Leopard Shark

The **leopard shark** (*Triakis semifasciata*) is a species of houndshark, in the family Triakidae. It is found along the Pacific coast of North America, from the U.S. state of Oregon to Mazatlán in Mexico. Typically measuring 1.2–1.5 m (3.9–4.9 ft) long, this slender-bodied shark is immediately identifiable by the striking pattern of black saddle-like markings and large spots over its back, from which it derives its common name. Large schools of leopard sharks are a common sight in bays and estuaries, swimming over sandy or muddy flats or rock-strewn areas near kelp beds and reefs. They are most common near the coast, in water less than 4 m (13 ft) deep.

Active-swimming predators, groups of leopard sharks often follow the tide onto intertidal mudflats to forage for food, mainly clams, spoon worms, crabs, shrimp, bony fish, and fish eggs. 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 uterus and are nourished by yolk. From March to June, the female gives birth to as many as 37 young after a gestation period 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 California 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 Least Concern, while noting that local stocksmay easily become overfished because of the shark's slow growth and limited migratory habits.

## Distribution and Habitat

The leopard shark occurs in the cool to warm-temperater continental waters of the northeastern Pacific Ocean, from Coos Bay, Oregon to Mazatlán, Mexico, including the Gulf of California. It favors muddy or sandy flats within enclosed bays and estuaries, and may also be encountered near kelp beds and rocky reefs, or along the open coast. Numbers have been known to gather near discharges of warm effluent from power plants. Leopard sharks generally swim close to the bottom and are most abundant from the intertidal zoneto a depth of 4 m (13 ft), though they may be found as deep as 91 m (299 ft). Many leopard sharks, particularly in the north, leave their coastal habitats in winter and return in early spring. A study in Tomales Bay in northern California determined that they depart when the water temperature drops below 10–12°C (50–54°F); one tagged shark was found to have swum some 140 km (87 mi) 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 gene pools have been identified along the Californian coast between Humboldt Bay and San Diego. 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 Los Angeles represents a genetic transitional zone between subpopulations whose boundaries are more diffuse. Off Baja California, 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 nares. The eyes are large and oval, with a nictitating membrane (a protective third eyelid). 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.

The large first dorsal fin is positioned about halfway between the pectoral and pelvic fins; the second is almost as large as the first and much larger than the anal fin. The pectoral fins are wide and triangular. The lower lobe of the caudal fin 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 1.2–1.5 m (3.9–4.9 ft). Rarely males may grow to 1.5 m (4.9 ft) and females 1.8 m (5.9 ft), and there is a record of an exceptional female that measured 2.1 m (6.9 ft) long. The heaviest known leopard shark weighed 18.4 kg (41 lb).

## 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 surf zone. It is more active at night than during the day, and sometimes lies still on the bottom. In Tomales Bay and elsewhere, the leopard shark follows the tide onto mudflats to forage for food, retreating just fast enough to prevent being stranded or trapped as the water recedes.[13] At Santa Catalina Island, resident sharks spend the day together in the shallows and at night disperse into deeper water, ranging up to 10 km (6.2 mi) away.

From birth, leopard sharks form large schools generally segregated by age and sex, which may intermingle with grey or brown smoothhounds (*Mustelus californicus* and *M. henlei*) or spiny dogfish (*Squalus acanthias*).[5] These schools are apparently nomadic, often appearing suddenly in an area for a few hours and then 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 3°C (5.4°F), they are speculated to be taking advantage of the heat to speed their growth and that of their gestating 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 estuary environments. Their eyes contain very few cone cells, 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 Humboldt Bay, striking with such momentum that the larger predator momentarily beached itself. Known parasites of this species include the tapeworms*Phyllobothrium riseri*, *Lacistorhynchus dollfusi* and *Paraorygmatobothrium barber*, as well as the copepods *Echthrogaleus eoleoptratus* and *Achtheinus oblongatus*.

### **Feeding**

The diet of the leopard shark consists of small benthic and littoral animals, most significantly crabs (Cancridae, Grapsidae, and Hippoidea), shrimp, bony fish (including anchovies, herring, topsmelt, croakers, surfperch, gobies, rockfish, sculpins, flatfish, and midshipmen), fish eggs, clams, and the echiurid fat innkeeper worm (*Urechis caupo*). This opportunistic hunter has also been known to eat ghost shrimp, polychaete worms, and the young of smoothhounds, shovelnose guitarfish (*Rhinobatos productus*), and bat rays (*Myliobatis californicus*). Eelgrass (*Zostera*) and algae may be swallowed incidentally.

The leopard shark captures prey by expanding its buccal cavity to create a suction force, which is facilitated by its labial cartilages 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 Elkhorn Slough at Monterey Bay, 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 70–80 cm (2.3–2.6 ft) long. The largest sharks are the ones that consume the most fish.

### **Life History**

The leopard shark is aplacental viviparous, with the developing embryos sustained by a yolk sac 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, Bodega Bay, San Francisco Bay, Elkhorn Slough, Morro Bay, 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. 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 1 m (3.3 ft) of water, their backs and dorsal fins exposed; the pups stay in even shallower water less than 30 cm (12 in) deep. Mating occurs in early summer following birth; the only known observation of this in the wild was in 2004 off La Jolla, amongst nine sharks gathered 19 m (62 ft) from shore in water 0.3–3 m (0.98–9.84 ft) deep.

Newborn leopard sharks measure around 20 cm (7.9 in) long. Relatively slow-growing after the first 3–4 years of life, male leopard sharks grow an average of 2.0 cm (0.79 in) per year, reaching maturity at an age of 7–13 years and a length of 0.7–1.2 m (2.3–3.9 ft), while females grow an average of 2.3 cm (0.91 in) per year, reaching maturity at an age of 10–15 years and a length of 1.1–1.3 m (3.6–4.3 ft). 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 4 cm (1.6 in) in length over 12 years. The maximum lifespan of this species is estimated to be 30 years.