mya to 13 mya during the Paleocene and Miocene epochs.[15][23] In this model, O. obliquus evolved into O. aksuaticus, which evolved into C. auriculatus, and then into C. angustidens, and then into C. chubutensis, and then finally into C. megalodon. The evolution of this lineage is characterized by the increase of serrations, the widening of the crown, the development of a more triangular shape, and the disappearance of the lateral cusps.[14]:28–31[23] The genus Otodus is ultimately derived from Cretolamna, a shark from the Cretaceous period.[20][24]

Another model of the evolution of Carcharocles, proposed in 2001 by paleontologist Michael Benton, is that the three other species are actually a single species of shark that gradually changed over time between the Paleocene and the Pliocene, making it a chronospecies.[14]:17[17][25] Some authors suggest that C. auriculatus, C. angustidens, and C. chubutensis should be classified as a single species in the genus Otodus, leaving C. megalodon the sole member of Carcharocles.[17][26]

The genus Carcharocles may be invalid, and the shark may actually belong in the genus Otodus, making it Otodus megalodon.[4] A 1974 study on Paleogene sharks by Henri Cappetta erected the subgenus Megaselachus, classifying the shark as Otodus (Megaselachus) megalodon, along with O. (M.) chubutensis. A 2006 review of Chondrichthyes elevated Megaselachus to genus, and classified the sharks as Megaselachus megalodon and M. chubutensis.[4] The discovery of fossils assigned to the genus Megalolamna in 2016 led to a re-evaluation of Otodus, which concluded that it is paraphyletic, that is, it consists of a last common ancestor but it does not include all of its descendants. The inclusion of the Carcharocles sharks in Otodus would make it monophyletic, with the sister clade being Megalolamna.[20] There was one apparent description of the shark in 1881 classifying it as Selache manzonii.[27]

Biology
Appearance
A drawing of a swimming shark showing the front left underside view
Restoration of megalodon with a similar appearance to the great white shark
One interpretation on how megalodon appeared was that it was a robust-looking shark, and may have had a similar build to the great white shark. The jaws may have been blunter and wider than the great white, and the fins would have also been similar in shape, though thicker due to its size. It may have had a pig-eyed appearance, in that it had small, deep-set eyes.[22]:64–65

Another interpretation is that megalodon bore a similarity to the whale shark (Rhincodon typus) or the basking shark (Cetorhinus maximus). The tail fin would have been crescent-shaped, the anal fin and second dorsal fin would have been small, and there would have been a caudal keel present on either side of the tail fin (on the caudal peduncle). This build is common in other large aquatic animals, such as whales, tuna, and other sharks, in order to reduce drag while swimming. The head shape can vary between species as most of the drag-reducing adaptations are toward the tail-end of the animal.[14]:35–36


Sculpture at the Museo de la Evolución de Puebla in Mexico
Since Carcharocles is derived from Otodus, and the two had teeth that bear a close similarity to those of the sand tiger shark (Carcharias taurus), megalodon may have had a build more similar to the sand tiger shark than to other sharks. This is unlikely since the sand tiger shark is a carangiform swimmer which require faster movement of the tail for propulsion through the water than the great white shark, a thunniform swimmer.[14]:35–36[28]

Size
Statistics
At the top of the picture is a line representing twenty meters. Below this is a gray megalodon silhouette that measures twenty meters, below is a red megalodon silhouette that measures fifteen meters, below is a violet whale shark silhouette that measures ten meters, below is a green great white shark that measures five meters. Standing next to this shark is a black human silhouette that stands two meters.
Megalodon (gray and red representing the largest and smallest estimates) with the whale shark (violet), great white shark (green), and a human (black) for scale
Due to fragmentary remains, there have been many contradictory size estimates for megalodon, as they can only be drawn from fossil teeth and vertebrae.[14]:87[29] Also because of this, the great white shark is the basis of its reconstruction and size estimation,[22]:57 as it is regarded as the best analogue to megalodon.[26] Various size estimates exist for megalodon; in 1973, Hawaiian ichthyologist John E. Randall estimated that the maximum length attained by C. megalodon was about 13 meters (43 ft),[30] while in the 1990s, marine biologists Patrick J. Schembri and Stephen Papson opined that C. megalodon may have approached a maximum of around 24 to 25 meters (79 to 82 ft) in total length;[31][32] Gottfried and colleagues asserted that C. megalodon could have reached a maximum of 20.3 meters (67 ft) in total length.[33][22][34] Nowadays, the commonly acknowledged maximum total length of C. megalodon is about 18 meters (59 ft), with the average size being 10.5 meters (34 ft),[18][19][21] compared to the maximum recorded sizes of the great white shark at 6.1 meters (20 ft) and the whale shark (the largest extant fish) at 12.65 m (42 ft).[35][36][37][38] It is possible that different populations of megalodon around the globe had different body sizes and behaviors due to different ecological pressures.[21] If it did attain a size of over 16 meters (52 ft), it would have been the largest known fish that has ever lived, surpassing the Jurassic fish Leedsichthys.[39]

Mature male megalodon may have had a body mass of 12.6 to 33.9 metric tons (13.9 to 37.4 short tons), and mature females may have been 27.4 to 59.4 metric tons (30.2 to 65.5 short tons), given that males could range in length from 10.5 to 14.3 meters (34 to 47 ft) and females 13.3 to 17 meters (44 to 56 ft).[22]:61[33] A 2015 study linking shark size and typical swimming speed estimated that megalodon would have typically swum at 18 kilometers per hour (11 mph), given that its body mass was typically 48 metric tons (53 short tons), which is consistent with other aquatic creatures of its size, such as the fin whale which typically cruises at speeds of 14.5 to 21.5 km/h (9.0 to 13.4 mph).[40]

Its large size may have been due to climatic factors and the abundance of large prey items, and it may have also been influenced by the evolution of regional endothermy (mesothermy) which would have increased its metabolic rate and swimming speed. Since the otodontid sharks are considered to have been ectotherms, and megalodon was a close relative to them, megalodon may have also been ectothermic. Contrary to this, the largest contemporary ectothermic sharks, such as the whale shark, are filter feeders, implying some metabolic constraints with a predatory lifestyle. That is to say, it is unlikely that megalodon was ectothermic.[41]

Estimations
Gordon Hubbell from Gainesville, Florida, possesses an upper anterior megalodon tooth whose maximum height is 18.4 centimeters (7.25 in), one of the largest known tooth specimens from the shark.[42] In addition, a 2.7-by-3.4-meter (9 by 11 ft) megalodon jaw reconstruction developed by fossil hunter Vito Bertucci contains a tooth whose maximum height is reportedly over 18 centimeters (7 in).[43]

The first attempt to reconstruct the jaw of megalodon was made by Bashford Dean in 1909, displayed at the American Museum of Natural History. From the dimensions of this jaw reconstruction, it was hypothesized that megalodon could have approached 30 meters (98 ft) in length. Dean had overestimated the size of the cartilage on both jaws, causing it to be too tall.[39][44]
Black-and-white photo of a man sitting inside a megalodon jaw reconstruction.
Reconstruction by Bashford Dean in 1909
A white megalodon tooth on the palms of a person. On the right side of the image is a ruler. The tip of the tooth starts at zero and ends at the seventeen centimeter marker on the ruler.
Tooth compared to hand
John E. Randall, an ichthyologist, used the enamel height (the vertical distance of the blade from the base of the enamel portion of the tooth to its tip) to measure the length of the shark, yielding a maximum length of about 13 meters (43 ft).[30] Tooth enamel height does not necessarily increase in proportion to the animal's total length.[22]:99

Shark researchers Michael D. Gottfried, Leonard Compagno, and S. Curtis Bowman proposed a linear relationship between a shark's total length and the height of the largest upper anterior tooth. The proposed relationship is: total length in meters = − (0.096) × [UA maximum height (mm)]-(0.22).[17][22]:60 They had estimated the average height, based on the slant height of the largest tooth discovered, for large female megalodon to be 15.6 meters (51 ft), though larger teeth may exist.[22]:55–60

In 2002, shark researcher Clifford Jeremiah proposed that total length was proportional to the root width of an upper anterior tooth. He claimed that for every 1 centimeter (0.39 in) of root width, there are approximately 1.4 meters (4.6 ft) of shark length. Jeremiah pointed out that the jaw perimeter of a shark is directly proportional to its total length, with the width of the roots of the largest teeth being a tool for estimating jaw perimeter. The largest tooth in Jeremiah's possession had a root width of about 12 centimeters (4.7 in), which yielded 16.5 meters (54 ft) in total length.[14]:88

In 2002, paleontologist Kenshu Shimada of DePaul University proposed a linear relationship between tooth crown height and total length after conducting anatomical analysis of several specimens, allowing any sized tooth to be used. Shimada stated that the previously proposed methods were based on a less-reliable evaluation of the dental homology between megalodon and the great white shark, and that the growth rate between the crown and root is not isometric, which he considered in his model. Using this model, the upper anterior tooth possessed by Gottfried and colleagues corresponded to a total length of 15 meters (49 ft).[45] Among the specimens found in the Gatún Formation of Panama, other shark researchers used this method and calculated a maximum height of 16.8 meters (55 ft) for a specimen,[26] and for another a total length of 17.9 meters (59 ft). This result appears to be an error within the matrix, and the length of this individual is actually 19.6 meters (64 ft).[46]

Teeth and bite force
A sideview of the inside of a megalodon jaw reconstruction showing five rows of teeth. Each row is more horizontal than the last, with the last row essentially resting on the jaw.
Reconstruction showing the position of the replacement teeth
The most common fossils of megalodon are its teeth. Diagnostic characteristics include a triangular shape, robust structure, large size, fine serrations, a lack of lateral denticles, and a visible V-shaped neck (where the root meets the crown).[22]:55[26] The tooth met the jaw at a steep angle, similar to the great white shark. The tooth was anchored by connective tissue fibers, and the roughness of the base may have added to mechanical strength.[47] The lingual side of the tooth, the part facing the tongue, was convex; and the labial side, the other side of the tooth, was slightly convex or flat. The anterior teeth were almost perpendicular to the jaw and symmetrical, whereas the posterior teeth were slanted and asymmetrical.[48]

Megalodon teeth can measure over 180 millimeters (7.1 in) in slant height (diagonal length) and are the largest of any known shark species.[14]:33 In 1989, a nearly complete set of megalodon teeth was discovered in Saitama, Japan. Another nearly complete associated megalodon dentition was excavated from the Yorktown Formations in the United States, and served as the basis of a jaw reconstruction of megalodon at the National Museum of Natural History (USNM). Based on these discoveries, an artificial dental formula was put together for megalodon in 1996.[22]:55[49]

The dental formula of megalodon is: 
2.1.7.4
3.0.8.4
. As evident from the formula, megalodon had four kinds of teeth in its jaws: anterior, intermediate, lateral, and posterior. Megalodon's intermediate tooth technically appears to be an upper anterior and is termed as "A3" because it is fairly symmetrical and does not point mesially (side of the tooth toward the midline of the jaws where the left and right jaws meet). Megalodon had a very robust dentition,[22]:20–21 and had over 250 teeth in its jaws, spanning 5 rows.[14]:iv It is possible that large megalodon individuals had jaws spanning roughly 2 meters (6.6 ft) across.[14]:129 The teeth were also serrated, which would have improved efficiency in cutting through flesh or bone.[9][14]:1 The shark may have been able to open its mouth to a 75° angle, though a reconstruction at the USNM approximates a 100° angle.[22]:63

A dark-yellow megalodon jaw reconstruction with two rows of white teeth stained black on the top.
Reconstructed jaws on display at the National Aquarium in Baltimore
In 2008, a team of scientists led by S. Wroe conducted an experiment to determine the bite force of the great white shark, using a 2.5-meter (8.2 ft) long specimen, and then isometrically scaling the results for its maximum confirmed size and the conservative minimum and maximum body mass of megalodon. They placed the bite force of the latter between 110,000 to 180,000 newtons (24,000 to 41,000 lbf) in a posterior bite, compared to the 18,000 N (4,100 lbf) bite force for the largest confirmed great white shark, and 7,400 N (1,700 lbf) for the placoderm fish Dunkleosteus. In addition, Wroe and colleagues pointed out that sharks shake sideways while feeding, amplifying the force generated, which would probably have caused the total force experienced by prey to be higher than the estimate.[33][50]

Internal anatomy
A skeletal reconstruction of megalodon. Visible are the jaws with two rows of teeth, eye sockets, a pointed snout, several long, straight spines protruding outwards in the gill area behind the head, and a long horizontal item representing the vertebral column
Reconstructed megalodon skeleton on display at the Calvert Marine Museum
Megalodon is represented in the fossil record by teeth, vertebral centra, and coprolites.[22]:57 As with all sharks, the skeleton of megalodon was formed of cartilage rather than bone; consequently most fossil specimens are poorly preserved.[51] To support its large dentition, the jaws of megalodon would have been more massive, stouter, and more strongly developed than those of the great white, which possesses a comparatively gracile dentition. Its chondrocranium, the cartilaginous skull, would have had a blockier and more robust appearance than that of the great white. Its fins were proportional to its larger size.[22]:64–65

Some fossil vertebrae have been found. The most notable example is a partially preserved vertebral column of a single specimen, excavated in the Antwerp Basin, Belgium, in 1926. It comprises 150 vertebral centra, with the centra ranging from 55 millimeters (2.2 in) to 155 millimeters (6 in) in diameter. The shark's vertebrae may have gotten much bigger, and scrutiny of the specimen revealed that it had a higher vertebral count than specimens of any known shark, possibly over 200 centra; only the great white approached it.[22]:63–65 Another partially preserved vertebral column of a megalodon was excavated from the Gram Formation in Denmark in 1983, which comprises 20 vertebral centra, with the centra ranging from 100 millimeters (4 in) to 230 millimeters (9 in) in diameter.[47]

Smmothly rounded dark brown rock-like coprolite
Coprolite attributed to megalodon
The coprolite remains of megalodon are spiral-shaped, indicating that the shark may have had a spiral valve, a corkscrew-shaped portion of the lower intestines, similar to extant lamniform sharks. Miocene coprolite remains were discovered in Beaufort County, South Carolina, with one measuring 14 cm (5.5 in).[52]

Gottfried and colleagues reconstructed the entire skeleton of megalodon, which was later put on display at the Calvert Marine Museum in the United States and the Iziko South African Museum.[22]:56[23] This reconstruction is 11.3 meters (37 ft) long and represents a mature male,[22]:61 based on the ontogenetic changes a great white shark experiences over the course of its life.[22]:65

Paleobiology
Range and habitat
Megalodon had a cosmopolitan distribution;[18][46] its fossils have been excavated from many parts of the world, including Europe, Africa, the Americas, and Australia.[22]:67[53] It most commonly occurred in subtropical to temperate latitudes.[18][22]:78 It has been found at latitudes up to 55° N; its inferred tolerated temperature range was 1–24 °C (34–75 °F). It arguably had the capacity to endure such low temperatures due to mesothermy, the physiological capability of large sharks to conserve metabolic heat by maintaining a higher body temperature than the surrounding water.[18]

Megalodon inhabited a wide range of marine environments (i.e., shallow coastal waters, areas of coastal upwelling, swampy coastal lagoons, sandy littorals, and offshore deep water environments), and exhibited a transient lifestyle. Adult megalodon were not abundant in shallow water environments, and mostly inhabited offshore areas. Megalodon may have moved between coastal and oceanic waters, particularly in different stages of its life cycle.[14]:33[54] Megalodon teeth have been excavated from regions far away from continental lands, such as the Marianas Trench in the Pacific Ocean.[14]:iv

Fossil remains show a trend for specimens to be larger on average in the southern hemisphere than in the northern, with mean lengths of 11.6 and 9.6 meters (38 and 31 ft), respectively; and also larger in the Pacific than the Atlantic, with mean lengths of 10.9 and 9.5 meters (36 and 31 ft) respectively. They do not suggest any trend of changing body size with absolute latitude, or of change in size over time (although the Carcharocles lineage in general is thought to display a trend of increasing size over time). The overall modal length has been estimated at 10.5 meters (34 ft), with the length distribution skewed towards larger individuals, suggesting an ecological or competitive advantage for larger body size.[21]

Locations of fossils
Megalodon had a global distribution and fossils of the shark have been found in many places around the world, bordering all oceans of the Neogene


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