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Overview

Brief Summary

Biology

The biology of the bull shark is still little known but it shows extraordinary physiological adaptations that allow it to persist in both freshwater and saltwater. Bull sharks have been captured in places you would never imagine a shark to be found; in the foothills of the Peruvian Andes, 3,700 kilometres up the Amazon River; and in Lake Nicaragua, the largest lake in Central America. However, the bull shark may not be able to complete its entire life cycle in freshwater, and all sharks in freshwater require access to saltwater through rivers and estuaries (2) (3). Its swims slowly and heavily, usually near the bottom, concealing its surprising agility and speed employed when attacking prey (2), and deceiving one into believing this may not be one of the most dangerous species of tropical shark, as it is frequently cited (2) (5) (6). Along with the great white and tiger shark the bull shark is responsible for the most accidents involving people (2); a result of its tendency to take large prey and the proximity of its habitat to the activities of humans (2). The bull shark's broad and varied diet includes bony fishes, other shark species (even occasionally young bull sharks), sea turtles, birds, dolphins, and terrestrial mammals (2). The bull shark is viviparous, giving birth to 1 to 13 young in each litter after a pregnancy of 10 to 11 months (2). The female gives birth in late spring and early summer in both hemispheres, in estuaries, river mouths, and very occasionally in freshwater lakes (2). Mating takes place at the same time of the year but it is unknown where exactly as it has never been directly observed (4).
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Description

The bull shark, named for its stocky body and ferocious reputation (3), is most famous for its remarkable ability to thrive in both saltwater and freshwater. It has a grey, robust body, a paler underside, and a blunt, rounded snout (2). The large, triangular first dorsal fin and moderately large second dorsal fin, as well as the other fins, have dusky tips but are not strikingly marked (4). Bull sharks have relatively small eyes, indicating that vision is not the most important sense required for hunting in its frequently turbid habitat (2).
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Comprehensive Description

Description

  Common names: shark (English), tiburón (Espanol)
 
Carcharhinus leucas (Muller & Henle, 1839)


Bull shark


Heavy-bodied; head thick; snout broadly rounded and very short (length before mouth 4.6-6.7% of TL);  eye small; nasal flaps broadly triangular; lip folds very short;  top teeth broadly triangular, straight, heavily serrated, lower teeth high, straight, narrow; no ridge on back between dorsal fins; origin of first dorsal fin usually over or just posterior to pectoral axil; first dorsal fin broadly triangular and moderately high (height 7.0-11.3% of TL), its apex fairly pointed, its origin a little before insertion point of pectoral; 2nd  dorsal fin relatively large, its origin slightly in front of anal origin, its free rear tip short; pectoral large, wide, pointed; inconspicuous keels on tail base.


Grey, becoming white ventrally, often with faint pale grey horizontal band extending into the white of the upper abdomen; fins of small individuals with dusky tips or edges, adults plain.

May reach 350 cm.

Habitat: continental coasts, estuarine, muddy areas, lagoons, often travels far up rivers.

Depth: 0-150 m.

All tropical and subtropical seas; southern California to the Gulf and to Peru, Malpelo and the Revillagigedos.
   
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Biology

A coastal and freshwater shark inhabiting shallow waters especially in bays, estuaries, rivers, and lakes. Readily penetrates far up rivers and hypersaline bays (Ref. 9997, 44894). Capable of covering great distances (up to 180 kilometers in 24 hours), moving between fresh and brackish water at random (Ref. 44894). Adults often found near estuaries and freshwater inflows to the sea; young enter rivers and may be found hundreds of km from the sea (Ref. 4967, 44894, 58304). Feeds on bony fishes, other sharks, rays, mantis shrimps, crabs, squid, sea snails, sea urchins, mammalian carrion, sea turtles, and occasionally garbage (Ref. 244, 5578, 44894). Very hardy and lives well in captivity, probably the most dangerous species of tropical shark (Ref. 244), it is repeatedly implicated in attacks on humans (Ref. 4967, 44894). Attacks in fresh water are rare (Ref. 44894). Viviparous (Ref. 50449). Gives birth to litters of up to 13 young (Ref. 26938, 44894). Size at birth is 56-81 cm TL (Ref. 81623). Sexual maturity is attained after 10-15 years (at a length between 160-200 centimeters) (Ref. 44894). Though not commercially important, this species is a good food fish (Ref. 12484). Utilized fresh, fresh-frozen or smoked for human consumption, fins for soup, hide for leather, liver for oil, and carcass for fishmeal (Ref. 244).
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Distribution

occurs (regularly, as a native taxon) in multiple nations

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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: Circumtropical. Gulf of Mexico, coast of eastern U.S. usually from Chesapeake Bay south. Migratory in Gulf of Mexico.

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Western Atlantic: Massachusetts, USA to southern Brazil
  • North-West Atlantic Ocean species (NWARMS)
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Range Description

The Bull Shark has a worldwide distribution in tropical and warm temperate areas, with seasonal appearances in cool, temperate waters (Garrick 1982, Compagno 1984, Last and Stevens 1994). This has resulted in multiple descriptions and numerous common names for the species (including Zambezi Shark, Swan River Shark and Lake Nicaragua Shark) from throughout its range (see Compagno (1984) for a full list of synonyms).

Primarily the Bull Shark is an inhabitant of continental shelf waters to a depth of about 150 m (but mostly less than 30 m), but it commonly moves into estuarine and fresh waters. It has been documented as travelling large distances up rivers (Thorson 1972), including the Amazon, Gambia, Ganges, Mississippi, San Juan (and Lake Nicaragua), Tigris and Zambezi. It also has been observed to tolerate hypersaline conditions up to 53 parts per thousand (ppt) (sea water is approximately 35 ppt). Although mostly a continental species, there are insular records from the Philippines and the South Pacific islands of New Caledonia, Fiji and Rangiroa (Compagno et al. 1989).
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Zoogeography

See Map (including site records) of Distribution in the Tropical Eastern Pacific 
 
Global Endemism: All species, TEP non-endemic, Circumtropical ( Indian + Pacific + Atlantic Oceans), "Transpacific" (East + Central &/or West Pacific), All Pacific (West + Central + East), East Pacific + Atlantic (East +/or West), Transisthmian (East Pacific + Atlantic of Central America), East Pacific + all Atlantic (East+West)

Regional Endemism: All species, Eastern Pacific non-endemic, Tropical Eastern Pacific (TEP) non-endemic, Continent + Island (s), Continent, Island (s)

Residency: Resident

Climate Zone: North Temperate (Californian Province &/or Northern Gulf of California), Northern Subtropical (Cortez Province + Sinaloan Gap), Northern Tropical (Mexican Province to Nicaragua + Revillagigedos), Equatorial (Costa Rica to Ecuador + Galapagos, Clipperton, Cocos, Malpelo), South Temperate (Peruvian Province )
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Widespread in warm oceans, rivers and lakes. Western Atlantic: Massachusetts, USA to Argentina (Ref. 58839). Eastern Atlantic: Morocco, Senegal to Angola (Ref. 81283). Indo-Pacific: Kenya and South Africa to India, then, Viet Nam to Australia; southern Baja California, Mexico to Ecuador and possibly occurring in Peru. Sympatric with Carcharhinus amboinensis, Glyphis gangeticus.
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Geographic Range

The Bull Shark inhabits coastal waters in tropical and subtropical seas worldwide. (Enchanted Learning 2000)

Biogeographic Regions: australian (Native ); oceanic islands (Native ); indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native )

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Cosmopolitan in tropical seas (including Madagascar, Mascarenes), enters freshwater rivers and lakes.
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Depth

Depth Range (m): 0 (S) - 150 (S)
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Range

Occurs along the continental coasts of all subtropical and tropical seas, and can often be found in freshwater rivers and lakes as well (2) (4).
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Physical Description

Morphology

Dorsal spines (total): 0; Dorsal soft rays (total): 0; Analspines: 0; Analsoft rays: 0
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Physical Description

The Bull Shark can be recognized by its unique body shape, which is much wider in comparison to its length than other sharks, and its snout, which is wider than it is long. These features give the Bull Shark an almost stout appearance. The shark is gray on the top half of its body and off white underneath. Several individuals have been found with pale stripes on the sides of their bodies. The Bull Shark also has two dorsal fins, the second of which is much smaller than the first. Males of the species are approximately 7 feet long and weich 90 kg while females grow to 11.4 feet on average and weigh 230 kg. The young sharks can be distinguished by the dark edges on their fins. (Australian Museum 1999, Enchanted Learning 2000)

Range mass: 90 to 230 kg.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: female larger

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Size

Length max (cm): 350.0 (S)
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Size

Length: 305 cm

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Maximum size: 3500 mm TL
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Max. size

400 cm TL (female); max. published weight: 316.5 kg (Ref. 40637); max. reported age: 32 years (Ref. 42004)
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Diagnostic Description

Description

Euryhaline, found in coastal waters including bays, estuaries, and rivers. Occasionally found in the surf zone (Ref. 9710). Feeds on small sharks, rays, and sea urchins. Viviparous, with a yolk-sac placenta, 1 to 13 young per litter. Very hardy and lives well in captivity. The most dangerous species of tropical shark. Has repeatedly been implicated in attacks on humans (Ref. 4967). Utilized for human consumption, fins for soup, hides for leather, and liver for oil. Sympatric with @C. amboinensis, G. gangeticus@.
  • Anon. (1996). FishBase 96 [CD-ROM]. ICLARM: Los Baños, Philippines. 1 cd-rom pp.
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A massive shark with a short, broad and blunt snout, small eyes and triangular saw-edged upper teeth; and lack of interdorsal ridge are sufficient to distinguish this species (Ref. 26938). Teeth in the upper jaw are upright, strongly serrated, triangular and broad at their bases, while in the lower jaw they are more slender and pointed (Ref. 12484, 81283, 81623). Symphysial teeth present on both jaws, tooth formula 12-14/12-13 on each side of jaws (Ref. 81623). Spiracles absent, posterior lateral gill-slits overlapping origin of pectoral fins (Ref. 81623). First dorsal fin broad and triangular and less than 3.2 times height of second dorsal fin; beginning slightly before anal-fin origin; no interdorsal ridge (Ref. 5578, 81253). Second dorsal fin much smaller than first, situated over anal fin (Ref. 81623). Pectoral fins rather large and broad (Ref. 81623). Precaudal vertebral centra 101-103, total vertebral centra 198-227 (Ref. 81623). The colour of the body is predominantly grey with a pale to white underside (Ref. 5578, 12484). Juveniles have black tips to their fins, which fade with age (Ref. 9997, 12484).
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Type Information

Type for Carcharhinus leucas
Catalog Number: USNM 16887
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Fishes
Preparation: Unknown
Collector(s): D. Bransford
Locality: Lake Nicaragua, Nicaragua, North America
  • Type: Gill, T. N. 1877. Proceedings of the Academy of Natural Sciences of Philadelphia. 29: 190.
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Ecology

Habitat

Habitat Type: Marine

Comments: Often in shallow coastal waters, low salinity situations; sometimes ascends rivers (only North American shark to do so).

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nektonic
  • North-West Atlantic Ocean species (NWARMS)
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Habitat and Ecology

Habitat and Ecology
The Bull Shark is one of the few shark species that can tolerate long periods of time in freshwater, often penetrating long distances up freshwater rivers that connect to the ocean. For example, Thorson (1971) reported the movement of Bull Sharks from the ocean through the Rio San Juan system and into Lake Nicaragua, a large freshwater lake. C. leucas move into estuarine or fresh waters to give birth to their young (Springer 1963), while neonates and young juveniles appear to use these habitats as nursery areas (Snelson et al. 1984).

Bull Sharks grow up to about 340 cm in total length. The young are born at between 56?81 cm. Males mature at 157?226 cm and females at 180?230 cm (Compagno 1984b).

The diet of this species is relatively diverse, including turtles, birds, dolphins, terrestrial mammals, crustaceans, echinoderms, teleost fishes and elasmobranchs (Last and Stevens 1994). However, the most diverse diet is restricted to larger individuals that are capable of consuming larger prey. The most commonly eaten prey items are teleost fishes and elasmobranchs. In the coastal lagoons of Florida, Snelson and Williams (1981) recorded a wide array of species in the diet including jacks, snook, tarpon, mullets, catfish, croaker, stingrays and sandbar sharks, but noted that saltwater catfish and stingrays (Dasyatis spp.) were most commonly eaten.

Reproduction is by placental viviparity. Litter sizes range from 1?13 (Compagno 1984b), with most between 6?8 (Pattillo et al. 1997). The gestation period is 10?11 months, with birth normally occurring in late spring and summer (Clark and von Schmidt 1965, Bass et al. 1973, Branstetter 1981). In warmer areas (e.g., Nicaragua) breeding (and hence parturition) may occur year?round (Castro 1983). The length of the reproductive cycle has not been published, but is probably biennial (Compagno in prep. b).

Thorson and Lacy (1982) and Branstetter and Stiles (1987) have provided age and growth data for this species. Thorson and Lacy (1982) used tag recapture information to estimate the growth rates of Bull Sharks in Lake Nicaragua. They estimated that the growth rates in the first two years of life were 18 and 16 cm per year, respectively. Growth subsequently slowed to 11?12 cm per year and finally to 9?10 cm per year. They estimated that females would live up to 16 years and males to 12 years.

Branstetter and Stiles (1987) used vertebral ageing techniques to estimate growth parameters for animals from the northern Gulf of Mexico. They estimated the von Bertalanffy parameters to be L?=285 cm, k=0.076 year-1 and t = -3.0 years. The oldest estimated male was 21.3 years and the oldest female 24.2 years. Their estimates of growth for early years were similar to those of Thorson and Lacy (1982), but growth in later years was thought to be much slower (4?5 cm per year), accounting for the larger maximum ages. Ages at maturity based on Branstetter and Stiles (1987) data are 14?15 years for males and 18+ years for females.

A number of migratory habits have been documented for this shark. Pregnant females migrate to estuarine areas to give birth. The juveniles remain in these areas until temperatures drop below optimum levels and then migrate to warmer offshore waters. A general migration along the United States east coast is also observed, with movement northwards during the summer as water temperatures rise and southwards again as temperatures cool in the north (Castro 1983).

Smith et al. (1998) have reported the results of demographic analysis for C. leucas. The technique that these authors used estimated the rebound potential (r2M, similar to the intrinsic rate of increase) from litter size, age at maturity, maximum age and natural mortality. They estimated natural mortality to be 0.166 year-1 based on a maximum age of 27 years. The estimated the rebound potential was 0.027?0.039 year-1.

Systems
  • Freshwater
  • Marine
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Environment

reef-associated; amphidromous (Ref. 51243); freshwater; brackish; marine; depth range 1 - 152 m (Ref. 30573), usually 1 - 30 m (Ref. 55183)
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Although Bull Sharks have been caught in considerably deeper water, they most often reside in water between 30 meters and waist deep. The sharks also seem to favor murky water for hunting. It is one of the only sharks that is able to survive in freshwater for extended periods of time. (Australian Museum 1999, Smith 1999)

Aquatic Biomes: reef ; lakes and ponds; rivers and streams; coastal

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Depth range based on 19 specimens in 1 taxon.
Water temperature and chemistry ranges based on 13 samples.

Environmental ranges
  Depth range (m): 1 - 1860
  Temperature range (°C): 4.216 - 26.358
  Nitrate (umol/L): 0.520 - 29.561
  Salinity (PPS): 34.899 - 36.025
  Oxygen (ml/l): 2.883 - 5.086
  Phosphate (umol/l): 0.131 - 1.950
  Silicate (umol/l): 0.567 - 28.497

Graphical representation

Depth range (m): 1 - 1860

Temperature range (°C): 4.216 - 26.358

Nitrate (umol/L): 0.520 - 29.561

Salinity (PPS): 34.899 - 36.025

Oxygen (ml/l): 2.883 - 5.086

Phosphate (umol/l): 0.131 - 1.950

Silicate (umol/l): 0.567 - 28.497
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Depth: 1 - 152m.
From 1 to 152 meters.

Habitat: demersal. Zambezi / Bull Shark.  (Müller & Henle 1839) Attains 3 Metres. Prominantly grey with a pale underside, a rounded blunt snout and sharp serrated triangular teeth. It feeds in muddy water and preys on bony fish such as mullet and spotted grunter which form 50% of the diet. Small sharks, dolphins, skates and turtles are also part of the diet. The remains of birds, land mammals, miscellaneous debris and crabs are also found in their stomachs. This shark is remarkable for its ability to live in fresh water for indefinite periods. It has been recorded hundreds of kilometres upstream in large rivers such as the Zambezi and lake Nicaragua in Central America. Litter contains 10 -12 pups born at 60 -70 cms. The juveniles are often seen spinning out of the water, apparently in an attempt to rid themselves  of external parasites. Adults occur in shallow inshore coastal waters in estuaries and rivers. The Zambezi shark has been responsible for several, sometimes fatal attacks on bathers along the East Coast of southern Africa. Worldwide distribution in warmer seas.
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Salinity: Marine, Brackish, Freshwater

Inshore/Offshore: Inshore, Inshore Only

Water Column Position: Surface, Near Surface, Mid Water, Near Bottom, Bottom + water column

Habitat: Reef (rock &/or coral), Rocks, Corals, Reef and soft bottom, Reef associated (reef + edges-water column & soft bottom), Soft bottom (mud, sand,gravel, beach, estuary & mangrove), Mud, Sand & gravel, Beach, Estuary, Mangrove, Freshwater, Water column

FishBase Habitat: Demersal
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The bull shark is most frequently found in coastal waters between 30 and 50 metres deep, but makes occasional deep dives to well over 100 metres; the maximum depth recorded is 204.4 metres for a bull shark from Fiji (4). It also commonly enters estuaries, bays and harbours, and penetrates far into freshwater, inhabiting rivers and lakes (2).
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Migration

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

Gulf of Mexico population migratory.

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Amphidromous. Refers to fishes that regularly migrate between freshwater and the sea (in both directions), but not for the purpose of breeding, as in anadromous and catadromous species. Sub-division of diadromous. Migrations should be cyclical and predictable and cover more than 100 km.Characteristic elements in amphidromy are: reproduction in fresh water, passage to sea by newly hatched larvae, a period of feeding and growing at sea usually a few months long, return to fresh water of well-grown juveniles, a further period of feeding and growing in fresh water, followed by reproduction there (Ref. 82692).
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Trophic Strategy

Comments: Feeds opportunistically on fishes, porpoises, sea turtles, mollusks, crustaceans, and refuse.

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Juveniles move upstream and may be found several km from the sea, adults found near estuaries (Ref. 4967).
  • Wetherbee, B.M., S.H. Gruber and E. Cortes 1990 Diet, feeding habits, digestion, and consumption in sharks, with special reference to the lemon shark, Negaprion brevirostris. p. 29-47. In H.L. Pratt, Jr., S.H. Gruber and T. Taniuchi (eds.) Elasmobranchs as living resources: advances in the biology, ecology, systematics, and the status of the fisheries. NOAA Tech. Rep. NMFS 90. 517 p. (Ref. 568)   http://www.fishbase.org/references/FBRefSummary.php?id=568&speccode=139 External link.
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Food Habits

The Bull Shark is an omnivorous animal. It routinely preys upon fish, sharks (especially young sandbar sharks), rays, turtles, echinoderms, birds, mollusks, dolphins, and almost anything else it can find. Remains of everything from humans to hippopotami have been found in Bull Sharks' stomachs. (Australian Museum 1999, Bilson and Bilson 1999)

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Feeding

Feeding Group: Carnivore

Diet: mobile benthic crustacea (shrimps/crabs), mobile benthic gastropods/bivalves, octopus/squid/cuttlefish, bony fishes, sharks/rays, sea snakes/mammals/turtles/birds
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Associations

Known prey organisms

Carcharhinus leucas preys on:
Actinopterygii
Aves
Zalophus californianus

Based on studies in:
USA: Florida, Everglades (Estuarine)

This list may not be complete but is based on published studies.
  • W. E. Odum and E. J. Heald, The detritus-based food web of an estuarine mangrove community, In Estuarine Research, Vol. 1, Chemistry, Biology and the Estuarine System, Academic Press, New York, pp. 265-286, from p. 281 (1975).
  • Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2006. The Animal Diversity Web (online). Accessed February 16, 2011 at http://animaldiversity.org. http://www.animaldiversity.org
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Life History and Behavior

Life Cycle

Viviparpous, with a yolk-sac placenta, 1-13 young in a litter. Size at birth about 60 cm TL (Ref. 9997). In the western North Atlantic off Florida and the Gulf of Mexico, and off South Africa, young are born in late spring or early summer. Off Nicaragua, females may have young throughout the year, with a peak in spring and early summer. Estimated gestation period is 10 to 11 months (Ref. 244). Distinct pairing with embrace (Ref. 205). Females often have courtship scars, but males are rarely seen with fighting scars (Ref. 244).
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Life Expectancy

Lifespan/Longevity

Average lifespan

Status: wild:
13.7 years.

Average lifespan

Status: wild:
28.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 32 years (wild)
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Reproduction

Gives birth to litter of 5-10 young, April-June along central Gulf coast of Florida.

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Bull Sharks are viviparous, meaning they give birth to live young that are nourished inside the mother shark. Sexual maturity is reached between the ages of 8 - 10. Bull Sharks breed in the summer months and the young sharks are born approximately one year later. The pups are born in litters of up to 13 and are around 28 inches at birth. A common breeding place for the Bull Shark is the brackish water where freshwater rivers meet the saltwater oceans. (Microsoft Encarta 1997, Enchanted Learning 2000)

Average age at sexual or reproductive maturity (male)

Sex: male:
6296 days.

Average age at sexual or reproductive maturity (female)

Sex: female:
6570 days.

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Egg Type: Live birth, No pelagic larva
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Carcharhinus leucas

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 26 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

CTTTATCTGATTTTTGGTGCATGAGCAGGTATAGTTGGTACAGCCCTA---AGTCTCCTAATTCGAGCTGAACTTGGACAACCTGGATCACTTTTAGGAGAT---GATCAGATCTATAATGTAATCGTAACTGCCCACGCTTTTGTAATAATCTTTTTCATAGTTATACCAATTATAATTGGTGGTTTCGGAAATTGATTAGTTCCTTTAATA---ATTGGTGCACCAGACATAGCCTTTCCACGAATAAATAACATAAGTTTCTGACTTCTTCCACCATCATTTCTTCTTCTCCTCGCCTCTGCTGGAGTAGAAGCTGGAGCAGGTACTGGTTGAACAGTCTATCCTCCATTAGCTAGCAATCTAGCACATGCTGGGCCATCTGTTGACTTA---GCTATCTTCTCCCTTCACTTAGCTGGTGTTTCATCAATTTTAGCTTCAATTAATTTTATTACAACTATTATTAATATAAAACCACCAGCCATTTCCCAATATCAAACACCATTATTTGTTTGATCCATTCTTGTAACCACTATTCTTCTTCTCCTCTCACTCCCAGTCCTTGCAGCA---GGGATTACAATATTACTTACAGATCGTAACCTTAATACTACATTCTTTGATCCTGCAGGTGGAGGAGACCCAATCCTTTATCAACACTTA------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------TTT
-- end --

Download FASTA File

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Statistics of barcoding coverage: Carcharhinus leucas

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 29
Specimens with Barcodes: 64
Species With Barcodes: 1
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Conservation

Conservation Status

National NatureServe Conservation Status

United States

Rounded National Status Rank: N5 - Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

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IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2009

Assessor/s
Simpfendorfer, C. & Burgess, G.H.

Reviewer/s
Musick, J.A. & Fowler, S.L. (Shark Red List Authority)

Contributor/s

Justification
This assessment is based on the information published in the 2005 sharks status survey (Fowler et al. 2005).

The Bull Shark (Carcharhinus leucas) is a common tropical and subtropical species that occurs in marine, estuarine and freshwater. It is the only species of shark that can exist for long periods in freshwater and penetrates long distances up large rivers. It is caught in fisheries throughout its range, but it is rarely a target species. Its occurrence in estuarine and freshwater areas makes it more vulnerable to human impacts and habitat modification.
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IUCN Red List of Threatened Species: near threatened

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IUCN Red List: Listed, Near threatened

CITES: Not listed
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Status

Classified as Lower Risk / Near Threatened (LR/nt) by the IUCN Red List (1).
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Population

Population Trend
Unknown
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Threats

Major Threats
The frequent use of estuarine and freshwater areas by the Bull Shark makes it more susceptible to deleterious human impacts than species of sharks occurring in other coastal or offshore areas. Bull sharks more frequently encounter humans while in waters of low salinity, and are thereby subjected to increased fishing pressure and environmental changes associated with habitat modification.

Bull Sharks are commonly caught in both commercial and recreational fisheries. Thorson (1982a) reported that a commercial fishery existed for C. leucas in Lake Nicaragua and the Rio San Juan river system in Central America. However, in most situations, Bull Sharks are not normally a fishery target species but are caught as bycatch or as part of a multi-species fishery. For example, in the US Atlantic region they are an important component of inshore ecosystems, but only comprise 1?6% of the large coastal shark catch for this area (Branstetter and Burgess 1997).

While the Bull Shark has been exploited commercially for its skin, liver oil and flesh, currently its fins are the major product driving demand for this and many other species. There are limited data on recreational catches of this species. The best data come from the Gulf of Mexico where Casey and Hoey (1985) reported that in 1978 C. leucas made up about 11% (by weight) of the recreational shark catch of around three million pounds (Casey and Hoey 1985). Recreational catches of large sharks in the Gulf of Mexico have decreased substantially since the 1970s, but Casey and Hoey?s results illustrate that recreational fishing may have a substantial impact on Bull Shark populations.

Beach protection programmes in KwaZulu-Natal, South Africa and Queensland, Australia also regularly catch Bull Sharks. Cliff and Dudley (1992) reported that between 1978 and 1990 the South African programme caught 59 Bull Sharks, 21% of which were released alive. Species identification problems occurred in the Queensland programme until the early 1990s, thus the importance of C. leucas can only be based on data from latter years. Gribble et al. (1998) reported that after identification was improved 16% of the sharks caught state-wide were Bull Sharks, with the majority caught in the central part of the state. The lack of historical data for the Queensland programme and the low abundance in the South African programme make it impossible to assess the impact of beach meshing on Bull Shark populations.

The location of nursery areas in estuarine and freshwater systems makes the species vulnerable to pollution and habitat modification, but there has been only limited study of these impacts on Bull Sharks. Canal developments have been prolific in some estuarine areas where the species is commonly found. It is not known whether these developments have negative impacts. In Florida, USA and the Gold Coast of Queensland, Australia, these developments have substantially altered the environment. Bull Sharks occur frequently in Gold Coast canals and the species has been responsible for a number of attacks on humans (Simpfendorfer unpubl.). The warm water effluent from power stations may also impact this shark. In Florida, USA, juveniles have been reported to be trapped in the warm water outfalls during winter when they would normally have migrated to warmer water areas (Snelson et al. 1984, C. Manire, Center for Shark Research, Mote Marine Laboratory pers. Comm.). The potential impacts of pollution and habitat modification need to be further investigated for this species.

This shark is also exploited by large aquariums. The species is good for public display, adapting well to life in a tank and providing a good example of a larger, aggressive shark. With the number of public aquaria rising worldwide there is an increasing demand for this and other species of sharks for display. While populations are healthy the needs of aquaria can probably be met without affecting the wild population. However, if a wild subpopulation is depleted this may not be the case and aquaria need to have responsible collection policies that will not result in further pressure being placed on a species. At present there is no evidence that collecting for aquariums has any impact on the wild population of Bull Shark.
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Near Threatened (NT)
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Although the bull shark is rarely the target species of commercial fisheries, it is often caught as by-catch throughout its range (1), and its abundance in inshore habitats make it a target of artisanal fisheries (2) (3). When captured, the meat is consumed by humans or used in fishmeal, the hide is used for leather, the fins are used in shark-fin soup and the liver is utilised for its vitamin-rich oil (2). In certain areas, such as the Gulf of Mexico and South Africa, the bull shark is also a popular game fish (2). The bull shark's inshore and freshwater habitat not only makes it an easier target of fisheries, but these are habitats that are particularly vulnerable to habitat modification and pollution caused by human activities (1).
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Management

Conservation Actions

Conservation Actions
No specific management or conservation programmes are known for this species. It is managed in the U.S. east coast shark fisheries as part of the ?large coastal? groups of species. The current quota for this group is 1,285 t/year (1997), but Bull Sharks make up only small percentage of this group (see above). Recent closures of coastal waters in several states in the southern US to gillnetting have removed pressure on the juveniles in estuarine and coastal nursery areas.
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Conservation

Whilst the IUCN consider there to be insufficient information to determine the extent to which the bull shark may be threatened, its proximity to human populations, with their hunting and environmentally damaging activities, is likely to be greatly impacting populations (1). Further research is evidently required, to determine the conservation status of the bull shark, and subsequently enable appropriate conservation measures to be implemented to ensure the future of this unique shark.
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Relevance to Humans and Ecosystems

Benefits

Importance

fisheries: commercial; gamefish: yes
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Economic Importance for Humans: Negative

Bull Sharks have an appetite for young Sandbar Sharks. Because many Sandbar Sharks do not reach maturity, this has a negative impact on the large commercial businesses that fish for them. (Smith 1999)

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Economic Importance for Humans: Positive

The Bull Shark is one of the most commonly caught sharks in the world. It is frequently used as food in coastal areas and its skin is used to make leather. (Smith 1999)

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Wikipedia

Bull shark

The bull shark (Carcharhinus leucas), also known as the Zambezi shark or, unofficially, as Zambi in Africa and Nicaragua shark in Nicaragua, is a requiem shark commonly found worldwide in warm, shallow waters along coasts and in rivers. The bull shark is known for its aggressive nature, predilection for warm shallow water, and presence in brackish and freshwater systems including estuaries and rivers.

Bull sharks can thrive in both saltwater and freshwater and can travel far up rivers. They have even been known to travel as far up the Mississippi River as Illinois,[2] although there have been few recorded freshwater attacks. They are probably responsible for the majority of near-shore shark attacks, including many attacks attributed to other species.[3]

Unlike the river sharks of the genus Glyphis, bull sharks are not true freshwater sharks, despite their ability to survive in freshwater habitats.

Etymology[edit]

The name "bull shark" comes from the shark's stocky shape, broad, flat snout, and aggressive, unpredictable behavior.[4] In India, the bull shark may be confused with the Sundarbans or Ganges shark. In Africa, it is also commonly called the Zambezi River shark or just Zambi. Its wide range and diverse habitats result in many other local names, including Ganges River shark, Fitzroy Creek whaler, van Rooyen’s shark, Lake Nicaragua shark,[5] river shark, freshwater whaler, estuary whaler, Swan River whaler,[6] cub shark, and shovelnose shark.[7]

Anatomy and appearance[edit]

A female Carcharhinus leucas at the Shark Reef Marine Reserve, Fiji
A sketch of a bull shark

Bull sharks are large and stout, with females being larger than males. The bull shark can be up to 81 cm (2.66 ft) in length at birth.[8] Adult female bull sharks average 2.4 m (7.9 ft) long and typically weigh 130 kg (290 lb), whereas the slightly smaller adult male averages 2.25 m (7.4 ft) and 95 kg (209 lb). While a maximum size of 3.5 m (11 ft) is commonly reported, there is a questionable record of a female specimen of exactly 4 m (13 ft). The maximum recorded weight of a bull shark was 315 kg (694 lb) but may be larger.[3][9][10] Bull sharks are wider and heavier than other requiem sharks of comparable length, and are grey on top and white below. The second dorsal fin is smaller than the first. The bull shark's caudal fin is longer and lower than that of the larger sharks, and it has a small snout, and lacks an interdorsal ridge.[8]

Bull sharks have a bite force of up to 600 kilograms (1,300 lb), pound for pound the highest among all investigated cartilaginous fishes.[11]

Distribution and habitat[edit]

The bull shark is commonly found worldwide in coastal areas of warm oceans, in rivers and lakes, and occasionally salt and freshwater streams if they are deep enough. It is found to a depth of 150 metres (490 ft), but does not usually swim deeper than 30 metres (98 ft).[12] In the Atlantic, it is found from Massachusetts to southern Brazil, and from Morocco to Angola. In the Indian Ocean, it is found from South Africa to Kenya, India, and Vietnam to Australia.

Populations of bull sharks are also found in several major rivers, with more than 500 bull sharks thought to be living in the Brisbane River. One was reportedly seen swimming the flooded streets of Brisbane, Queensland, Australia, during the Queensland floods of late 2010/early 2011.[13] Several were sighted in one of the main streets of Goodna, Queensland, shortly after the peak of the January 2011, floods.[14] There are greater numbers still in the canals of the Gold Coast, also in Queensland. A large bull shark was caught in the canals of Scarborough, two hours north of the Gold Coast.[15] In the Pacific Ocean, it can be found from Baja California to Ecuador. The shark has traveled 4,000 kilometres (2,500 mi) up the Amazon River to Iquitos in Peru.[16] It also lives in fresh water Lake Nicaragua, in the Ganges and Brahmaputra rivers of West Bengal and Assam in eastern India and adjoining Bangladesh. It can live in water with a high salt content as in St. Lucia Estuary in South Africa. The bull shark is generally prolific in the warm coastal waters and estuarine systems of the Mozambique Channel and southward, including Kwa-Zulu Natal and Mozambique. The species has a distinct preference for warm currents.

After Hurricane Katrina, many bull sharks were sighted in Lake Pontchartrain.[17] Bull sharks have occasionally gone up the Mississippi River as far upstream as Alton, Illinois.[18] They have also been found in the Potomac River in Maryland.[19][20] A golf course lake in Queensland, Australia is the home to several bull sharks. They are believed to have become trapped following a flood in the 1990s. The golf course has capitalized on the novelty and now hosts a monthly tournament called the "Shark Lake Challenge."[21]

Behavior[edit]

Freshwater tolerance[edit]

The bull shark is the best known of 43 species of elasmobranch in ten genera and four families to have been reported in fresh water.[22] Other species that enter rivers include the stingrays (Dasyatidae, Potamotrygonidae and others) and sawfish (Pristidae). Some skates (Rajidae), smooth dogfishes (Triakidae), and sandbar sharks (Carcharhinus plumbeus) regularly enter estuaries. Elasmobranchs' ability to enter fresh water is limited because their blood is normally at least as salty (in terms of osmotic strength) as seawater through the accumulation of urea and trimethylamine oxide, but bull sharks living in fresh water show a significantly reduced concentration of urea within their blood.[23] Despite this, the solute composition (i.e. osmolarity) of a bull shark in freshwater is still much higher than that of the external environment. This results in a large influx of water across the gills due to osmosis and loss of sodium and chloride from the shark's body. However, bull sharks in freshwater possess several organs with which to maintain appropriate salt and water balance; these are the rectal gland, kidneys, liver and gills. All elasmobranchs have a rectal gland which functions in the excretion of excess salts accumulated as a consequence of living in seawater. Bull sharks in freshwater environments decrease the salt-excretory activity of the rectal gland, thereby conserving sodium and chloride.[24] The kidneys produce large amounts of dilute urine, but also play an important role in the active reabsorption of solutes into the blood.[24] The gills of bull sharks are likely to be involved in the uptake of sodium and chloride from the surrounding freshwater,[25] whereas urea is produced in the liver as required with changes in environmental salinity.[26]

Initially, scientists thought the sharks in Lake Nicaragua belonged to an endemic species, the Lake Nicaragua shark (Carcharhinus nicaraguensis). In 1961, following specimens comparisons, taxonomists synonymized them.[27] They can jump along the rapids of the San Juan River (which connects Lake Nicaragua and the Caribbean Sea), almost like salmon.[12] Bull sharks tagged inside the lake have later been caught in the open ocean (and vice versa), with some taking as little as seven to eleven days to complete the journey.[27]

Bull sharks are able to regulate themselves to live in either fresh water or salt water. It is possible for the bull shark to live in fresh water for its entire life, but it has been observed that this does not happen for certain reasons, mostly due to reproduction. Young bull sharks will leave the brackish water in which they are born and move out into the sea in order to breed with bull sharks of the other sex. While theoretically, it may be possible for bull sharks to live in purely freshwater, it was observed that the bull sharks that were being experimented on had died within four years. The stomach was opened and all that was found were 2 small fishes that were unidentifiable. The cause of death could have been starvation since the primary food source for bull sharks resides in salt water.[28]

The bull shark is known to be able to live in both saltwater and freshwater. It was found in a research experiment that, the majority of the time, the bull sharks were found to be at the mouth of an estuary.[29] The water salinity was tested every month to determine if the difference in salinity is what drove bull sharks into the river. It was found that the bull shark stayed at the mouth of the river independent of the salinity of the water.[29] The driving factor for a bull shark to be in freshwater or saltwater is its age; as the bull shark ages the tolerance for very low or high salinity increases.[29] It was found that the majority of the newborn or very young bull sharks were found in the freshwater area, whereas the much older bull sharks were found to be in the saltwater, as they had developed a much better tolerance for the salinity.[29] This is one of the reasons why adult bull sharks will travel into the river (in order to reproduce). The newborns are not born with a high tolerance for high salinity, so they are born in freshwater and stay there until they are able to travel out.

Diet[edit]

The bull shark's diet consists mainly of bony fish and small sharks, including other bull sharks,[3] but can also include turtles, birds, dolphins, terrestrial mammals, crustaceans, echinoderms, and stingrays. They hunt in murky waters because it is harder for the prey to see the shark coming.[1][30][31] Bull sharks have been known to use the bump-and-bite technique to attack their prey. After the first initial contact the bull shark continues to bite and tackle it's prey until they are unable to flee.[32] Bull sharks are believed to have attacked hippopotamuses in Africa's Zambezi River, based on bite marks on the hippos' feet.[33]

The bull shark is known to be a solitary hunter, although there are brief moments in which the bull sharks will team up with another bull shark in order to make it easier to hunt and to trick prey.[34][35]

Sharks are known to be opportunistic feeders,[32] and the bull shark is no exception to this, as it is part of the Carcharhinus family of sharks. Normally, sharks eat in short bursts, and when food is scarce, sharks digest for a much longer period of time in order to avoid starvation.[32] As part of their survival mechanism, bull sharks will regurgitate the food in their stomachs in order to escape from a predator. This is a distraction tactic, if the predator moves to eat the regurgitated food the bull shark can use the opportunity to escape.[36]

Reproduction[edit]

Bull sharks mate during late summer and early autumn,[37] often in the brackish water of river mouths. After gestating for 12 months, a bull shark may give birth to four to ten live young.[37] They are viviparous; they are born live and free-swimming. The young are about 70 cm (27.6 in) at birth and take 10 years to reach maturity. Coastal lagoons, river mouths, and other low-salinity estuaries are common nursery habitats.[3]

175 cm to 235 cm seems to be the size of a fully matured female bull shark that produce viable eggs for fertilization. The courting routine between bull sharks has not been observed in detail as of yet. It is speculated that the male bites the females on the tail until they turn upside down and they male can copulate at that point. At some points, the harassment of the male can become violent. It is not uncommon to see scratches and other marks on a mature female from the mating ritual.[38]

Bull sharks have an unusual migratory pattern in comparison to other sharks. They are found in rivers all over the world. They have the ability to go from seawater and freshwater. They give birth in the freshwater of rivers. The young bull sharks are free from predators while they grow up in the river before they go out to the sea in order to find mates.[39]

The ability to be able to survive in both freshwater and saltwater also gives another benefit that has been driven by evolution. Because the majority of sharks are only able to survive in saltwater, the bull shark has evolved to have their offspring in the freshwater where other sharks cannot enter.[40] The freshwater acts as a protective area where the young are able to grow and mature without the threat of larger sharks preying on the younger bull sharks.[40] This is an explanation for the behavior that is observed from the bull sharks as to why there would be any reason for the adult bull shark to ever travel into a freshwater area despite being able to tolerate the high salinity of marine water.

Bull sharks are born alive in freshwater which is in contrast to most sharks since they lay eggs. The size range of a litter for a female bull shark is around 1 to 13 pups.[41] The average time span for a female bull shark to be pregnant is around 10 to 11 months.[42] The male bull shark is able to begin reproducing around the age of 15 years while the female cannot begin reproducing until the age of 18 years.[42] Unlike most sharks though, the bull shark does not rear its young like other sharks, the young bull sharks are born into flat, protected areas.[42] Freshwater presents a natural defense against most larger predators, and the flat land is an added defense as most large predators will not swim in shallow areas. This increases their chance of survival since the parents do not rear the young in the traditional manner. This is also the reason why there is a high mortality rate in young bull sharks. Since the parents do not rear and protect the young, any predator that is able to attack a young bull shark is easily able to kill and eat the young bull shark without much resistance.[43]

Interactions with humans[edit]

Photo of bull shark in shallow water
Bull shark (Bahamas)

Since bull sharks often dwell in very shallow waters, they may be more dangerous to humans than any other species of shark,[12] and along with the tiger shark, oceanic whitetip and great white shark, are among the four shark species most likely to attack humans.[4]

One or several bull sharks may have been responsible for the Jersey Shore shark attacks of 1916, which were the inspiration for Peter Benchley's novel Jaws.[44] The speculation of bull sharks possibly being responsible is based on some attacks occurring in brackish and freshwater.

The bull shark is responsible for attacks around the Sydney Harbour inlets.[45] Most of these attacks were previously attributed to great white sharks. In India, bull sharks swim up the Ganges River and have attacked people. Many of these attacks have been attributed to the Ganges shark, Glyphis gangeticus, a critically endangered river shark species that is probably the only other shark in India able to survive in fresh water, although the sand tiger shark was also blamed during the 1960s and 1970s.

The bull shark prefers coastal water which is less than 100 feet in depth. This is mostly due to their feeding patterns since they prefer murky waters. This is also a problem since this gives the most interaction with humans. It is known that bull sharks inhabit areas off the coast of Florida, and there have been reports of bull sharks getting close enough to the coast to attack humans since the bull shark is a territorial animal.[46]

Visual cues[edit]

Behavioral studies have confirmed that sharks can take visual cues in order to discriminate between different objects.[47] The bull shark is able to discriminate between colors of mesh netting that is present underwater.[47] It was found that bull sharks tended to avoid mesh netting of bright colors rather than colors that blended in with the water. Bright yellow mesh netting was found to be easily avoided when it was placed in the path of the bull shark. This was found to be the reason that sharks are attracted to bright yellow survival gear rather than ones that were painted black.[47] This is very important because it gives an insight into how bull sharks are able to pick up certain visual keys underwater that might give them an advantage when seeking out certain prey.

Energy conservation[edit]

In 2008, researchers tagged and recorded the movements of young bull sharks in the Caloosahatchee River estuary. Specifically, they were testing to find out what determined the movement of the young bull sharks.[48] It was found that the young bull sharks synchronously moved downriver when the environmental conditions changed.[48] This large movement of young bull sharks were found to be moving as a response rather than other external factors such as predators. An interesting find was that the movement was directly related to the bull shark conserving energy for itself. One way the bull shark is able to conserve energy is that when the tidal flow changes, the bull shark uses the tidal flow in order to conserve energy as it moves downriver.[48] Another way for the bull shark to conserve energy is to decrease the amount of energy needed to osmoregulate the surrounding environment.[48]

Ecology[edit]

Bull sharks are apex predators and seldom have to fear being attacked by other animals. Humans are their biggest threat. Larger sharks, such as the tiger shark and great white shark, may attack them.[3] There have been stories documented in the media of saltwater crocodiles preying on young bull sharks in the rivers and estuaries of Northern Australia.[49]

See also[edit]

Notes and references[edit]

  1. ^ a b Simpfendorfer, C. & Burgess, G.H. (2005). "Carcharhinus leucas". IUCN Red List of Threatened Species. Version 2011.1. International Union for Conservation of Nature. Retrieved 18 August 2011. 
  2. ^ Sharks In Illinois. In-Fisherman (16 July 2012). Retrieved on 30 November 2013.
  3. ^ a b c d e "Bull shark". Florida Museum of Natural History. Retrieved 8 September 2006. 
  4. ^ a b "Bull shark". National Geographic. Retrieved 3 April 2011. 
  5. ^ "Biology of Sharks and Rays". ReefQuest Centre for Shark Research. Retrieved 19 August 2010. 
  6. ^ McGrouther, Mark (12 May 2010). "Bull Shark, Carcharhinus leucas Valenciennes, 1839 – Australian Museum". Australian Museum. Retrieved 19 August 2010. 
  7. ^ Allen, Thomas B. (1999). The Shark Almanac. New York: The Lyons Press. ISBN 1-55821-582-4. 
  8. ^ a b "Shark Species; Bull Sharks". Shark Diver Magazine 17: 34. 2003. 
  9. ^ "The Rosenstiel School of Marine & Atmospheric Science". 
  10. ^ "9 Biggest Sharks Ever Caught". Total Pro Sports.com. 
  11. ^ Habegger, M. L.; Motta, P. J.; Huber, D. R.; Dean, M. N. (2012). "Feeding biomechanics and theoretical calculations of bite force in bull sharks (Carcharhinus leucas) during ontogeny". Zoology. doi:10.1016/j.zool.2012.04.007.  edit; for a popular summary, see Walker, Matt (12 October 2012). "Bull sharks have strongest bite of all shark species". BBC News. Retrieved 12 October 2012. 
  12. ^ a b c Crist, Rick. "Carcharhinus leucas". University of Michigan Museum of Zoology, Animal Diversity Web. Retrieved 8 September 2006. 
  13. ^ "Queensland rebuilding 'huge task'". BBC News. 12 January 2011. 
  14. ^ Bull sharks seen in flooded streets | Offbeat | Weird News, Odd and Freaky Stories in Northern Rivers | Clarence Valley Daily Examiner. Dailyexaminer.com.au (14 January 2011). Retrieved on 4 May 2012.
  15. ^ Berrett, Nick (14 November 2008). "Canal shark shock". Redcliffe & Bayside Herald. Quest Community Newspapers. Retrieved 26 March 2009. 
  16. ^ Shark Gallery. Bull shark (Carcharhinus leucas). sharks-med.netfirms.com
  17. ^ High number of sharks reported in Lake Pontchartrain. wwltv.com. 16 September 2006
  18. ^ "Sharks in Illinois". In-Fisherman. Retrieved 26 July 2010. 
  19. ^ 8-Foot Shark Caught In Potomac River. Nbcwashington.com. Retrieved on 4 May 2012.
  20. ^ Zauzmer, Julie (22 August 2013). "Man catches 2 bull sharks in Potomac". Washington Post. 
  21. ^ "Shark-Infested Australian Golf Course Believed to Be World's First". Fox News. October 11, 2011. 
  22. ^ Compagno, Leonard I.V. and Cook, Sid F. (March 1995). "Freshwater elasmobranchs; a questionable future". Florida Museum of Natural History Ichthyology Department. Retrieved 27 April 2011. 
  23. ^ Pillans, R.D.; Franklin, C.E. (2004). "Plasma osmolyte concentrations and rectal gland mass of bull sharks Carcharhinus leucas, captured along a salinity gradient.". Comparative Biochemistry and Physiology A: Molecular and Integrative Physiology 138 (3): 363–371. doi:10.1016/j.cbpb.2004.05.006. PMID 15313492. 
  24. ^ a b Pillans, R.D.; Good, J.P., Anderson, W.G., Hazon, N and Franklin, C.E. (2005). "Freshwater to seawater acclimation of juvenile bull sharks (Carcharhinus leucas): plasma osmolytes and Na+/K+-ATPase activity in gill, rectal gland, kidney and intestine". Journal of Comparative Physiology B: Biochemical, Systemic, and Environmental Physiology 175 (1): 37–44. doi:10.1007/s00360-004-0460-2. PMID 15565307. 
  25. ^ Reilly, B.D.; Cramp, R.L., Wilson, J.M., Campbell, H.A and Franklin, C.E. (2011). "Branchial osmoregulation in the euryhaline bull shark, Carcharhinus leucas: a molecular analysis of ion transporters". The Journal of Experimental Biology 214 (17): 2883–2895. doi:10.1242/jeb.058156. PMID 21832131. 
  26. ^ Anderson, W.G.; Good, J.P., Pillans, R.D., Hazon, N and Franklin, C.E. (2005). "Hepatic urea biosynthesis in the euryhaline elasmobranch Carcharhinus leucas". Journal of Experimental Zoology Part A: Comparative Experimental Biology 303A (10): 917–921. doi:10.1002/jez.a.199. PMID 16161010. 
  27. ^ a b Fresh Waters: Unexpected Haunts. elasmo-research.org. Accessed 6 April 2008.
  28. ^ Montoya, Rafael Vasquez; Thorson, Thomas B. (1982). "The bull shark and largetooth sawfish in Lake Bayano, a tropical man-made impoundment in Panama". Env. Biol. Fish 7 (4): 341–347. doi:10.1007/BF00005568. 
  29. ^ a b c d Heupel, Michelle R.; Colin A. Simpfendorfer (2008). "Movement and distribution of young bull sharks Carcharhinus leucas in a variable estuarine environment". Aqutic Biology 1: 277–289. doi:10.3354/ab00030. 
  30. ^ Kindersley, Dorling (2001) in Animal, David Burnie and Don E. Wilson (eds.) London & New York: Smithsonian Institution, ISBN 0789477645.
  31. ^ Snelson, Franklin F; Mulligan, Timothy J; Williams, Sherry E. (1 January 1984). "Food Habits, Occurrence, and Population Structure of the Bull Shark, Carcharhinus Leucas, in Florida Coastal Lagoons". Bulletin of Marine Science 1: 71–80. 
  32. ^ a b c Motta, Philip J; Wilga, Cheryl D. (2001). "Advances in the study of feeding behaviors, mechanisms, and mechanics or sharks". Environmental Biology of Fishes 60 (60): 131–156. doi:10.1023/A:1007649900712. 
  33. ^ International Masters Publishers. Wildlife Explorer. Group 4, cards 33 and 4. ISBN 1-886614-77-6.
  34. ^ Bull Sharks, Carcharhinus leucas. Marinebio.org (14 January 2013). Retrieved on 30 November 2013.
  35. ^ Life of Bull Shark | Life of Sea. Life-sea.blogspot.com (15 November 2011).
  36. ^ Tuma, Robert E (1976). Investigations of the ichthyofauna of Nicaraguan lakes. American Society of Ichthyologists and Herpetologists. 
  37. ^ a b McAuley, R. B.; Simpfendorfer, C. A.; Hyndes, G. A. and Lenanton, R. C. J. (2007). "Distribution and reproductive biology of the sandbar shark, Carcharhinus plumbeus (Nardo), in Western Australian waters". Mar. Freshwater Res. 58 (1): 116–126. doi:10.1071/MF05234. "The proportion of mature males with running spermatozoa increased from 7.1% in October to 79 and 80% in January and March, respectively, suggesting that mating activity peaks during late summer and early autumn." 
  38. ^ Jenson, Norman H (1976). Investigation of the Icthyofauna of Nicaraguan Lakes. American Society of Ichthyologists and Herpetologists. 
  39. ^ Bres, M (1993). "The Behaviour of Sharks". Reviews in Fish Biology and Fisheries 3 (2): 133–159. doi:10.1007/BF00045229. 
  40. ^ a b Heupel, Michelle R.; Carlson, John K. and Simpfendorfer, Colin A. (14 May 2007). "Shark nursery areas: concepts, definitoin characterization and assumptions". Marine: Ecology Progress Series 337: 289–297. doi:10.3354/meps337287. 
  41. ^ Pacific Shark Research Center » Featured Elasmobranch – Bull Shark. Psrc.mlml.calstate.edu (16 February 2009). Retrieved on 30 November 2013.
  42. ^ a b c Fact Sheet: Bull Sharks. Sharkinfo.ch (15 October 1999). Retrieved on 30 November 2013.
  43. ^ Bull Shark – Animal Facts and Information. Bioexpedition.com. Retrieved on 30 November 2013.
  44. ^ Handwerk, Brian. "Great Whites May Be Taking the Rap for Bull Shark Attacks". National Geographic News. Retrieved 1 February 2007. 
  45. ^ Quinn, Ben (15 March 2009). "Shark attacks bring panic to Sydney's shore". The Guardian (London). Retrieved November 2009. 
  46. ^ Frantz, Vickie (18 July 2011). "Bull Sharks Attacks Comonly in Warm, Shallow Waters". accuweather. 
  47. ^ a b c Bres, M. (1993). "The behaviour of sharks". Reviews in Fish Bviology and Fisheries 3 (2): 133–159. doi:10.1007/BF00045229. 
  48. ^ a b c d Ortega, Lori A.; Heupel, Michelle R.; van Beynen, Philip and Motta, Philip J. (2009). "Movement patterns and water quality preferences of juvenile bull sharks (Carcharhinus lecuas) in a Florida estuary". Evionrmen Biol Fish 84 (4): 361–373. doi:10.1007/s10641-009-9442-2. 
  49. ^ "No Bull: Saltwater Crocodile Eats Shark". UnderwaterTimes.com. 13 August 2007. Retrieved 15 June 2008. 

General references[edit]

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Comments: Revision of LEUCUS-GANGETICUS group soon to be (or already) published by Garrick. C. AZUREUS is a synonym (Lee et al. 1980).

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