dcsimg

Lifespan, longevity, and ageing

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Maximum longevity: 12 years (wild) Observations: Unverified estimates suggest these animals may live up to 23 years (http://www.fishbase.org/).
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Life Cycle

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Young Raja clavata hatch at 11 to 13 cm in length and 8 to 9 g in weight with a male:female sex ratio of 1:1. Juveniles remain in shallow water. As they increase in size, juveniles grow large button like structures called bucklers. When R. clavata reaches sexual maturity, usually around 8.8 years, it begins seasonally migrating. During summer, adults move in shore during the summer and migrate into deeper waters during the winter.

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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John Berini, Special Projects
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Associations

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Raja clavata eggs are preyed upon by some fish and mollusks, such as necklace shells. Juveniles are commonly preyed upon by larger fish, specifically common skates and adult R. clavata. The coloration of R. clavata likely helps reduce risk of predation.

Known Predators:

  • necklace shells, Polinices
  • common skate, Dipturus batis
  • thornback rays, Raja clavata

Anti-predator Adaptations: cryptic

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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Morphology

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Raja clavata shares its general body shape with all rays in that it is a flattened, disc shape with broad pectoral fins connected to the head and body with triangular pelvic fins at the base of the body. Raja clavata has a long, narrow tail that is roughly equivalent to the length of the body. As the common name denotes (thornback ray), the upper portion of the body and tail are covered with thorn-like projections. When R. clavata reaches sexual maturity the bases of the thorns thicken to resemble small button like projections called buckler. The snout and small portions of the body are covered in thorns in sub adults and the underside may be thorny in large females. Adults typically have between 21 and 25 large thorns running from the nape to its first pectoral fin. It is sexually dimorphic, as females have a maximum length of 118 cm and males have a maximum length of 98 cm. The dorsal surface of R. clavata exhibits a wide range of coloration from light brown to grey and is sprinkled with light and dark spots of varying size. Its underside is solid white except for the snout which is grey. The largest specimen ever recorded weighed 18 kg.

Range mass: 18 (high) kg.

Range length: 98 to 120 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: female larger

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Life Expectancy

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In the wild, Raja clavata lives between 12 and 15 years. There is no information available regarding the average lifespan of captive individuals.

Typical lifespan
Status: wild:
12 to 15 years.

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Habitat

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Raja clavata lives at the bottom of the continental shelf and upper slope of the coastal sea floor. It inhabits a wide range of sea floor habitats including mud, sand, shingle, gravel and rocky areas. Raja clavata has been found in coastal waters as deep as 300 m but is most abundant in waters between 10 and 60 meters. Juveniles are often found further inshore in more shallow waters than their adult counterparts.

Range depth: 10 to 300 m.

Average depth: 60 m.

Habitat Regions: saltwater or marine

Aquatic Biomes: coastal

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Distribution

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Raja clavata is wide spread in coastal waters from Iceland to Norway. Its geographic ranges extends into the North Sea, the Mediterranean Sea, the western Black Sea, Madeira Island, the Atlantic coasts of Africa, as well the waters off of South Africa and the south-western Indian Ocean. Adult R. clavata remains inshore during summer and move off shore into deeper waters during winter.

Biogeographic Regions: palearctic (Native ); ethiopian (Native ); mediterranean sea (Native )

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Trophic Strategy

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Raja clavata is carnivorous. Juveniles feed predominantly on bottom dwelling aquatic crustaceans such as amphipods, mysids and crangonid shrimps. Larger adults feed on larger crustaceans such as swimming crabs and fish, such as sandeels, small gadoids and dragonets.

Animal Foods: fish; aquatic crustaceans

Primary Diet: carnivore (Piscivore , Eats non-insect arthropods)

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Associations

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Raja clavata feeds on a variety of benthic animals. Juveniles feed on small crustaceans such as amphipods, mysids and crangonid shrimps, and adults feed on fish and larger crustaceans such as swimming crabs and fish such as sandeels, small gadoids and dragonets. This species is also known to feed on conspecifics as well. The eggs of R. clavata act as prey for some fish and mollusks such as necklace shells. Juveniles are preyed upon by common skate and adult R. clavata. Due to its high placement in the costal sea floor food web, R. clavata has a significant effect on trophic levels of the benthos. Parasites of this species have not been documented.

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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Benefits

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Raja clavata is more highly valued for its meat than other skates and rays. Thus, R. clavata plays an important economic role in fisheries, particularly those off the coast of Portugal.

Positive Impacts: food

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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Benefits

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There are no known adverse effects of Raja clavata on humans.

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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John Berini, Special Projects
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Conservation Status

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Raja clavata is classified as near threatened on to the ICUN's Red List of Threatened Species. Declines in R. clavata landings are thought to be due to over exploitation and poor species identification. It is caught primarily as by-catch in both otter and beam trawl fisheries, and is targeted with gillnets, long-lines, and recreational anglers. Landings of rays and skates are often lumped together under one generic category (i.e., Raja spp.), but market samplings suggest that R. clavata is the most dominant species landed in the North Sea and Skagerrak. Over exploitation in this species is especially relevant as it has slow growth, long life span, and long period before individuals become reproductively mature make. Research suggests that R. clavata may be extinct from the North Sea due to over fishing. In an effort to help R. clavata recover (as well as other batids), total allowable catch (TAC) for all skates and rays was reduced by 47% from 1999 to 2005. Some fisheries in the United Kingdom have also implemented a minimum landing size. Unfortunately, localized management efforts are not expected to have an impact on regional population conservation.

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

IUCN Red List of Threatened Species: near threatened

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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John Berini, Special Projects
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Behavior

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Like most other elasmobranchs, Raja clavata uses a diverse array of senses that allow it to perceive orientation as well as locate potential predators and prey. In order locate predators and prey, it depends on its acute sense of smell and a lateral line system that allows it to detect changes in pressure in the local environment. The lateral line system is made up of individual mechano-receptors called neuromasts which run the length of the body. These neuromasts are hair filled structures surrounded with a jelly like substance. These are connected to lateral line canals which interact with the environment. Pressure changes in the water displace the hair and fires impulses to the organism’s brain alerting it to the direction and strength of the disturbance. Raja clavata also depends on electrosensory perception organs known as Ampullae of Lorenzini. The ampullae, which consists of a series of gelatin filled pits that create an electrically sensitive network along the ventral surface of the animal, help sense minor electrical pulses created by the muscle contractions of organisms in the local environment. The ampullary organs interact with local electric fields through an opening in the skin on the snout of the animal that leads to a jelly filled canal. The electrical fields produced by animals dissipate rather quickly so R. clavata can only perceive the weak current at short range. Within that short range the thornback ray can accurately determine the location of the prey, even if buried in the sand. Some studies have indicated that R. clavata as well as other sharks and rays can use their Ampullae of Lorenzini to detect the strength and direction of the Earth’s magnetic field as a sort of rudimentary positioning system. Studies have shown that the thornback ray does not rely sight for locating prey.

Raja clavata acquires information from its three otilith organs (the utriculus, the sacculus, and the lagena) to maintain spatial equilibrium in its marine habitat. These organs are located in the inner ear vestibular system where the semicircular canals converge next to the cochlea. The three organs respond to tilts in both the longitudinal and transverse axes. The utriculus and sacculus produce similar responses and thus create mutually reinforcing signals. The macula lagena is made up of gravity receptors which provide a marker for R. clavata upright position. The macula utriculus produces an increase in discharge to signal side-up and nose-up displacements while the macula sacculus signals side-up and nose-down displacements. All three maculae act as an "out-of-position receptor" when the individual is subjected to a constant speed spatial deviation.

Thornback rays have a very acute sense of smell that they use for locating prey. They have two openings called nares that are located ventrally on the organism’s snout. Water enters the nares and passes through a structure known as the olfactory sac. The olfactory sac is lined with a series of folded tissue called the olfactory lamellae which provide increased surface area for molecule-receptor interactions. Molecules dissolved in the water bind to neuroreceptors in the olfactory lamellae to provide chemosensory information to the brain.

Perception Channels: visual ; acoustic ; vibrations ; chemical ; electric ; magnetic

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Jason Dooney, The College of New Jersey
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Matthew Wund, The College of New Jersey
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Reproduction

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Little is known of the mating behavior of many skates, including Raja clavata. Some speculate that this is because the animals mate in the cover of night. Studies of sting rays may provide insight into the mating behaviors of elasmobranchs. During mating, females swim in shore and spend the morning either buried in the sand or lay atop each other in large groups. Those females buried are attempting to avoid males that already mated or are not yet ready for reproduction whilst those in the aggregate groups are prepared for mating. Meanwhile, males swim up and down the beach searching for potential mates. In the afternoon, they switch roles and the males bury themselves in the sand while the females become mobile to search for food in the grassy beds. This cycle can continue for multiple weeks.

Thornback rays are oviparous with females migrating further in shore to lay their eggs. Egg-laying season for Raja clavata occurs between March and September. Mature females deposit one egg at a time in sandy or muddy substrate close to shore. Eggs range in length from 5 to 9 cm long and 3.8 to 6.4 cm wide. Mature females can lay between 140 and 160 eggs in a single year. Eggs are rectangular and oblong in shape and are marked by rigid horns on each corner. During development, embryos feed solely on their yolk. Embryonic development typically takes between 4 and 6 months and is largely dependent on ambient water temperature.

Breeding season: Thornback rays breed from March to September.

Range number of offspring: 140 to 160.

Range gestation period: 4 to 6 months.

Average age at sexual or reproductive maturity (female): 8.8 years.

Average age at sexual or reproductive maturity (male): 7.1 years.

Key Reproductive Features: iteroparous ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous

There is no information available regarding parental care in Raja clavata; however, studies on closely related elasmobranches indicate a high level of parental investment.

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Dooney, J. 2011. "Raja clavata" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Raja_clavata.html
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Biology

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All skates are oviparous (8) (egg-laying), with large female thornbacks laying up to 140 to 160 egg capsules per year (6). The species moves into shallower, inshore waters in spring when ready to spawn, laying eggs between March and September, which take a further four to six months to hatch (6). Young feed on small crustaceans and other bottom-dwelling creatures, predating more on fish and larger crustaceans such as crabs as they develop (5) (6). Maturity is reached at around eight years of age (5), and the maximum lifespan is at least 15 years (6).
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Conservation

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Fishing of the thornback skate is controlled through the authorities limiting the Total Allowable Catch (TAC), although the quota is set for all rajids combined, covering EU waters in the North Sea and Norwegian Sea. This measure helps ensure continuity of economic activity in the fisheries concerned, while giving depleted stocks a reasonable chance of recovery. The TAC for skates and rays has been reduced by approximately 47 % between 1999 and 2005, providing greater protection to the species by reducing the quota allowed to be caught. However, records show that the actual numbers caught each year have in fact been below what is permissible through the TAC (6). Thus, these skate are not currently considered to be particularly over-fished and are not seen as a high conservation priority.
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Description

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The thornback skate is probably one of the commonest skates encountered by divers (3), being among the most abundant rajids in the north-eastern Atlantic and Mediterranean (1). As with all skates, the body is flattened and disc-shaped, with the pectoral fins broadly expanded and joined to the head and body (4). The tail is distinctly demarcated from the disc-like body, relatively narrow, and about as long as body length (3) (4). The upper surface of the disc and tail are covered with numerous thorns, which become thickened with button-like bases (known as bucklers) once the skate is sexually mature, hence the species' common name (3) (5). Only the snout and margins of the disc are prickly in young, and the underside is only prickly in large, mature females, which also possess more developed 'bucklers' on their back and tail (2) (3). The colour varies from light brown to grey on the upper surface, variegated with dark and light spots and blotches that camouflage the skate in the dappled light of the sea bed, while the underside is creamy-white (2) (3).
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Habitat

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A bottom-dwelling species occupying continental shelf and upper slope waters to depths of 300 m, but mainly from 10 to 60 m (6). However, off South Africa this species occurs to 1000 m (4). Usually found on sediment type sea beds such as mud, sand or gravel, occasionally adjacent to rocky reefs (3) (7).
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Range

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Found in the eastern Atlantic from Iceland, Norway, North Sea and the western Baltic southward to Morocco and Namibia, including the Mediterranean and the Black Sea (2). Also reported from South Africa eastward into the south-western Indian Ocean (4).
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Status

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Classified as Lower Risk/near threatened (LR/nt) on the IUCN Red List 2004 (1).
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Threats

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The thornback skate is an important component of mixed trawl fisheries, has been targeted by recreational anglers (1), and is also caught incidentally as bycatch in beam trawl fisheries (9). The thorniness of this species adds to their chance of being entangled in nets (9). There is some evidence of a decline in catch rates in north-west European waters, which could imply a decline in population numbers, but more data is required to confirm this (1). The impact of the fishing industry on rays and skates is the subject of increasing concern due to their slow growth rate, late maturity and low fecundity, making them particularly vulnerable to population collapse due to fishing activity (10).
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Brief Summary

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The thornback ray owes its name to the numerous thorns on its back and tail. Nevertheless, the animal is not dangerous because the thorns are not poisonous. In order to catch its food, it hides under a layer of sand on the sea floor. Together with the spots on its back, it is well camouflaged. Any worms, crustaceans, sea snails or small benthic fish that happen to swim past are caught from its ambush. Thornbacks have rows of strong blunt teeth. When a tooth is worn down, the next one in line appears. Adult thornback rays have 36 to 44 rows of teeth.
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Diagnostic Description

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Sub-rhomboid in shape with pointed wing-tips; disc-width 1,25 to 1,36 times in its length, its length 1,70 to 1,83 times in total length. Short rostrum, rounded at his extremity; pectoral fins with clear angles on lateral side; triangular pelvic fins (Ref. 39246). Dorsally prickly; large females also prickly throughout their ventral surface; juveniles and large males prickly along the margins of their discs and the underside of their snout. 30-50 thorns form a median row from the nape to the first dorsal fin; additional large 'buckler' thorns with swollen bases scattered on upper surface of disc in adults (Ref. 3167). Coloration is highly variable, with the dorsal surface having all shades of brown with dark and light spots and blotches; ventral surface white (Ref. 3167, 58137).
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Recorder
Gert Boden
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Life Cycle

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Oviparous. Distinct pairing with embrace. Polyandrous species (Ref. 89024). Paired eggs are laid and deposited on shallow sand, mud, pebble or gravel bottoms (Ref. 205, 89025). Up to 170 egg cases can be laid by a single female in a year (Ref. 3167), but average fecundity is much lower (around 48-74 eggs) (Ref. 3603, 31302). In northwestern Europe, egg cases are laid during spring (Ref. 3167) and in the Mediterranean during winter and spring (Ref. 3167). Egg cases are oblong capsules with stiff pointed horns at the corners, each containing one embryo (Ref. 205). Capsules are 5.0-9.0 cm long without the horns (Ref. 41250, 88187) and 3.4-6.8 cm wide (Ref. 41250). Egg cases are anchored with an adhesive film (Ref. 82399). Embryos feed solely on yolk (Ref. 50449). Egg cases hatch after about 4-5 months and pups are about 11-13 cm TL (Ref. 88864). Mating season from February to September, peaking in June (Ref. 74501). Adults observed to form same-sex aggregations during the mating season with females moving to shallower inshore waters approximately a month before the males (Ref 3603, 58137, 74501). Mating does not occur in the Baltic Sea (Ref. 82311).
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Cristina V. Garilao
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Morphology

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Dorsal spines (total): 0; Analspines: 0; Analsoft rays: 0
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Gert Boden
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Trophic Strategy

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Nocturnal predator. Feeds on all kinds of bottom animals (Ref. 3167). Polychaetes and crustaceans are important prey items of smaller individuals, bony fishes become more important prey items of larger individuals (Ref. 28070).
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Pascualita Sa-a
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Biology

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Inhabits shelf and upper slope waters. Reported depth ranges varies, from 10-300 m (Ref. 4426, 82319, 88187), and from 300-577 m in the eastern Ionian Sea (Ref. 56504); most common in coastal waters between 10-60 m depth (Ref. 58137). Tolerates low salinities (Ref. 88171). Found on mud, sand and gravel bottoms, rarely on rougher bottoms (Ref. 6808, 58137). Nocturnal species. Feeds on all kinds of bottom animals, preferably crustaceans (Ref. 3167) and fishes (Ref. 114953). Undertakes migrations with mean distances of 54-117 km per month; shows a clear annual migration cycle (Ref. 89017, 89018), moves from deeper offshore waters (10-30 m) in autumn and winter to shallower areas (
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Susan M. Luna
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Importance

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fisheries: commercial; gamefish: yes
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Thornback ray

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The thornback ray (Raja clavata), or thornback skate, is a species of ray fish in the family Rajidae.[2]

Distribution

The Thornback ray is found in the Atlantic coastal waters of Europe and western Africa. It is also present from south Africa to the southwestern Indian Ocean [2] and in the Mediterranean and Black Seas.[3] It is native possibly as far south as Namibia and South Africa.[2]

Its natural habitats are open large seas and shallow seas. It is sometimes seen trapped in large estuarine pools at low tide.[2]

The thornback ray is probably one of the most common rays encountered by divers.

Habitat

The thornback ray is usually found on sedimentary seabeds such as mud, sand or gravel at depths between 10-60m. Juvenile fish feed on small crustaceans, particularly amphipods and bottom-living shrimps; adults feed on crabs, shrimps and small fish.

Description

Like all rays, the thornback ray has a flattened body with broad, wing-like pectoral fins. The body is kite-shaped with a long, thorny tail. The back is covered in numerous thorny spines, as is the underside in older females.[2][4]

Adult fish can grow to 1 metre (3.3 ft) in length, although most are less than 85 centimetres (33 in). This ray can weigh from 4.5 to 8.75 lb (2 to 4 kg).[5]

Their colours vary from light brown to grey with darker blotches and numerous small darker spots and yellow patches. Sometimes the yellow patches are surrounded by small dark spots. The underside is creamy-white with a greyish margin. When threatened they can appear black.[6][7]

In sexually mature fish, some of the spines are thickened with button-like bases (known as bucklers). These are particularly well developed on the tails and backs of sexually mature females.[7]

Personificated face of a Raja clavata .

Taxonomy

Raja clavata, the thornback ray (or thornback skate, roker), was named by Carl Linnaeus in the 10th edition of Systema Naturae in 1758, in the genus Raja of the Order Rajiformes in the Family Rajidae.[2]

It is one of about 13 species of skate (family Rajidae) that are known from the North Sea and adjacent Atlantic waters.

Common names

Raja clavata at the Aquarium de Vannes.

Common names include:[8]

  • Danish — Sømrokke.
  • Dutch — Stekelrog.
  • English — Roker / Thornback ray / Dorn.
  • Estonian — Astelrai.
  • Faeroese — Naglaskøta.
  • Finnish — Okarausku.
  • French — Raie bouclée.
  • German — Nagelrochen.
  • Icelandic — Dröfnuskata.
  • Latvian — Dzelkņraja.
  • Norwegian — Piggskate.
  • Polish — Raja nabijana a. ciernista.
  • Portuguese — Raia lenga / Raia brocheada
  • Russian — Колючий скат.
  • Spanish — Raya de clavos.
  • Swedish — Knaggrocka.

Morphometrics

Dorsal spines : 0; Anal spines: 0; Anal soft rays: 0. ; disc-width 1,25 to 1,36 times in its length, its length 1,70 to 1,83 times in total length; pectoral fins with clear angles on lateral side; triangular pelvic fins . Dorsally prickly; large females also prickly throughout their ventral surface; young and large males prickly along the borders of their discs and the underside of their snout. 30-50 thorns form a median row from the nape to the first dorsal fin; additional large 'buckler' thorns with swollen bases scattered on upper surface of disc in adults . Max length : 105 cm male/unsexed; 139.0 cm (female); common length : 85.0 cm ; max. weight: 18.0 kg.

Top side view of Thornback ray on a white background.

Life cycle

Oviparous. Polyandrous species. Paired eggs are laid and deposited on shallow sand, mud, pebble or gravel bottoms . Up to 170 egg cases can be laid by a single female in a year, average fecundity around 48-74 eggs. In northwestern Europe, egg cases are laid during spring, and in the Mediterranean during winter and spring. Egg cases are oblong capsules with stiff pointed horns at the corners, each containing one embryo. Capsules are 5.0-9.0 cm long without the horns and 3.4-6.8 cm wide. Egg cases are anchored with an adhesive film.

Embryos feed solely on yolk. Egg cases hatch after about 4–5 months and pups are about 11–13 cm.

Mating season is from February to September, peaking in June. Adults observed to form same-sex aggregations during the mating season with females moving to shallower inshore waters approximately a month before the males. Mating does not occur in the Baltic Sea.[2] This is one of the fish used by the marine leech Pontobdella muricata as a host.[9]

British Isles population
Skeletal mount

A search about the growth and maturation of Raja clavata in the Solway Firth (part of the border between Cumbria, England and Dumfries and Galloway, Scotland) shows that the males and females appear to mature at 42 and 45 cm in disc width respectively. The Solway population is heavily exploited by an unrestricted commercial fishery and a considerable proportion (48.6%) of the retained catch is immature. It is suggested that fishing pressure has brought about a reduction in the size at which female fish mature.[10]

See also

References

  1. ^ Ellis, J. (2016). "Raja clavata". IUCN Red List of Threatened Species. 2016: e.T39399A103110667. doi:10.2305/IUCN.UK.2016-3.RLTS.T39399A103110667.en. Retrieved 15 November 2021.
  2. ^ a b c d e f g Fishbase.org: Raja clavata
  3. ^ Guide of Mediterranean Skates and Rays (Raja clavata). Oct. 2022. Mendez L., Bacquet A. and F. Briand. http://www.ciesm.org/Guide/skatesandrays/raja-clavata
  4. ^ Filaman.ifm: Morphology of Raja clavata
  5. ^ Kindersley, Dorling (2005) [2001]. Animal. New York City: DK Publishing. ISBN 0-7894-7764-5.
  6. ^ Shark-references.com: Lists/literature by V.A. Vanov
  7. ^ a b Oceanario.pt: Raja clavata
  8. ^ http://www.ices.dk/explore-us/projects/EU-RFP/EU Repository/ICES ICES Fish Map: species factsheet for Raja clavata
  9. ^ "Pontobdella muricata Linnaeus, 1758". SeaLifeBase. Retrieved 27 December 2019.
  10. ^ Nottage, A. S. (1983). "Growth and maturation of roker Raja clavata L. in the Solway Firth". Journal of Fish Biology. 23: 43–48. doi:10.1111/j.1095-8649.1983.tb02880.x.

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Thornback ray: Brief Summary

provided by wikipedia EN

The thornback ray (Raja clavata), or thornback skate, is a species of ray fish in the family Rajidae.

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