Overview

Brief Summary

Biology

The lemon shark has been the subject of one of the most long-term and intensive studies on a shark species. Much of what is known about this shark is due to the work of Dr Samuel Gruber and his colleagues. This predator is most active at dawn and dusk, and occurs singly or in loose aggregations of up to 20 individuals. It feeds primarily on fishes, including sea catfishes, mullet, stingrays and eagle rays, but also on crustaceans and molluscs. During the day they often lie quietly on the seabed, apparently resting, but in reality this behaviour uses up more energy than when swimming, due to the extra effort required to pump water over the gills (2). Therefore, they may be lying motionless waiting for wrasses or other small reef fishes to clean them of any parasites (3). The lemon shark is viviparous; the embryos develop inside the mother and receive nutrients via a yolk sac placenta. After a gestation period of 10 to 12 months, pregnant females enter shallow nursery areas in spring and summer to give birth to litters of 4 to 17 pups. The pups have a very slow growth rate and remain within nursery grounds for a considerable length of time, where they are less vulnerable to predation by larger sharks (2). The mangroves that the young frequently inhabit are highly productive waters, creating a marvellous site for feeding, but also an area of very low oxygen content. Luckily, the lemon shark has numerous adaptations that enhance oxygen uptake, such as blood with an unusually high affinity for oxygen, and thus the pups can lie feeding in the rich waters, protected from any large potential predators by the mangrove's tangled roots (2) (6). As they grow, their range expands dramatically, from six to eight kilometres up to around 300 kilometres. Maturity is reached at about six and a half years of age, and it is believed that the lemon shark lives for up to 27 years (2).
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Description

This stocky, powerful shark is named for its pale yellow-brown to grey skin, which lacks any distinctive markings. This provides perfect camouflage when swimming over the sandy seafloor in its coastal habitat (3). It has a flattened head with a short, broad snout, and the second dorsal fin is almost as large as the first (2). The lemon shark's retina has a specialized horizontal band across the middle, which is disproportionately rich in cones that discern fine detail and colour in well-illuminated conditions. This 'visual streak' is thought to provide the shark with a particularly clear view of its underwater world (4). This shark is potentially dangerous to humans due to its large size and powerful bite, and though there have been some unprovoked attacks, many were the result of provocation from divers and swimmers (2) (3).
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Comprehensive Description

Description

  Common names: shark (English), tiburón (Espanol), cazón (Espanol)
 
Negaprion brevirostris (Poey, 1868)


Lemon shark


A large stocky shark; snout short and broad, somewhat rounded; eyes small, with lower eyelid;  no spiracle; lip furrows at corners of mouth; mouth arched, ends well past eye; upper and lower front teeth with a single narrowly triangular straight point, mainly smooth, large fish have serrations on bases of upper teeth; 5 gill slits, last 2 over pectoral; spiracles usually absent (occasionally with very small spiracles); no ridge on back between dorsal fins; origin of first dorsal fin behind rear margin of pectoral fins; second dorsal fin 80-100% size of first; pectoral fins broad and slightly curved; anal fin large, a little smaller than second dorsal, with strongly notched rear margin;  pit on top of tail base is longitudinal, straight; tail asymmetric, with well developed lower lobe.


Pale yellow brown on back and sides, yellowish or whitish below.


Attains about 340 cm; size at birth 60-65 cm.

A common inshore shark seen in a variety of habitats including rocky reefs, estuaries, and river mouths (may enter fresh water).

Depth: 0-90 m.

Tropical waters of the Atlantic and eastern Pacific (southern Baja and the Gulf of California to Peru.   
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Biology

Occurs on continental and insular shelves, frequenting mangrove fringes, coral keys, docks, sand or coral mud bottoms, saline creeks, enclosed bays or sounds, and river mouths. May enter fresh water. Occasionally moves into the open ocean, near or at the surface, apparently for purposes of migration. May rest motionless on the bottom (Ref. 9710). May occur singly or in small groups. Feeds mainly on fish but also takes crustaceans and mollusks. Viviparous, with 4 to 17 young in a litter. Size at birth 60 to 65 cm. Has been involved in several attacks on people. Meat is utilized for human consumption, hides for leather, fins for shark-fin soup base, liver oil for vitamins, and carcasses for fish meal. Marketed fresh, dried-salted and frozen (Ref. 9987).
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Distribution

Western Atlantic: New Jersey, USA to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Caribbean; also in Gulf of Mexico
  • North-West Atlantic Ocean species (NWARMS)
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Range Description

This inshore species is common along the coasts in the Atlantic Ocean ranging from the US in the north down to southern Brazil and possibly in some areas on the West African Coast. It is not known whether these populations are the same species (Compagno 1984). Lemon Sharks also occur in the Pacific Ocean from Baja California in the north to Ecuador in the south.
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Zoogeography

See Map (including site records) of Distribution in the Tropical Eastern Pacific 
 
Global Endemism: All species, TEP non-endemic, East Pacific + Atlantic (East +/or West), East Pacific + all Atlantic (East+West)

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

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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Western Atlantic: New Jersey, USA to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Caribbean; also in Gulf of Mexico (Ref. 26938). Northeast Atlantic: Senegal, Côte d'Ivoire and probably wide-ranging off West Africa, but this requires confirmation. Eastern Pacific: southern Baja California, Mexico and the Gulf of California to Ecuador.
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Atlantic; Eastern Pacific.
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Depth

Depth Range (m): 0 (S) - 90 (S)
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Range

Occurs in the tropical western Atlantic, from New Jersey to southern Brazil; and in the north eastern Atlantic, off west Africa. It is also occasionally found in the eastern Pacific, from southern Baja California and the Gulf of California to Ecuador (2) (5).
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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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Physical Description

Morphology

Size

Length max (cm): 340.0 (S)
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Size

Maximum size: 3400 mm TL
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Max. size

340 cm TL (male/unsexed; (Ref. 26938)); max. published weight: 183.7 kg (Ref. 40637); max. reported age: 25 years (Ref. 31395)
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Diagnostic Description

A brownish shark with yellow overtones but no conspicuous markings. Large second dorsal fin nearly same size as first dorsal (Ref. 26938).
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Type Information

Type for Negaprion brevirostris
Catalog Number: USNM 28167
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Fishes
Collector(s): C. Gilbert
Year Collected: 1880
Locality: Mexico: Mazatlan., Sinaloa, Mexico, Pacific
  • : Jordan, D. S. & Gilbert, C. H. 1882. Proceedings of the United States National Museum. 5 (268): 102.; Type: Jordan, D. S. & Gilbert, C. H. 1882. Proceedings of the United States National Museum. 5 (268): 102.
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Ecology

Habitat

Habitat Type: Marine

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

Habitat and Ecology
The Lemon Shark inhabits shallow waters around coral keys, mangrove fringes, around docks, on sand or coral mud bottoms, in saline creeks, in enclosed sounds or bays and in river mouths. It may enter fresh water but has not been found far up in rivers (Compagno 1984b). Occasionally it ventures into the open ocean and has been found down at depths of at least 90 m (Springer 1950).

Mating occurs during spring and summer with parturition in shallow nursery grounds the following year after a 10?12 month gestation period (Bigelow and Schroeder 1948, Springer 1950, Clark and von Schmidt 1965). The female Lemon Shark gives birth to 4?17 young (Clark and von Schmidt 1965, Compagno 1984b) of 50?60 cm TL (Gruber and Stout 1983, Brown and Gruber 1988). Maturity is reached at 225 cm (males) and 235 cm (females) or at an age of 12 and 13 years, respectively (Compagno 1984b, Brown and Gruber 1988). Growth follows the von Bertalanffy equation (Brown and Gruber 1988):

PCL = 317.65 × (1-e -0.057 (t + 2.302)), n = 110, r2 = 0.99 , where PCL is precaudal length (m) at time t (yrs).

This equation assumes the maximum length to be 317 cm, but the lemon shark can become bigger. Hueter and Gruber (1982) examined a 368 cm large male. The normal size range of the adult is 250?290 cm with females being slightly bigger than males (Brown and Gruber 1988) but sizes of up to 3 m or more are not unusual (Clark and von Schmidt 1965). At this size the shark would have a weight of approximately 250 kg (Gruber 1984) and is probably more than 30 years old.

Activity space ranges from a few km² in the highly site-attached juveniles (Morrissey and Gruber 1993a) up to several hundred km² in the more active adults (Compagno 1984b). Juvenile Lemon Sharks appear to select shallow (0?50 cm) and warmer water (30°C or more). They also prefer rocky or sandy substrate (Morrissey and Gruber 1993b). Almost all field research on the Lemon Shark originates from the waters in and around the Bimini Lagoon, Bahamas where a high annual, density-dependent mortality rate (35?62%) for young-of-the-year Lemon Sharks was found (Gruber et al. 2001). This is probably due to predation by larger sharks (Manire and Gruber 1993). Jacobsen (1987) suggested that the same area could support about 250 juveniles while Henningsen and Gruber (1985) estimated the population to be around 500 specimens with a density of five sharks per km². The annual production of these Lemon Sharks was 320 kg corresponding to about 0.3 g of new Lemon Shark tissue for every m² of lagoon (Henningsen 1989). Later, Gruber et al. (2001) estimated that the maximum number of juveniles that could survive each year in the Bimini Lagoon was 30. Young Lemon Sharks feed mainly on teleosts, crustaceans (small portunid crabs and panaeid shrimp) and octopods. As they grow the diet becomes dominated by teleost and cartilaginous fishes and the adults even eat sea birds (Springer 1950, Cortés and Gruber 1990). The energy consumed and later used for growth depends on the daily feeding rate but maximum conversion rate is probably close to 25% (Cortés and Gruber 1994).

Systems
  • Freshwater
  • Marine
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Environment

reef-associated; oceanodromous (Ref. 51243); brackish; marine; depth range 0 - 92 m (Ref. 244)
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Depth range based on 10 specimens in 1 taxon.
Water temperature and chemistry ranges based on 8 samples.

Environmental ranges
  Depth range (m): 1 - 60
  Temperature range (°C): 22.006 - 23.236
  Nitrate (umol/L): 0.951 - 1.990
  Salinity (PPS): 36.017 - 36.251
  Oxygen (ml/l): 4.658 - 4.984
  Phosphate (umol/l): 0.090 - 0.424
  Silicate (umol/l): 1.391 - 2.629

Graphical representation

Depth range (m): 1 - 60

Temperature range (°C): 22.006 - 23.236

Nitrate (umol/L): 0.951 - 1.990

Salinity (PPS): 36.017 - 36.251

Oxygen (ml/l): 4.658 - 4.984

Phosphate (umol/l): 0.090 - 0.424

Silicate (umol/l): 1.391 - 2.629
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Depth: 0 - 92m.
Recorded at 92 meters.

Habitat: reef-associated.
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Salinity: Marine, Brackish

Inshore/Offshore: Inshore, Inshore Only

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

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

FishBase Habitat: Reef Associated
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The lemon shark occurs over continental shelves, from the surface down to a depth of least 92 meters. It usually occurs around corals keys, at the fringes of mangroves, around docks, in saline creeks, in enclosed bays and at river mouths. It occasionally travels short distances upriver, entering freshwater, and also ventures into the open ocean when migrating (2) (5).
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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.

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Oceanodromous. Migrating within oceans typically between spawning and different feeding areas, as tunas do. Migrations should be cyclical and predictable and cover more than 100 km.
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Trophic Strategy

Exhibits no pattern of periodicity. Feeding is asynchronous and intermittent. Daily ration is estimated at 1.5 - 2.1% body weight per day; meal completely evacuated from stomach 25-41 hr after feeding, depending on meal type and temperature; fecal production continues for 68-82 hr after feeding (Wetherbee 1990).
  • 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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Feeding

Feeding Group: Carnivore

Diet: mobile benthic crustacea (shrimps/crabs), mobile benthic gastropods/bivalves, octopus/squid/cuttlefish, bony fishes, sea snakes/mammals/turtles/birds
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Life History and Behavior

Life Cycle

Distinct pairing with embrace (Ref. 205). Viviparous, placental (Ref. 50449), with 5 to 17 embryos (Ref. 9253). Both male and female during precopulatory and courtship swim with body axes in parallel (Ref. 49562, 51112). During copulation, the pair performs coordinated swimming (Ref. 49562, 51112).
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Life Expectancy

Lifespan, longevity, and ageing

Maximum longevity: 20 years (wild)
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Reproduction

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

Molecular Biology

Barcode data: Negaprion brevirostris

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


There are 11 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.

CCTTTACTTGATTTTTGGTGCATGAGCAGGTATAGTTGGAACAGCCCTTAGTCTTTTGATTCGGGCTGAACTTGGCCAACCTGGATCACTTTTAGGGGATGACCAAATTTATAATGTAATCGTAACTGCCCACGCTTTTGTAATGATCTTTTTTATAGTTATGCCAATCATAATTGGTGGTTTCGGAAACTGACTAGTTCCATTAATAATTGGTGCACCAGATATAGCCTTCCCACGAATAAACAACATAAGTTTCTGACTTCTTCCACCATCATTTCTTCTTCTCCTCGCCTCTGCTGGAGTAGAGGCTGGAGCAGGTACTGGCTGAACAGTATATCCTCCATTAGCTAGTAACCTAGCCCATGCTGGACCATCTGTTGATTTAGCTATCTTCTCTCTTCACTTAGCCGGTGTTTCATCAATTTTAGCTTCAATTAACTTCATTACAACCATTATTAATATAAAACCACCAGCTATCTCCCAATATCAAACACCATTATTTGTTTGATCCATCCTTGTAACTACTATTCTCCTTCTCCTTTCACTTCCAGTTCTTGCAGCAGGGATTACAATATTACTTACAGATCGTAACCTTAATACTACATTCTTTGACCCTGCAGGTGGAGGAGATCCAATCCTTTNNNNNNNNNNN
-- end --

Download FASTA File
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Statistics of barcoding coverage: Negaprion brevirostris

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 10
Specimens with Barcodes: 14
Species With Barcodes: 1
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Genomic DNA is available from 2 specimens with morphological vouchers housed at British Antarctic Survey
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Conservation

Conservation Status

National NatureServe Conservation Status

United States

Rounded National Status Rank: NNR - Unranked

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NatureServe Conservation Status

Rounded Global Status Rank: GNR - Not Yet Ranked

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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
Sundström, L.F.

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 shark status survey (Fowler et al. 2005).

The Lemon Shark (Negaprion brevirostris) is a large coastal shark that is common in the Atlantic Ocean along the coasts of the United States to Brazil and possibly in some areas on the West African coast, as well as in the Pacific from Baja California to Ecuador. Young sharks are highly site attached but adults may undertake long migrations, possibly to deeper waters at the onset of winter. The species is caught both in commercial and recreational fisheries, but no management plans are implemented.
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IUCN Red List: Listed, Near threatened

CITES: Not listed
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Status

Classified as Near Threatened (NT) on the IUCN Red List (1).
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Population

Population Trend
Unknown
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Threats

Major Threats
Lemon Sharks are caught commercially on longlines and the meat is dried, salted, or smoked. The fins fetch a very high price. The Lemon Shark is consumed in the United States and in Central and South America (Rose 1996). The rough and heavy skin has made the lemon shark preferable among tanneries for the production of leather. However, it is not included in TRAFFIC Network?s list of species frequently appearing in available information on worldwide shark fisheries (Rose 1996). It is a target species in Belize, Mexico and USA and reported as bycatch in St Lucia (Oliver 1996, Anon. 1997). Lemon Sharks were seen at a fish market in Cameroon in 1991, but not since then (C. Grist pers. comm.). The species is also caught in recreational fishing and was reported as the 13th most common shark species in the US recreational fishery (Casey and Hoey 1985). A decrease in the number of juvenile Lemon Sharks between 1986?1989 in the lower Florida Keys may have been caused by several years of shark fishing tournaments and 20 years of targeting with gillnets affecting the return of females to bear new litters (Manire and Gruber 1990). The Lemon Shark is a popular aquarium species and it is also used extensively for research purposes. Lemon Sharks used to be common in the western Atlantic, from New Jersey, USA to Brazil, but lately their numbers have been depleted, especially around Florida (S.H. Gruber pers. comm.).
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Near Threatened (NT)
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The lemon shark is caught in commercial and recreational fisheries; for their meat, skin, fins for soup, and liver for vitamin-rich oil (2). There is some evidence indicating that this exploitation is depleting populations in the eastern Pacific and western Atlantic (1) (5). The shallow, coastal nursery grounds are susceptible to human-induced habitat degradation (1) (5), particularly mangrove swamps which are being destroyed to make way for shore front hotels and shops throughout the Gulf of Mexico, Bahamas and Caribbean region (6).
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Management

Conservation Actions

Conservation Actions
There are no management plans in place for the Lemon Shark. Some research, however, has dealt with related issues so there is a base of knowledge should a plan ever be implemented.
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Conservation

The lemon shark is not currently considered to be at risk from extinction, and there are no known specific conservation measures in place. However, it may gain some protection from the United Nations International Plan of Action for the Conservation and Management of Sharks. This requires that signatory states, such as Ecuador, Mexico and the United States, implement a national programme for the conservation and management of shark stocks, and carry out regular assessments of stocks (7).
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Relevance to Humans and Ecosystems

Benefits

Importance

fisheries: commercial; gamefish: yes
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Wikipedia

Lemon shark

The lemon shark (Negaprion brevirostris) is a stocky and powerful shark named for its yellow color.[2] A member of the family Carcharhinidae, lemon sharks can grow to 10 feet (3.0 m) in length.[3] They are often found in shallow subtropical waters and are known to inhabit and return to specific nursery sites for breeding.[4] Often feeding at night, these sharks use electroreceptors to find their main source of prey, fish.[5] Lemon sharks use the many benefits of group living such as enhanced communication, courtship, predatory behavior, and protection.[6] This species of shark is viviparous, and the females are polyandrous and have a biennial reproductive cycle.[4] Lemon sharks are not thought to be a large threat to humans.[7]

Taxonomy[edit]

The lemon shark was first named and described in 1868 by Felipe Poey.[7] He originally named it Hypoprion brevirostris, but later renamed it Negaprion brevirostris. [7] The lemon shark has also appeared in literature as Negaprion fronto and Carcharias fronto (Jordan and Gilbert, 1882), Carcharias brevirostris (Gunther, 1870), and Carcharhinus brevirostris (Henshall, 1891).[7]

Description[edit]

The shark's yellow colouring serves as a perfect camouflage when swimming over the sandy seafloor in its coastal habitat.[8] The lemon shark commonly attains a length of 2.4 to 3.1 m (7.9 to 10.2 ft) and a weight of up to 90 kg (200 lb) by adulthood, although sexual maturity is attained at 2.24 m (7.3 ft) in males and 2.4 m (7.9 ft) in females.[9] The maximum recorded length and weight is 3.43 m (11.3 ft) and 183.7 kg (405 lb).[10] It has a flattened head with a short, broad snout, and the second dorsal fin is almost as large as the first. Lemon sharks have electroreceptors concentrated in their heads, called the ampullae of Lorenzini.[11] These receptors detect electrical pulses emitted by potential prey and allow these nocturnal feeders to sense their prey in the dark.[11]

Distribution[edit]

Lemon sharks are found from New Jersey to southern Brazil in the tropical western Atlantic Ocean. They also live off the coast of west Africa in the southeastern Atlantic.[2] In addition, lemon sharks have been found in the eastern Pacific, from southern Baja California to Ecuador.[2] This species of shark often occupies the subtropical shallow waters of coral reefs, mangroves, enclosed bays, and river mouths; however, lemon sharks have also been found in the open ocean down to depths of 92 m.[12] Although lemon sharks do swim up rivers, they never seem to travel very far into fresh water. They are found in open water primarily during migrations, and tend to stay along the continental and insular[disambiguation needed] shelves for most of their lives.[13]

Habitat selection[edit]

A lemon shark with many remoras clinging to its body.

Information about activity patterns and the utilization of space is important in understanding a species’ behavioral ecology.[14] Animals often make decisions about habitat use by evaluating their environment’s abiotic conditions that serve as valuable indicators of good foraging sites or predator-safe locations.[15] Lemon sharks select habitats in water that is warm and shallow with a rocky or sandy bottom.[14]

The environmental temperature influences an individual’s body temperature, which ultimately affects physiological processes such as growth and metabolism.[15] Lemon sharks, therefore, select warm-water habitats to maintain optimal metabolic levels. They are believed to avoid areas with thick seagrasses because it makes finding prey more difficult.[15] Lemon sharks tend to live in or near shallow-water mangroves, which are often the nursery areas of several species of fish. One theory is that lemon sharks select mangrove habitats due to the abundance of prey that resides there, while another theory posits that mangroves provide a safe haven from adult lemon sharks that occasionally feed on juvenile sharks and are unable to enter the shallow waters.[16] Ontogenetic niche shifts, or changes in an animal’s niche breadth or position, to deeper waters are known to occur in relation to a lemon shark’s size. These changes occur due to the dramatic decrease in the risk of predation as body size increases.[15] Habitat selection clearly depends on a variety of biological and environmental variables.

The mangrove areas that lemon sharks inhabit are often referred to as their nursery sites. A nursery site is best defined as the most common area sharks are encountered, the location sharks tend remain after birth or frequently return to, and the habitat used by shark groups repeatedly for several years.[17] The nursery ground concept has been known and studied for at least a century. In addition, fossil evidence from 320 million years ago suggests the use of shallow, costal areas as pupping grounds is primitive.[17]

Feeding behavior[edit]

Solitary lemon shark at Tiger Beach, Bahamas

Lemon sharks have proven to be an ideal model species to challenge the belief that all sharks are asynchronous opportunistic predators due to their tendency to use nursery areas for an extended period of time.[18] Lemon shark feeding behaviors are easy to determine because their well-defined home ranges are conducive to accurate calculations of both the amount and types of prey in the environment and diet of a lemon shark.

Lemon sharks feed at night and are mainly piscivorous; however, they have been known to feed on crustaceans and benthic organisms.[5] Intraspecific predation, or cannibalism, of juvenile lemon sharks by larger conspecifics has also been documented.[15] Rather than feeding randomly, lemon sharks display a high degree of preference for certain species and size of prey when environmental conditions are favorable.[19] They also tend to prefer a prey when it is more abundant and available. Lemon sharks feed selectively on species that are slower and more easily captured by using a stalking technique.[20] For example, parrotfish and mojarras are common prey in the Bahamas because they use camouflage rather than an escape response and are vulnerable due to their stationary foraging behavior. Lemon sharks feed on prey that are intermediate in size compared to other available prey.[5] This tendency can be explained by the tradeoff between the probability of capture and the profitability when it comes to prey size. The general trend in the foraging behavior of lemon sharks conforms to the optimal foraging theory, which suggests a positive relationship between prey selectivity and availability.[18]

Rather than rolling on their sides to rip off chunks of prey, lemon sharks approach their victim with speed only to brake suddenly using their pectoral fins upon contact.[5] The animal then jabs forward multiple times until it has a good grasp of its prey in its jaw and proceeds to shake its head from side to side until it tears off a chunk of flesh. A feeding frenzy, or large swarm of other sharks, then forms as the individuals sense the blood and bodily fluids released from the prey.[5] Sounds of struggling prey also attract groups of sharks, suggesting they use sound detection for predation.[20] Group feeding behavior such as pack hunting or communal scavenging was observed in a study in which pieces of the same stingray were found in the stomachs of several lemon shark individuals that were caught and examined.

Social behavior[edit]

Lemon Shark off the Coast of Naples, Florida, May 1987

Many species of shark, including the lemon shark, are known to actively prefer to be social and live in groups or loose aggregations.[6] A few benefits of group living are enhanced communication, courtship, predatory behavior, and protection. Group living and a preference for social interaction is thought to be important for the survival and success of juvenile lemon sharks.[6] Group living, though, comes with its costs. A few include increased risk of disease, ease of parasite transmission, and competition for resources.[21]

Lemon sharks are found in groups based on similar size. Passive sorting mechanisms such as the ontogenetic habitat shift discussed above have been postulated to contribute to the formation of groups organized based on size or sex.[22] One exception to this behavior is that sharks aged 0–1 years show no preference for groups of matched or unmatched size.[6] One hypothesis for this finding is that it is beneficial for the small young lemon sharks to associate with the larger individuals because they have an easier time gathering information about the habitat regarding elements such as predators and local prey.[6] Lemon shark groups form due to an active desire to be social rather than a simple attraction to the same limited resources such as the mangrove habitat and prey associated with such a habitat.[22]

Many studies have related brain size with complex social behaviors in mammals and birds.[22] The brain of a lemon shark, being comparable in relative mass to that of a mammal or bird, suggests they have the ability to learn from social interactions, cooperate with other individuals, and have the potential to establish dominance hierarchies and stable social bonds.[6]

Reproduction[edit]

Lemon sharks congregate for reproduction at special mating grounds.[4] Females give birth to their young in shallow nursery waters to which they are philopatric. Lemon shark young are known as pups and they tend to remain in the nursery area for several years before venturing into deeper waters.[23] Lemon sharks are viviparous, meaning that the mother directly transfers nutrients to her young via a yolk-sac placenta and the young are born alive.[4] Fertilization is internal and occurs after a male lemon shark holds a female, bites her, and inserts his clasper into her cloaca.[4] Female lemon sharks are polyandrous and sperm competition occurs due to their ability to store sperm in an oviducal gland for several months.[4] Several studies suggest that polyandry in female lemon sharks has adapted out of convenience, rather than indirect genetic benefits to offspring.[24] This type of polyandry is termed as convenience polyandry because females are believed to mate multiple times to avoid harassment by males.[24] Females have a biennial reproductive cycle, requiring a year for gestation and another year for oogenesis and vitellogenesis after parturition. Lemon sharks reach sexual maturity around 12–16 years of age and have low fecundity.[4] The maximum number of pups recorded in a litter is 18.[4]

Importance to humans[edit]

This species of shark is best known in its behavior and ecology. This is mainly due to the work of Samuel Gruber at the University of Miami, who has been studying the lemon shark both in the field and in the laboratory since 1967.[14] The population around the Bimini Islands in the western Bahamas, where Gruber's Bimini Biological Field Station is situated, is probably the best known of all shark populations.[14] As of 2007, this population was experiencing a severe decline and may disappear altogether as a result of destruction of the mangroves for construction of a golf resort.[14]

The lemon shark is targeted by commercial and recreational fishermen along the US Atlantic Ocean, Caribbean, and in the eastern Pacific Ocean due to its prized meat, fins, and skin.[13] Lemon shark skin may be used for leather and its meat can be consumed and is believed to be a delicacy in many cultures.[13] Concern exists that over-fishing has led the lemon shark populations in the western north Atlantic and eastern Pacific Ocean to decline.[7]

Lemon sharks do not represent a large threat to humans. The International Shark Attack File shows 10 unprovoked lemon shark attacks have occurred, none of which has been fatal.[7]

See also[edit]

References[edit]

This article incorporates text from the ARKive fact-file "Lemon shark" under the Creative Commons Attribution-ShareAlike 3.0 Unported License and the GFDL.

  1. ^ Sundström, L.F. (2005). "Negaprion brevirostris". IUCN Red List of Threatened Species. Version 2011.1. International Union for Conservation of Nature. Retrieved 20 August 2011. 
  2. ^ a b c "Lemon shark". Retrieved 15 November 2013. 
  3. ^ Lemon Shark - SharkSurvivor.com
  4. ^ a b c d e f g h Feldheim, K. A.; Gruber, S. H.; Ashley, M. V. (22 August 2002). "The breeding biology of lemon sharks at a tropical nursery lagoon". Proceedings of the Royal Society B: Biological Sciences 269 (1501): 1655–1661. doi:10.1098/rspb.2002.2051. 
  5. ^ a b c d e Bright, Michael (2000). The private life of sharks : the truth behind the myth. Mechanicsburg, PA: Stackpole Books. ISBN 0-8117-2875-7. 
  6. ^ a b c d e f Guttridge, T (August 2009). "Social preferences of juvenile lemon sharks, Negaprion brevirostris". Animal Behaviour 78 (2). doi:10.1016/j.anbehav.2009.06.009. Retrieved 25 October 2013. 
  7. ^ a b c d e f "Florida Museum of Natural History". Retrieved 15 November 2013. 
  8. ^ 3.Carwardine, M. and Watterson, K. (2002) The Shark Watcher’s Handbook. BBC Worldwide Ltd, London.
  9. ^ "FLMNH Ichthyology Department: Lemon Shark". flmnh.ufl.edu. Retrieved 2014-01-25. 
  10. ^ Negaprion brevirostris, Lemon shark - FishBase
  11. ^ a b "Evaluation of a Three-Dimensional Magnetic Barrier on Juvenile Negaprion brevirostris". Retrieved 15 November 2013. 
  12. ^ "Marinebio". Retrieved 15 November 2013. 
  13. ^ a b c "IUCN Red List of Threatened Species". Retrieved 15 November 2013. 
  14. ^ a b c d e Samuel H. Gruber, John F. Morrissey (1993). "Habitat selection by juvenile lemon sharks, Negaprion brevirostris". Environmental Biology of Fishes 38 (4). Retrieved 25 October 2013. 
  15. ^ a b c d e Guttridge, TL; Gruber, SH; Franks, BR; Kessel, ST; Gledhill, KS; Uphill, J; Krause, J; Sims, DW (20 January 2012). "Deep danger: intra-specific predation risk influences habitat use and aggregation formation of juvenile lemon sharks Negaprion brevirostris". Marine Ecology Progress Series 445: 279–291. doi:10.3354/meps09423. 
  16. ^ Wetherbee, BM; Gruber, SH; Rosa, RS (7 August 2007). "Movement patterns of juvenile lemon sharks Negaprion brevirostris within Atol das Rocas, Brazil: a nursery characterized by tidal extremes". Marine Ecology Progress Series 343: 283–293. doi:10.3354/meps06920. 
  17. ^ a b Franks, Bryan (October 2007). "The Spatial Ecology and Resource Selection of Juvenile Lemon Sharks (Negaprion brevirostris) in their Primary Nursery Areas". Drexel University. Retrieved 25 October 2013. 
  18. ^ a b Cortés, Enric; Samuel H. Grube (March 1990). "Diet, Feeding Habits and Estimates of Daily Ration of Young Lemon Sharks, Negaprion brevirostris (Poey)". Copeia 1. JSTOR 1445836. 
  19. ^ Newman, SP; Handy, RD; Gruber, SH (5 January 2010). "Diet and prey preference of juvenile lemon sharks Negaprion brevirostris". Marine Ecology Progress Series 398: 221–234. doi:10.3354/meps08334. 
  20. ^ a b Banner, A (June 1972). "Use of Sound in Predation by Young Lemon Sharks, Negaprion Brevirostris (Poey)". Bulletin of Marine Science 22 (2). Retrieved 25 October 2013. 
  21. ^ Alexander, R D (November 1974). "The Evolution of Social Behavior". Annual Review of Ecology and Systematics 5 (1): 325–383. doi:10.1146/annurev.es.05.110174.001545. 
  22. ^ a b c Jacoby, David M P; Croft, Darren P; Sims, David W (1 December 2012). "Social behaviour in sharks and rays: analysis, patterns and implications for conservation". Fish and Fisheries 13 (4): 399–417. doi:10.1111/j.1467-2979.2011.00436.x. 
  23. ^ "BBC Nature". Retrieved 15 November 2013. 
  24. ^ a b DiBATTISTA, JOSEPH D.; FELDHEIM, KEVIN A.; GRUBER, SAMUEL H.; HENDRY, ANDREW P. (9 January 2008). "Are indirect genetic benefits associated with polyandry? Testing predictions in a natural population of lemon sharks". Molecular Ecology 17 (3): 783–795. doi:10.1111/j.1365-294X.2007.03623.x. 
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