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The leopard shark ( Triakis semifasciata) is a species of , in the family . It is found along the coast of , from the U.S. state of to Mazatlán in . Typically measuring long, this slender-bodied is immediately identifiable by the striking pattern of black saddle-like markings and large spots over its back, from which it derives its . Large schools of leopard sharks are a common sight in and , swimming over sandy or muddy flats or rock-strewn areas near and . They are most common near the coast, in water less than deep.

Active-swimming , groups of leopard sharks often follow the onto mudflats to forage for food, mainly , , , , , and . Most leopard sharks tend to remain within a particular area rather than undertaking long movements elsewhere, which has led to genetic divergence between populations of sharks living in different regions. This species is aplacental viviparous, meaning that the young hatch inside the and are nourished by . From March to June, the female gives birth to as many as 37 young after a of 10–12 months. It is relatively slow-growing and takes many years to mature.

Harmless to humans, the leopard shark is caught by commercial and recreational fisheries for food and the aquarium trade. This species is mostly fished in the waters off where, after a period of population decline in the 1980s, new fishing regulations in the early 1990s reduced harvesting to sustainable levels. The International Union for Conservation of Nature has assessed this species as of , while noting that local may easily become because of the shark's slow growth and limited migratory habits.


Taxonomy and phylogeny
The first applied to the leopard shark was Triakis californica, coined by John Edward Gray in the 1851 List of the specimens of fish in the collection of the British Museum. However, Gray did not furnish the name with a proper description, rendering it a . In December 1854, American ichthyologist William Orville Ayres gave a lecture describing the species as Mustelus felis, which included the first scientific description of the species. His lecture was reprinted first in The Pacific, a newspaper, and then in the journal Proceedings of the California Academy of Natural Sciences. In April 1855, French biologist Charles Frédéric Girard published another description of this species, naming it Triakis semifasciata.

Despite M. felis being the , an error in recording the dates of publication resulted in the widespread use of T. semifasciata as the leopard shark's scientific name. As a result of this long-standing error, Triakis semifasciata came to be recognized as the valid name (as a ) and Mustelis felis was invalidated (as a ). The specific epithet semifasciata comes from the words semi ("half") and fasciatus ("banded"), describing this shark's dorsal pattern of saddle-like markings. In older literature, this species may be referred to as "" or "".

(2025). 9780520222656, University of California Press.

The genus Triakis contains two , Triakis and Cazon. The leopard shark is placed within the subgenus Triakis along with the banded houndshark ( T. (Triakis) scyllium).

(1984). 9789251013847, Food and Agricultural Organization.
A 2006 analysis by J. Andrés López and colleagues, based on -coding , revealed that Triakis and Cazon are in fact not closely related, and additionally that the leopard shark may be the most basal member of its family.


Distribution and habitat
The leopard shark occurs in the Northeastern Pacific Ocean, from the temperate continental waters of , to the tropical waters of Mazatlán, , including the Gulf of California. It favors muddy or sandy flats within enclosed and , and may also be encountered near beds and rocky , or along the open coast. Numbers have been known to gather near discharges of warm from .
(2025). 9782831707006, International Union for Conservation of Nature and Natural Resources.
Leopard sharks generally swim close to the bottom and are most abundant from the to a depth of , though they may be found as deep as . Many leopard sharks, particularly in the north, leave their coastal habitats in winter and return in early spring. A study in in northern California determined that they depart when the water temperature drops below ; one tagged shark was found to have swum some south.

While a few leopard sharks have been documented traveling hundreds of kilometers, most individuals tend to remain in a localized area for much of their lives. This low level of dispersal has led to genetic divergence across its range. Seven discrete have been identified along the coast between and . Of these, the Humboldt Bay subpopulation is perhaps the most isolated, with the sharks there maturing at a larger size and producing fewer offspring than those from other areas. By contrast, the area around represents a genetic transitional zone between subpopulations whose boundaries are more diffuse. Off , the leopard sharks on the Pacific side are probably distinct from those in the northern Gulf of California. Although there is equivocal evidence for natal philopatry (returning to one's birthplace to breed) in this species, proximity to established breeding grounds likely contributes to the structuring of these different subpopulations.


Description
The leopard shark has a moderately stout body, with a short, rounded snout. There are well-developed, triangular flaps of skin in front of the . The eyes are large and oval, with a nictitating membrane (a protective third ). The line of the mouth is strongly curved. There are furrows at the corners of the mouth extending onto both jaws, with those on the lower jaw almost long enough to meet at the midline. The tooth rows number 41–55 in the upper jaw and 34–45 in the lower jaw; each tooth has a slightly oblique, smooth-edged cusp in the center and 1–2 small cusplets on either side. These teeth are arranged into a flat, "pavement"-like surface with overlapping ridges.Delius, B. Biological Profiles: Leopard Shark. Florida Museum of Natural History Ichthyology Department. Retrieved on August 9, 2009.

The large first is positioned about halfway between the and ; the second is almost as large as the first and much larger than the . The pectoral fins are wide and triangular. The lower lobe of the is well-developed in adults but less than half the length of the upper lobe, which has a strong ventral notch near the tip. The coloration is unique, consisting of prominent black "saddles" and large black spots running along the back, on a silvery to bronzy gray background. Adult sharks often have more spots and saddles with lighter centers compared to juveniles. The underside is whitish and plain. The average length of a leopard shark is . Rarely males may grow to and females , and there is a record of an exceptional female that measured long. The heaviest known leopard shark weighed . Triakis semifasciata monterey.jpg|The dorsal color pattern of the leopard shark gives it its common name. Triakis semifasciata jaws.jpg|Jaws Triakis semifasciata lower teeth.jpg|Lower teeth


Biology and ecology
An active species that swims with a strong undulating motion, the leopard shark is commonly spotted cruising in or just beyond the . It is more active at night than during the day, and sometimes lies still on the bottom. In (among other similar regions), the leopard shark follows the onto to forage for food, retreating just fast enough to prevent being stranded or trapped as the water recedes. At Santa Catalina Island, resident sharks spend the day together in the shallows and at night, they disperse into deeper water, up to away.Manley, J.F. (1995). Diel movement patterns of leopard sharks, Triakis semifasciata, at Santa Catalina Island, California. MS Thesis. California State University, Long Beach, CA.

From birth, leopard sharks form large schools, generally segregated by age and sex; these groups may intermingle with or brown smoothhounds ( Mustelus californicus and M. henlei) and ( Squalus acanthias). These schools are apparently nomadic, often appearing suddenly in an area for a few hours before vanishing just as quickly. In captivity, larger sharks have been observed establishing their dominance over smaller individuals via light nips to the pectoral fins. During summer days, large aggregations of mature females assemble in shallow bays and estuaries, dispersing at night. As these females follow the warmest patches of water (allowing them to raise their internal body temperatures by up to , they are speculated to be taking advantage of the heat to speed their own growth and that of their young. The membership of these aggregations is inconstant, with individual females periodically switching between different sites scattered over several kilometers.

Compared to the related grey and brown smoothhounds that share its range, the leopard shark has smaller and more numerous red blood cells, allowing it to process oxygen more efficiently. This may be an adaptation for foraging in deoxygenated environments.Martin, R.A. Estuaries: Leopard Shark. ReefQuest Centre for Shark Research. Retrieved on August 9, 2009. Their eyes contain very few , likely due to the murky water they inhabit. Small leopard sharks fall prey to larger sharks such as the great white shark ( Carcharodon carcharias) and the broadnose sevengill shark ( Notorynchus cepedianus). On one occasion, a sevengill shark was seen ambushing a leopard shark on a tidal mudflat in , striking with such momentum that the larger predator momentarily beached itself.Martin, R.A. Sandy Plains: Broadnose Sevengill Shark. ReefQuest Centre for Shark Research. Retrieved on August 9, 2009. Known parasites of this species include the Phyllobothrium riseri, Lacistorhynchus dollfusi and Paraorygmatobothrium barber, as well as the Echthrogaleus eoleoptratus and Achtheinus oblongatus.


Feeding
The diet of the leopard shark consists of small and animals, most significantly (, , and ), , (including , , , , , , , , , and ), , , and the ( Urechis caupo). This opportunistic hunter has also been known to eat , , and the young of smoothhounds, shovelnose guitarfish ( Rhinobatos productus), and ( Myliobatis californicus). ( Zostera) and may be swallowed incidentally.

The leopard shark captures prey by expanding its to create a suction force, which is facilitated by its swinging forward to form the mouth into a tube. Simultaneously, the shark protrudes its jaws forward to grip the prey between its teeth. As with other sharks, the teeth of the leopard shark are periodically shed and replaced; it takes 9–12 days for a replacement tooth to move into position. Leopard sharks have been caught with stomachs filled with clam siphons, which the sharks seize before the clams can retract and break off with a levering motion of their bodies. On occasion, the shark tears the entire clam body out of its shell this way. Other sharks examined have had stomachs containing whole innkeeper worms with no bite marks, suggesting that the sharks sucked them out of their burrows. Under a hollow bridge support in San Francisco Bay, a group of leopard sharks and spiny dogfish have been observed feeding on a dense school of anchovies by slowly swimming counterclockwise through the clockwise-swimming school, and swallowing any anchovies that accidentally entered their open mouths.

In some places, this species feeds only on a few prey types and little else (e.g. innkeeper worms and cancrid crabs in Tomales Bay, jack silverside ( Atherinopsis californiensis) eggs and the crabs Romaleon antennarium and Metacarcinus magister in Humboldt Bay). The predominant prey taken depends on location, time of year, and age. For example, in the at , cancrid crabs and innkeeper worms are mostly eaten in winter and spring, fish eggs from winter to early summer, bony fish in summer, and grapsid crabs and clams in fall. Young sharks feed mostly on crabs and transition to clam siphons, fish eggs, and innkeeper worms once they reach long. The largest sharks are the ones that consume the most fish.


Life history
The leopard shark is , with the developing sustained by a until birth. In northern areas, females use bays and sloughs as nursery areas, while to the south they give birth in more open areas. Known breeding grounds along the coast of California include Humboldt Bay, Tomales Bay, , San Francisco Bay, Elkhorn Slough, , Santa Monica Bay (Los Angeles), Catalina Harbor (Santa Catalina Island), and San Diego Bay.

Females give birth to 1–37 young annually from March and July (most in April or May), with the number of pups increasing with female size. Multiple males may father a litter from a single mother In Humboldt and San Francisco Bays, females drop their pups in beds of eelgrass that provide both shelter and food. In Catalina Harbor, females give birth on flats in of water, their backs and dorsal fins exposed; the pups stay in even shallower water less than deep. occurs in early summer following birth; the only known observation of this in the wild was in 2004 off , amongst nine sharks gathered from shore in water deep.Smith, S.E. (2004). "Leopard Shark ( Triakis semifasciata) Mating Activity Observed off La Jolla, California". Southwest Fisheries Science Center, San Diego. Note.

Newborn leopard sharks measure around long. Relatively slow-growing after the first 3–4 years of life, male leopard sharks grow an average of per year, reaching maturity at an age of 7–13 years and a length of , while females grow an average of per year, reaching maturity at an age of 10–15 years and a length of . Individuals of the same age can vary significantly in size and large sharks are especially slow-growing: one specimen was documented to have gained only in length over 12 years. The maximum lifespan of this species is estimated to be 30 years.


Human interactions
Wary and quick to flee, leopard sharks pose almost no danger to humans. There was only one recorded incident which was in 1955 of a leopard shark harassing a skin diver with a nosebleed, though no injuries resulted. This species is caught by commercial fisheries using and , and by recreational anglers and ; it is also caught as in . The meat is considered excellent eating and is sold fresh or frozen. However, leopard sharks living alongside human development may accumulate such as mercury, , and polychlorinated biphenyls (PCBs) within their bodies, and the California Department of Fish and Wildlife warns against eating them regularly. The leopard shark is prized by the aquarium trade for its attractive appearance and hardiness; this led to many newborn pups being captured with hook-and-line off southern California in the late 1980s and early 1990s. It may live for over 20 years in captivity.

Most of the fishing for leopard sharks occurs in the waters off California, where commercial landings peaked at in 1983 and has averaged annually since 1991. These figures are believed to be underestimates, as an unknown portion of the leopard shark catch is recorded simply as "shark". In recent decades the number of sharks taken recreationally has exceeded those taken commercially, with sport fishers catching an average of 52,000 sharks per year from 1980 to 1988, and 45,000 sharks per year since 1993.

(2025). 9781879906570, ANR Publications.
This species contributes negligibly to fishery landings off Oregon. The status of leopard sharks off Mexico is uncertain, although Mexican shark fisheries tend to focus more on larger species.

The International Union for Conservation of Nature (IUCN) has assessed the leopard shark as of . It is highly susceptible to local depletion due to its slow growth rate and low level of exchange between regional . It was considered for, but is not managed under, the U.S. Pacific Fishery Management Council's (PMC) 1982 Groundfish Management Plan. In 1992, the State of California imposed a recreational minimum size limit of and a possession limit of three fish, a move that was strongly supported and promoted by the sport fishing community. In 1993, the State also instituted a minimum size limit of for the commercial harvesting of all sharks and rays, including for the aquarium trade. In addition, Californian restrictions on the use of certain types of fishing gear in near-shore waters confer good protection to this shark's core population. These management measures seem to have successfully reduced fishing mortality and halted the population decline from the 1980s; the status of the leopard shark off California is regarded as secure, with demographic models estimating an annual population increase of 3–6% with sustainable fishing.


See also
  • List of sharks


External links

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