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Goblin shark

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Goblin shark
Temporal range:Miocene–Holocene[1]
Diagram showing a goblin and scuba diver from the side: the shark is not quite twice as long as the human
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Subdivision: Selachimorpha
Order: Lamniformes
Family: Mitsukurinidae
Genus: Mitsukurina
Species:
M. owstoni
Binomial name
Mitsukurina owstoni
Range of the goblin shark[2]
Synonyms
  • Odontaspis nasutusBragança, 1904
  • Scapanorhynchus dofleiniEngelhardt, 1912
  • Scapanorhynchus jordaniHussakof, 1909
  • Scapanorhynchus mitsukuriiWhite,1937

Thegoblin shark(Mitsukurina owstoni) is a rarespeciesof deep-seashark.Sometimes called a "living fossil",it is the onlyextantrepresentative of thefamilyMitsukurinidae,a lineage some 125 million years old. This pink-skinned animal has a distinctive profile with an elongated, flat snout, and highly protrusible jaws containing prominent nail-like teeth. It is usually between 3 and 4 m (10 and 13 ft) long when mature, though it can grow considerably larger such as one captured in 2000 that is thought to have measured 6 m (20 ft).[3]Goblin sharks arebenthopelagiccreatures that inhabit uppercontinental slopes,submarine canyons,andseamountsthroughout the world at depths greater than 100 m (330 ft), with adults found deeper than juveniles. Some researchers believe that these sharks could also dive to depths of up to 1,300 m (4,270 ft), for short periods of time.[3]

Various anatomical features of the goblin shark, such as its flabby body and small fins, suggest that it is sluggish in nature. This species hunts forteleostfishes,cephalopods,andcrustaceansboth near thesea floorand in the middle of thewater column.Its long snout is covered withampullae of Lorenzinithat enable it to sense minuteelectric fieldsproduced by nearby prey, which it can snatch up by rapidly extending its jaws. Small numbers of goblin sharks areunintentionally caughtby deepwaterfisheries.TheInternational Union for Conservation of Nature(IUCN) has assessed it asLeast Concern,despite its rarity, citing its wide distribution and low incidence of capture.

Taxonomy

Differing jaw positions in preserved goblin sharks caused several specimens to be described erroneously as distinct species.

AmericanichthyologistDavid Starr Jordandescribed the goblin shark in an 1898 issue ofProceedings of the California Academy of Sciences,recognizing the peculiar fish not only as a new species, but also a newgenusand family. He based his account on an immature male 107 cm (42 in) caught inSagami BaynearYokohama,Japan. The specimen had been acquired byshipmasterandnaturalistAlan Owston,who had given it to ProfessorKakichi Mitsukuriat theUniversity of Tokyo,who in turn brought it to Jordan. Jordan named the sharkMitsukurina owstoniin honor of these two men.[4]

Thecommon name"goblin shark" is acalqueof its traditional Japanese nametenguzame,atengubeing a Japanese mythical creature often depicted with a long nose and red face.[5]Another name for this species is elfin shark.[2]

Soon after Jordan's description was published, several scientists noted the similarity betweenMitsukurinaand the extinctMesozoicsharkScapanorhynchus.[6]For a time, the prevailing opinion was to treatMitsukurinaas ajunior synonymofScapanorhynchus.Eventually, more completefossilsrevealed many anatomical differences betweenScapanorhynchusandMitsukurina,causing modern authors to again regard them as distinct genera.[7]Several goblin shark specimens were described as separate species from 1904 to 1937, none of which are now considered valid. Thistaxonomicconfusion began because the specimens' jaws were fixed at varying degrees of protrusion during preservation, giving the appearance of proportional differences among the heads.[8]

Phylogeny and evolution

Phylogeneticstudies based onmorphologyhave classified the goblin shark as the mostbasalmember of theorderLamniformes,known as mackerel sharks.[9][10]Studies usinggenetic datahave also confirmed a basal classification for this species.[11][12]The family Mitsukurinidae, represented byMitsukurina,Scapanorhynchus,andAnomotodon,dates back to theAptianage of theCretaceousperiod (c.125–113Ma).Mitsukurinaitself first appears in thefossil recordduring the periodMiddle Eocene(c.49–37 Ma);[13][14]extinct species includeM. lineataandM. maslinensis.[15][16]Striatolamia macrota,which lived in warm shallow waters during thePaleogeneperiod (c.66–23 Ma), may also be aMitsukurinaspecies.[17]As the last member of an ancient lineage, and one that retains several "primitive"traits, the goblin shark has been described as a"living fossil".[18]

Description

The goblin shark's jaws extend dramatically when feeding.

The goblin shark has a distinctively long and flat snout, resembling a blade. The proportional length of the snout decreases with age.[19]The eyes are small and lack protectivenictitating membranes;behind the eyes arespiracles.The large mouth isparabolicin shape. The jaws are very protrusible and can be extended almost to the end of the snout, though normally they are held flush against the underside of the head. It has 35–53 upper and 31–62 lower tooth rows. The teeth in the main part of the jaws are long and narrow, particularly those near the symphysis (jaw midpoint), and are finely grooved lengthwise. The rear teeth near the corners of the jaw are small and have a flattened shape for crushing. Much individual variation of tooth length and width occurs, as for whether the teeth have a smaller cusplet on each side of the maincusp,and regarding the presence of toothless gaps at the symphysis or between the main and rear teeth. The five pairs ofgill slitsare short, with thegill filamentsinside partly exposed; the fifth pair is above the origin of thepectoral fins.[5][7][20]

The body is fairly slender and flabby. The twodorsal finsare similar in size and shape, both being small and rounded. The pectoral fins are also rather small and rounded. Thepelvicandanal finshave long bases and are larger than the dorsal fins. Thecaudal peduncleis flattened from side-to-side and lacks keels or notches. The asymmetriccaudal finhas a long upper lobe with a shallow ventral notch near the tip, and an indistinct lower lobe.[7][20]The soft, semitranslucent skin has a rough texture from a covering ofdermal denticles,each shaped like a short upright spine with lengthwise ridges. Living sharks of this species are pink or tan due to visibleblood vesselsbeneath the skin; the color deepens with age, and young sharks may be almost white. The fins' margins are translucent gray or blue, and the eyes are black with bluish streaks in theirises.After death, the coloration fades quickly to dull gray or brown.[5][8]Adult sharks usually measure between 3 and 4 m (9.8 and 13.1 ft) long.[5]However, the capture of an enormous female estimated at 5.4–6.2 m (18–20 ft) long during 2000 showed this species can grow far larger than suspected previously.[19]A 2019 study suggested that it would have reached 7 m (23 ft) in maximum length.[21]Until 2022, the maximum weight recorded was 210 kg (460 lb) for a shark of 3.8 m (12.5 ft) in length.[7]In 2023, a heavily pregnant, 4.7-metre (15 ft) individual weighing 800 kg (1,800 lb), was landed inTaiwan.The enormous individual sparked criticism of the fishing method ofbottom trawlingwhich was used to catch it.[22]

Distribution and habitat

The goblin shark has been caught in all three major oceans, indicating a wide global distribution. In the Atlantic Ocean, it has been recorded from the northernGulf of Mexico,Suriname, French Guiana, and southern Brazil in the west, and France, Portugal,Madeira,and Senegal in the east.[2][5][23]It has also been collected fromseamountsalong theMid-Atlantic Ridge.[24][25]In theIndo-PacificandOceania,it has been found off South Africa, Mozambique, Japan, Taiwan, Australia and New Zealand. This species has been recorded from offEast CapetoKaikōura Canyonand from theChallenger Plateaunear New Zealand.[26]A single eastern Pacific specimen is known, collected off southernCalifornia.[2][5]This species is most often found over the uppercontinental slopeat depths of 270–960 m (890–3,150 ft).[7]It has been caught as deep as 1,300 m (4,300 ft), and a tooth has been found lodged in an undersea cable at a depth of 1,370 m (4,490 ft).[27]Adults inhabit greater depths than juveniles. Immature goblin sharks frequent thesubmarine canyonsoff southern Japan at depths of 100–350 m (330–1,150 ft), with individuals occasionally wandering intoinshorewaters as shallow as 40 m (130 ft).[2][28]On 19 April 2014, fishermen inKey West,Florida,while fishing in theGulf of Mexico,caught a goblin shark in their fishing net, only the second one ever to be caught in the Gulf.[29]The shark was photographed and released back into the water.[29]The first shark found in the Gulf was caught by commercial fisherman on 25 July 2000 at a depth of approximately 919-1,099 m (3,016–3606 ft) and is thought to have been about 20 ft long.[3]

During July 2014, a goblin shark was found in a fishery net inSri Lanka,near the eastern coast of Sri Lanka. The shark was about 4 ft (1.2 m) long and weighed about 7.5 kg (17 lb). The shark was given to the NARA (National Aquatic Resource Research & Development Agency) for further research.[30]

Biology and ecology

Although observations of living goblin sharks are scant, its anatomy suggests its lifestyle is inactive and sluggish.[31]Its skeleton is reduced and poorlycalcified,the muscle blocks along its sides (myomeres) are weakly developed, and its fins are soft and small. Its long caudal fin, held at a low angle, is also typical of a slow-swimming shark. The long snout appears to have a sensory function, as it bears numerousampullae of Lorenzinithat candetectthe weakelectric fieldsproduced by other animals. Due to the snout's softness, it is unlikely to be used for stirring up prey from the bottom as has been proposed.[8]Vision seems to be less important than other senses, considering the relatively smalloptic tectumin the shark's brain.[5]Yet unlike most deep-sea sharks, it can change the size of itspupils,thus probably does use its sight in some situations.[27]Goblin sharks may be the prey ofblue sharks(Prionace glauca).[28]Parasitesdocumented from this species include thecopepodEchthrogaleus mitsukurinae,[32]and thetapewormsLitobothrium amsichensisandMarsupiobothrium gobelinus.[33]

Feeding

Rattails and other deep-living teleosts are the main food of the goblin shark.

The goblin shark feeds mainly onteleostfishes such asrattailsanddragonfishes.It also consumescephalopodsandcrustaceans,includingdecapodsandisopods.Garbage has been recorded from the stomachs of some specimens.[34]Its known prey includes bottom-dwelling species such as theblackbelly rosefish(Helicolenus dactylopterus), and midwater species such as thesquidTeuthowenia pellucidaand theostracodMacrocypridina castanea rotunda.Thus, the goblin shark appears to forage for food both near the sea floor and far above it.[8][35]

Since it is not a fast swimmer, the goblin shark may be anambush predator.Its low-density flesh and largeoilylivermake itneutrally buoyant,allowing it to drift towards its prey with minimal motions so as to avoid detection.[28]Once prey comes into range, the shark's specialized jaws can snap forward to capture it. The protrusion of the jaw is assisted by two pairs of elasticligamentsassociated with themandibularjoint, which are pulled taut when the jaws are in their normal retracted position; when the shark bites, the ligaments release their tension and essentially "catapult" the jaws forward.[7]At the same time, the well-developed basihyal (analogous to a tongue) on the floor of the mouth drops, expanding the oral cavity and sucking in water and prey.[8]Striking and prey capture events were videotaped and recorded for the first time during 2008 and 2011 and helped to confirm the use and systematics of the protrusible jaws of goblin sharks. The video evidence suggests that while the jaws are definitely unique, goblin sharks use ram feeding, a type of prey capture that is typical of many mackerel sharks.[36]What makes the goblin shark unique is the kinematics of their jaw when feeding. The lower jaw seems to undergo more complex movements and is important in capturing the prey. The measured protrusions of the upper and lower jaw combined put the goblin shark jaws at 2.1–9.5 times more protrusible than other sharks. The lower jaw has a velocity about two times greater than the upper jaw because it not only protrudes forward, but also swings upward to capture the prey, and the maximum velocity of the jaws is 3.14 m/s. The goblin shark has a re-opening and re-closing pattern during the strike, a behavior that has never been seen in other sharks before and could be related to the extent with which the goblin shark protrudes its jaws.[36]

Life history

Little is known about goblin shark reproduction because a pregnant female has yet to be found and studied. It likely shares the reproductive characteristics of other mackerel sharks, which areviviparouswith small litter sizes andembryosthat grow duringgestationby eating undevelopedeggs(oophagy).[2]The birth size is probably close to 82 cm (32 in), the length of the smallest known specimen.[5]Malesmature sexuallyat about 2.6 m (8.5 ft) long, while female maturation size is unknown.[2]No data is available concerning growth and aging.[5]Some researchers have estimated, based on their own research and prior findings, that male goblin sharks mature at approximately 16 years old and can live up to 60 years.[37]

Human interactions

Some of the first known findings pertaining to the goblin shark were published in 1910, and the researcher wrote that, "the new shark is certainly grotesque," and that, "the most remarkable feature is the curiously elongated nose."[38]Given the depths at which it lives, the goblin shark poses little danger to humans. A few specimens have been collected alive and brought topublic aquariums,though they only survived briefly. One was kept atTokai Universityand lived for a week, while another was kept at Tokyo Sea Life Park and lived for two days.[7][39]Its economic significance is minimal; the meat may be dried and salted, while the jaws fetch high prices from collectors.[2][7]At one time, the Japanese also used it for liver oil andfertilizer.[40]This shark is not targeted by anyfisheries,but is occasionally found asbycatchin bottomgillnetsandtrawls,hooked onlonglines,or entangled in fishing gear. Most captures are isolated incidents; one of the few areas where it is caught regularly is off southern Japan, where around 30 individuals (mostly juveniles) are taken each year. Ablack scabbardfish(Aphanopus carbo) fishery offMadeiraalso takes two or three goblin sharks annually. During April 2003, more than a hundred goblin sharks were caught off northwestern Taiwan; the cause of the event was unknown, though observers noted it was preceded by a majorearthquake.The species had never been recorded in the area before, nor has it been found in such numbers since.[2][5]TheInternational Union for Conservation of Nature(IUCN) has categorized the goblin shark asLeast Concern.[2]In addition to its wide range, most of its population is thought to reside in unfished environments because few adults are caught. Therefore, it is not believed to be threatened by human activity.[2]However, during June 2018 the New ZealandDepartment of Conservationclassified the goblin shark as "At Risk – Naturally Uncommon" with the qualifiers "Data Poor" and "Secure Overseas" using theNew Zealand Threat Classification System.[41]

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