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The lemon shark (''Negaprion brevirostris'') is a species of
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
from the family Carcharhinidae and is classified as a Vulnerable species by the International Union for the Conservation of Nature. Lemon sharks can grow to in length. They are often found in shallow subtropical waters and are known to inhabit and return to specific nursery sites for breeding. Often feeding at night, these sharks use electroreceptors to find their main source of prey:
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
. Lemon sharks enjoy the many benefits of group living such as enhanced communication, courtship, predatory behavior, and protection. This species of shark gives birth to live young, and the females are
polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wive ...
and have a
biennial Biennial means (an event) lasting for two years or occurring every two years. The related term biennium is used in reference to a period of two years. In particular, it can refer to: * Biennial plant, a plant which blooms in its second year and th ...
reproductive cycle. Lemon sharks are not thought to be a large threat to humans; there have been 10 recorded bites, none of which were life-threatening. The lemon shark's life span is unknown, but the average shark is 25 to 30 years old.


Taxonomy

The lemon shark was first named and described in 1868 by
Felipe Poey Felipe Poey (May 26, 1799 – January 28, 1891) was a Cuban zoologist. Biography Poey was born in Havana, the son of French and Spanish parents. He spent several years (1804 to 1807) of his life in Pau then studied law in Madrid. He became a ...
. He originally named it ''Hypoprion brevirostris'', but later renamed it ''Negaprion brevirostris.'' 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).


Description

The shark's yellow colouring serves as an excellent
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
when swimming over the sandy seafloor in its coastal habitat.3.Carwardine, M. and Watterson, K. (2002) The Shark Watcher’s Handbook. BBC Worldwide Ltd., London. The lemon shark commonly attains a length of and a weight up to by adulthood, although sexual maturity is attained at in males and in females. The maximum recorded length and weight is and , respectively. It has a flattened head with a short, broad snout, and the second
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through c ...
is almost as large as the first. Lemon sharks, as any other species of shark, have electroreceptors concentrated in their heads, called the ampullae of Lorenzini. These receptors detect electrical pulses emitted by potential prey and allow these nocturnal feeders to sense their prey in the dark.


Distribution

Lemon sharks are found from
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
to southern
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
in the tropical western
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
. They also live off the coast of west
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
in the southeastern
Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe an ...
. In addition, lemon sharks have been found in the eastern
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contine ...
, from southern
Baja California Baja California (; 'Lower California'), officially the Free and Sovereign State of Baja California ( es, Estado Libre y Soberano de Baja California), is a state in Mexico. It is the northernmost and westernmost of the 32 federal entities of Mex ...
to
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ' ...
and in Cape Verde in Sal Island. 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 meters (301 feet.) 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
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
s, and tend to stay along the continental and insular shelves for most of their lives.


Habitat selection

Information about activity patterns and the use of space is important in understanding a species’ behavioral ecology. Animals often make decisions about habitat use by evaluating their environment's
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
conditions that serve as valuable indicators of good foraging sites or predator-safe locations. Lemon sharks select habitats in warm and shallow water with a rocky or sandy bottom. The environmental temperature influences an individual's body temperature, which ultimately affects physiological processes such as growth and
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run c ...
. Lemon sharks, therefore, select warm-water habitats to maintain optimal metabolic levels. They are believed to avoid areas with thick sea grasses because they make finding prey more difficult. Lemon sharks tend to live in or near shallow-water mangroves, which are often the nursery areas of several species of fish. The data gathered about the characteristics of shark nursery areas is mostly based on coastal species, this is due to their occurrence in bays, estuaries, river deltas, and shallow coastal waters. 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. 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. 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 to remain at after birth or frequently return to, and the habitat used by shark groups repeatedly for several years. 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, coastal areas as pupping grounds is primitive. Lemon sharks have proven to be an ideal model species to challenge the belief that all sharks are asynchronous
opportunistic Opportunism is the practice of taking advantage of circumstances – with little regard for principles or with what the consequences are for others. Opportunist actions are expedient actions guided primarily by self-interested motives. The term ...
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s due to their tendency to use nursery areas for an extended period of time. 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 A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evoluti ...
; however, they have been known to feed on crustaceans and benthic organisms. Intraspecific predation, or cannibalism, of juvenile lemon sharks by larger conspecifics has also been documented. Rather than feeding randomly, lemon sharks display a high degree of preference for certain species and size of prey when environmental conditions are favorable. 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. For example, parrotfish and mojarras are common prey in the
Bahamas The Bahamas (), officially the Commonwealth of The Bahamas, is an island country within the Lucayan Archipelago of the West Indies in the North Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the ar ...
because they use
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
rather than an escape response and are vulnerable due to their stationary
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
behavior. Lemon sharks feed on prey that are intermediate in size compared to other available prey. 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 Optimal foraging theory (OFT) is a behavioral ecology model that helps predict how an animal behaves when searching for food. Although obtaining food provides the animal with energy, searching for and capturing the food require both energy and t ...
, which suggests a positive relationship between prey selectivity and availability. 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 Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as see ...
upon contact. 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 In ecology, a feeding frenzy occurs when predators are overwhelmed by the amount of prey available. The term is also used as an idiom in the English language. Examples in nature For example, a large school of fish can cause nearby sharks, such a ...
, or large swarm of other sharks, then forms as the individuals sense the blood and bodily fluids released from the prey. Sounds of struggling prey also attract groups of sharks, suggesting they use sound detection for predation. Group feeding behavior such as pack hunting or communal
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding ...
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

Many species of sharks, including the lemon shark, are known to actively prefer to be social and live in groups or loose aggregations. 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. Group living, though, comes with its costs. A few include increased risk of disease, ease of parasite transmission, and competition for resources. Lemon sharks are found in groups based on similar size. Passive sorting mechanisms such as its ontogenetic habitat shift have been postulated to contribute to the formation of groups organized based on size or sex. One exception to this behavior is that sharks up to 1 year old show no preference for groups of matched or unmatched size. 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. 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. Many studies have related brain size with complex social behaviors in
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur o ...
and birds. 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 A social relation or also described as a social interaction or social experience is the fundamental unit of analysis within the social sciences, and describes any voluntary or involuntary interpersonal relationship between two or more individuals ...
, cooperate with other individuals, and have the potential to establish dominance hierarchies and stable social bonds.


Reproduction

Lemon sharks congregate for reproduction at special mating grounds. Females give birth to their young in shallow nursery waters to which they are
philopatric Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derives ...
. Lemon shark young are known as pups and they tend to remain in the nursery area for several years before venturing into deeper waters. 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. Fertilization is internal and occurs after a male lemon shark holds a female, bites her, and inserts his
clasper In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's c ...
into her cloaca. Female lemon sharks are
polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wive ...
and sperm competition occurs due to their ability to store sperm in an oviducal gland for several months. Several studies suggest that polyandry in female lemon sharks has adapted out of convenience, rather than indirect genetic benefits to offspring. This type of polyandry is termed as convenience polyandry because females are believed to mate multiple times to avoid harassment by males. Females have a biennial reproductive cycle, requiring a year for gestation and another year for oogenesis and
vitellogenesis Vitellogenesis is the process of yolk protein formation in the oocytes of non mammalian vertebrates during sexual maturation. The term ''vitellogenesis'' comes from the Latin ''vitellus'' ("egg yolk"). Yolk proteins, such as Lipovitellin and P ...
after parturition. Lemon sharks reach sexual maturity around 12–16 years of age and have low
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
. Males tend to mature earlier than females. The maximum number of pups recorded in a litter is 18.


Relationship with humans

This species of
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
is best known in its behavior and ecology, mainly due to the work of Samuel Gruber at the
University of Miami The University of Miami (UM, UMiami, Miami, U of M, and The U) is a private research university in Coral Gables, Florida. , the university enrolled 19,096 students in 12 colleges and schools across nearly 350 academic majors and programs, i ...
, who studied the lemon shark both in the field and in the laboratory from 1967. The population around the Bimini Islands in the western
Bahamas The Bahamas (), officially the Commonwealth of The Bahamas, is an island country within the Lucayan Archipelago of the West Indies in the North Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the ar ...
, where Gruber's Bimini Biological Field Station is situated, is probably the best known of all shark populations. The lemon shark is targeted by commercial and recreational fishers along the U.S.
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
, Caribbean, and in the eastern
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contin ...
due to its prized meat,
fins A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fin ...
, and skin. Lemon shark skin may be used for leather and its meat can be consumed and is believed to be a
delicacy A delicacy is usually a rare and expensive food item that is considered highly desirable, sophisticated, or peculiarly distinctive within a given culture. Irrespective of local preferences, such a label is typically pervasive throughout a r ...
in many cultures. Concern exists that over-fishing has led the lemon shark populations in the western north Atlantic and eastern Pacific Ocean to decline. It is considered vulnerable. Lemon sharks do not represent a large threat to humans. The
International Shark Attack File The International Shark Attack File' is a global database of shark attacks. It began as an attempt to catalogue shark attacks on servicemen during World War II. The Office of Naval Research funded it from 1958 until 1968. During that time, a p ...
lists 10 unprovoked lemon shark bites, none of which were fatal.


See also

* List of sharks


References


Further reading

* * *''Washington Post'', 2005, Aug. 22nd: "Scientists Fear Oceans on the Cusp Of a Wave of Marine Extinctions"


External links

*
Lemon shark facts and pictures
{{Authority control lemon shark Viviparous fish Fish of the Atlantic Ocean Fish of the Americas Fish of the Dominican Republic Fish of the Caribbean Marine fauna of West Africa lemon shark