The dark shark [1] [2] , or the white-shark [2] , or the twilight shark [2] [3] , or the dark shark-dog [2] ( Latin Carcharhinus obscurus ) is a predatory fish of the family of gray sharks of the Karhariformes order. These sharks live in tropical waters and many temperate regions of all oceans. They are found at a depth of up to 400 m. They make seasonal migrations, moving in the summer closer to the poles, and in the winter to the equator. They have a slender, streamlined body with a wide and rounded snout. The second dorsal fin is much smaller than the first. The pectoral fins are large, sickle-shaped. There is a crest between the dorsal fins. Coloring of the dorsal surface of the body from bronze to bluish-gray. The maximum recorded length is 420 cm and the mass is 347 kg.
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Carcharhinus obscurus Lesueur , 1818 | |||||||||||||||||||||||||||||||||||||||||
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Carcharhinus iranzae Fourmanoir, 1961
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The diet of dark sharks is very diverse. They prey on a variety of bony and cartilaginous fish, as well as marine invertebrates. Marine mammals are rarely attacked. Like other representatives of the genus of gray sharks, dark sharks are viviparous. In the litter from 3 to 16 newborns. Pregnancy lasts up to 24 months. The reproduction cycle is three-year. Dark sharks grow very slowly and reach puberty late. These sharks are potentially dangerous to humans. They are the subject of commercial fishing. The slow reproduction cycle makes them very sensitive to the negative effects caused by human factors [4] [5] .
Content
- 1 Taxonomy and phylogenesis
- 2 Description
- 3 Area
- 4 Biology and Ecology
- 4.1 Diet and eating behavior
- 4.2 Reproduction and life cycle
- 5 Human interaction
- 6 Conservation measures
- 7 Notes
- 8 Literature
- 9 References
Taxonomy and Phylogenesis
French naturalist Charles Alexander Lesueur published the first scientific description of a dark shark in 1818 in the issue of the journal of the Academy of Natural Sciences of Philadelphia. He carried it to the genus Squalus and gave a specific epithet of lat. obscurus , which means “dark” or “dull”, referring to the color of the fish [6] . Subsequent authors attributed this species to the genus Carcharhinus . The holotype was not assigned, although it is known that Lesjuyer investigated an individual, probably caught in North American waters [7] .
Many early sources used the scientific name for the dark shark Carcharias (later Carcharhinus ) lamiella , given to this species in 1882 by David Starr Jordanan and Charles Henry Gilbert . Later it turned out that the jaws described by scientists did not belong to the dark, but toothed shark ( Archarhinus brachyurus ). Thus, Сarcharhinus lamiella is considered a synonym not for Сarcharhinus obscurus , but Сarcharhinus brachyurus [8] [9] .
In the paleontological literature, petrified teeth of dark sharks are often mentioned, although the species identification of Carcharhinus teeth is rather problematic [10] . These findings, dating back to the Miocene epoch (25-5.3 Ma), were discovered in the Kendis and Grand Bay formations in Carriacou , Grenadines [11] , Mohra ( Egypt ) [12] , Polk (Florida County), and probably , Cerro la Cruz in northern Venezuela [13] . In addition, teeth from the Late Miocene and Early Pliocene (11.6–3.6 Ma) are often found in the Yorktown Formation and the Pungo River , North Carolina , as well as in the Chesapeake Bay . teeth of modern dark sharks and are mistaken for the teeth of long-winged sharks [10] . The teeth of dark sharks were extracted from the fossilized remains of two baleen whales , of which one was preserved in the Guz Creek Limestone formation, North Carolina, and dates from the Early Pliocene (about 3.5 million years ago), and the other was found in mud sediments of the Pleistocene - Holocene (about 12,000 years ago) [14]
In 1982, New Zealand scientist Jack Garrick published the results of a phylogenetic analysis of Carcharhinus based on morphology and placed the dark and Galapagos sharks ( Carcharhinus galapagensis ) in the center of the obscurus group . The group included large sharks with triangular teeth and a crest between the dorsal fins , such as Carcharhinus altimus , Carcharhinus perezi , gray-blue and long-winged shark [15] . This interpretation was widely supported by the results of phenetic studies [16] and analysis of the allozyme sequence. The latter made it possible to establish relationships within the branch of Carcharhinus , the members of which are characterized by the presence of an inter-fin dorsal ridge. It was found that the Galapagos , gray-blue and long-winged sharks are a derivative of the clade [17] .
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| Phylogenetic relationships of the dark shark established on the basis of the allozyme sequence [17] |
Description
One of the largest members of the genus of gray sharks , the dark shark usually reaches a length of 3.2 m with a mass of 160-180 kg. The maximum recorded length and weight are 4.2 m and 347 kg, respectively [4] [18] . Females grow larger than males [19] . This shark has a slender, streamlined body with a wide rounded snout. Before the nostrils there are barely developed skin folds. Medium-sized round eyes are equipped with a blinking membrane . The mouth is very short; thin furrows are located in the corners. In the mouth, 13-15 (usually 14) dentitions on each side of both jaws. Upper teeth broad, triangular, slightly inclined, edges very serrated; the lower teeth are narrower and stand upright, the notches on the edges are smaller. The dark shark has five pairs of rather long gill slits [18] . Large pectoral fins are about 1/5 of the length of the shark's body, have a crescent shape, the tips are pointed. The first dorsal fin is of moderate size and fuzzy sickle-shaped, it has a pointed apex and a strongly concave hind margin. The fin base lies above the free posterior ends of the pectoral fins. The second dorsal fin is much smaller than the first and is located opposite the anal fin. There is a ridge between the first and second dorsal fins. The caudal fin is large and high, with a well developed lower lobe and ventral notch under the tip of the upper lobe [20] . Placoid scales have the shape of a rhombus, the scales are closely set, each has five horizontal teeth [18] . Coloring from bronze to bluish-gray, belly white, white color spreads on the sides, where weak light streaks can be seen. The fins, especially the lower pectoral and the lower lobe of the caudal fin, are darker along the edges. This characteristic coloration is more pronounced in young sharks [21] .
Range
Dark shark is found all over the world in tropical and warm temperate waters. Its range is vast, albeit intermittent. In the western Atlantic, these sharks inhabit from the state of Massachusetts and the George Bank to the Gulf of Mexico and from Nicaragua to the southern part of the Brazilian coast. In the eastern Atlantic, they are found in the west of the Mediterranean Sea , off the coast of the Canary Islands , Cape Verde , Senegal , Sierra Leone and, possibly, in other places. In the Indian Ocean, they inhabit the coastal waters of South Africa , Mozambique and Madagascar . Occasionally, there is evidence of the presence of these sharks in the Arabian Sea , the Bay of Bengal and, possibly, the Red Sea . In the western Pacific Ocean, they are found from Japan to the states of Queensland and New South Wales , Australia . In the eastern Pacific, the range covers the coast of the Americas from southern California to Chile [5] . Information about the presence of dark sharks in the north-eastern and central-eastern parts of the Atlantic Ocean is probably erroneous and refers to the Galapagos sharks [5] [22] . Studies of mitochondrial DNA and microsatellites showed that sharks living in the waters of Indonesia and Australia belong to separate populations [23] .
Dark sharks live on the continental coast from the surf zone to the outer edge of the continental shelf and adjacent oceanic waters, dividing the space with more specialized relatives, such as gray-blue sharks , pelagic silk sharks ( Сarcharhinus falciformis ), long-winged ocean sharks , species Carcharhinus altimus and the island Galapagos and white-feathered gray sharks ( Carcharhinus albimarginatus ) [7] . In a study conducted by tagging in the northern part of the Gulf of Mexico , it was found that dark sharks stay most of their time at a depth of 10-80 m, periodically dropping below 200 m. It is known that this species can reach a depth of 400 m. prefers a water temperature of 19–28 ° C and avoids areas with low salinity , such as estuaries [8] [24] .
Dark sharks migrate up to 3800 km. Off the coast of North America in the summer they move to the north, and in the winter to the equator [5] . In the waters of South Africa, young females and males, having reached a size of 0.9 m, swim south and north of the place of birth of Kwazulu-Natal and after a few years join unknown adults along unknown routes. In addition, growing individuals spend spring and summer in the surf zone, and autumn and winter in the open sea. Upon reaching a length of 2.2 m, they begin to migrate from north to south between KwaZulu-Natal in winter and Western Cape in summer. Large sharks, whose size exceeds 2.8 m, migrate south up to Mozambique [5] [7] [25] . Along the shores of Western Australia, adult and young dark sharks swim in the summer and autumn to the coast, without going into shallow bays and bays where newborns are born [5] .
Biology and Ecology
As predators occupying the top of the food pyramid, dark sharks are usually smaller in number compared to other sharks that share their range [7] . However, there are places where clusters of these sharks are found, especially young ones [8] . Adult sharks often accompany ships far from the earth, for example, along the course of Cape Igolny [25] . Studies conducted at the mouth of the Cape Fear River in North Carolina reported that their average speed is 0.8 km / h [26] . Dark sharks are often accompanied by sticking fish ( Echeneis naucrates ) [27] . Adult dark sharks have no enemies in nature [18] . Young sharks are hunted by the common sand shark ( Carcharias taurus ), the great white shark ( Carcharodon Carcharias ), the bull shark ( Сarcharhinus leucas ) and the tiger shark ( Galeocerdo cuvier ). Off the coast of KwaZulu-Natal, the use of nets to protect beaches from sharks has led to a reduction in the population of large predators. This caused a sharp increase in the number of immature dark sharks, which, in turn, destroyed small bony fish, which generally had negative consequences for the biodiversity of the local ecosystem [7] [28] .
On dark sharks, tapeworms Anthobothrium laciniatum [29] , Dasyrhynchus pacificus [30] , Platybothrium kirstenae [31] , Floriceps saccatus [32] , Tentacularia coryphaenae [33], and Triloculatum trilocheris trilochemata salpinggoides [36] , leeches of Stibarobdella macrothela [37] , copepods Alebion sp. , Pandarus cranchii [38] , P. sinuatus [18] and lamprey [39]
Diet and Eating Behavior
The dark shark is a universal predator, the diet of which includes a wide range of marine animals at all levels of the water column [21] [40] . Large individuals can consume more than 1/10 of their own weight in one meal [41] . The bite force of a 2-meter dark shark is 60 kg per 2 mm² (tooth apex area). This is the highest indicator reliably measured in any shark [42] . In the Indian Ocean, dense accumulations of young dark sharks were recorded in areas with a high concentration of food [5] .
The diet of the dark shark includes pelagic bony fish , such as herring and anchovies , tuna and mackerel , marlin , horse mackerel , needle fish , flying fish , saber fish ; bottom fish, including mullet , crucian carp , croaker , scorpion , eel and flounder ; reef fish, including barracudas , urchin fish , red mullet , paddlefish and sea bass ; cartilaginous fish , including sharks , sawfish , angelfish ( Squalus squatina ), marten sharks , hammer fish , fox sharks , fish guitars , stingrays , and invertebrates , including cephalopods , crustaceans , mollusks and starfish . Very rarely, large individuals eat sea turtles , marine mammals (mainly carrion) and human litter [7] [8] [40] [43] .
In the northwestern Atlantic, about 60% of the diet of the dark shark is made up of different fish belonging to more than 10 families, of which the main role is played by lufari ( Pomatomus saltatrix ) and summer paralytes ( Paralichthys dentatus ). The second component of the diet are stingrays, their egg capsules . Crabs Ovalipes ocellatus is another significant food source [40] . In South African and Australian waters, the basis of the diet is bony fish. Newborn and young sharks prey mainly on small pelagic fish and cephalopods such as sardines and squids .
Sharks longer than 2 m extend their diet due to large bony and cartilaginous fish [44] [45] . Every winter, South African sardines ( Sardinops sagax ) appear on the eastern coast of South Africa, which are hunted by medium and large sized dark sharks. Pregnant and giving birth to females do not join the hunt, possibly because pregnancy does not allow them to spend a lot of energy pursuing such a quick prey [41] . One study showed that 0.2% of the observed dark sharks hunted bottlenose dolphins [46] .
Reproduction and Life Cycle
Like other representatives of the genus of gray sharks , dark sharks are viviparous fish; developing embryos receive nutrition through placental communication with a mother formed by an empty yolk sac . In the northwestern Atlantic, mating occurs in spring, while in other areas, such as off the coast of South Africa, the reproductive cycle is not seasonal [7] [8] . Females are able to store sperm, probably of several males, for a long period from several months to several years in the non-essential gland (the organ that secretes the germ bags). This is an important factor for successful reproduction, given the long migrations performed by dark sharks and the low concentration of the population, which make intraspecific encounters rare and random [47] .
Pregnancy lasts up to 22-24 months, the female brings posterity once every three years [5] . In the litter from 3 to 16 newborns, the average number is 6-12, the number of offspring does not depend on the size of the female [8] [41] . In the western part of the Atlantic Ocean, as a rule, the litter is less than in the southeastern part of the Atlantic Ocean (on average 8 out of 10 newborns) [18] . Depending on the region, labor takes place all year round or seasonally: in the northwestern Atlantic, newborns were born at the end of winter, and offshore of Western Australia in summer and autumn, in the waters of South Africa, dark sharks are born in any season [7] [21] . As the term of labor approaches, pregnant females move to small coastal lagoons , where there is a lot of food and there are no large predators (including their species), and immediately after the offspring are born [8] [45] . Such natural nurseries exist on the coast of Kwazulu-Natal, in southwestern Australia, in Western California and in the eastern United States in New Jersey and North Carolina [5] [8] .
The length of newborns is 0.7-1.0 m [5] ; their size is directly related to the size of the mother and decreases with the number of offspring. Available data indicate that females can determine the optimal size at which their cubs should be born in order to increase their chances of survival depending on environmental conditions. In addition, at birth, the weight of the liver of sharks is 1/5 of their weight, and they use the nutrients stored in the liver until they learn how to hunt on their own [41] . Dark sharks are one of the slowest growing species of sharks. They reach puberty only by becoming large [48] : depending on the region of habitation, males reach puberty with a length of 2.8-3 m at the age of 18-23 years, for females it is 2.6-3.1 m and 17-32 years respectively [49] [50] [51] . Annual growth is 8–11 cm during the first five years of life [52] . The maximum life expectancy of 40-50 years or more [48] .
Human Interaction
Due to their large size, dark sharks are considered to be potentially dangerous to humans, although little is known about how they behave towards people under water [7] . By 2014, the list was International Shark Attack File 2 attacks on people and boats were mentioned, one of which was fatal [53] . Вероятно, нападения, приписываемые этому виду у Бермудских островов , на самом деле были совершены галапагосскими акулами . Для защиты пляжей в Южной Африке и Австралии активно используют противоакульи сети, в которые попадаются в основном взрослые особи. С 1978 по 1999 год у берегов Квазулу-Наталь в среднем ежегодно отлавливали 256 тёмных акул [48] . Молодые тёмные акулы хорошо приспосабливаются к жизни в неволе в аквариумах [7] . Тёмная акула является одним из наиболее востребованных видов акул на рынке, так как у неё крупные плавники с большим количеством внутренних лучей ( ceratotrichia ), которые используются для приготовления популярного супа [48] . Кроме того, мясо поступает в продажу свежим, замороженным, сушёным, солёным или копчёным, используют кожу и жир [4] . Тёмные акулы являются предметом целевого промышленного рыболовства у берегов восточной части Северной Америки, юго-западной Австралии и восточной Южной Африки. В водах юго-западной Австралии промысел начался в 1940 году и был расширен в 1970-е годы, когда вылавливалось по 500—600 тонн акул в год. Для добычи тёмных акул используются донные жаберные сети , в которые попадаются почти исключительно молодые акулы в возрасте до трёх лет, из них 18—28 % от всех новорождённых в первый год жизни. Демографические модели показывают, что промысел будет оставаться устойчивым при условии, что уровень смертности среди акул более 2 м в длину составит менее 4 % [48] . В дополнение к промышленному рыболовству тёмные акулы попадаются в качестве прилова в ярусы, предназначенные для тунца и рыбы-меч (в таких случаях туши акул обычно выбрасывают, сохраняя лишь ценные плавники). Этот вид ценится в спортивном рыболовстве. Большое количество тёмных акул, в основном неполовозрелых, вылавливают рыболовы-любители у берегов Южной Африки и Восточной Австралии. До резкого сокращения численности эта акула была одним из важнейших видов в турнирах Florida trophy shark [48] .
Меры по сохранению вида
Международный союз охраны природы (МСОП) оценил статус сохранности этого вида как «Близкий к уязвимому положению» (NT) во всём мире и «Уязвимый» (VU) в северо-западной Атлантике и Мексиканском заливе. Американское общество рыболовов оценило статус североамериканской популяции тёмных акул как «Уязвимый» [18] . Очень низкий уровень репродуктивного цикла делает этот вид чувствительным к перелову. Проведённая в 2006 году американской Национальной службой морского рыболовства оценка запасов показала, что численность сократилась до 15—20 % от уровня 1970-х годов. В 1998 году была запрещена коммерческая и любительская ловля тёмной акулы, но эти меры имели ограниченную эффективность в силу высокой смертности рыб из-за прилова. Кроме того, несмотря на запрет, в 2003 году рыболовами-любителями были пойманы около 2000 тёмных акул. В 2005 году Северная Каролина объявила о временном прекращении любительского промысла [54] . Для облегчения усилий по сохранению разработан метод молекулярной идентификации , позволяющий определить видовую принадлежность акульих плавников, поступивших на рынок, чтобы выявлять случаи браконьерства [55] .
Notes
- ↑ Линдберг Г. У. , Герд А. С. , Расс Т. С. Словарь названий морских промысловых рыб мировой фауны. — Ленинград: Наука, 1980. — С. 39. — 562 с.
- ↑ 1 2 3 4 Решетников Ю. С. , Котляр А. Н. , Расс Т. С. , Шатуновский М. И. Пятиязычный словарь названий животных. Рыбы. Latin, Russian, English, German, French. / edited by Acad. V. E. Sokolova . - M .: Rus. яз., 1989. — С. 31. — 12 500 экз. — ISBN 5-200-00237-0 .
- ↑ Губанов Е. П., Кондюрин В. В., Мягков Н. А. Акулы Мирового океана: Справочник-определитель. — М.: Агропромиздат, 1986. — С. 155. — 272 с.
- ↑ 1 2 3 Тёмная акула (англ.) в базе данных FishBase .
- ↑ 1 2 3 4 5 6 7 8 9 10 Carcharhinus obscurus (англ.) . The IUCN Red List of Threatened Species .
- ↑ Lesueur CA Description of several new species of North American fishes (part 1) // Journal of the Academy of Natural Sciences of Philadelphia. — 1818. — Vol. 1, № (2) . — P. 222—235.
- ↑ 1 2 3 4 5 6 7 8 9 10 Compagno LJV Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. — Rome: Food and Agricultural Organization, 1984. — P. 489—491. — ISBN 92-5-101384-5 .
- ↑ 1 2 3 4 5 6 7 8 Ebert DA Sharks, Rays, and Chimaeras of California. — California: University of California Press, 2003. — P. 149—152. — ISBN 0520234847 .
- ↑ Jordan DS, Gilbert CH Description of a new shark ( Carcharias lamiella ) from San Diego, California // Proceedings of the United States National Museum. — 1882. — Vol. 5, № 269 . — P. 110—111.
- ↑ 1 2 Heim, B. and Bourdon, J. Species from the Fossil Record: Carcharhinus obscurus . The Life and Times of Long Dead Sharks (April 20, 2009). Дата обращения 18 мая 2015.
- ↑ Portell RW, Hubbell G., Donovan SK, Green JK, Harpe DAT, Pickerill R. Miocene sharks in the Kendeace and Grand Bay formations of Carriacou, The Grenadines, Lesser Antilles // Caribbean Journal of Science. - 2008 .-- Vol. 44, № (3) . — P. 279—286.
- ↑ Cook TD, Murray AM, Simons EL, Attia YS, Chatrath PA Miocene selachian fauna from Moghra, Egypt // Historical Biology: an International Journal of Paleobiolog. — 2010. — P. 1029—2381.
- ↑ Sanchez-Villagra MR, Burnham RJ, Campbell DC, Feldmann RM, Gaffney ES, Kay RF, Lozsan R., Purdy R., Thewissen JGM A New Near-Shore Marine Fauna and Flora from the Early Neogene of Northwestern Venezuela // Journal of Paleontology. — 2000. — Vol. 74, № 5 . — P. 957—968. — DOI : 10.1666/0022-3360(2000)074<0957:ANNSMF>2.0.CO;2 .
- ↑ Cicimurri DJ, Knight JL Two Shark-bitten Whale Skeletons from Coastal Plain Deposits of South Carolina // Southeastern Naturalist. - 2009. - Vol. 8, № (1) . — P. 71—82. — DOI : 10.1656/058.008.0107 .
- ↑ Garrick JAF Sharks of the genus Carcharhinus // NOAA Technical Report, NMFS Circ.. — 1982. — Vol. 445.
- ↑ Compagno LJV Sharks of the Order Carcharhiniformes. — Princeton University Press. — P. 319—320. — ISBN 0-691-08453-X .
- ↑ 1 2 Naylor GJP The phylogenetic relationships among requiem and hammerhead sharks: inferring phylogeny when thousands of equally most parsimonious trees result // Cladistics. — 1999. — Vol. 8, № (4) . — P. 295—318. — DOI : 10.1111/j.1096-0031.1992.tb00073.x .
- ↑ 1 2 3 4 5 6 7 Knickle Craig. Biological report: duscky shark . Florida Museum of Natural History Ichthyology Department. Дата обращения 18 мая 2015.
- ↑ Natanson LJ, Casey JG, Kohler NE Age and growth estimates for the dusky shark, Carcharhinus obscurus , in the western North Atlantic Ocean // Fishery Bulletin. — 1995. — Vol. 93, № 1 . — P. 116—126.
- ↑ McEachran JD, Fechhelm JD Fishes of the Gulf of Mexico: Myxinformes to Gasterosteiformes. — University of Texas Press, 1998. — P. 87. — ISBN 0-292-75206-7 .
- ↑ 1 2 3 Last PR, Stevens JD Sharks and Rays of Australia. — 3-е. — Harvard University Press, 1993. — P. 254—255. — ISBN 0674034112 .
- ↑ Compagno L., Dando M., Fowler S. Sharks of the World. — Princeton University Press, 2005. — P. 302—303. — ISBN 978-0-691-12072-0 .
- ↑ Ovenden JR, Kashiwagi T., Broderick D., Giles J., Salini J. The extent of population genetic subdivision differs among four co-distributed shark species in the Indo-Australian archipelago // BMC Evolutionary Biology. - 2009. - Vol. 9, № 40 . — DOI : 10.1186/1471-2148-9-40 . — PMID 19216767 .
- ↑ Hoffmayer ER, Franks JS, Driggers WB (III), Grace MA Movements and Habitat Preferences of Dusky ( Carcharhinus obscurus ) and Silky ( Carcharhinus falciformis ) Sharks in the Northern Gulf of Mexico: Preliminary Results // 2009 MTI Bird and Fish Tracking Conference Proceedings. — 2009.
- ↑ 1 2 Van der Elst R. A Guide to the Common Sea Fishes of Southern Africa. — 3-е. — Struik, 1993. — P. 35. — ISBN 1868253945 .
- ↑ Huish MT, Benedict C. Sonic tracking of dusky sharks in Cape Fear River, North Carolina // Journal of the Elisha Mitchell Scientific Society. — 1977. — Vol. 93, № (1) . — P. 21—26.
- ↑ Schwartz FJ Five species of sharksuckers (family Echeneidae) in North Carolina // Journal of the North Carolina Academy of Science. — 2004. — Vol. 120, № (2) . — P. 44—49.
- ↑ Martin RA A Place For Sharks . ReefQuest Centre for Shark Research. Date of treatment May 22, 2015.
- ↑ Ruhnke TR, Caira JN Two new species of Anthobothrium van Beneden, 1850 (Tetraphyllidea: Phyllobothriidae) from carcharhinid sharks, with a redescription of Anthobothrium laciniatum Linton, 1890 // Systematic Parasitology. - 2009. - Vol. 72, № (3) . — P. 217—227. — DOI : 10.1007/s11230-008-9168-0 . — PMID 19189232 .
- ↑ Beveridge I., Campbell RA A revision of Dasyrhynchus Pintner (Cestoda, Trypanorhyncha), parasitic in elasmobranch and teleost fishes // Systematic Parasitology. — 1993. — Vol. 24, № 2 . — P. 129—157. — DOI : 10.1007/BF00009597 .
- ↑ Healy CJ A revision of Platybothrium Linton, 1890 (Tetraphyllidea : Onchobothriidae), with a phylogenetic analysis and comments on host-parasite associations // Systematic Parasitology. — 2003. — Vol. 56, № 2 . — P. 85—139. — DOI : 10.1023/A:1026135528505 . — PMID 14574090 .
- ↑ Linton E. Rhynchobothrium ingens spec. nov., a parasite of the dusky shark ( Carcharhinus obscurus ) // Journal of Parasitology. — 1921. — Vol. 8, № 1 . — P. 22—32. — DOI : 10.2307/3270938 .
- ↑ Knoff M., De Sao CSC, Pinto RM, Lanfredi RM, Gomes DC New records and expanded descriptions of Tentacularia coryphaenae and Hepatoxylon trichiuri homeacanth trypanorhynchs (Eucestoda) from carcharhinid sharks from the State of Santa Catarina off-shore, Brazil // Revista Brasileira de Parasitologia Veterinaria. — 2004. — Vol. 13, № 2 . — P. 72—80.
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- ↑ Bullard SA, Dippenaar SM, Hoffmayer ER, Benz GW New locality records for Dermophthirius carcharhini (Monogenea : Microbothriidae) and Dermophthirius maccallumi and a list of hosts and localities for species of Dermophthirius // Comparative Parasitology. — 2004. — Vol. 71, № 1 . — P. 78—80. — DOI : 10.1654/4093 .
- ↑ MacCullum GA Some new forms of parasitic worms // Zoopathologica; scientific contributions of the New York Zoological Society. — 1917. — Vol. 1, № 2 . — P. 1—75.
- ↑ Yamauchi T., Ota Y., Nagasawa K. Stibarobdella macrothela (Annelida, Hirudinida, Piscicolidae) from Elasmobranchs in Japanese Waters, with New Host Records // Biogeography. - 2008 .-- Vol. 10. — P. 53—57.
- ↑ Newbound DR, Knott B. Parasitic copepods from pelagic sharks in Western Australia // Bulletin of Marine Science. — 1999. — Vol. 65, № 3 . — P. 715—724.
- ↑ Jensen C., Schwartz FJ Atlantic Ocean occurrences of the sea lamprey, Petromyzon marinus (Petromyzontiformes: Petromyzontidae), parasitizing sandbar, Carcharhinus plumbeus , and dusky, C. obscurus (Carcharhiniformes: Carcharhinidae), sharks off North and South Carolina // Brimleyana. — 1994. — Vol. 21. — P. 69—76.
- ↑ 1 2 3 Gelsleichter J., Musick JA, Nichols S. Food habits of the smooth dogfish, Mustelus canis , dusky shark, Carcharhinus obscurus , Atlantic sharpnose shark, Rhizoprionodon terraenovae , and the sand tiger, Carcharias taurus , from the northwest Atlantic Ocean // Environmental Biology of Fishes. — 1999. — Vol. 54, № 2 . — P. 205—217.
- ↑ 1 2 3 4 Hussey NE, Cocks DT, Dudley SFJ, McCarthy ID, Wintner SP The condition conundrum: application of multiple condition indices to the dusky shark Carcharhinus obscurus // Marine Ecology Progress Series. - 2009. - Vol. 380. — P. 199—212. — DOI : 10.3354/meps07918 .
- ↑ Marin A. The Power of Shark Bites . ReefQuest Centre for Shark Research. Дата обращения 18 мая 2015.
- ↑ Gubanov EP Morphological characteristics of the requiem shark, Carcharinus obscurus , of the Indian Ocean // Journal of Ichthyology. - 1988. - Vol. 28, № (6) . — P. 68—73.
- ↑ Simpfendorfer CA, Goodreid A., McAuley RB Diet of three commercially important shark species from Western Australian waters // Marine and Freshwater Research. — 2001. — Vol. 52, № (7) . — DOI : 10.1071/MF01017 .
- ↑ 1 2 Smale MJ Occurrence and feeding of three shark species, Carcharhinus brachyurus , C. obscurus and Sphyrna zygaena , on the eastern Cape coast of South Africa // South African Journal of Marine Science. — 1991. — P. 31—42.
- ↑ Cockcroft VG, Cliff G., Ross GJB Shark predation on Indian Ocean bottlenose dolphins Tursiops truncatus off Natal, South Africa // South African Journal of Zoology. — 1989. — Vol. 24, № (4) . — P. 305—310.
- ↑ Pratt HL (Jr.). The storage of spermatozoa in the oviducal glands of western North Atlantic sharks. — 1993. — Vol. 38. — P. 139—149. — DOI : 10.1007/BF00842910 .
- ↑ 1 2 3 4 5 6 Fowler SL, Cavanagh RD, Camhi M., Burgess GH, Cailliet GM, Fordham SV, Simpfendorfer CA, Musick JA Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes. — International Union for Conservation of Nature and Natural Resources, 2005. — P. 106—109, 287—288. — ISBN 2831707005 .
- ↑ Natanson LJ, Kohler NE A preliminary estimate of age and growth of the dusky shark Carcharhinus obscurus from the South-West Indian Ocean, with comparisons to the western North Atlantic population // South African Journal of Marine Science. — 1996. — Vol. 17. — P. 217—224. — DOI : 10.2989/025776196784158572 .
- ↑ White WT Catch composition and reproductive biology of whaler sharks (Carcharhiniformes: Carcharhinidae) caught by fisheries in Indonesia // Journal of Fish Biology. - 2007. - Vol. 71, № (5) . — P. 1512—1540. — DOI : 10.1111/j.1095-8649.2007.01623.x. .
- ↑ Simpfendorfer CA, McAuley RB, Chidlow J., Unsworth P. Validated age and growth of the dusky shark, Carcharhinus obscurus, from Western Australian waters // Marine and Freshwater Research. - 2002. - Vol. 53, № (2) . — P. 567—573. — DOI : 10.1071/MF01131 .
- ↑ Simpfendorfer CA Growth rates of juvenile dusky sharks, Carcharhinus obscurus (Lesueur, 1818), from southwestern Australia estimated from tag-recapture data // Fishery Bulletin. — 2000. — Vol. 98, № (4) . — P. 811—822.
- ↑ ISAF Statistics on Attacking Species of Shark . Florida Museum of Natural History Ichthyology Department. Дата обращения 23 мая 2015.
- ↑ Species of Concern: Dusky Shark. (Jan. 6, 2009). NMFS Office of Protected Resources. Retrieved on May 18, 2009.
- ↑ Pank M., Stanhope M., Natanson L., Kohler N., Shivji M. Rapid and Simultaneous Identification of Body Parts from the Morphologically Similar Sharks Carcharhinus obscurus and Carcharhinus plumbeus (Carcharhinidae) Using Multiplex PCR // Marine Biotechnology. — 2001. — Vol. 3, № (3) . — P. 231—240. — DOI : 10.1007/s101260000071 . — PMID 14961360 .
Literature
- Ebert DA Sharks, Rays, and Chimaeras of California. — California: University of California Press, 2003. — P. 149—152. — ISBN 0520234847 .
- Compagno LJV Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. — Rome: Food and Agricultural Organization, 1984. — P. 489—491. — ISBN 92-5-101384-5 .
- Fowler SL, Cavanagh RD, Camhi M., Burgess GH, Cailliet GM, Fordham SV, Simpfendorfer CA, Musick JA Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes. — International Union for Conservation of Nature and Natural Resources, 2005. — P. 106—109, 287—288. — ISBN 2831707005 .
Links
- Вид Тёмная акула (англ.) в Мировом реестре морских видов ( World Register of Marine Species ).