View source for Skate (fish)

{{Short description|Family of fishes}}
{{More citations needed|date=October 2024}}
{{Automatic taxobox
| name = Skates
| image = Amblyraja hyperborea1.jpg
| image_caption = [[Arctic skate]], ''Amblyraja hyperborea''
| fossil_range = {{Fossil range|Late Cretaceous | Recent}}<ref name=FB>{{FishBase_family|family=Rajidae|year=2009|month=January}}</ref>
| taxon = Rajidae
| authority = [[Charles Lucien Bonaparte|Bonaparte]], 1831
}}

'''Skates''' are [[Chondrichthyes|cartilaginous fish]] belonging to the [[family (biology)|family]] '''Rajidae''' in the superorder [[Batoidea]] of [[Ray (fish)|rays]]. More than 150 species have been described, in 17 genera.<ref name=Last2016>{{cite book |first1=P.R. |last1=Last |first2=B. |last2=Séret |first3=M.F.W. |last3=Stehmann |first4=S. |last4=Weigmann |chapter=Skates: Family Rajidae |pages=204–363 |editor1-last=Last |editor1-first=Peter |editor2-last=Naylor |editor2-first=Gavin |editor3-last=Séret |editor3-first=Bernard |editor4-last=White |editor4-first=William |editor5-last=de Carvalho |editor5-first=Marcelo |editor6-last=Stehmann |editor6-first=Matthias |title=Rays of the World |date=2016 |publisher=Csiro Publishing |isbn=978-0-643-10914-8 }}</ref> [[Arhynchobatidae|Softnose skates]] and [[Gurgesiellidae|pygmy skates]] were previously treated as subfamilies of Rajidae (Arhynchobatinae and Gurgesiellinae), but are now considered as distinct families.<ref name=Last2016/> Alternatively, the name "skate" is used to refer to the entire order of [[Rajiformes]] (families [[Anacanthobatidae]], [[Arhynchobatidae]], [[Gurgesiellidae]] and [[Rajidae]]).<ref name=Last2016/>

Members of Rajidae are distinguished by a stiff snout and a [[rostrum (anatomy)|rostrum]] that is not reduced.<ref>EBERT, D.A. (2003) The Sharks, Rays and Chimaeras of California. University California Press, Berkeley, CA. 284 pp.{{pn|date=June 2024}}</ref><ref>{{cite book |last1=Compagno |first1=Leonard J |chapter=Systematics and Body Form |pages=1–42 |chapter-url=https://books.google.com/books?id=ON-sPP5rowwC&pg=PA1 |editor1-last=Hamlett |editor1-first=William C. |title=Sharks, Skates, and Rays: The Biology of Elasmobranch Fishes |date=1999 |publisher=JHU Press |isbn=978-0-8018-6048-5 }}</ref>

== Taxonomy and systematics==
===Evolution===
Skates belong to the ancient lineage of cartilaginous fishes. Fossil [[Dermal denticle|denticle]]s (tooth-like scales in the skin) resembling those of today's [[chondrichthyan]]s 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 [[shark]]s, [[sawfish]], [[ray (fish)|ray]]s and skates.<ref>[http://www.ucmp.berkeley.edu/vertebrates/basalfish/chondrofr.html UCMP Berkeley "Chondrichthyes: Fossil Record"]</ref>

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.<ref name='nature2023'>{{cite journal |last1=Marlétaz |first1=Ferdinand |last2=de la Calle-Mustienes |first2=Elisa |last3=Acemel |first3=Rafael D. |last4=Paliou |first4=Christina |last5=Naranjo |first5=Silvia |last6=Martínez-García |first6=Pedro Manuel |last7=Cases |first7=Ildefonso |last8=Sleight |first8=Victoria A. |last9=Hirschberger |first9=Christine |last10=Marcet-Houben |first10=Marina |last11=Navon |first11=Dina |last12=Andrescavage |first12=Ali |last13=Skvortsova |first13=Ksenia |last14=Duckett |first14=Paul Edward |last15=González-Rajal |first15=Álvaro |last16=Bogdanovic |first16=Ozren |last17=Gibcus |first17=Johan H. |last18=Yang |first18=Liyan |last19=Gallardo-Fuentes |first19=Lourdes |last20=Sospedra |first20=Ismael |last21=Lopez-Rios |first21=Javier |last22=Darbellay |first22=Fabrice |last23=Visel |first23=Axel |last24=Dekker |first24=Job |last25=Shubin |first25=Neil |last26=Gabaldón |first26=Toni |last27=Nakamura |first27=Tetsuya |last28=Tena |first28=Juan J. |last29=Lupiáñez |first29=Darío G. |last30=Rokhsar |first30=Daniel S. |last31=Gómez-Skarmeta |first31=José Luis |title=The little skate genome and the evolutionary emergence of wing-like fins |journal=Nature |date=20 April 2023 |volume=616 |issue=7957 |pages=495–503 |doi=10.1038/s41586-023-05868-1 |pmid=37046085 |pmc=10115646 |bibcode=2023Natur.616..495M }}</ref>

===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.<ref name="Ebert Davis 2007"/> The following genera recognized in the family Rajidae:<ref name = CofF>{{Cof family|family=Rajidae|access-date=31 October 2024}}</ref>

{{Linked genus list
| Amblyraja | [[August Wilhelm Malm|Malm]], 1877
| Beringraja | [[Hajimi Ishihara|Ishihara]], [[Michelle Treloar|Treloar]], [[Peter H. F. Bor|Bor]], [[Hiroshi Senou|Senou]] & [[Choon H. Jeong|Jeong]], 2012
| Breviraja | [[Henry Bryant Bigelow|Bigelow]] & [[William Charles Schroeder|Schroeder]], 1948
| Caliraja  | [[David A. Ebert|Ebert]], 2022
| Dactylobatus | [[Barton Appler Bean|Bean]] & [[Alfred Cleveland Weed|Weed]], 1909
| Dentiraja | [[Gilbert Percy Whitley|Whitley]], 1940
| Dipturus | [[Constantine Samuel Rafinesque|Rafinesque]], 1810
| ''[[Korean skate|Hongeo]]'' | Jeong & [[Tetsuji Nakabo|Nakabo]], 2009
| Leucoraja | Malm, 1877
| Malacoraja | [[Matthias Stehmann|Stehmann]], 1970
| Neoraja | [[John D. McEachran|McEachran]] & [[Leonard Compagno|Compagno]], 1982
| Okamejei | [[Reizo Ishiyama|Ishiyama]], 1958
| Orbiraja | [[Peter R. Last|Last]], [[Simon Weigmann|Weigmann]] & [[Don Dumale|Dumale]], 2016
| ''[[Raja (fish)|Raja]]'' | [[Carl Linnaeus|Linnaeus]], 1758
| Rajella | Stehmann, 1970
| Rostroraja | [[P. Alexander Hulley|Hulley]], 1972
| Spiniraja | Whitley, 1939
| Zearaja | Whitley 1939
}}

Skates have more valid species than any other group of cartilaginous fishes.{{cn|date=October 2024}} Since 1950, 126 new species of skates have been discovered. Five scientists take credit for the rapid increase of findings.{{cn|date=October 2024}} The [[Rajiformes|Rajidae]] are considered monophyletic because of their similarity in appearance. There are 18 genera<ref name = CofF/> and about 250 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.<ref name=":1" />

== 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.<ref name=":02">{{cite web|url=https://www.floridamuseum.ufl.edu/fish/discover/rays-skates/anatomy/|title=Ray & Skate Anatomy :: Florida Museum of Natural History|website=www.floridamuseum.ufl.edu|date=5 May 2017|language=en|access-date=2018-03-10}}</ref> This structure creates power for forward propulsion, providing the emergence swimming capabilities that enabled skates to colonize the sea floor.<ref name='nature2023'/>

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.<ref name=":8">{{Cite book|last1=Berkovitz|first1=Barry|last2=Shellis|first2=Peter|pages=5–27|doi=10.1016/b978-0-12-802850-6.00002-3|title = The Teeth of Non-Mammalian Vertebrates|year = 2017|isbn = 9780128028506}}</ref> Skate's [[gill slit]]s are located ventrally as well, but dorsal [[Spiracle (vertebrates)|spiracles]] allow the skate to be partially buried in floor sediment and still complete respiratory exchange.<ref name=":5">{{cite journal |author1=Carlos Bustamente |author2=Julio Lamilla |author3=Francisco Concha |author4=David A. Ebert |author5=Michael B. Bennett |author-link2=Julio Lamilla |title=Morphological Characters of the Thickbody Skate Amblyraja frerichsi (Krefft 1968) (Rajiformes: Rajidae), with Notes on Its Biology |journal=[[PLOS One]] |date=29 June 2012 |volume=7 |issue=8 |pages=e39963 |doi=10.1371/journal.pone.0039963 |pmid=22768186 |pmc=3386969 |bibcode=2012PLoSO...739963B |doi-access=free }}</ref> Also located on the dorsal side of the skate are their two eyes which allow for predator awareness.<ref name=":02" /> In addition to their pectoral fins, skates have a first and second [[dorsal fin]], [[Fish fin|caudal fin]] and paired [[pelvic fin]]s. 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 (biology)|electric organ]] and [[mermaid's purse]] create clear distinctions.
[[File:Mermaidpurse.jpg|thumb|Full view of a skate's mermaid's purse. Roughly 125 mm in length.]]

=== Skate specific characteristics ===
==== Mermaid's purse ====
{{Full article|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.<ref name=":03">{{cite journal |last1=Mabragaña |first1=E. |last2=Vazquez |first2=D. M. |last3=Gabbanelli |first3=V. |last4=Sabadin |first4=D. |last5=Barbini |first5=S. A. |last6=Lucifora |first6=L. O. |title=Egg cases of the graytail skate Bathyraja griseocauda and the cuphead skate Bathyraja scaphiops from the south-west Atlantic Ocean |journal=Journal of Fish Biology |date=September 2017 |volume=91 |issue=3 |pages=968–974 |doi=10.1111/jfb.13380 |pmid=28868748 |bibcode=2017JFBio..91..968M |hdl=11336/45691|hdl-access=free}}</ref><ref name="Ebert Davis 2007" /> 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 ====
[[File:Skate EO cross section .png|thumb|The fibers of the skate electric organ are embedded in the muscles located lateral to the notochord in the tail.]]

The [[Electric organ (biology)|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.<ref name=":22">{{cite journal |last1=Macesic |first1=Laura J. |last2=Kajiura |first2=Stephen M. |title=Electric organ morphology and function in the lesser electric ray, Narcine brasiliensis |journal=Zoology |date=November 2009 |volume=112 |issue=6 |pages=442–450 |doi=10.1016/j.zool.2009.02.002 |pmid=19651501 |bibcode=2009Zool..112..442M }}</ref> The impulses put out by the electric organs of the skate are considered to be weak, asynchronous, long-lasting signals.<ref name=":4">{{Cite journal|last=Koester|first=David M.|title=Anatomy and motor pathways of the electric organ of skates|journal=The Anatomical Record|volume=273A|issue=1|pages=648–662|doi=10.1002/ar.a.10076|pmid=12808649|year=2003|doi-access=free}}</ref> 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.<ref name=":4"/> 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.<ref name=":4"/>

== Distribution and habitats ==
Skates are primarily found from the intertidal down to depths greater than {{cvt|3,000.|m}}.<ref name=":1">{{cite journal |last1=Ebert |first1=David A. |last2=Compagno |first2=Leonard J. V. |title=Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei) |journal=Environmental Biology of Fishes |date=October 2007 |volume=80 |issue=2–3 |pages=111–124 |doi=10.1007/s10641-007-9247-0 |bibcode=2007EnvBF..80..111E |s2cid=22875855 }}</ref> They are most commonly found along outer continental shelves and upper slopes.<ref>{{cite journal |last1=Ebert |first1=David A. |last2=Bizzarro |first2=Joseph J. |title=Standardized diet compositions and trophic levels of skates (Chondrichthyes: Rajiformes: Rajoidei) |journal=Environmental Biology of Fishes |date=October 2007 |volume=80 |issue=2–3 |pages=221–237 |doi=10.1007/s10641-007-9227-4 |bibcode=2007EnvBF..80..221E |s2cid=1382600 }}</ref> 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.<ref name="Ebert Davis 2007">{{cite journal |last1=Ebert |first1=David A. |last2=Davis |first2=Chante D. |title=Descriptions of skate egg cases (Chondrichthyes: Rajiformes: Rajoidei) from the eastern North Pacific |journal=Zootaxa |date=18 January 2007 |volume=1393 |issue=1 |doi=10.11646/ZOOTAXA.1393.1.1 }}</ref> 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.<ref>{{Cite web |url=https://www.ct.gov/deep/lib/deep/fishing/fisheries_management/trawl_show.pdf |title=Archived copy |access-date=2019-01-16 |archive-date=2017-01-28 |archive-url=https://web.archive.org/web/20170128113229/http://ct.gov/deep/lib/deep/fishing/fisheries_management/trawl_show.pdf |url-status=dead }}</ref> Some skate fauna have been found inhabiting areas of rock cobble and high rocky relief.

== Behavior and ecology ==
=== Reproduction ===
[[File:Dorron - 16197242004.jpg|thumb|A skate laying an [[Egg case (Chondrichthyes)|egg case (mermaid's purse)]] at the [[San Sebastian Aquarium]]]]
[[File:Big skate 4 embryos.jpg|thumb|Four developing embryos located in an opened [[big skate]] [[Egg case (Chondrichthyes)|egg case (mermaid's purse)]]]]
Skates mate at the same nursery ground each year. In order to fertilize the egg, males use [[clasper]]s, a structure attached to the pelvic fins. The claspers allow them to direct the flow of semen into the female's [[cloaca]]. Skates are [[Oviparity|oviparous]], meaning they lay eggs with very little development in the mother. This is one major difference from rays, which are [[Viviparity|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 (Chondrichthyes)|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 web]]s 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 [[Batoid locomotion#Rajiform swimming|Rajiform locomotion]] to glide along the water floor.<ref>{{cite book |editor1-last=Hall |editor1-first=Brian K. |title=Fins into Limbs: Evolution, Development, and Transformation |date=2008 |publisher=University of Chicago Press |isbn=978-0-226-31340-5 |oclc=308649613 }}{{pn|date=June 2024}}</ref> Current research suggests that some species of skates, in addition to their Rajiform locomotion, use their pelvic fins to perform ambulatory locomotion.<ref>{{cite journal |last1=Lucifora |first1=Luis O. |last2=Vassallo |first2=Aldo I. |title=Walking in skates (Chondrichthyes, Rajidae): anatomy, behaviour and analogies to tetrapod locomotion |journal=Biological Journal of the Linnean Society |date=28 August 2002 |volume=77 |issue=1 |pages=35–41 |doi=10.1046/j.1095-8312.2002.00085.x |doi-access=free |hdl=11336/128800 |hdl-access=free }}</ref> 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.<ref>{{cite journal |last1=Jung |first1=Heekyung |last2=Baek |first2=Myungin |last3=D’Elia |first3=Kristen P. |last4=Boisvert |first4=Catherine |last5=Currie |first5=Peter D. |last6=Tay |first6=Boon-Hui |last7=Venkatesh |first7=Byrappa |last8=Brown |first8=Stuart M. |last9=Heguy |first9=Adriana |last10=Schoppik |first10=David |last11=Dasen |first11=Jeremy S. |title=The Ancient Origins of Neural Substrates for Land Walking |journal=Cell |date=February 2018 |volume=172 |issue=4 |pages=667–682.e15 |doi=10.1016/j.cell.2018.01.013 |pmid=29425489 |pmc=5808577 }}</ref>

==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.<ref name=":0">{{Cite book|title=Marine Biology 10th Edition|last1=Castro|first1=Peter|last2=Huber|first2=Michael|publisher=McGraw-Hill Education|year=2017|isbn=978-1-259-25199-3|location=Singapore|pages=155–158}}</ref> [[File:Rajidae swmimming.jpg|thumb|right|Skates swim with their pectoral fins]]

{{clear left}}

{| class="wikitable collapsible expanded"
|-
! style="width:360px;" colspan="4"| Comparison of skates and [[Myliobatiformes|stingrays]]
|-
| style="width:360px;" colspan="4"| {{center|Superficially, skates and [[Myliobatiformes|stingrays]] appear somewhat similar.}} 
{{multiple image
 | align = center
 | direction = horizontal
 | image1    = Amblyraja hyperborea4.jpg
 | width1    = 158
 | caption1  = {{center|Skate}}
 | image2    = Dasyatis brevicaudata 4x3.jpg
 | width2    = 233
 | caption2  = {{center|Stingray}}
}}
{{Clear}}
{{center|However, there are fundamental differences.}} 
|-
! style="width:110px;"| Characteristic
! Skates
! Rays
! style="width:70px;"| Sources
|-
| Reproduction
| Skates are [[oviparous]], that is they lay eggs. Their fertilized eggs are laid in a protective hard case called a [[mermaid's purse]].
| valign=top | Rays are [[viviparous]], that is, they bear their young inside their bodies and give birth to them alive. 
| align=center | <ref name=fmnh>[http://www.flmnh.ufl.edu/fish/education/questions/raybasics.html#large Ichthyology: Ray and Skate Basics] ''Florida Museum of Natural History''. Retrieved 21 March 2013.</ref>
|-
| [[Dorsal fin]]
| Distinct
| Missing or vestigial
| align=center | <ref name=fmnh />
|-
| [[Pelvic fin]]s
| Fins are divided into two lobes
| Fins have one lobe
| align=center | <ref name=ReefQuest>[http://www.elasmo-research.org/education/shark_profiles/skate_or_ray.htm Skate or Ray?] ''ReefQuest Centre for Shark Research''. Retrieved 212 March 2013.</ref>
|-
| Tail
| Fleshy tails which lack spines
| Whip-like with one or two stinging spines
| align=center | <ref name=fmnh /><ref name=ReefQuest />
|-
| Protection
| Rely 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)
| align=center | <ref name=fmnh />
|-
| Teeth
| Small
| "Plate-like teeth adapted for crushing prey"
| align=center | <ref name=fmnh />
|-
| Size
| Usually smaller than rays
| Usually larger than skates
| align=center | <ref name=fmnh />
|-
| Colour
| Often drab, brown or gray (but not always)
| Often boldly patterned (but not always)
| align=center | <ref name=ReefQuest />
|-
| Habitat
| Often deep water (but not always)
| Often shallow water (but not always)
| align=center | <ref name=ReefQuest />
|}

==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.

== See also ==
* [[Jenny Haniver]], a fake sea monster created from a skate carcass.
* [[Hongeohoe]], a Korean dish made from fermented skate. 
* [[Mokpo]], a South Korean city famous for its skate cuisine.

==References==
{{Reflist}}

==External links==
{{Commons category|Rajidae|<br />Rajidae}}
{{Wikispecies|Rajidae}}
{{Wiktionary|skate}}
{{cookbook}}
* ARKive – [https://web.archive.org/web/20070405020743/http://www.arkive.org/species/GES/fish/Dipturus_batis/ images and movies of the common skate ''(Dipturus batis)'']
* Kliman, Todd. "[http://www.washingtonian.com/articles/bestbites/302.html Skate Goes From Trash Fish to Treasure] {{Webarchive|url=https://web.archive.org/web/20120319234004/http://www.washingtonian.com/articles/bestbites/302.html |date=2012-03-19 }}", ''Washingtonian'', May 1, 2006.

{{Taxonbar|from=Q837104}}
{{Authority control}}

[[Category:Rajidae|.]]
[[Category:Ray families]]
[[Category:Cretaceous cartilaginous fish]]
[[Category:Extant Late Cretaceous first appearances]]
[[Category:Taxa named by Charles Lucien Bonaparte]]