Skate (fish)

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Skates
Temporal range: Late Cretaceous–Recent
Amblyraja hyperborea1.jpg
Arctic skate, Amblyraja hyperborea
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Superorder: Batoidea
Order: Rajiformes
Family: Rajidae
Bonaparte, 1831

Skates are cartilaginous fish belonging to the family Rajidae in the superorder Batoidea of rays. More than 150 species have been described, in 17 genera. [2] Softnose skates and pygmy skates were previously treated as subfamilies of Rajidae (Arhynchobatinae and Gurgesiellinae), but are now considered as distinct families. [2] Alternatively, the name "skate" is used to refer to the entire order of Rajiformes (families Anacanthobatidae, Arhynchobatidae, Gurgesiellidae and Rajidae). [2]

Contents

Members of Rajidae are distinguished by a stiff snout and a rostrum that is not reduced. [3] [4]

Taxonomy and systematics

Evolution

Skates belong to the ancient lineage of cartilaginous fishes. Fossil denticles (tooth-like scales in the skin) resembling those of today's chondrichthyans date at least as far back as the Ordovician, with the oldest unambiguous fossils of cartilaginous fish dating from the middle Devonian. A clade within this diverse family, the Neoselachii, emerged by the Triassic, with the best-understood neoselachian fossils dating from the Jurassic. This clade is represented today by sharks, sawfish, rays and skates. [5]

The body plan of skates is caused by skate-specific genomic rearrangements that have altered the three-dimensional regulatory landscape of genes. These changes arose about 286–221  million years ago when skates diverged from sharks. [6]

Classification

The skate belongs to the class Chondrichthyes. This class consists of all the cartilaginous fishes, including sharks and stingrays. Chondrichthyes is divided into two subclasses; of which Elasmobranchii includes skates, rays, and sharks. Skates are the most diverse elasmobranch group, comprising over 20% of the known species. The number of species is likely to increase as taxonomic issues are resolved and new species are identified. [7] There are 17 genera recognized in the family Rajidae: [2]

Skates have more valid species (574) than any other group of cartilaginous fishes. Since 1950, 126 new species of skates have been discovered. Five scientists take credit for the rapid increase of findings. The Rajidae are considered monophyletic because of their similarity in appearance. There are 13 genera and about 245 valid species. However, there is little information about the diets of about 24% of these species. There are at least 45 dubious species of skates worldwide. [8]

Description

General Batoidea characteristics

Skates are cartilaginous fishes like other Chondrichthyes, however, skates, like rays and other Rajiformes, have a flat body shape with flat pectoral fins that extend the length of their body. [9] This structure creates power for forward propulsion, providing the emergence swimming capabilities that enabled skates to colonize the sea floor. [6]

A large portion of the skate's dorsal body is covered by rough skin made of placoid scales. Placoid scales have a pointed tip that is oriented caudally and are homologous to teeth. Their mouths are located on the underside of the body, with a jaw suspension common to Batoids known as euhyostyly. [10] Skate's gill slits are located ventrally as well, but dorsal spiracles allow the skate to be partially buried in floor sediment and still complete respiratory exchange. [11] Also located on the dorsal side of the skate are their two eyes which allow for predator awareness. [9] In addition to their pectoral fins, skates have a first and second dorsal fin, caudal fin and paired pelvic fins. Distinct from their rhomboidal shape is a long fleshy slender tail. While skate anatomy is similar to other Batoidea, features such as their electric organ and mermaid's purse create clear distinctions.

Full view of a skate's mermaid's purse. Roughly 125 mm in length. Mermaidpurse.jpg
Full view of a skate's mermaid's purse. Roughly 125 mm in length.

Skate specific characteristics

Mermaid's purse

Skates produce their young in an egg case called a mermaid's purse. These egg cases have distinct characteristics that are individualized to each species. [12] [13] This makes a great tool for identifying different species of skates. One of these identifiable structures is the keel. The keel is a flexible ridge that runs along the outside of the structure. Another characteristic is the number of embryos in the egg case. Some species contain only one embryo while others can have up to seven. The size of the fibrous shell around the case is another characteristic. Some species have thick layers on the exterior.

Electric organ

The fibers of the skate electric organ are embedded in the muscles located lateral to the notochord in the tail. Skate EO cross section .png
The fibers of the skate electric organ are embedded in the muscles located lateral to the notochord in the tail.

The electric organ is a characteristic exclusive to aquatic species. Among the Chondrichthyes, the only groups to possess electric organs are the electric ray and the skates. Unlike many other electrogenic fishes, skates are unique in having paired electric organs which run longitudinally through the tail in the lateral musculature of the notochord. [14] The impulses put out by the electric organs of the skate are considered to be weak, asynchronous, long-lasting signals. [15] Although the anatomy of the skate's electric organ is well described, its function is poorly understood. Some research suggests the electric impulses are too weak to be a mechanism used for defense or hunting. It is also too irregular to be useful for electrolocation purposes. [15] The most reasonable explanation in the literature suggests that the electric organ discharges may be used as a form of communication used for reproduction purposes. [15]

Distribution and habitats

Skates are primarily found from the intertidal down to depths greater than 3,000 m (9,843 ft). [8] They are most commonly found along outer continental shelves and upper slopes. [16] They are typically more diverse at higher latitudes and in deep-water. In fact, skates are the only cartilaginous fish taxon to exhibit more diversity of species at higher latitudes. A cool, temperate to polar water in the deep sea can be a favorable environment for skates. [7] As the water becomes more shallow and warmer, skates are seen to be replaced by stingrays. Skates are absent from brackish and freshwater environments. However, there is a single estuarine species that has been found in Tasmania, Australia. Also, the Connecticut Department of Environmental Protection has caught and studied skates within the Long Island Sound estuary. [17] Some skate fauna have been found inhabiting areas of rock cobble and high rocky relief.

Behavior and ecology

Reproduction

A skate laying an egg case (mermaid's purse) at the San Sebastian Aquarium Dorron - 16197242004.jpg
A skate laying an egg case (mermaid's purse) at the San Sebastian Aquarium
Four developing embryos located in an opened big skate egg case (mermaid's purse) Big skate 4 embryos.jpg
Four developing embryos located in an opened big skate egg case (mermaid's purse)

Skates mate at the same nursery ground each year. In order to fertilize the egg, males use claspers, a structure attached to the pelvic fins. The claspers allow them to direct the flow of semen into the female's cloaca. Skates are oviparous, meaning they lay eggs with very little development in the mother. This is one major difference from rays, which are viviparous, meaning they give birth to live young. When a female skate is fertilized, a protected case forms around the embryo called an egg case, or more commonly mermaid's purse. This egg case is then deposited out of the mother's body onto the ocean floor where the skates develop for up to 15 months before they enter the external environment.

Diet and feeding

The majority of skates feed on bottom dwelling animals, such as shrimp, crab, oyster, clams, and other invertebrates. To feed on these animals they have grinding plates in their mouths. Skates are an influential part of the food webs of demersal marine communities. They utilize similar resources to those of other upper trophic-level marine predators, such as seabirds, marine mammals, and sharks. The flattened body shape, ventral eyes and well developed spiracles of the skate allows them to live benthically, buried in the sediment or using a longitudinal undulation of the pectoral fins known as Rajiform locomotion to glide along the water floor. [18] Current research suggests that some species of skates, in addition to their Rajiform locomotion, use their pelvic fins to perform ambulatory locomotion. [19] This form of locomotion performed by the skate is being explored as a possible origin for our own development of walking by looking for similar neural pathways used for movement between skates and animals walking on land. [20]

Skates versus stingrays

Skates are like stingrays in that they have five pairs of gill slits that are located ventrally, which means on the underside of their body (unlike sharks that have their gills located on their sides). Skates and rays both have pectoral fins that are flat and expanded, which are typically fused to the head. Both skates and stingrays typically have their eyes on top of their head. Skates also share similar feeding habits with rays.

Skates are different from rays in that they lack a whip-like tail and stinging spines. However, some skates have electric organs located in their tail. The main difference between skates and rays is that skates lay eggs, whereas rays give birth to live young.

Moreover, skates can be more abundant than rays, and are fished for food in some parts of the world. [21]

Skates swim with their pectoral fins Rajidae swmimming.jpg
Skates swim with their pectoral fins
Comparison of skates and stingrays
Superficially, skates and stingrays appear somewhat similar.
Amblyraja hyperborea4.jpg
Skate
Dasyatis brevicaudata 4x3.jpg
Stingray
However, there are fundamental differences.
CharacteristicSkatesRaysSources
ReproductionSkates are oviparous, that is they lay eggs. Their fertilized eggs are laid in a protective hard case called a mermaid's purse.Rays are viviparous, that is, they bear their young inside their bodies and give birth to them alive. [22]
Dorsal fin DistinctMissing or vestigial [22]
Pelvic fins Fins are divided into two lobesFins have one lobe [23]
TailFleshy tails which lack spinesWhip-like with one or two stinging spines [22] [23]
ProtectionRely on "thorny projections on their backs and tails for protection from predators"Rely on stinging spines or barbs for protection (though some, such as manta rays lack these) [22]
TeethSmall"Plate-like teeth adapted for crushing prey" [22]
SizeUsually smaller than raysUsually larger than skates [22]
ColourOften drab, brown or gray (but not always)Often boldly patterned (but not always) [23]
HabitatOften deep water (but not always)Often shallow water (but not always) [23]

Conservation

Skates have slow growth rates and, since they mature late, low reproductive rates. As a result, skates are vulnerable to overfishing and appear to have been overfished and are suffering reduced population levels in many parts of the world.

In 2010, Greenpeace International added the barndoor skate, bottlenose skate, and maltese skate to its seafood red list. "The Greenpeace International seafood red list is a list of fish that are commonly sold in supermarkets around the world, and which have a very high risk of being sourced from unsustainable fisheries." [24]

See also

Related Research Articles

<span class="mw-page-title-main">Chondrichthyes</span> Class of jawed cartilaginous fishes

Chondrichthyes is a class of jawed fish that contains the cartilaginous fish or chondrichthyians, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or bony fish, which have skeletons primarily composed of bone tissue. Chondrichthyes are aquatic vertebrates with paired fins, paired nares, placoid scales, conus arteriosus in the heart, and a lack of opecula and swim bladders. Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.

<span class="mw-page-title-main">Stingray</span> Suborder of fishes

Stingrays are a group of sea rays, which are cartilaginous fish related to sharks. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae, Plesiobatidae, Urolophidae (stingarees), Urotrygonidae, Dasyatidae, Potamotrygonidae, Gymnuridae and Myliobatidae . There are about 220 known stingray species organized into 29 genera.

<span class="mw-page-title-main">Elasmobranchii</span> Subclass of fishes

Elasmobranchii is a subclass of Chondrichthyes or cartilaginous fish, including modern sharks, rays, skates, and sawfish. Members of this subclass are characterised by having five to seven pairs of gill clefts opening individually to the exterior, rigid dorsal fins and small placoid scales on the skin. The teeth are in several series; the upper jaw is not fused to the cranium, and the lower jaw is articulated with the upper. The details of this jaw anatomy vary between species, and help distinguish the different elasmobranch clades. The pelvic fins in males are modified to create claspers for the transfer of sperm. There is no swim bladder; instead, these fish maintain buoyancy with large livers rich in oil.

<span class="mw-page-title-main">Egg case (Chondrichthyes)</span> Natural collagen casing found encompassing some aquatic lifeforms fertilized eggs

An egg case or egg capsule, often colloquially called a mermaid's purse, is the casing that surrounds the eggs of oviparous chondrichthyans. Living chondricthyans that produce egg cases include some sharks, skates and chimaeras. Egg cases typically contain one embryo, except for big skate and mottled skate egg cases, which contain up to 7 embryos. Oviparity is completely absent in the superorder Squalomorphii. Egg cases are also thought to have been produced by some extinct chondricthyan groups, such as hybodonts and xenacanths.

<span class="mw-page-title-main">Rajiformes</span> Order of fishes in the superorder Batoidea

Rajiformes is one of the four orders in the superorder Batoidea, flattened cartilaginous fishes related to sharks. Rajiforms are distinguished by the presence of greatly enlarged pectoral fins, which reach as far forward as the sides of the head, with a generally flattened body. The undulatory pectoral fin motion diagnostic to this taxon is known as rajiform locomotion. The eyes and spiracles are located on the upper surface of the head and the gill slits are on the underside of the body. Most species give birth to live young, although some lay eggs enclosed in a horny capsule.

<span class="mw-page-title-main">Longhead catshark</span> Species of shark

The longhead catshark or smoothbelly catshark is a species of catshark, family Scyliorhinidae, with a patchy distribution in the Indo-Pacific from Mozambique to southern Japan to northern Australia. It is found in water between 500 and 1,140 m deep. This species grows to 59 cm (23 in) long and is characterized by its extremely long and narrow snout, short abdomen, and long anal and caudal fins. In addition, a large area of the anterior ventral portion of its body lacks dermal denticles. The longhead catshark is oviparous and the only known cartilaginous fish that is normally hermaphroditic, with the majority of individuals having both the functional reproductive organs of one sex and the undeveloped reproductive organs of the opposite sex.

<i>Raja</i> (fish) Genus of cartilaginous fishes

Raja, also known as raia, is a genus of skates in the family Rajidae containing 16 species. Formerly a wastebasket genus, many species historically categorized here have been moved to other genera in the family, such as Amblyraja, Beringraja, Dipturus, Leucoraja and Rostroraja. Raja are flat-bodied, cartilaginous fish with a rhombic shapes due to their large pectoral fins extending from or near from the snouts to the bases of their tails. Their sharp snouts are produced by a cranial projection of rostral cartilage. The mouth and gills are located on the underside of the body. They may be either solid-coloured or patterned, and most skates have spiny or thorn-like structures on the upper surface, while some species contain weak electrical organs within their tails. Mating typically occurs in the spring and the female lays numerous eggs per clutch which are encapsulated in leathery cases, commonly known as "mermaid’s purses". Species vary in size, ranging from about 40 to 140 cm (1.3–4.6 ft) in length. These bottom-dwellers are active during both day and night, and typically feed on molluscs, crustaceans and fish. Raja skates are found in the East Atlantic, also in the Mediterranean, and western Indian Ocean, ranging from relatively shallow water to a depth of 800 m (2,600 ft). Skates and related species have fossil records dating from the Upper Cretaceous period, thus this well-adapted species is quite ancient.

<span class="mw-page-title-main">Thornback ray</span> Species of cartilaginous fish

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

<span class="mw-page-title-main">Big skate</span> Species of cartilaginous fish

The big skate is the largest species of skate in the waters off North America. They are found along the Pacific Coast from Alaska to Baja California, typically from the intertidal zone to a depth of 120 m (390 ft), and feed on benthic invertebrates and small fishes. They are unusual among skates in that their egg cases may contain up to seven eggs each. This species is one of the most commercially important skates off California and is sold for food.

<span class="mw-page-title-main">Arhynchobatidae</span> Family of fishes

Arhynchobatidae is a family of skates whose members are commonly known as the softnose skates. It belongs to the order Rajiformes in the superorder Batoidea of rays. At least 104 species have been described, in 13 genera. Softnose skates have at times been placed in the same family as hardnose skates, but most recent authors recognize them as a distinct family. Members of the Arhynchobatidae can be distinguished from hardnose skates in having a soft and flexible snout, as well as a more or less reduced rostrum.

<span class="mw-page-title-main">Batoidea</span> Superorder of cartilaginous fishes

Batoidea is a superorder of cartilaginous fishes, commonly known as rays. They and their close relatives, the sharks, comprise the subclass Elasmobranchii. Rays are the largest group of cartilaginous fishes, with well over 600 species in 26 families. Rays are distinguished by their flattened bodies, enlarged pectoral fins that are fused to the head, and gill slits that are placed on their ventral surfaces.

<span class="mw-page-title-main">Mottled skate</span> Species of cartilaginous fish

The mottled skate is a species of skate in the family Rajidae. An inhabitant of shallow coastal waters, it is found in the northwestern Pacific Ocean off Korea, Japan, and China. This species grows to 1.12 m (3.7 ft) long and has a diamond-shaped pectoral fin disc with a long snout. It is characterized by a covering of prickles above and below its snout, but not elsewhere on its body, and a dark ring in the middle of each "wing".

<span class="mw-page-title-main">Clearnose skate</span> Species of cartilaginous fish

The clearnose skate is a species of cartilaginous fish in the family Rajidae. R. eglanteria is also known by other common names such as the brier skate and summer skate. Clearnose skates are easily identified by the translucent patches on either side of their snouts and their mottled dorsal surface. They are found along the Atlantic and Gulf coasts of the United States in shallow waters of the continental shelf.

<span class="mw-page-title-main">Fish fin</span> Bony skin-covered spines or rays protruding from the body of a fish

Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the spine and are supported only by muscles.

<i>Raja stellulata</i> Species of cartilaginous fish

Raja stellulata, commonly known as the Pacific starry skate, rock skate, prickly skate, or starry skate, is a species of cartilaginous fish in the family Rajidae. It is found on rocky bottoms at 18–982 m depths in the Northeast and Eastern Central Pacific, from Coronado Bank in northern Baja California in Mexico to Barkley Sound in British Columbia, Canada. Females reach a maximum total length of 76.1 cm and a maximum age of 15 years, while males can be up to 71.7 cm long and live up to 14 years; the total length at birth is 15.5–22.5 cm. This skate prefers cold water with a temperature of 4.1–11.6°C.

Batoids are a superorder of cartilaginous fish consisting of skates, rays and other fish all characterized by dorsoventrally flattened bodies and large pectoral fins fused to the head. This distinctive morphology has resulted in several unique forms of locomotion. Most Batoids exhibit median paired fin swimming, utilizing their enlarged pectoral fins. Batoids that exhibit median paired fin swimming fall somewhere along a spectrum of swimming modes from mobuliform to rajiform based on the number of waves present on their fin at once. Of the four orders of Batoidae this holds truest for the Myliobatiformes (rays) and the Rajiformes (skates). The two other orders: Rhinopristiformes and Torpediniformes exhibit a greater degree of body caudal fin swimming.

<i>Raja radula</i> Ray in the genus Raja

The rough ray is a Mediterranean ray species of the Rajidae family described by Delaroche in 1809.

The deepwater ray, also called the deepwater skate or abyssal skate, is a species of skate in the family Rajidae.

<i>Rajella fyllae</i> Species of ray fish

Rajella fyllae is a species of skate in the family Rajidae.

Okamejei panayensis is a demersal fish belonging to the skate family, occurring in nearshore temperate environments and deep-water tropical and boreal regions.

References

  1. Froese, Rainer, and Daniel Pauly, eds. (2009). "Rajidae" in FishBase. January 2009 version.
  2. 1 2 3 4 LAST, P.R. & SÉRET, B. & STEHMANN, M.F.W. & WEIGMANN, S. (2016) Skates, Family Rajidae. In: Last, P.R., White, W.T., Carvalho, M.R. de, Séret, B., Stehmann, M.F.W & Naylor, G.J.P (Eds.) Rays of the World. CSIRO Publishing, Melbourne: 204–363
  3. EBERT, D.A. (2003) The Sharks, Rays and Chimaeras of California. University California Press, Berkeley, CA. 284 pp.
  4. COMPAGNO, L.J.V. (1999) Chapter 1. Systematics and body form. pp. 1–42. In: W.C. Hamlett, ed. Sharks, skates and rays. The biology of elasmobranch fishes. Johns Hopkins Press, Maryland.
  5. UCMP Berkeley "Chondrichthyes: Fossil Record"
  6. 1 2 Marlétaz, F., de la Calle-Mustienes, E., Acemel, R.D. et al. The little skate genome and the evolutionary emergence of wing-like fins. Nature 616, 495–503 (2023). https://doi.org/10.1038/s41586-023-05868-1
  7. 1 2 Ebert, David; Davis, Chante (2003). "Descriptions of skate egg cases (Chondrichthyes: Rajiformes: Rajoidei) from the eastern North Pacific". Zootaxa.
  8. 1 2 Ebert, David A.; Compagno, Leonard J. V. (2007-10-01). "Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei)". Environmental Biology of Fishes. 80 (2–3): 111–124. doi:10.1007/s10641-007-9247-0. ISSN   0378-1909. S2CID   22875855.
  9. 1 2 "Ray & Skate Anatomy :: Florida Museum of Natural History". www.floridamuseum.ufl.edu. 5 May 2017. Retrieved 2018-03-10.
  10. Berkovitz, Barry; Shellis, Peter (2017). The Teeth of Non-Mammalian Vertebrates. pp. 5–27. doi:10.1016/b978-0-12-802850-6.00002-3. ISBN   9780128028506.
  11. Carlos Bustamente; Julio Lamilla; Francisco Concha; David A. Ebert; Michael B. Bennett (29 June 2012). "Morphological Characters of the Thickbody Skate Amblyraja frerichsi (Krefft 1968) (Rajiformes: Rajidae), with Notes on Its Biology". PLOS One . 7 (8): e39963. Bibcode:2012PLoSO...739963B. doi: 10.1371/journal.pone.0039963 . ISSN   1932-6203. PMC   3386969 . PMID   22768186.
  12. Mabragaña, E.; Vazquez, D. M.; Gabbanelli, V.; Sabadin, D.; Barbini, S. A.; Lucifora, L. O. (2017-09-01). "Egg cases of the graytail skate Bathyraja griseocauda and the cuphead skate Bathyraja scaphiops from the south-west Atlantic Ocean". Journal of Fish Biology. 91 (3): 968–974. doi:10.1111/jfb.13380. ISSN   1095-8649. PMID   28868748.
  13. Ebert, David; Davis, Chante (2007-01-18). Descriptions Of Skate Egg Cases (Chondrichthyes: Rajiformes: Rajoidei) From The Eastern North Pacific. Vol. 1393.
  14. Macesic, Laura J.; Kajiura. "Electric organ morphology and function in the lesser electric ray, Narcine brasiliensis" (PDF). Zoology. 112: 442–450 – via ScienceDirect.
  15. 1 2 3 Koester, David M. (2003). "Anatomy and motor pathways of the electric organ of skates". The Anatomical Record. 273A (1): 648–662. doi: 10.1002/ar.a.10076 . PMID   12808649.
  16. Ebert, David A.; Bizzarro, Joseph J. (2007-10-01). "Standardized diet compositions and trophic levels of skates (Chondrichthyes: Rajiformes: Rajoidei)". Environmental Biology of Fishes. 80 (2–3): 221–237. doi:10.1007/s10641-007-9227-4. ISSN   0378-1909. S2CID   1382600.
  17. "Web_Slide_Show04.indd" (PDF). Retrieved 2019-11-29.
  18. Hall, Brian K., ed. (2007). Fins into limbs : evolution, development, and transformation. Chicago: University of Chicago Press. ISBN   9780226313405. OCLC   308649613.
  19. Lucifora, Luis O.; Vassallo, Aldo I. (2002-09-01). "Walking in skates (Chondrichthyes, Rajidae): anatomy, behaviour and analogies to tetrapod locomotion". Biological Journal of the Linnean Society. 77 (1): 35–41. doi: 10.1046/j.1095-8312.2002.00085.x . hdl: 11336/128800 . ISSN   0024-4066.
  20. Jung, Heekyung; Baek, Myungin; D’Elia, Kristen P.; Boisvert, Catherine; Currie, Peter D.; Tay, Boon-Hui; Venkatesh, Byrappa; Brown, Stuart M.; Heguy, Adriana (2018-02-08). "The Ancient Origins of Neural Substrates for Land Walking 3". Cell. 172 (4): 667–682.e15. doi:10.1016/j.cell.2018.01.013. PMC   5808577 . PMID   29425489. (embargo until 2019-02-08, but abstract available)
  21. Castro, Peter; Huber, Michael (2017). Marine Biology 10th Edition. Singapore: McGraw-Hill Education. pp. 155–158. ISBN   978-1-259-25199-3.
  22. 1 2 3 4 5 6 Ichthyology: Ray and Skate Basics Florida Museum of Natural History. Retrieved 21 March 2013.
  23. 1 2 3 4 Skate or Ray? ReefQuest Centre for Shark Research. Retrieved 212 March 2013.
  24. "Greenpeace Seafood Red list". Greenpeace International.