Dusky shark


The dusky shark is a species of requiem shark, in the family Carcharhinidae, occurring in tropical and warm-temperate continental seas worldwide. A generalist apex predator, the dusky shark can be found from the coast to the outer continental shelf and adjacent pelagic waters, and has been recorded from a depth of 400 m. Populations migrate seasonally towards the poles in the summer and towards the equator in the winter, traveling hundreds to thousands of kilometers. One of the largest members of its genus, the dusky shark reaches more than in length and in weight. It has a slender, streamlined body and can be identified by its short round snout, long sickle-shaped pectoral fins, ridge between the first and second dorsal fins, and faintly marked fins.
Adult dusky sharks have a broad and varied diet, consisting mostly of bony fishes, sharks and rays, and cephalopods, but also occasionally crustaceans, sea stars, bryozoans, sea turtles, marine mammals, carrion, and humans. This species is viviparous with a three-year reproductive cycle; females bear litters of 3-14 young after a gestation period of 22-24 months, after which there is a year of rest before they become pregnant again. This shark, tied with the Spiny dogfish as a result is the animal with the longest gestation period. Females are capable of storing sperm for long periods, as their encounters with suitable mates may be few and far between due to their nomadic lifestyle and low overall abundance. Dusky sharks are one of the slowest-growing and latest-maturing sharks, not reaching adulthood until around 20 years of age.
Because of its slow reproductive rate, the dusky shark is very vulnerable to human-caused population depletion. This species is highly valued by commercial fisheries for its fins, used in shark fin soup, and for its meat, skin, and liver oil. It is also esteemed by recreational fishers. The International Union for Conservation of Nature has assessed this species as Endangered worldwide and Vulnerable off the eastern United States, where populations have dropped to 15-20% of 1970s levels. The dusky shark is regarded as potentially dangerous to humans due to its large size, but there are few attacks attributable to it.

Taxonomy

French naturalist Charles Alexandre Lesueur published the first scientific description of the dusky shark in an 1818 issue of Journal of the Academy of Natural Sciences of Philadelphia. He placed it in the genus Squalus and gave it the specific epithet obscurus, referring to its coloration. Subsequent authors have recognized this species as belonging to the genus Carcharhinus. Lesueur did not designate a type specimen, though he was presumably working from a shark caught in North American waters.
Many early sources gave the scientific name of the dusky shark as Carcharias ''lamiella, which originated from an 1882 account by David Starr Jordan and Charles Henry Gilbert. Although Jordan and Gilbert referred to a set of jaws that came from a dusky shark, the type specimen they designated was later discovered to be a copper shark. Therefore, C. lamiella is not considered a synonym of C. obscurus but rather of C. brachyurus''. Other common names for this species include bay shark, black whaler, brown common gray shark, brown dusky shark, brown shark, common whaler, dusky ground shark, dusky whaler, river whaler, shovelnose, and slender whaler shark.

Phylogeny and evolution

Teeth belonging to the dusky shark are fairly well represented in the fossil record, though assigning Carcharhinus teeth to species can be problematic. Dusky shark teeth dating to the Miocene have been recovered from the Kendeace and Grand Bay formations in Carriacou, the Grenadines, the Moghra Formation in Egypt, Polk County, Florida, and possibly Cerro La Cruz in northern Venezuela. Teeth dating to the Late Miocene or Early Pliocene are abundant in the Yorktown Formation and the Pungo River, North Carolina, and from the Chesapeake Bay region; these teeth differ slightly from the modern dusky shark, and have often been misidentified as belonging to the oceanic whitetip shark. Dusky shark teeth have also been recovered from the vicinity of two baleen whales in North Carolina, one preserved in Goose Creek Limestone dating to the Late Pliocene, and the other in mud dating to the Pleistocene-Holocene.
In 1982, Jack Garrick published a phylogenetic analysis of Carcharhinus based on morphology, in which he placed the dusky shark and the Galapagos shark at the center of the "obscurus group". The group consisted of large, triangular-toothed sharks with a ridge between the dorsal fins, and also included the bignose shark, the Caribbean reef shark, the sandbar shark, and the oceanic whitetip shark. This interpretation was largely upheld by Leonard Compagno in his 1988 phenetic study, and by Gavin Naylor in his 1992 allozyme sequence study. Naylor was able to further resolve the interrelationships of the "ridge-backed" branch of Carcharhinus, finding that the dusky shark, Galapagos shark, oceanic whitetip shark, and blue shark comprise its most derived clade.

Distribution and habitat

The range of the dusky shark extends worldwide, albeit discontinuously, in tropical and warm-temperate waters. In the western Atlantic Ocean, it is found from Massachusetts and the Georges Bank to southern Brazil, including the Bahamas and Cuba. In the eastern Atlantic Ocean, it has been reported from the western and central Mediterranean Sea, the Canary Islands, Cape Verde, Senegal, Sierra Leone, and possibly elsewhere including Portugal, Spain, Morocco, and Madeira. In the Indian Ocean, it is found off South Africa, Mozambique, and Madagascar, with sporadic records in the Arabian Sea, the Bay of Bengal, and perhaps the Red Sea. In the Pacific Ocean, it occurs off Japan, mainland China and Taiwan, Vietnam, Australia, and New Caledonia in the west, and from southern California to the Gulf of California, around Revillagigedo, and possibly off northern Chile in the east. Records of dusky sharks from the northeastern and eastern central Atlantic, and around tropical islands, may in fact be of Galapagos sharks. Mitochondrial DNA and microsatellite evidence suggest that Indonesian and Australian sharks represent distinct populations.
Residing off continental coastlines from the surf zone to the outer continental shelf and adjacent oceanic waters, the dusky shark occupies an intermediate habitat that overlaps with its more specialized relatives, such as the inshore sandbar shark, the pelagic silky shark and oceanic whitetip shark, the deepwater bignose shark, and the islandic Galapagos shark and silvertip shark. One tracking study in the northern Gulf of Mexico found that it spends most of its time at depths of, while making occasional forays below ; this species has been known to dive as deep as. It prefers water temperatures of, and avoids areas of low salinity such as estuaries.
The dusky shark is nomadic and strongly migratory, undertaking recorded movements of up to ; adults generally move longer distances than juveniles. Sharks along both coasts of North America shift northward with warmer summer temperatures, and retreat back towards the equator in winter. Off South Africa, young males and females over long disperse southward and northward respectively from the nursery area off KwaZulu-Natal; they join the adults several years later by a yet-unidentified route. In addition, juveniles spend spring and summer in the surf zone and fall and winter in offshore waters, and as they approach in length begin to conduct a north–south migration between KwaZulu-Natal in the winter and the Western Cape in summer. Still-larger sharks, over long, migrate as far as southern Mozambique. Off Western Australia, adult and juvenile dusky sharks migrate towards the coast in summer and fall, though not to the inshore nurseries occupied by newborns.

Description

Alongside the bull shark, the dusky shark is the largest members of its genus, commonly reaching a length of and a weight of ; the maximum recorded length and weight are and respectively. However, the maximum reported size of the species is, while the maximum weight is reported to reach up to. Females grow larger than males. This shark has a slender, streamlined body with a broadly rounded snout no longer than the width of the mouth. The nostrils are preceded by barely developed flaps of skin. The medium-sized, circular eyes are equipped with nictitating membranes. The mouth has very short, subtle furrows at the corners and contains 13–15 tooth rows on either side of both jaws. The upper teeth are distinctively broad, triangular, and slightly oblique with strong, coarse serrations, while the lower teeth are narrower and upright, with finer serrations. The five pairs of gill slits are fairly long.
The large pectoral fins measure around one-fifth as long as the body, and have a falcate darken towards the tips; this is more obvious in juveniles. Dusky sharks can be found at Redondo Beach, southern California to the Gulf of California, and to Ecuador. But sometimes rarely off southern California; common in tropics. Dusky sharks have a total length of at least 3.6 m or possibly to 4.2 m. At birth, dusky sharks are about a length of 70–100 cm. In the surf zone, dusky sharks swim to a depth of 573 m. Dusky sharks have a color of Gray or beige.

Biology and ecology

As an apex predator positioned at the highest level of the trophic web, the dusky shark is generally less abundant than other sharks that share its range. However, high concentrations of individuals, especially juveniles, can be found at particular locations. Adults are often found following ships far from land, such as in the Agulhas Current. A tracking study off the mouth of the Cape Fear River in North Carolina reported an average swimming speed of. The dusky shark is one of the hosts of the sharksucker. Known parasites of this species include the tapeworms Anthobothrium laciniatum, Dasyrhynchus pacificus, Platybothrium kirstenae, Floriceps saccatus, Tentacularia coryphaenae, and Triloculatum triloculatum, the monogeneans Dermophthirius carcharhini and Loimos salpinggoides, the leech Stibarobdella macrothela, the copepods Alebion sp., Pandarus cranchii, P. sinuatus, and P. smithii, the praniza larvae of gnathiid isopods, and the sea lamprey.
Full-grown dusky sharks have no significant natural predators. Major predators of young sharks include the ragged tooth shark, the great white shark, the bull shark, and the tiger shark. Off KwaZulu-Natal, the use of shark nets to protect beaches has reduced the populations of these large predators, leading to a dramatic increase in the number of juvenile dusky sharks. In turn, the juvenile sharks have decimated populations of small bony fishes, with negative consequences for the biodiversity of the local ecosystem.