Centroscyllium Fabricii
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The black dogfish (''Centroscyllium fabricii'') is a species of
dogfish shark Squalidae, more commonly known as dogfish, dog sharks, or spiny dogfish, are one of several families of sharks categorized under Squaliformes, making it the second largest order of sharks, numbering 119 species across 7 families. Having earned t ...
in the family Etmopteridae. It is common over the outer
continental shelf A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
and continental slope at depths of . Females generally inhabit deeper water than males, and depending on the region, smaller sharks may occur at different depths than larger ones. This species is distributed widely in the Atlantic Ocean, from Greenland and Iceland to Virginia and West Africa in the north, and off southwestern Africa and Argentina in the south. The largest member of its family, the black dogfish, typically measures long. It has a stocky, dark brown body that is darker below than above, and bears scattered, minute
bioluminescent Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorganisms including ...
organs. Its two dorsal fins are preceded by stout spines, and the
anal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
is absent. Active and
schooling A school is an educational institution designed to provide learning spaces and learning environments for the teaching of students under the direction of teachers. Most countries have systems of formal education, which is sometimes compuls ...
, the black dogfish is an opportunistic predator and
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
that mainly consumes
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
es, crustaceans, and
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s. It is
aplacental viviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
, with females producing litters of four to 40 pups that are sustained to term by a yolk sac. There is no well-defined
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
, and mating and birthing take place year-round. The black dogfish contributes significantly to the
bycatch Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
of deep-sea commercial fisheries operating in the
North Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe and ...
; it is of little commercial value and is usually discarded. As large portions of its range see little deepwater fishing activity and its northwestern Atlantic population seems to be stable, the International Union for Conservation of Nature has listed this species under Least Concern overall. It has been assessed as Near Threatened in the northeastern Atlantic, where its numbers may have declined from heavy fishing pressure.


Taxonomy and phylogeny

The first known specimen of the black dogfish was collected near
Julianehåb Qaqortoq, formerly Julianehåb, is a city in the Kujalleq municipalities of Greenland, municipality in southern Greenland, located near Cape Thorvaldsen. With a population of 3,050 in 2020, it is the most populous town and the municipal capital in ...
in Greenland and described by Danish zoologist
Johannes Reinhardt Johannes Theodor Reinhardt (3 December 1816, in Copenhagen – 23 October 1882, in Frederiksberg) was a Danish zoologist and herpetologist. The son of Johannes Christopher Hagemann Reinhardt. Biography He participated as botanist in the first ...
in his 1825 ''Ichthyologiske bidrag''. Reinhardt gave it the name ''Spinax fabricii'' in honor of missionary and naturalist Otto Fabricius, who pioneered the study of Greenlandic fishes. German biologists Johannes Müller and Jakob Henle, in their 1839–41 ''Systematische Beschreibung der Plagiostomen'', created the new genus ''
Centroscyllium ''Centroscyllium'' is a genus of big-eyed, deepwater dogfishes with no anal fin, a grey or black-brown body, and dorsal spines, with the second one being much larger than the first. Seven extant species are described. Species * '' Centroscyll ...
'' for this species. According to the IUCN, further
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
investigation is required to confirm the black dogfish in the
North Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe and ...
and the southeastern Atlantic represent the same species. A 2010 phylogenetic study by Nicolas Straube and colleagues, based on
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: * Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics *Nuclear space *Nuclear ...
and
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
, found the black dogfish is the sister species of the whitefin dogfish (''C. ritteri''), and the two are, in turn, sister to the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
formed by the granular dogfish (''C. granulatum'') and
combtooth dogfish The combtooth dogfish (''Centroscyllium nigrum'') is a little-known, deepwater dogfish shark, named after its teeth that are comb-shaped. Description The combtooth dogfish has no anal fin, grooved dorsal spines, two dorsal fins of about same ...
(''C. nigrum'').


Distribution and habitat

The black dogfish is a common species, with a wide but discontinuous distribution in the temperate waters of the Atlantic Ocean; its range abuts, but does not extend into, the Arctic Ocean. In the northeast, it occurs from Iceland to Sierra Leone, including the Faroe Islands, southern Norway, and the Rockall Trough and Porcupine Seabight off Ireland. In the northwest, it is found from southern Greenland and
Baffin Island Baffin Island (formerly Baffin Land), in the Canadian territory of Nunavut, is the largest island in Canada and the fifth-largest island in the world. Its area is , slightly larger than Spain; its population was 13,039 as of the 2021 Canadia ...
to Virginia, being particularly abundant in the Laurentian Channel, and may occur further south to the Gulf of Mexico off Alabama. In the southeast, it is found off Namibia and South Africa as far as Cape Agulhas. In the southwest, it has been recorded from the Beagle Channel at the southern tip of Argentina. Inhabiting the outer
continental shelf A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
and continental slope, the black dogfish is found mostly near the bottom in water deep. It is most common at depths of off Iceland, in the Rockall Trough, off Greenland, off northern Canada, and below off southern Africa. The species may venture closer to the surface in the northern extreme of its range, particularly during the dark, cold winter months. Depth segregation by sex has been documented in the North Atlantic, with females outnumbering males at depths greater than . Depth segregation by size varies by region: larger sharks are generally found in deeper water off western Greenland, in shallower water off western Iceland, and without pattern with respect to depth off eastern Iceland. The black dogfish prefers water temperatures of , though off northern Canada, it is most abundant in water of ; it can tolerate temperatures down to . There is some evidence that this species conducts seasonal migrations, spending winter and spring in shallower water. Sharks off northern Canada perform development-related movements (see below) not observed off western Greenland, suggesting the presence of two distinct
stocks Stocks are feet restraining devices that were used as a form of corporal punishment and public humiliation. The use of stocks is seen as early as Ancient Greece, where they are described as being in use in Solon's law code. The law describing ...
in the northwestern Atlantic.


Description

Adult black dogfish typically measure in length and can reach , making it the largest member of its family. Females attain a larger ultimate size than males. The shark has a rather stocky and laterally compressed body, with a moderately long, thick, and flattened snout that forms a very broad arch at the front. The sizable, horizontally oval eyes are a reflective green in life and lack
nictitating membrane The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
s (protective third eyelids); they are followed a short distance behind by much smaller
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
s (accessory respiratory openings). The nostrils are anteriorly placed and preceded by short flaps of skin. The mouth is wide and evenly arched, with thin lips and short but deep furrows around the corners. There are around 34 tooth rows in either side of both jaws; each tooth has three (occasionally up to five) slender cusps, with the central one the longest. Both dorsal fins are immediately preceded by stout, grooved spines, with the second much longer than the first. The small first dorsal fin has a rounded apex and a nearly straight trailing margin, with its origin lying behind the pectoral fins. The second dorsal fin is rather angular and has about double the area of the first, with its origin located opposite the midpoint of the
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods. Structure and function Structure In actinopterygians, the pelvic fin consists of two en ...
bases. The pectoral fins are small and rounded. The pelvic fins are about as large as the second dorsal fin, with rounded tips and nearly straight trailing margins. The caudal peduncle is short and leads to a broad caudal fin comprising less than a quarter of the total length; the upper lobe has a convex upper margin leading to a squared-off tip, while the lower lobe is indistinct. The skin is densely covered by tiny
dermal denticle A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as we ...
s; each one is recurved and thorn-like, rising from an irregular star-shaped base. This species is a plain dark brown above, darkening to almost black below, with white dorsal fin spines. Juvenile sharks have white edges on the dorsal, pectoral, and pelvic fins. Minute, luminescent dots are scattered about the skin without a regular pattern.


Biology and ecology

The black dogfish forms shoals or schools that tend to be larger during the winter and spring. Though fairly active, its swimming muscles exhibit lower activity of
glycolytic Glycolysis is the metabolic pathway that converts glucose () into pyruvate (). The free energy released in this process is used to form the high-energy molecules adenosine triphosphate (ATP) and reduced nicotinamide adenine dinucleotide (NADH ...
enzymes and higher activity of creatine phosphokinase compared to the shallow-water spiny dogfish (''Squalus acanthias''), suggesting a lesser capacity for bursts of speed. The lipid-filled liver comprises about one-fifth of its total weight and functions in maintaining neutral buoyancy. Potential predators of the black dogfish are larger sharks and bony fishes. It is one of several deep-sea sharks
parasitized Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
by the
barnacle A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in eros ...
''Anelasma squalicola'', which attaches in front of the second dorsal fin and impairs the reproductive development of its host. Other known parasites of this species include the
fluke Fluke may refer to: Biology * Fluke (fish), a species of marine flatfish * Fluke (tail), the lobes of the tail of a cetacean, such as dolphins or whales, ichthyosaurs, mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ...
''Otodistomum cestoides'', the
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s ''Neoalbionella fabricii'' and ''Neoalbionella centroscyllii'', and the
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
ns ''Haemogregarina delagei'' and ''Trypanosoma rajae''. Apparently opportunistic in feeding habits, the black dogfish typically hunts in open water, but also scavenges off the bottom. The bulk of its diet consists of a variety of
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
es, including
rattail Grenadiers or rattails are generally large, brown to black gadiform marine fish of the subfamily Macrourinae, the largest subfamily of the family Macrouridae. Found at great depths from the Arctic to Antarctic, members of this subfamily are am ...
s, whitings, rockfishes, lanternfishes, and barracudinas, as well as pelagic crustaceans such as krill and
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are refer ...
, and
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s. Fish become a progressively more important food source as the shark ages, while crustaceans become less important. Infrequently, polychaete worms and jellyfish are also eaten. In the northwestern Atlantic, Greenland halibut (''Reinhardtius hippoglossoides'') and rattail offal discarded from fishing vessels have become a major source of food for this species, particularly for older sharks that are capable of consuming larger pieces such as heads.


Life history

Reproduction in the black dogfish occurs year-round, with no well-defined seasonal pattern. This species is
aplacental viviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
, in which the developing
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
s are retained inside the uterus and are sustained to term solely by yolk. Mature females have two functional ovaries and two functional uteruses. Fertilized eggs are ovulated into the uterus at a diameter of , though a few may be retained in the ovary; the eggs are not enclosed in a capsule. The external yolk sac is fully resorbed when the embryo is close to term, with the remaining yolk having been transferred to an internal yolk sac attached to the
intestine The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
. The internal yolk sac serves to provision the newborn shark until it learns to feed. The litter size ranges from four to 40. Newborns measure long. Various authors have reported the size at maturity as between for males and for females, reflecting differences between geographical areas. Off northern Canada, females give birth in the portion of the Laurentian Channel less than deep. As the young grow, they migrate into the deeper parts of the Channel, and eventually a long distance northward over the Grand Banks or the
Labrador Shelf , nickname = "The Big Land" , etymology = , subdivision_type = Country , subdivision_name = Canada , subdivision_type1 = Province , subdivision_name1 ...
, to the deep continental slope. This movement pattern has not been observed in black dogfish inhabiting adjacent waters off western Greenland. A number of anomalous hermaphroditic specimens have been documented.


Human interactions

The black dogfish is harmless to humans and of little commercial value. Substantial numbers are caught incidentally by commercial deep-sea trawl,
gillnet Gillnetting is a fishing method that uses gillnets: vertical panels of netting that hang from a line with regularly spaced floaters that hold the line on the surface of the water. The floats are sometimes called "corks" and the line with corks is ...
, and
longline Long line or longline may refer to: *''Long Line'', an album by Peter Wolf *Long line (topology), or Alexandroff line, a topological space *Long line (telecommunications), a transmission line in a long-distance communications network *Longline fish ...
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, both ...
operating throughout the North Atlantic, including the Icelandic Greenland halibut fishery, the French mixed-species trawl fishery, and the Canadian Greenland halibut,
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
, redfish, monkfish, and witch fisheries. Captured sharks are usually discarded, though in recent years this and other small deepwater sharks have been increasingly retained and used for fishmeal. Reported catches by European countries, of which France made the largest contribution, have followed a declining trend from 486
ton Ton is the name of any one of several units of measure. It has a long history and has acquired several meanings and uses. Mainly it describes units of weight. Confusion can arise because ''ton'' can mean * the long ton, which is 2,240 pounds ...
s in 2001 to 35 tons in 2006. The average catch by Canadian fisheries was 68 tons per year from 1996 to 2005. The black dogfish occurs mostly too deep for fisheries off southern Africa; in the remainder of its range, little information is available on fishery impact. The International Union for Conservation of Nature (IUCN) has listed the black dogfish under Least Concern worldwide; it is minimally affected by fishing activity across many parts of its range, while its population in the northwestern Atlantic presently seems to be stable and may have increased from 1978 to 1995. By contrast, the intensity of deepwater fisheries in the northeastern Atlantic has led the IUCN to give this species a regional assessment of Near Threatened. The reproductive characteristics of the black dogfish, such as a large female maturation size, may render it susceptible to
overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in th ...
, though it is more fecund than other deep-sea dogfish sharks. In the northeastern Atlantic, catches of this species are managed as part of the total allowable catch for deep-sea sharks set by the International Council for the Exploration of the Sea (ICES).


References


External links


''Centroscyllium fabricii'', Black dogfish
a
FishBase


a
Florida Museum of Natural History Ichthyology Department
{{DEFAULTSORT:Black dogfish Centroscyllium Fish of the Atlantic Ocean Fish described in 1825 Taxa named by Johan Reinhardt