The zebra shark was first described as Squalus varius by Seba in 1758. No type specimen was designated, though Seba included a comprehensive description in Latin and an accurate illustration of a juvenile. Müller and Henle placed this species in the genus Stegostoma in 1837, using the specific epithet fasciatus (or the neuter form fasciatum, as Stegostoma is neuter while Squalus is masculine) from an 1801 work by Bloch and Schneider. In 1984, Compagno rejected the name "varius/m" in favor of "fasciatus/m" for the zebra shark, because Seba did not consistently use binomial nomenclature in his species descriptions (though Squalus varius is one that can be construed as a binomial name). In Compagno's view, the first proper usage of "varius/m" was by Garman in 1913, making it a junior synonym. Both S. fasciatum and S. varium are currently in wide usage for this species.
Early taxonomists thought that juvenile zebra sharks were a different species because of their different appearance from adults. The genus name is derived from the Greek stego meaning "covered", and stoma meaning "mouth". The specific epithet fasciatum means "banded", referring to the striped pattern of the juvenile. The juvenile coloration is also the origin of the common name "zebra shark". The name "leopard shark" is sometimes applied to the spotted adult, but that name usually refers to the houndshark Triakis semifasciata, and is also sometimes used for the tiger shark (Galeocerdo cuvier). Due to their different color patterns and body proportions, both juveniles and subadults have historically been described as separate species (Squalus tigrinus and S. longicaudatus respectively).
There is robust morphological support for the placement of the zebra shark, the whale shark (Rhincodon typus), and the nurse sharks (Ginglymostoma cirratum, Nebrius ferrugineus, and Pseudoginglymostoma brevicaudatum) in a single clade. However, the interrelationships between these taxa are disputed by various authors. Dingerkus (1986) suggested that the whale shark is the closest relative of the zebra shark, and proposed a single family encompassing all five species in the clade. Compagno (1988) suggested affinity between this species and either Pseudoginglymostoma or a clade containing Rhincodon, Ginglymostoma, and Nebrius. Goto (2001) placed the zebra shark as the sister group to a clade containing Rhincodon and Ginglymostoma.
Distribution and habitat
The zebra shark occurs in the tropical waters of the Indo-Pacific region, from South Africa to the Red Sea and the Persian Gulf (including Madagascar and the Maldives), to India and Southeast Asia (including Indonesia, the Philippines, and Palau), northward to Taiwan and Japan, eastward to New Caledonia and Tonga, and southward to northern Australia.
Bottom-dwelling in nature, the zebra shark is found from the intertidal zone to a depth of 62 m (210 ft) over the continental and insular shelves. Adults and large juveniles frequent coral reefs, rubble, and sandy areas. There are unsubstantiated reports of this species from fresh water in the Philippines. Zebra sharks sometimes cross oceanic waters to reach isolated seamounts. Movements of up to 140 km (87 mi) have been recorded for individual sharks. However, genetic data indicates that there is little exchange between populations of zebra sharks, even if their ranges are contiguous.
Adult zebra sharks have longitudinal ridges on the body and a spotted pattern.
The zebra shark has a cylindrical body with a large, slightly flattened head and a short, blunt snout. The eyes are small and placed on the sides of the head; the spiracles are located behind them and are as large or larger. The last 3 of the 5 short gill slits are situated over the pectoral fin bases, and the fourth and fifth slits are much closer together than the others. Each nostril has a short barbel and a groove running from it to the mouth. The mouth is nearly straight, with three lobes on the lower lip and furrows at the corners. There are 28–33 tooth rows in the upper jaw and 22–32 tooth rows in the lower jaw; each tooth has a large central cusp flanked by two smaller ones.
There are five distinctive ridges running along the body in adults, one along the dorsal midline and two on the sides. The dorsal midline ridge merges into the first dorsal fin, placed about halfway along the body and twice the size of the second dorsal fin. The pectoral fins are large and broad; the pelvic and anal fins are much smaller but larger than the second dorsal fin. The caudal fin is almost as long as the rest of the body, with a barely developed lower lobe and a strong ventral notch near the tip of the upper lobe. The zebra shark attains a length of 2.5 m (8.2 ft), with an unsubstantiated record of 3.5 m (11.5 ft). Males and females are not dimorphic in size.
The color pattern in young sharks is dark brown above and light yellow below, with vertical yellow stripes and spots. As the shark grows to 50–90 cm (20–35 in) long, the dark areas begin to break up, changing the general pattern from light-on-dark stripes to dark-on-light spots. There is substantial variation in pattern amongst adults, which can be used to identify particular individuals. In 1964, a partially albino zebra shark completely lacking spots was discovered in the Indian Ocean. The shark, a 1.9 m (6.2 ft) long mature female, was unusual in that albino animals rarely survive long in the wild due to their lack of crypsis.
Biology and ecology
The zebra shark swims with an eel-like motion.
During the day, zebra sharks are sluggish and usually found resting on the sea bottom, sometimes using their pectoral fins to prop up the front part of their bodies and facing into the current with their mouths open to facilitate respiration. Reef channels are favored resting spots, since the tightened space yields faster, more oxygenated water. They become more active at night or when food becomes available. Zebra sharks are strong and agile swimmers, propelling themselves with pronounced anguilliform (eel-like) undulations of the body and tail. In a steady current, they have been seen hovering in place with sinuous waves of their tails.
The zebra shark feeds primarily on shelled molluscs, though it also takes crustaceans, small bony fishes, and possibly sea snakes. The slender, flexible body of this shark allows it to wriggle into narrow holes and crevices in search of food, while its small mouth and thickly muscled buccal cavity allow it to create a powerful suction force with which to extract prey. This species may be preyed upon by larger fishes and marine mammals. Known parasites of the zebra shark include four species of tapeworms in the genus Pedibothrium.
A juvenile zebra shark with a color pattern intermediate between that of young and adults. The courtship behavior of the zebra shark consists of the male following the female and biting vigorously at her pectoral fins and tail, with periods in which he holds onto her pectoral fin and both sharks lie still on the bottom. On occasion this leads to mating, in which the male curls his body around the female and inserts one of his claspers into her cloaca. Copulation lasts for two to five minutes. The zebra shark is oviparous, with females laying large egg capsules measuring 17 cm (7 in) long, 8 cm (3 in) wide, and 5 cm (2 in) thick. The egg case is dark brown to purple in color, and has hair-like fibers along the sides that secure it to the substrate. The adhesive fibers emerge first from the female's vent; the female circles vertical structures such as reef outcroppings to entangle the fibers, so as to anchor the eggs. Females have been documented laying up to 46 eggs over a 112-day period. Eggs are deposited in batches of around four. Reproductive seasonality in the wild is unknown.
In captivity, the eggs hatch after two to four weeks. The hatchlings measure 20–36 cm (8–14 in) long and have proportionately longer tails than adults. The habitat preferences of juveniles are unclear; one report places them at depths greater than 50 m (160 ft), while another report from India suggests they inhabit shallower water than adults. The stripes of the juveniles may have an anti-predator function, making each individual in a group harder to target. Males attain sexual maturity at 1.5–1.8 m (4.9–5.9 ft) long, and females at 1.7 m (5.6 ft) long. The lifespan has been estimated to be 25–30 years in the wild.