Velvet belly lanternshark: Difference between revisions
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==Taxonomy== |
==Taxonomy== |
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The velvet belly was originally described as ''Squalus spinax'' by Swedish natural historian [[Karl Linnaeus]], known as the "father of [[taxonomy]]", in the 1758 tenth edition of ''[[Systema Naturae]]''. He did not designate a [[type specimen]]; the [[specific epithet]] ''spinax'' is in reference to the spiny [[dorsal fin]]s. This species was later moved to the genus ''Etmopterus'' via the [[synonym (biology)|synonymy]] of [[Constantine Samuel Rafinesque]]'s ''Etmopterus aculeatus'' with ''Squalus spinax''.<ref name="compagno">{{cite book |author=Compagno, L.J.V. |date=1984 |title=Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date |place=Rome |publisher=Food and Agricultural Organization |isbn=9251013845. |page=85}}</ref> The velvet belly is grouped with the [[Caribbean lanternshark]] (''E. hillianus''), [[fringefin lanternshark]] (''E. schultzi''), [[brown lanternshark]] (''E. unicolor''), [[broadbanded lanternshark]] (''E. gracilispinis''), [[combtooth lanternshark]] (''E. decacuspidatus''), and [[dwarf lanternshark]] (''E. perryi'') in having irregularly arranged, needle-shaped [[dermal denticle]]s.<ref name="springer and burgess">{{cite journal |title=Two New Dwarf Dogsharks (''Etmopterus'', Squalidae), Found off the Caribbean Coast of Colombia |author=Springer, S. and G.H. Burgess |journal=Copeia |volume=1985 |issue=3 |date=August 5, 1985 |pages=584–591}}</ref> Its [[common name]] comes from this shark's black ventral surface, which is sharply delineated from the rest of its body like a patch of [[velvet]].<ref name="ellis">{{cite book |author=Ellis, R. |title=Deep Atlantic: Life, Death, and Exploration in the Abyss |publisher=The Lyons Press |date=1996 |isbn=1558216634 |pages=195–196}}</ref> |
The velvet belly was originally described as ''Squalus spinax'' by Swedish natural historian [[Karl Linnaeus]], known as the "father of [[taxonomy]]", in the 1758 tenth edition of ''[[Systema Naturae]]''. He did not designate a [[type specimen]]; the [[specific epithet]] ''spinax'' is in reference to the spiny [[dorsal fin]]s. This species was later moved to the genus ''Etmopterus'' via the [[synonym (biology)|synonymy]] of [[Constantine Samuel Rafinesque]]'s ''Etmopterus aculeatus'' with ''Squalus spinax''.<ref name="compagno">{{cite book |author=Compagno, L.J.V. |date=1984 |title=Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date |place=Rome |publisher=Food and Agricultural Organization |isbn=9251013845. |page=85}}</ref> The velvet belly is grouped with the [[Caribbean lanternshark]] (''E. hillianus''), [[fringefin lanternshark]] (''E. schultzi''), [[brown lanternshark]] (''E. unicolor''), [[broadbanded lanternshark]] (''E. gracilispinis''), [[combtooth lanternshark]] (''E. decacuspidatus''), and [[dwarf lanternshark]] (''E. perryi'') in having irregularly arranged, needle-shaped [[dermal denticle]]s.<ref name="springer and burgess">{{cite journal |title=Two New Dwarf Dogsharks (''Etmopterus'', Squalidae), Found off the Caribbean Coast of Colombia |author=Springer, S. and G.H. Burgess |journal=Copeia |volume=1985 |issue=3 |date=August 5, 1985 |pages=584–591 |doi=10.2307/1444748}}</ref> Its [[common name]] comes from this shark's black ventral surface, which is sharply delineated from the rest of its body like a patch of [[velvet]].<ref name="ellis">{{cite book |author=Ellis, R. |title=Deep Atlantic: Life, Death, and Exploration in the Abyss |publisher=The Lyons Press |date=1996 |isbn=1558216634 |pages=195–196}}</ref> |
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==Distribution and habitat== |
==Distribution and habitat== |
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The range of the velvet belly is in the eastern Atlantic, extending from Iceland and Norway to Gabon, including the [[Mediterranean Sea]], the [[Azores]], the [[Canary Islands]], and [[Cape Verde]]. It has also been reported off [[Cape Province]], South Africa. This shark mainly inhabits the outer [[Continental shelf|continental]] and insular shelves and upper slopes over mud or clay, from close to the bottom to the middle of the water column.<ref name="compagno"/><ref name="sion et al">{{cite journal |author=Sion, L., Bozzano, A., D’Onghia, G., Capezzuto, F. and Panza, M. |title=Chondrichthyes species in deep waters of the Mediterranean Sea |journal=Scientia Marina |volume=68 |issue=S3 |date=2004 |pages=153–162}}</ref> It is most common at a depth of {{convert|200|-|500|m|ft|abbr=on}}, though in the [[Rockall Trough]] it is only found at a depth of {{convert|500|-|750|m|ft|abbr=on}}.<ref name="martin"/><ref name="mauchline and gordon">{{cite journal |author=Mauchline, J. and Gordon, J.D.M. |title=Diets of the sharks and chimaeroids of the Rockall Trough, northeastern Atlantic Ocean |journal=Marine Biology |volume=75 |pages=269–278 |date=1983}}</ref> This species has been reported from as shallow as {{convert|70|m|ft|abbr=on}}, and as deep as {{convert|2490|m|ft|abbr=on}}.<ref name="jones et al"/> |
The range of the velvet belly is in the eastern Atlantic, extending from Iceland and Norway to Gabon, including the [[Mediterranean Sea]], the [[Azores]], the [[Canary Islands]], and [[Cape Verde]]. It has also been reported off [[Cape Province]], South Africa. This shark mainly inhabits the outer [[Continental shelf|continental]] and insular shelves and upper slopes over mud or clay, from close to the bottom to the middle of the water column.<ref name="compagno"/><ref name="sion et al">{{cite journal |author=Sion, L., Bozzano, A., D’Onghia, G., Capezzuto, F. and Panza, M. |title=Chondrichthyes species in deep waters of the Mediterranean Sea |journal=Scientia Marina |volume=68 |issue=S3 |date=2004 |pages=153–162}}</ref> It is most common at a depth of {{convert|200|-|500|m|ft|abbr=on}}, though in the [[Rockall Trough]] it is only found at a depth of {{convert|500|-|750|m|ft|abbr=on}}.<ref name="martin"/><ref name="mauchline and gordon">{{cite journal |author=Mauchline, J. and Gordon, J.D.M. |title=Diets of the sharks and chimaeroids of the Rockall Trough, northeastern Atlantic Ocean |journal=Marine Biology |volume=75 |pages=269–278 |date=1983 |doi=10.1007/BF00406012}}</ref> This species has been reported from as shallow as {{convert|70|m|ft|abbr=on}}, and as deep as {{convert|2490|m|ft|abbr=on}}.<ref name="jones et al"/> |
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==Description== |
==Description== |
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The velvet belly is a robustly built shark with a moderately long, broad, flattened snout. The mouth has thin, smooth lips. The upper teeth are small, with a narrow central [[Cusp (dentistry)|cusp]] and usually fewer than three pairs of lateral cusplets. The lower teeth are much larger, with a strongly slanted blade-like cusp at the top and interlocking bases. The five pairs of [[gill slit]]s are tiny, comparable in size to the [[spiracle]]s. Both dorsal fins bear stout, grooved spines at the front, with the second much longer than the first and curved. The first dorsal fin originates behind the short and rounded [[pectoral fin]]s; the second dorsal fin is twice the size of the first and originates behind the [[pelvic fin]]s. The [[anal fin]] is absent. The tail is slender, leading to a long [[caudal fin]] with a small lower lobe and a low upper lobe with a prominent ventral notch near the tip.<ref name="compagno"/> |
The velvet belly is a robustly built shark with a moderately long, broad, flattened snout. The mouth has thin, smooth lips. The upper teeth are small, with a narrow central [[Cusp (dentistry)|cusp]] and usually fewer than three pairs of lateral cusplets. The lower teeth are much larger, with a strongly slanted blade-like cusp at the top and interlocking bases. The five pairs of [[gill slit]]s are tiny, comparable in size to the [[spiracle]]s. Both dorsal fins bear stout, grooved spines at the front, with the second much longer than the first and curved. The first dorsal fin originates behind the short and rounded [[pectoral fin]]s; the second dorsal fin is twice the size of the first and originates behind the [[pelvic fin]]s. The [[anal fin]] is absent. The tail is slender, leading to a long [[caudal fin]] with a small lower lobe and a low upper lobe with a prominent ventral notch near the tip.<ref name="compagno"/> |
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The [[dermal denticle]]s are thin with hooked tips, arranged without a regular pattern well-separated from one another. The coloration is brown above, abruptly transitioning to black below. There are thin black marks above and behind the pelvic fins, and along the caudal fin.<ref name="compagno"/> The velvet belly possesses numerous photophores that emit a blue-green light visible from {{convert|3|-|4|m|ft|abbr=on}} away.<ref name="ellis"/> Varying densities of photophores are arranged in nine patches on the shark's sides and belly, creating a pattern unique to this species: photophores are present along the [[lateral line]], scattered beneath the head but excluding the mouth, evenly on the belly, and concentrated around the pectoral fins and beneath the [[caudal peduncle]].<ref name="martin">Martin, R.A. [http://www.elasmo-research.org/education/ecology/deepsea-velvetbelly.htm Deep Sea: Velvetbelly Lanternshark]. ReefQuest Centre for Shark Research. Retrieved on June 24, 2009.</ref><ref name="claes and mallefet">{{cite journal |author=Claes, J.M. and Mallefet, J. |title=Early development of bioluminescence suggests camouflage by counter-illumination in the velvet belly lantern shark ''Etmopterus spinax'' (Squaloidea: Etmopteridae) |journal=Journal of Fish Biology |volume=73 |issue=6 |pages=1337–1350 |date=2008}}</ref> The maximum reported length is {{convert|60|cm|in|abbr=on}}, although few are longer than 45 cm (18 in).<ref name="martin"/> Females are larger than males.<ref name="hickling"/> |
The [[dermal denticle]]s are thin with hooked tips, arranged without a regular pattern well-separated from one another. The coloration is brown above, abruptly transitioning to black below. There are thin black marks above and behind the pelvic fins, and along the caudal fin.<ref name="compagno"/> The velvet belly possesses numerous photophores that emit a blue-green light visible from {{convert|3|-|4|m|ft|abbr=on}} away.<ref name="ellis"/> Varying densities of photophores are arranged in nine patches on the shark's sides and belly, creating a pattern unique to this species: photophores are present along the [[lateral line]], scattered beneath the head but excluding the mouth, evenly on the belly, and concentrated around the pectoral fins and beneath the [[caudal peduncle]].<ref name="martin">Martin, R.A. [http://www.elasmo-research.org/education/ecology/deepsea-velvetbelly.htm Deep Sea: Velvetbelly Lanternshark]. ReefQuest Centre for Shark Research. Retrieved on June 24, 2009.</ref><ref name="claes and mallefet">{{cite journal |author=Claes, J.M. and Mallefet, J. |title=Early development of bioluminescence suggests camouflage by counter-illumination in the velvet belly lantern shark ''Etmopterus spinax'' (Squaloidea: Etmopteridae) |journal=Journal of Fish Biology |volume=73 |issue=6 |pages=1337–1350 |date=2008 |doi=10.1111/j.1095-8649.2008.02006.x}}</ref> The maximum reported length is {{convert|60|cm|in|abbr=on}}, although few are longer than 45 cm (18 in).<ref name="martin"/> Females are larger than males.<ref name="hickling"/> |
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==Biology and ecology== |
==Biology and ecology== |
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Along with the [[blackmouth catshark]] (''Galeus melastomus'') and the [[Portuguese dogfish]] (''Centroscymnus coelolepis''), the velvet belly is one of the most abundant deep-sea sharks in the northeastern Atlantic.<ref name="carrasson et al">{{cite journal |author=Carrassón, M., Stefanescu, C. and Cartes, J.E. |title=Diets and bathymetric distributions of two bathyal sharks of the Catalan deep sea (western Mediterranean) |journal=Marine Ecology Progress Series |volume=82 |issue=1 |pages=21–30 |date=1992}}</ref> It is found individually or in small [[shoaling and schooling|shoals]].<ref name="klimpel">{{cite journal |author=Klimpel, S., Palm, H.W. and Seehagen, A. |title=Metazoan parasites and food composition of juvenile ''Etmopterus spinax'' (L., 1758) (Dalatiidae, Squaliformes) from the Norwegian Deep |journal=Parasitology Research |date=2003 |volume=89 |pages=245–251}}</ref> Samplings in the Mediterranean have found females outnumbering males across all ages; this imbalance increases in the older age classes.<ref name="gennari and scacco">{{cite journal |author=Gennari, E. and Scacco, U. |title=First age and growth estimates in the deep water shark, ''Etmopterus Spinax'' (Linnaeus, 1758), by deep coned vertebral analysis |journal=Marine Biology |volume=152 |issue=5 |pages=1207–1214}}</ref> In the [[Rockall Trough]] and the [[Catalan Sea]], large adults are found in deeper waters than juveniles, which may serve to reduce competition between the two groups.<ref name="carrasson et al"/> However, this pattern has not been observed at other sites in the eastern Mediterranean.<ref name="jones et al">{{cite journal |author=Jones, E.G., Tselepides, E., Bagley, P.M., Collins, M.A. and Priede, I.G. |title=Bathymetric distribution of some benthic and benthopelagic species attracted to baited cameras and traps in the deep eastern Mediterranean |journal=Marine Ecology Progress Series |volume=251 |pages=75–86 |date=2003}}</ref> |
Along with the [[blackmouth catshark]] (''Galeus melastomus'') and the [[Portuguese dogfish]] (''Centroscymnus coelolepis''), the velvet belly is one of the most abundant deep-sea sharks in the northeastern Atlantic.<ref name="carrasson et al">{{cite journal |author=Carrassón, M., Stefanescu, C. and Cartes, J.E. |title=Diets and bathymetric distributions of two bathyal sharks of the Catalan deep sea (western Mediterranean) |journal=Marine Ecology Progress Series |volume=82 |issue=1 |pages=21–30 |date=1992 |doi=10.3354/meps082021}}</ref> It is found individually or in small [[shoaling and schooling|shoals]].<ref name="klimpel">{{cite journal |author=Klimpel, S., Palm, H.W. and Seehagen, A. |title=Metazoan parasites and food composition of juvenile ''Etmopterus spinax'' (L., 1758) (Dalatiidae, Squaliformes) from the Norwegian Deep |journal=Parasitology Research |date=2003 |volume=89 |pages=245–251 |pmid=12632160 |issue=4 |doi=10.1007/s00436-002-0741-1}}</ref> Samplings in the Mediterranean have found females outnumbering males across all ages; this imbalance increases in the older age classes.<ref name="gennari and scacco">{{cite journal |author=Gennari, E. and Scacco, U. |title=First age and growth estimates in the deep water shark, ''Etmopterus Spinax'' (Linnaeus, 1758), by deep coned vertebral analysis |journal=Marine Biology |volume=152 |issue=5 |pages=1207–1214 |doi=10.1007/s00227-007-0769-y |year=2007}}</ref> In the [[Rockall Trough]] and the [[Catalan Sea]], large adults are found in deeper waters than juveniles, which may serve to reduce competition between the two groups.<ref name="carrasson et al"/> However, this pattern has not been observed at other sites in the eastern Mediterranean.<ref name="jones et al">{{cite journal |author=Jones, E.G., Tselepides, E., Bagley, P.M., Collins, M.A. and Priede, I.G. |title=Bathymetric distribution of some benthic and benthopelagic species attracted to baited cameras and traps in the deep eastern Mediterranean |journal=Marine Ecology Progress Series |volume=251 |pages=75–86 |date=2003 |doi=10.3354/meps251075}}</ref> |
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The velvet belly's liver accounts for 17% of its body mass, three-quarters of which is [[oil]], making it nearly [[neutral buoyancy|neutrally buoyant]].<ref>{{cite journal |author=Schmidt-Nielsen, S., Flood, A. and Stene, J. |date=1934 |title=On the size of the liver of some gristly fishes, their content of fat and vitamin A |journal=Kongeleige Norske Videnskabers Selskab Forhandlinger |volume=7 |pages=47–50}}</ref> To deal with the higher concentrations of heavy metals in the deep sea, the velvet belly has [[T-cell]]s in its [[bloodstream]] that can identify and mark toxic compounds for elimination. These T-cells are produced by a [[lymphatic system|lymphomyeloid]] [[gland]] in its [[esophagus]] called a "[[Leydig's organ]]", which is also found in some other sharks and rays. In its liver, there are also specialized proteins capable of detoxifying [[cadmium]], [[copper]], [[mercury (element)|mercury]], [[zinc]], and other toxic contaminants.<ref name="martin"/> The velvet belly's bioluminescence is thought to function in counter-illumination, which eliminates the shark's silhouette and camouflages it from upward-looking predators.<ref name="claes and mallefet"/> Its bioluminescence may also serve a social function, such as finding mates or coordinating groups, as the pattern is species-specific. The velvet belly is an important food of larger fishes such as other sharks; a major predator of this species is the [[longnosed skate]] (''Dipturus oxyrinchus'').<ref name="martin"/><ref name="klimpel"/> |
The velvet belly's liver accounts for 17% of its body mass, three-quarters of which is [[oil]], making it nearly [[neutral buoyancy|neutrally buoyant]].<ref>{{cite journal |author=Schmidt-Nielsen, S., Flood, A. and Stene, J. |date=1934 |title=On the size of the liver of some gristly fishes, their content of fat and vitamin A |journal=Kongeleige Norske Videnskabers Selskab Forhandlinger |volume=7 |pages=47–50}}</ref> To deal with the higher concentrations of heavy metals in the deep sea, the velvet belly has [[T-cell]]s in its [[bloodstream]] that can identify and mark toxic compounds for elimination. These T-cells are produced by a [[lymphatic system|lymphomyeloid]] [[gland]] in its [[esophagus]] called a "[[Leydig's organ]]", which is also found in some other sharks and rays. In its liver, there are also specialized proteins capable of detoxifying [[cadmium]], [[copper]], [[mercury (element)|mercury]], [[zinc]], and other toxic contaminants.<ref name="martin"/> The velvet belly's bioluminescence is thought to function in counter-illumination, which eliminates the shark's silhouette and camouflages it from upward-looking predators.<ref name="claes and mallefet"/> Its bioluminescence may also serve a social function, such as finding mates or coordinating groups, as the pattern is species-specific. The velvet belly is an important food of larger fishes such as other sharks; a major predator of this species is the [[longnosed skate]] (''Dipturus oxyrinchus'').<ref name="martin"/><ref name="klimpel"/> |
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[[Image:Etmopterus spinax 010.jpg|thumb|Velvet bellies are often heavily parasitized; this shark has an ''Anelasma squalicola'' barnacle attached near the fin spine.]] |
[[Image:Etmopterus spinax 010.jpg|thumb|Velvet bellies are often heavily parasitized; this shark has an ''Anelasma squalicola'' barnacle attached near the fin spine.]] |
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Numerous [[parasite]]s are known for this species, and both juveniles and adults often carry heavy parasite loads. Known internal parasites include the [[monogenea]]n ''Squalonchocotyle spinacis'', the [[tapeworm]]s ''Aporhynchus norvegicus'', ''Lacistorhynchus tenuis'', and ''Phyllobothrium squali'', and the [[nematode]]s ''Anisakis simplex'' and ''Hysterothylacium aduncum''. Some of these parasites utilize the velvet belly's prey as intermediate hosts and are acquired via ingestion, while others use the shark itself as an intermediate host.<ref name="klimpel">{{cite journal |author=Klimpel, S., Palm, H.W. and Seehagen, A. |title=Metazoan parasites and food composition of juvenile ''Etmopterus spinax'' (L., 1758) (Dalatiidae, Squaliformes) from the Norwegian Deep |journal=Parasitology Research |date=2003 |volume=89 |pages=245–251}}</ref> The [[barnacle]] ''Anelasma squalicola'' is an external parasite that attaches to the shark's spine socket and penetrates deeply into the muscle, in the process often providing an attachment site for a second (and rarely a third) barnacle. Infestation by this barnacle reduces the host's [[fecundity]] by impairing the development of the [[reproductive organ]]s.<ref name="hickling">{{cite journal |title=On the small deep-sea shark Etmopterus spinax L., and its cirripede parasite Anelasma squalicola (Lovén) |author=Hickling, C.F. |journal=Journal of the Linnean Society of London, Zoology |volume=45 |issue=303 |pages=17–24}}</ref> |
Numerous [[parasite]]s are known for this species, and both juveniles and adults often carry heavy parasite loads. Known internal parasites include the [[monogenea]]n ''Squalonchocotyle spinacis'', the [[tapeworm]]s ''Aporhynchus norvegicus'', ''Lacistorhynchus tenuis'', and ''Phyllobothrium squali'', and the [[nematode]]s ''Anisakis simplex'' and ''Hysterothylacium aduncum''. Some of these parasites utilize the velvet belly's prey as intermediate hosts and are acquired via ingestion, while others use the shark itself as an intermediate host.<ref name="klimpel">{{cite journal |author=Klimpel, S., Palm, H.W. and Seehagen, A. |title=Metazoan parasites and food composition of juvenile ''Etmopterus spinax'' (L., 1758) (Dalatiidae, Squaliformes) from the Norwegian Deep |journal=Parasitology Research |date=2003 |volume=89 |pages=245–251 |pmid=12632160 |issue=4 |doi=10.1007/s00436-002-0741-1}}</ref> The [[barnacle]] ''Anelasma squalicola'' is an external parasite that attaches to the shark's spine socket and penetrates deeply into the muscle, in the process often providing an attachment site for a second (and rarely a third) barnacle. Infestation by this barnacle reduces the host's [[fecundity]] by impairing the development of the [[reproductive organ]]s.<ref name="hickling">{{cite journal |title=On the small deep-sea shark Etmopterus spinax L., and its cirripede parasite Anelasma squalicola (Lovén) |author=Hickling, C.F. |journal=Journal of the Linnean Society of London, Zoology |volume=45 |issue=303 |pages=17–24 |doi=10.1111/j.1096-3642.1963.tb00484.x |year=1963}}</ref> |
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===Feeding=== |
===Feeding=== |
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A generalist predator, velvet bellies feed on crustaceans (e.g. [[Pasiphaeidae|pasiphaeid shrimp]] and [[krill]]), cephalopods (e.g. [[Ommastrephidae|ommastrephid squid]] and [[Sepiolidae|sepiolid]]s), and bony fishes (e.g. [[Sternoptychidae|shads]], [[barracudina]]s, [[lanternfish]]es, and [[Gadidae|pouts]]).<ref name="martin"/> Sharks off [[Italy]] also eat small amounts of nematodes, [[polychaete worm]]s, and other cartilaginous fishes.<ref>{{cite book |author=Serena, F., Cecchi, E., Mancusi, C. and Pajetta, R. |chapter=Contribution to the knowledge of the biology of ''Etmopterus spinax'' (Linnaeus 1758) (Chondrichthyes, Etmopteridae) |pages=388–394 |title=Deep Sea 2003: Conference on the Governance and Management of Deep-sea Fisheries |publisher=Food and Agricultural Organization |date=2006 |isbn=9251054576 |editor=FAO}}</ref> Studies of velvet bellies off Norway and [[Portugal]], and in the Rockall Trough, have found that small sharks under {{convert|27|cm|in|abbr=on}} long feed mainly on the krill ''[[Meganyctiphanes norvegica]]'' and the small fish ''[[Maurolicus muelleri]]''. As the sharks grow larger, their diet becomes more varied, consisting mainly of squid and the shrimp ''[[Pasiphaea tarda]]'', as well fishes other than ''M. muelleri''.<ref name="klimpel"/><ref name="mauchline and gordon"/><ref>{{cite journal |author=Neiva, J., Coelho, R. and Erzini, K. |title=Feeding habits of the velvet belly lanternshark ''Etmopterus spinax'' (Chondrichthyes: Etmopteridae) off the Algarve, southern Portugal |journal=Journal of the Marine Biological Association of the United Kingdom |date=2006 |volume=86 |issue=4 |pages=835–841}}</ref> It has been speculated that smaller velvet bellies may be too slow to catch fast-moving cephalopods.<ref name="klimpel"/> The cephalopod diet of adults overlaps with that of the Portuguese dogfish; the latter species may avoid competition with the velvet belly by living in deeper water.<ref name="carrasson et al"/> The bite force exerted by the velvet belly is only around 1 [[Newton (unit)|N]].<ref>{{cite journal |author=Huber, D.R., Claes, J.M., Mallefet, J. and Herrel, A. |title=Is Extreme Bite Performance Associated with Extreme Morphologies in Sharks? |journal=Physiological and Biochemical Zoology |volume=82 |issue=1 |pages=20–28 |date=2009}}</ref> |
A generalist predator, velvet bellies feed on crustaceans (e.g. [[Pasiphaeidae|pasiphaeid shrimp]] and [[krill]]), cephalopods (e.g. [[Ommastrephidae|ommastrephid squid]] and [[Sepiolidae|sepiolid]]s), and bony fishes (e.g. [[Sternoptychidae|shads]], [[barracudina]]s, [[lanternfish]]es, and [[Gadidae|pouts]]).<ref name="martin"/> Sharks off [[Italy]] also eat small amounts of nematodes, [[polychaete worm]]s, and other cartilaginous fishes.<ref>{{cite book |author=Serena, F., Cecchi, E., Mancusi, C. and Pajetta, R. |chapter=Contribution to the knowledge of the biology of ''Etmopterus spinax'' (Linnaeus 1758) (Chondrichthyes, Etmopteridae) |pages=388–394 |title=Deep Sea 2003: Conference on the Governance and Management of Deep-sea Fisheries |publisher=Food and Agricultural Organization |date=2006 |isbn=9251054576 |editor=FAO}}</ref> Studies of velvet bellies off Norway and [[Portugal]], and in the Rockall Trough, have found that small sharks under {{convert|27|cm|in|abbr=on}} long feed mainly on the krill ''[[Meganyctiphanes norvegica]]'' and the small fish ''[[Maurolicus muelleri]]''. As the sharks grow larger, their diet becomes more varied, consisting mainly of squid and the shrimp ''[[Pasiphaea tarda]]'', as well fishes other than ''M. muelleri''.<ref name="klimpel"/><ref name="mauchline and gordon"/><ref>{{cite journal |author=Neiva, J., Coelho, R. and Erzini, K. |title=Feeding habits of the velvet belly lanternshark ''Etmopterus spinax'' (Chondrichthyes: Etmopteridae) off the Algarve, southern Portugal |journal=Journal of the Marine Biological Association of the United Kingdom |date=2006 |volume=86 |issue=4 |pages=835–841 |doi=10.1017/S0025315406013762}}</ref> It has been speculated that smaller velvet bellies may be too slow to catch fast-moving cephalopods.<ref name="klimpel"/> The cephalopod diet of adults overlaps with that of the Portuguese dogfish; the latter species may avoid competition with the velvet belly by living in deeper water.<ref name="carrasson et al"/> The bite force exerted by the velvet belly is only around 1 [[Newton (unit)|N]].<ref>{{cite journal |author=Huber, D.R., Claes, J.M., Mallefet, J. and Herrel, A. |title=Is Extreme Bite Performance Associated with Extreme Morphologies in Sharks? |journal=Physiological and Biochemical Zoology |volume=82 |issue=1 |pages=20–28 |date=2009 |doi=10.1086/588177 |pmid=19006469}}</ref> |
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===Life history=== |
===Life history=== |
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The velvet belly is [[ovoviviparous]], with the [[embryo]]s hatching inside the [[uterus]] and being sustained by a [[yolk sac]]. The reproductive cycle may be 2–3 years long, with [[ovulation]] occurring in early autumn, [[fertilization]] in the summer (or possibly in the winter if females are capable of storing sperm), and parturition in late winter or early spring. The [[gestation period]] is under one year.<ref name="hickling"/><ref name="coelho and erzini">{{cite journal |author=Coelho, R. and Erzini, K. |title=Life history of a wide-ranging deepwater lantern shark in the north-east Atlantic, ''Etmopterus spinax'' (Chondrichthyes: Etmopteridae), with implications for conservation |journal=Journal of Fish Biology |volume=73 |issue=6 |pages=1419–1443 |date=2008}}</ref> The litter size is 6–20, with the number of young increasing with female size. At birth the young measure {{convert|12|-|14|cm|in|abbr=on}} long.<ref name="compagno"/><ref name="coelho and erzini"/> The shark's bioluminescence develops before birth; the yolk sac is [[fluorescence|fluorescent]] before any photophores have formed, suggesting that the mother transfers luminescent materials to her offspring. The first luminous tissue appears when the embryo is {{convert|55|mm|in|abbr=on}} long, and the complete pattern is laid down by the time it is {{convert|95|mm|in|abbr=on}} long. At birth, the young shark is already capable of counter-illumination with 80% of its ventral surface luminescent.<ref name="claes and mallefet"/> |
The velvet belly is [[ovoviviparous]], with the [[embryo]]s hatching inside the [[uterus]] and being sustained by a [[yolk sac]]. The reproductive cycle may be 2–3 years long, with [[ovulation]] occurring in early autumn, [[fertilization]] in the summer (or possibly in the winter if females are capable of storing sperm), and parturition in late winter or early spring. The [[gestation period]] is under one year.<ref name="hickling"/><ref name="coelho and erzini">{{cite journal |author=Coelho, R. and Erzini, K. |title=Life history of a wide-ranging deepwater lantern shark in the north-east Atlantic, ''Etmopterus spinax'' (Chondrichthyes: Etmopteridae), with implications for conservation |journal=Journal of Fish Biology |volume=73 |issue=6 |pages=1419–1443 |date=2008 |doi=10.1111/j.1095-8649.2008.02021.x}}</ref> The litter size is 6–20, with the number of young increasing with female size. At birth the young measure {{convert|12|-|14|cm|in|abbr=on}} long.<ref name="compagno"/><ref name="coelho and erzini"/> The shark's bioluminescence develops before birth; the yolk sac is [[fluorescence|fluorescent]] before any photophores have formed, suggesting that the mother transfers luminescent materials to her offspring. The first luminous tissue appears when the embryo is {{convert|55|mm|in|abbr=on}} long, and the complete pattern is laid down by the time it is {{convert|95|mm|in|abbr=on}} long. At birth, the young shark is already capable of counter-illumination with 80% of its ventral surface luminescent.<ref name="claes and mallefet"/> |
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The growth rate of the velvet belly is slow, though faster than some other deep-sea sharks such as the [[leafscale gulper shark]] (''Centrophorus squamosus'') or the [[shortspine spurdog]] (''Squalus mitsukurii''). Males [[sexual maturation|mature sexually]] at {{convert|28|-|33|cm|in|abbr=on}} long and females at {{convert|34|-|36|cm|in|abbr=on}} long.<ref name="martin"/><ref name="gennari and scacco"/> The average age at maturity is 4.0 years for males and 4.7 years for females, though 4-year-old mature individuals of both sexes have been caught in the wild, along with immature females over 8 years old.<ref name="coelho and erzini"/> Males and females 8 and 11 years old respectively have been caught in the wild; the potential lifespan of this species has been estimated at 18 years for males and 22 years for females.<ref name="gennari and scacco"/><ref name="coelho and erzini"/> |
The growth rate of the velvet belly is slow, though faster than some other deep-sea sharks such as the [[leafscale gulper shark]] (''Centrophorus squamosus'') or the [[shortspine spurdog]] (''Squalus mitsukurii''). Males [[sexual maturation|mature sexually]] at {{convert|28|-|33|cm|in|abbr=on}} long and females at {{convert|34|-|36|cm|in|abbr=on}} long.<ref name="martin"/><ref name="gennari and scacco"/> The average age at maturity is 4.0 years for males and 4.7 years for females, though 4-year-old mature individuals of both sexes have been caught in the wild, along with immature females over 8 years old.<ref name="coelho and erzini"/> Males and females 8 and 11 years old respectively have been caught in the wild; the potential lifespan of this species has been estimated at 18 years for males and 22 years for females.<ref name="gennari and scacco"/><ref name="coelho and erzini"/> |
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Revision as of 08:15, 25 February 2010
| Velvet belly lantern shark | |
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| Scientific classification | |
| Kingdom: | |
| Phylum: | |
| Class: | |
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| Family: | |
| Genus: | |
| Species: | E. spinax
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| Binomial name | |
| Etmopterus spinax (Linnaeus, 1758)
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| Range of the velvet belly lantern shark | |
| Synonyms | |
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Etmopterus aculeatus Rafinesque, 1810 * ambiguous synonym | |
Template:FixHTML Template:Sharksportal Template:FixHTML The velvet belly lantern shark (or simply velvet belly), Etmopterus spinax, is a species of dogfish shark in the family Etmopteridae. One of the most common deepwater sharks in the northeastern Atlantic Ocean, the velvet belly is found from Iceland and Norway to Gabon and South Africa at a depth of 70–2,490 m (230–8,170 ft). A small shark generally no more than 45 cm (18 in) long, the velvet belly is so named because its black underside is abruptly distinct from the brown coloration on the rest of its body. The body of this species is fairly stout, with a moderately long snout and tail, and very small gill slits. Like other lantern sharks, the velvet belly is bioluminescent, with light-emitting photophores forming a species-specific pattern over its flanks and abdomen. These photophores are thought to function in counter-illumination, which camouflages the shark against predators. They may also play a role in social interactions.
Young velvet bellies feed mainly on krill and small bony fish, transitioning to squid and shrimp as they grow larger. There is evidence that individuals also move into deeper water as they age. This species exhibits a number of adaptations to living in the deep sea, such as specialized T-cells and liver proteins for dealing with the higher concentrations of heavy metals found there. Velvet bellies often carry a heavy parasite load. It is ovoviviparous, giving birth to litters of 6–20 young every 2–3 years. This species has virtually no commercial value but large numbers are caught as bycatch in deepwater commercial fisheries. Although its population status has not been assessed, the heavy fishing pressure throughout its range and its slow reproductive rate are raising conservation concerns.
Taxonomy
The velvet belly was originally described as Squalus spinax by Swedish natural historian Karl Linnaeus, known as the "father of taxonomy", in the 1758 tenth edition of Systema Naturae. He did not designate a type specimen; the specific epithet spinax is in reference to the spiny dorsal fins. This species was later moved to the genus Etmopterus via the synonymy of Constantine Samuel Rafinesque's Etmopterus aculeatus with Squalus spinax.[1] The velvet belly is grouped with the Caribbean lanternshark (E. hillianus), fringefin lanternshark (E. schultzi), brown lanternshark (E. unicolor), broadbanded lanternshark (E. gracilispinis), combtooth lanternshark (E. decacuspidatus), and dwarf lanternshark (E. perryi) in having irregularly arranged, needle-shaped dermal denticles.[2] Its common name comes from this shark's black ventral surface, which is sharply delineated from the rest of its body like a patch of velvet.[3]
Distribution and habitat
The range of the velvet belly is in the eastern Atlantic, extending from Iceland and Norway to Gabon, including the Mediterranean Sea, the Azores, the Canary Islands, and Cape Verde. It has also been reported off Cape Province, South Africa. This shark mainly inhabits the outer continental and insular shelves and upper slopes over mud or clay, from close to the bottom to the middle of the water column.[1][4] It is most common at a depth of 200–500 m (660–1,640 ft), though in the Rockall Trough it is only found at a depth of 500–750 m (1,640–2,460 ft).[5][6] This species has been reported from as shallow as 70 m (230 ft), and as deep as 2,490 m (8,170 ft).[7]
Description
The velvet belly is a robustly built shark with a moderately long, broad, flattened snout. The mouth has thin, smooth lips. The upper teeth are small, with a narrow central cusp and usually fewer than three pairs of lateral cusplets. The lower teeth are much larger, with a strongly slanted blade-like cusp at the top and interlocking bases. The five pairs of gill slits are tiny, comparable in size to the spiracles. Both dorsal fins bear stout, grooved spines at the front, with the second much longer than the first and curved. The first dorsal fin originates behind the short and rounded pectoral fins; the second dorsal fin is twice the size of the first and originates behind the pelvic fins. The anal fin is absent. The tail is slender, leading to a long caudal fin with a small lower lobe and a low upper lobe with a prominent ventral notch near the tip.[1]
The dermal denticles are thin with hooked tips, arranged without a regular pattern well-separated from one another. The coloration is brown above, abruptly transitioning to black below. There are thin black marks above and behind the pelvic fins, and along the caudal fin.[1] The velvet belly possesses numerous photophores that emit a blue-green light visible from 3–4 m (9.8–13.1 ft) away.[3] Varying densities of photophores are arranged in nine patches on the shark's sides and belly, creating a pattern unique to this species: photophores are present along the lateral line, scattered beneath the head but excluding the mouth, evenly on the belly, and concentrated around the pectoral fins and beneath the caudal peduncle.[5][8] The maximum reported length is 60 cm (24 in), although few are longer than 45 cm (18 in).[5] Females are larger than males.[9]
Biology and ecology
Along with the blackmouth catshark (Galeus melastomus) and the Portuguese dogfish (Centroscymnus coelolepis), the velvet belly is one of the most abundant deep-sea sharks in the northeastern Atlantic.[10] It is found individually or in small shoals.[11] Samplings in the Mediterranean have found females outnumbering males across all ages; this imbalance increases in the older age classes.[12] In the Rockall Trough and the Catalan Sea, large adults are found in deeper waters than juveniles, which may serve to reduce competition between the two groups.[10] However, this pattern has not been observed at other sites in the eastern Mediterranean.[7]
The velvet belly's liver accounts for 17% of its body mass, three-quarters of which is oil, making it nearly neutrally buoyant.[13] To deal with the higher concentrations of heavy metals in the deep sea, the velvet belly has T-cells in its bloodstream that can identify and mark toxic compounds for elimination. These T-cells are produced by a lymphomyeloid gland in its esophagus called a "Leydig's organ", which is also found in some other sharks and rays. In its liver, there are also specialized proteins capable of detoxifying cadmium, copper, mercury, zinc, and other toxic contaminants.[5] The velvet belly's bioluminescence is thought to function in counter-illumination, which eliminates the shark's silhouette and camouflages it from upward-looking predators.[8] Its bioluminescence may also serve a social function, such as finding mates or coordinating groups, as the pattern is species-specific. The velvet belly is an important food of larger fishes such as other sharks; a major predator of this species is the longnosed skate (Dipturus oxyrinchus).[5][11]
Numerous parasites are known for this species, and both juveniles and adults often carry heavy parasite loads. Known internal parasites include the monogenean Squalonchocotyle spinacis, the tapeworms Aporhynchus norvegicus, Lacistorhynchus tenuis, and Phyllobothrium squali, and the nematodes Anisakis simplex and Hysterothylacium aduncum. Some of these parasites utilize the velvet belly's prey as intermediate hosts and are acquired via ingestion, while others use the shark itself as an intermediate host.[11] The barnacle Anelasma squalicola is an external parasite that attaches to the shark's spine socket and penetrates deeply into the muscle, in the process often providing an attachment site for a second (and rarely a third) barnacle. Infestation by this barnacle reduces the host's fecundity by impairing the development of the reproductive organs.[9]
Feeding
A generalist predator, velvet bellies feed on crustaceans (e.g. pasiphaeid shrimp and krill), cephalopods (e.g. ommastrephid squid and sepiolids), and bony fishes (e.g. shads, barracudinas, lanternfishes, and pouts).[5] Sharks off Italy also eat small amounts of nematodes, polychaete worms, and other cartilaginous fishes.[14] Studies of velvet bellies off Norway and Portugal, and in the Rockall Trough, have found that small sharks under 27 cm (11 in) long feed mainly on the krill Meganyctiphanes norvegica and the small fish Maurolicus muelleri. As the sharks grow larger, their diet becomes more varied, consisting mainly of squid and the shrimp Pasiphaea tarda, as well fishes other than M. muelleri.[11][6][15] It has been speculated that smaller velvet bellies may be too slow to catch fast-moving cephalopods.[11] The cephalopod diet of adults overlaps with that of the Portuguese dogfish; the latter species may avoid competition with the velvet belly by living in deeper water.[10] The bite force exerted by the velvet belly is only around 1 N.[16]
Life history
The velvet belly is ovoviviparous, with the embryos hatching inside the uterus and being sustained by a yolk sac. The reproductive cycle may be 2–3 years long, with ovulation occurring in early autumn, fertilization in the summer (or possibly in the winter if females are capable of storing sperm), and parturition in late winter or early spring. The gestation period is under one year.[9][17] The litter size is 6–20, with the number of young increasing with female size. At birth the young measure 12–14 cm (4.7–5.5 in) long.[1][17] The shark's bioluminescence develops before birth; the yolk sac is fluorescent before any photophores have formed, suggesting that the mother transfers luminescent materials to her offspring. The first luminous tissue appears when the embryo is 55 mm (2.2 in) long, and the complete pattern is laid down by the time it is 95 mm (3.7 in) long. At birth, the young shark is already capable of counter-illumination with 80% of its ventral surface luminescent.[8]
The growth rate of the velvet belly is slow, though faster than some other deep-sea sharks such as the leafscale gulper shark (Centrophorus squamosus) or the shortspine spurdog (Squalus mitsukurii). Males mature sexually at 28–33 cm (11–13 in) long and females at 34–36 cm (13–14 in) long.[5][12] The average age at maturity is 4.0 years for males and 4.7 years for females, though 4-year-old mature individuals of both sexes have been caught in the wild, along with immature females over 8 years old.[17] Males and females 8 and 11 years old respectively have been caught in the wild; the potential lifespan of this species has been estimated at 18 years for males and 22 years for females.[12][17]
Human interactions
Throughout its range, large numbers of velvet bellies are caught as bycatch in bottom trawls meant for shrimp and lobsters, and deepwater longlines meant for other fish. Lacking commercial value, these sharks are almost always discarded with extremely high mortality, though occasionally they are dried and salted or made into fishmeal. The impact of these fisheries on velvet belly numbers has not been measured but is likely to be heavy. The slow reproductive rate of this species gives cause for concern, as it cannot easily recover from population depletion.[1][12][17]
References
- ^ a b c d e f Compagno, L.J.V. (1984). Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. Rome: Food and Agricultural Organization. p. 85. ISBN 9251013845..
{{cite book}}: Check|isbn=value: invalid character (help) - ^ Springer, S. and G.H. Burgess (August 5, 1985). "Two New Dwarf Dogsharks (Etmopterus, Squalidae), Found off the Caribbean Coast of Colombia". Copeia. 1985 (3): 584–591. doi:10.2307/1444748.
- ^ a b Ellis, R. (1996). Deep Atlantic: Life, Death, and Exploration in the Abyss. The Lyons Press. pp. 195–196. ISBN 1558216634.
- ^ Sion, L., Bozzano, A., D’Onghia, G., Capezzuto, F. and Panza, M. (2004). "Chondrichthyes species in deep waters of the Mediterranean Sea". Scientia Marina. 68 (S3): 153–162.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c d e f g Martin, R.A. Deep Sea: Velvetbelly Lanternshark. ReefQuest Centre for Shark Research. Retrieved on June 24, 2009.
- ^ a b Mauchline, J. and Gordon, J.D.M. (1983). "Diets of the sharks and chimaeroids of the Rockall Trough, northeastern Atlantic Ocean". Marine Biology. 75: 269–278. doi:10.1007/BF00406012.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b Jones, E.G., Tselepides, E., Bagley, P.M., Collins, M.A. and Priede, I.G. (2003). "Bathymetric distribution of some benthic and benthopelagic species attracted to baited cameras and traps in the deep eastern Mediterranean". Marine Ecology Progress Series. 251: 75–86. doi:10.3354/meps251075.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c Claes, J.M. and Mallefet, J. (2008). "Early development of bioluminescence suggests camouflage by counter-illumination in the velvet belly lantern shark Etmopterus spinax (Squaloidea: Etmopteridae)". Journal of Fish Biology. 73 (6): 1337–1350. doi:10.1111/j.1095-8649.2008.02006.x.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c Hickling, C.F. (1963). "On the small deep-sea shark Etmopterus spinax L., and its cirripede parasite Anelasma squalicola (Lovén)". Journal of the Linnean Society of London, Zoology. 45 (303): 17–24. doi:10.1111/j.1096-3642.1963.tb00484.x.
- ^ a b c Carrassón, M., Stefanescu, C. and Cartes, J.E. (1992). "Diets and bathymetric distributions of two bathyal sharks of the Catalan deep sea (western Mediterranean)". Marine Ecology Progress Series. 82 (1): 21–30. doi:10.3354/meps082021.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c d e Klimpel, S., Palm, H.W. and Seehagen, A. (2003). "Metazoan parasites and food composition of juvenile Etmopterus spinax (L., 1758) (Dalatiidae, Squaliformes) from the Norwegian Deep". Parasitology Research. 89 (4): 245–251. doi:10.1007/s00436-002-0741-1. PMID 12632160.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c d Gennari, E. and Scacco, U. (2007). "First age and growth estimates in the deep water shark, Etmopterus Spinax (Linnaeus, 1758), by deep coned vertebral analysis". Marine Biology. 152 (5): 1207–1214. doi:10.1007/s00227-007-0769-y.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Schmidt-Nielsen, S., Flood, A. and Stene, J. (1934). "On the size of the liver of some gristly fishes, their content of fat and vitamin A". Kongeleige Norske Videnskabers Selskab Forhandlinger. 7: 47–50.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Serena, F., Cecchi, E., Mancusi, C. and Pajetta, R. (2006). "Contribution to the knowledge of the biology of Etmopterus spinax (Linnaeus 1758) (Chondrichthyes, Etmopteridae)". In FAO (ed.). Deep Sea 2003: Conference on the Governance and Management of Deep-sea Fisheries. Food and Agricultural Organization. pp. 388–394. ISBN 9251054576.
{{cite book}}: CS1 maint: multiple names: authors list (link) - ^ Neiva, J., Coelho, R. and Erzini, K. (2006). "Feeding habits of the velvet belly lanternshark Etmopterus spinax (Chondrichthyes: Etmopteridae) off the Algarve, southern Portugal". Journal of the Marine Biological Association of the United Kingdom. 86 (4): 835–841. doi:10.1017/S0025315406013762.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Huber, D.R., Claes, J.M., Mallefet, J. and Herrel, A. (2009). "Is Extreme Bite Performance Associated with Extreme Morphologies in Sharks?". Physiological and Biochemical Zoology. 82 (1): 20–28. doi:10.1086/588177. PMID 19006469.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c d e Coelho, R. and Erzini, K. (2008). "Life history of a wide-ranging deepwater lantern shark in the north-east Atlantic, Etmopterus spinax (Chondrichthyes: Etmopteridae), with implications for conservation". Journal of Fish Biology. 73 (6): 1419–1443. doi:10.1111/j.1095-8649.2008.02021.x.
{{cite journal}}: CS1 maint: multiple names: authors list (link)
External links
