Skip to main content
Log in

Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei)

  • Special Issue-Skates
  • Published:
Environmental Biology of Fishes Aims and scope Submit manuscript

Abstract

Skates (Rajiformes: Rajoidei) are a highly diverse fish group, comprising more valid species than any other group of cartilaginous fishes. The high degree of endemism exhibited by the skates is somewhat enigmatic given their relatively conserved body morphology and apparent restrictive habitat, e.g. soft bottom substrates. Skates are primarily marine benthic dwellers found from the intertidal down to depths in excess of 3,000 m. They are most diverse at higher latitudes and in deepwater, but are replaced in shallower, warm temperate to tropical waters by stingrays (Myliobatodei). The number of valid skate species has increased exponentially, with more species having been described since 1950 (n = 126) than had been described in the previous 200 years (n = 119). Much of the renaissance in skate systematics has largely been through the efforts of a few individuals who through author–coauthor collaboration have accounted for 78 of the 131 species described since 1948 and for nine of 13 genera named since 1950. Furthermore, detailed regional surveys and accounts of skate biodiversity have also contributed to a better understanding of the diversity of the skates. A checklist of the living valid skate species is presented.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5

Similar content being viewed by others

References

  • Bigelow HB, Schroeder WC (1948) New genera and species of batoid fishes. J Mar Res 7:543–566

    Google Scholar 

  • Bigelow HB, Schroeder WC (1950) New and little known cartilaginous fishes from the Atlantic. Bull Mus Comp Zool Harvard 103:385–408

    Google Scholar 

  • Bigelow HB, Schroeder WC (1951a) A new genus and species of anacanthobatid skate from the Gulf of Mexico. J Wash Acad Sci 41:110–113

    Google Scholar 

  • Bigelow HB, Schroeder WC (1951b) Three new skates and a new chimaerid fish from the Gulf of Mexico. J Wash Acad Sci 41:383–392

    Google Scholar 

  • Bigelow HB, Schroeder WC (1953) Fishes of the western North Atlantic. Mem Sears Found Mar Res 1(pt 2):588

    Google Scholar 

  • Bigelow HB, Schroeder WC (1954) A new family, a new genus, and two new species of batoid fishes from the Gulf of Mexico. Breviora, Mus Comp Zool 24:1–16

    Google Scholar 

  • Bigelow HB, Schroeder WC (1958) Four new rajids from the Gulf of Mexico. Bull Mus Comp Zool 119:201–233

    Google Scholar 

  • Bigelow HB, Schroeder WC (1962) New and little known batoid fishes from the western Atlantic. Bull Mus Comp Zool 128:159–244

    Google Scholar 

  • Bigelow HB, Schroeder WC (1964) A new skate Raja cervigoni, from Venezuela and the Guianas. Breviora Mus Comp Zool 209:1–5

    Google Scholar 

  • Bigelow HB, Schroeder WC (1965) A further account of batoid fishes from the western Atlantic. Bull Mus Comp Zool 132:446–475

    Google Scholar 

  • Carvalho MR de (1996) Higher-level elasmobranch phylogeny, basal squaleans, and paraphyly. In: Stiassny MLJ, Parenti LR, Johnson GD (eds) Interrelationships of fishes. Academic Press, San Diego, California, USA, pp 35–62

    Google Scholar 

  • Carvalho MR de, Gomes UL, Gadig OBF (2005) Description of a new species of skate of the genus Malacoraja Stehmann, 1970: the first species from the southwestern Atlantic Ocean with notes on generic monophyly and composition (Chondrichthyes: Rajidae). Neotrop Ichthy 3:239–258

    Google Scholar 

  • Chen CT, Joung SJ (1989) Fishes of the genus Raja (Rajiformes: Rajidae) from Taiwan. J Taiwan Mus 42:1–12

    Google Scholar 

  • Compagno LJV (1999) Checklist of living elasmobranchs. In: Hamlett WC (ed) Sharks, skates, and rays: the biology of elasmobranch fishes. The Johns Hopkins University Press, Baltimore, MD, pp 471–498

    Google Scholar 

  • Compagno LJV (2001) Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Vol. 2. Bullhead, mackerel, and carpet sharks (Heterodontiformes, Lamniformes, and Orectolobiformes). FAO Species Catalogue for Fishery Purposes. No. 1, vol 2. Rome, FAO, 2001, 269 p

  • Compagno LJVC (2005) Checklist of living chondrichthyes. In: Hamlett WC (ed) Reproductive biology and phylogeny of chondrichthyes: sharks, batoids, and chimaeras. Science Publishers, Inc, Enfield, New Hampshire, USA, pp 501–548

    Google Scholar 

  • Compagno LJV, Ebert DA (2007) Southern African skate biodiversity. Environ Biol Fish (this volume)

  • Compagno LJV, Ebert DA, Cowley PD (1991) Distribution of offshore demersal cartilaginous fishes (Class Chondrichthyes) off the west coast of southern Africa, with notes on their systematics. S Afr J Mar Sci 11:43–139

    Google Scholar 

  • Compagno LJV, Ebert DA, Smale MJ (1989) Guide to the sharks and rays of southern Africa. Struik Pub, Cape Town, 160 pp

    Google Scholar 

  • Cousseau MB, Figueroa DE, Diaz de Astarloa JM (2000) Clave de identificacion de las rayas del litoral maritime de Argentina y Uruguay (Chondrichthyes, Familia Rajidae). Publicaciones especiales, INIDEP, Mar del Plata, 35 pp

  • Dolganov VN (1983) Preliminary review of skates Rajidae family (Pacific coast of North America). Izv Tikhookean Nauchno-Issled Inst Rybn Khoz Okeanogr 107:56–72

    Google Scholar 

  • Dolganov VN (1985) New aspects of the ray family Rajidae. Russian version of “New species of skates of the family Rajidae from the northwestern Pacific Ocean”. J Ichthyol 25:121–132

    Google Scholar 

  • Ebert DA (2003) The sharks, rays and chimaeras of California. University California Press, Berkeley, California, 284 pp

  • Eschmeyer WN, Herald ES, Hammond H (1983) A field guide to Pacific coast fishes of North America, vol 28. Houghton Mifflin Co Field Guide, 336 pp

  • Fowler HW (1941) The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostariophysi obtained by United States Bureau of Fisheries Steamer Albatross in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bull US Natn Mus (100) 13:i–x, 1–879

  • Gomes UL (2002) Revisão taxonômica da família Rajidae no Brazil (Chondrichthyes, Elasmobranchii, Rajiformes). Unpublished doctoral thesis, Universidade Federal do Rio de Janeiro, 286 pp

  • Hulley PA (1970) An investigation of the Rajidae of the west and south coasts of southern Africa. Ann S Afr Mus 55:151–220

    Google Scholar 

  • Hulley PA (1972a) The origin, interrelationships and distribution of Southern African Rajidae (Chondrichthyes, Batoidei). Ann S Afr Mus 60:1–103

    Google Scholar 

  • Hulley PA (1972b) A new species of southern African Brevirajid skate (Chondrichthyes, Batoidei, Rajidae). Ann S Afr Mus 60:253–263

    Google Scholar 

  • Ishihara H (1987) Revision of the Western North Pacific species of the genus Raja. Japan J Ichthyol 34:241–285

    Google Scholar 

  • Ishihara H, Ishiyama R (1985) Two new North Pacific skates (Rajidae) and a revised key to Bathyraja in the area. Japan J Ichthyol 32:143–179

    Google Scholar 

  • Ishihara H, Ishiyama R (1986) Systematics and distribution of the skates of the North Pacific (Chondrichthyes, Rajoidei). Indo-Pacific Fish Biology: Proc 2nd Int Conf Indo-Pacific Fishes 269–280

  • Ishiyama R (1952) Studies on the rays and skates belonging to the family Rajidae, found in Japan and adjacent regions. 4. A revision of three genera of Japanese rajids, with descriptions of one new genus and four new species mostly occurred in northern Japan. J Shimonoseki Coll Fish 1:1–34

    Google Scholar 

  • Ishiyama R (1955) Studies on the rays and skates belonging to the family Rajidae, found in Japan and adjacent regions. 6. Raja macrocauda, a new skate. J Shimonoseki Coll Fish 4:43–51

    Google Scholar 

  • Ishiyama R (1958) Studies on the Rajid fishes (Rajidae) found in the waters around Japan. J Shimonoseki Coll Fish 7:193–394

    Google Scholar 

  • Ishiyama R (1967) Fauna Japonica. Rajidae (Pisces). Biogeogr Soc Japan, 84 pp

  • Ishiyama R, Ishihara H (1977) Five new species of skates in the genus Bathyraja from the western North Pacific, with reference to their interspecific relationships. Japan J Ichthy 24:71–90

    Google Scholar 

  • Kuhnz LA, Bizzarro JJ, Chaney L, Ebert DA (2006) In situ video observations of deep-living skates and rays in the central and eastern North Pacific. 22nd Annual Meeting of the American Elasmobranch Society, abstract

  • Lamilla J, Sáez S (2003) Clave taxonomica para el reconocimiento de especies de rayas chilenas (Chondrichthyes, Batoidei). Invest Mar, Valparaiso 31:3–16

    Google Scholar 

  • Last PR, McEachran JD (2006) New softnose skate genus Brochiraja from New Zealand (Rajidae: Arhynchobatinae) with description of four new species. NZ J Mar Freshwater Res 40:65–90

    Article  Google Scholar 

  • Last PR, Stevens JD (1994) Sharks and rays of Australia. CSIRO Division of Fisheries, Melbourne, Australia, 605 pp

  • Last PR, Yearsley GK (2002) Zoogeography and relationships of Australasian skates (Chondrichthyes: Rajidae). J Biogeogr 29:1627–1641

    Article  Google Scholar 

  • Leigh-Sharpe WH (1920) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir I J Morph 34:245–265

    Google Scholar 

  • Leigh-Sharpe WH (1921) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir II J Morph 35:359–380

    Article  Google Scholar 

  • Leigh-Sharpe WH (1922a) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir III J Morph 36:191–198

    Article  Google Scholar 

  • Leigh-Sharpe WH (1922b) The comparative morphology of the secondary sexual characters of Holocephali and elasmobranch fishes. Memoir IV J Morph 36:199–220

    Article  Google Scholar 

  • Leigh-Sharpe WH (1922c) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir V J Morph 39:221–243

    Article  Google Scholar 

  • Leigh-Sharpe WH (1924a) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir VI J Morph 39:553–566

    Article  Google Scholar 

  • Leigh-Sharpe WH (1924b) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir VII J Morph 42:567–577

    Article  Google Scholar 

  • Leigh-Sharpe WH (1926a) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir VIII J Morph 42:307–320

    Google Scholar 

  • Leigh-Sharpe WH (1926b) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir IX J Morph 42:321–334

    Article  Google Scholar 

  • Leigh-Sharpe WH (1926c) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir X J Morph 42:335–348

    Article  Google Scholar 

  • Leigh-Sharpe WH (1926d) The comparative morphology of the secondary sexual characters of elasmobranch fishes. Memoir XI J Morph 42:349–358

    Article  Google Scholar 

  • Malm AW (1877) Goteborgs och Bohusläns fauna, Ryggradjuren. Göteborg. Göteborgs Bohusläns Fauna, 674 pp

  • McEachran JD (1982) Revision of the South American skate genus Sympterygia (Elasmobranchii: Rajiformes). Copeia 4:867–890

    Article  Google Scholar 

  • McEachran JD (1983) Results of the research cruises of FRV “Walther Herwig” to South America. LXI. Revision of the South American skate genus Psammobatis Günther, 1870 (Elasmobranchii: Rajiformes, Rajidae). Arch FischWiss 34:23–80

    Google Scholar 

  • McEachran JD, Aschliman N (2004) Phylogeny of Batoidea. In: Carrier JC, Musick JA, Heithaus MR (eds) Biology of sharks and their relatives. CRC Press, Boca Raton, FL, pp 79–114

    Google Scholar 

  • McEachran JD, Compagno LJV (1982) Interrelationships of and within Breviraja based on anatomical structures (Pisces: Rajoidei). Bull Mar Sci 33:399–425

    Google Scholar 

  • McEachran JD, Dunn KA (1998) Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs (Chondrichthyes: Rajidae). Copeia 2:271–290

    Article  Google Scholar 

  • Menni RC, Stehmann MFW (2000) Distribution, environment, and biology of batoid fishes off Argentina, Uruguay, and Brazil. A review Rev Mus Argentino Cienc Nat 2:69–109

    Google Scholar 

  • Seret B (1986) Deep water skates from Madagascar. Part 1. Anacanthobatidae (Pisces, Chondirchthyes, Batoidea), second record of the skate Anacanthobatis ori (Wallace, 1967) from off Madagascar. Cybium 10:307–326

    Google Scholar 

  • Seret B (1989a) Deep water skates of Madagascar. Part 2. Rajidae. (Pisces, Chondrichthyes, Batoidea) Gurgesiella (Fenestraja) maceachrani sp. n. Cybium 13:55–64

    Google Scholar 

  • Seret B (1989b) Deep water skates of Madagascar. Part 3. Rajidae. (Pisces, Chondrichthyes, Batoidea) Raja (Dipturus) crosnieri sp. n. Cybium 13:115–130

    Google Scholar 

  • Stehmann M (1970) A taxonomic rearrangement of the northeastern Atlantic Rajidae (Chondrichthyes, Batoidea) based on comparative morphological and anatomical studies. Arch FischWiss 21:73–164

    Google Scholar 

  • Stehmann M (1995) First and new records of skates (Chondrichthyes, Rajiformes, Rajidae) from the West African contintental slope (Morocco to South Africa), with descriptions of two new species. Arch Fish Mar Res 43:1–119

    Google Scholar 

  • Stehmann M, Bürkel DL (1984) Rajidae. In: Whitehead PJP, Bauchot M-L, Hureau JC, Tortonese E (eds) Fishes of the Northeastern Atlantic and Mediterranean. UNESCO, Paris, pp 163–196

    Google Scholar 

  • Stevenson DE (2004) Identification of skates, sculpins, and smelts by observers in North Pacific groundfish fisheries (2002–2003). NOAA Tech Memo NMFS-AFSC-142

  • Stevenson DE, Orr JW (2005) Records of two deepwater skate species from the eastern Bering Sea. Northwest Nat 86:71–81

    Article  Google Scholar 

  • Wallace JH (1967) The batoid fishes of the east coast of southern Africa. Part III: skates and electric rays. S Afr Assoc Mar Biol Res, Oceanogr Res Inst, Invest Rep 17:1–62

    Google Scholar 

  • Whitley GP (1931) Studies in Ichthyology No 4. Rec Aust Mus v 18:96–133

    Google Scholar 

  • Whitley GP (1939) Taxonomic notes on sharks and rays. Aust Zool 9:227–262

    Google Scholar 

  • Zorzi GD, Anderson ME (1988) Records of the deep-sea skates, Raja (Amblyraja) badia Garman, 1899 and Bathyraja abyssicola (Gilbert, 1896) in the Eastern North Pacific, with a new key to California skates. Calif Fish Game 74:87–105

    Google Scholar 

Download references

Acknowledgements

Funding for this research was provided to DAE by NOAA/NMFS to the National Shark Research Consortium and Pacific Shark Research Center and by the American Elasmobranch Society. LJVC’s research was supported by grants from the South African Foundation for Research Development (FRD), National Science Foundation (NRF) and Fisheries Development Corporation, and by the South African Museum and the former J.L.B. Smith Institute of Ichthyology (now South African Institute of Aquatic Biodiversity).

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to David A. Ebert.

Appendix 1

Appendix 1

A checklist of the known skate species. Names of doubtful species are preceeded by a question mark; valid species that are doubtfully placed in agences have a question mark at the end of the generic name.

Class Chondrichthyes—Cartilaginous fishes

       Subclass Elasmobranchii—Sharks and Rays

           Superorder Batoidea—Rays

                Order Rajiformes—Skates

                      Suborder Rajoidei

Family Arhynchobatidae Fowler, 1934—Softnose Skates

This group in an alternate arrangement is ranked as subfamily Arhynchobatinae (e.g., McEachran and Dunn 1998) within a single family, Rajidae, for all skates. Worldwide this family has 12 genera and approximately 89 valid species and 25 undescribed and dubious species.

Genus Arhynchobatis Waite, 1909—Longtailed skates. Found only in New Zealand waters on insular shelves. A single species.

  • Arhynchobatis asperrimus Waite, 1909—Longtailed skate

Genus Atlantoraja Menni, 1972—La Plata skates. Endemic to the western South Atlantic coast of South America, mostly on continental shelves. Three species, plus possibly one undescribed species.

  • Atlantoraja castelnaui (Ribero, 1904)—Spotback skate

  • Atlantoraja cyclophora (Regan, 1903)—Eyespot skate

  • Atlantoraja platana (Günther, 1880)—La Plata skate

Genus Bathyraja Ishiyama 1958—Softnose skates. This wide-ranging speciose and primarily deep-water genus is most diverse at high latitudes, on continental insular shelves and slopes. Forty-six species and at least four, possibly more, undescribed species.

  • Bathyraja abyssicola (Gilbert, 1896)—Deepsea skate

  • Bathyraja aguja (Kendall and Radcliffe, 1912)—Aguja skate

  • Bathyraja aleutica (Gilbert, 1895)—Aleutian skate

  • Bathyraja andriashevi Dolganov, 1985—Little-eye skate

  • Bathyraja bergi Dolganov, 1985—Bottom skate

  • Bathyraja brachyurops (Fowler, 1910)—Broadnose skate

  • Bathyraja caerulonigricans Ishiyama and Ishihara, 1977—Purpleblack skate

  • Bathyraja cousseauae de Astarloa and Mabragaña, 2004—Joined fins skate

  • Bathyraja diplotaenia (Ishiyama, 1952)—Duskypink skate

  • Bathyraja eatoni (Günther, 1876)—Eaton’s skate

  • Bathyraja fedorovi Dolganov, 1985—Cinnamon skate

  • Bathyraja griseocauda (Norman, 1937)—Graytail skate

  • Bathyraja hesperafricana Stehmann, 1995—West African skate

  • Bathyraja irrasa Hureau and Ozouf-Costaz, 1980—Kerguelen sandpaper skate

  • Bathyraja ishiharai Stehmann, 2005—Abyssal skate

  • Bathyraja isotrachys (Günther, 1877)—Challenger skate

  • Bathyraja kincaidii (Garman, 1908)—Sandpaper skate

  • Bathyraja lindbergi Ishiyama and Ishihara, 1977—Commander Skate

  • Bathyraja longicauda (de Buen, 1959)—Slimtail skate

  • Bathyraja maccaini Springer, 1971—McCain’s skate

  • Bathyraja maculata Ishiyama and Ishihara, 1977—Whiteblotched skate

  • Bathyraja mariposa Stevenson, Orr, Hoff, and McEachran, 2004—Butterfly skate

  • Bathyraja matsubarai (Ishiyama, 1952)—Duskypurple skate

  • Bathyraja meridionalis Stehmann, 1987—Darkbelly skate

  • Bathyraja microtrachys (Osburn and Nichols, 1916)—Fine-spine skate

  • Bathyraja minispinosa Ishiyama and Ishihara, 1977—Smallthorn skate

  • Bathyraja notoroensis Ishiyama and Ishihara, 1977–Notoro skate

  • ?Bathyraja ogilbyi (Whitley, 1939)—Great skate (=‘Raja’ whitleyi?)

  • Bathyraja pallida (Forster, 1967)—Pale skate

  • Bathyraja papilionifera Stehmann, 1985—Butterfly skate

  • Bathyraja parmifera (Bean, 1881)—Alaska skate

  • Bathyraja peruana McEachran and Miyake, 1984—Peruvian skate

  • Bathyraja richardsoni (Garrick, 1961)—Richardson’s skate

  • Bathyraja scaphiops (Norman, 1937)—Cuphead skate

  • Bathyraja schroederi (Krefft, 1968)—Whitemouth skate

  • Bathyraja shuntovi Dolganov, 1985—Narrownose skate

  • Bathyraja simoterus (Ishiyama, 1967)—Hokkaido skate

  • Bathyraja smirnovi (Soldatov and Pavlenko, 1915)—Golden skate

  • Bathyraja smithii (Müller and Henle, 1841)—African softnose skate

  • Bathyraja spinicauda (Jensen, 1914)—Spinetail skate

  • Bathyraja spinosissima (Beebe and Tee-Van, 1941)—White skate

  • Bathyraja trachouros (Ishiyama, 1958)—Eremo skate

  • Bathyraja trachura (Gilbert, 1892)—Roughtail skate

  • Bathyraja tunae Stehmann, 2005—Cristina’s skate

  • Bathyraja tzinovskii Dolganov, 1985—Creamback skate

  • Bathyraja violacea (Suvorov, 1935)—Okhotsk skate

Genus Brochiraja Last and McEachran, 2006—Thornsnout skates. An endemic genus found in the deepseas surrounding New Zealand. Six described species.

  • Brochiraja aenigma Last and McEachran, 2006—Enigma skate

  • Brochiraja albilabiata Last and McEachran, 2006—Whitelipped skate

  • Brochiraja asperula (Garrick and Paul, 1974)—Prickly deepsea skate

  • Brochiraja leviveneta Last and McEachran, 2006—Smooth blue skate

  • Brochiraja microspinifera Last and McEachran, 2006—Small prickly skate

  • Brochirajaspinifera (Garrick and Paul, 1974)—Spiny deepsea skate

Genus Irolita Whitley, 1931—Round skates. Endemic to Australia, found on the outer shelves and upper slopes. One described and one undescribed species.

  • Irolita waitii (McCulloch, 1911)—Southern round skate

Genus Notoraja Ishiyama, 1958—Velvet skates. Western Pacific, found mostly on continental and insular slopes. Four described and one undescribed species.

  • Notoraja laxipella (Yearsley and Last, 1992)—Eastern looseskin skate

  • Notoraja ochroderma McEachran and Last, 1994—Yellow skate

  • Notoraja subtilispinosa Stehmann, 1989—Deep-water skate

  • Notoraja tobitukai (Hiyama, 1940)—Leadhued skate

Genus Pavoraja Whitley 1939—Peacock skates. Western South Pacific, mainly on outer continental shelves and upper slopes. Two described species with at least four or more undescribed species.

  • Pavoraja alleni McEachran and Fechhelm, 1982—Allen’s skate

  • Pavoraja nitida (Günther, 1880)—Peacock skate

Genus Psammobatis Günther, 1870—Sandskates. Endemic to the eastern South Pacific and western South Atlantic coasts of South America, mostly found on continental shelves. Eight described species.

  • Psammobatis bergi Marini, 1932—Blotched sandskate

  • Psammobatis extenta Garman, 1913—Zipper skate

  • Psammobatis lentiginosa (Bigelow and Schroeder, 1951)—Freckled sandskate

  • Psammobatis normani McEachran, 1983—Shortfin sandskate

  • Psammobatis parvacauda McEachran 1983—Smalltail sandskate

  • Psammobatis rudis Günther, 1870—Smallthorn sandskate

  • Psammobatis rutrum Jordan, 1890—Spade sandskate

  • Psammobatis scobina (Philippi, 1857)—Raspthorn sandskate

Genus Pseudoraja Bigelow and Schroeder, 1954—Fanfin skates. A single endemic species to the western North Atlantic, found on continental slopes.

  • Pseudoraja fischeri Bigelow and Schroeder, 1954—Fanfin skate

Genus Rhinoraja Ishiyama, 1952—Jointnose skates. A wide-ranging, but patchily distributed, group found in the North Pacific, western South Atlantic coast of South America, and Antarctica, found on continental and insular shelves and slopes. This genus differs from Bathyraja primarily in having a basal joint in the rostral cartilage and may not be separable from that genus. Twelve species.

  • Rhinoraja albomaculata (Norman, 1937)—White-dotted skate

  • Rhinoraja interrupta (Gill and Townsend, 1897)—Bering skate

  • Rhinoraja kujiensis (Tanaka, 1916)—Dapple-bellied softnose skate

  • Rhinoraja longicauda Ishiyama, 1952—Whitebellied softnose skate

  • Rhinoraja macloviana (Norman, 1937)—Patagonian skate

  • Rhinoraja magellanica (Philippi, 1902)—Magellan skate

  • Rhinoraja multispinis (Norman, 1937)—Multispine skate

  • Rhinoraja murrayi (Günther, 1880)—Murray’s skate

  • ?Rhinoraja obtusa (Gill and Townsend, 1897)—Blunt skate.

  • Rhinoraja odai Ishiyama, 1958—Oda’s skate

  • ?Rhinoraja rosispinis (Gill and Townsend, 1897)—Flathead skate

  • Rhinoraja taranetzi Dolganov, 1985—Mud skate

Genus Rioraja Whitley, 1939—Rio skates. Endemic to the western South Atlantic coast of South America, found on continental shelves. One species.

  • Rioraja agassizi (Müller and Henle, 1841)—Rio Skate

Genus Sympterygia Müller and Henle, 1837—Fanskates. Endemic to the Atlantic and Pacific coasts of South America, found primarily on continental shelves. Four valid species.

  • Sympterygia acuta Garman, 1877—Bignose fanskate

  • Sympterygia bonapartei Müller and Henle, 1841—Smallnose fanskate

  • Sympterygia brevicaudata Cope, 1877—Shorttail fanskate

  • Sympterygia lima (Poeppig, 1835)—Filetail skate

Family Rajidae Blainville, 1816—Hardnose Skates

The Rajidae, in some arrangements (e.g. McEachran and Dunn 1998; McEachran and Aschliman 2004), are considered monophyletic, with all species occurring within this family due largely to their similarity in appearance. In the present arrangement this family has 13 genera, approximately 138 valid species, and at least 45 undescribed and dubious species world-wide.

Genus Amblyraja Malm, 1877—Stout skates. A wide-ranging, circumglobal, genus mostly found at higher latitudes and in deep tropical waters, mostly on continental and insular shelves and slopes, and around seamounts. Ten valid species and two undescribed species.

  • Amblyraja badia (Garman, 1899)—Broad skate

  • Amblyraja doellojuradoi (Pozzi, 1935)—Southern thorny skate

  • Amblyraja frerichsi (Krefft, 1968)—Thickbody skate

  • Amblyraja georgiana (Norman, 1938)—Antarctic starry skate

  • Amblyraja hyperborea (Collette, 1879)—Arctic skate

  • Amblyraja jenseni (Bigelow and Schroeder, 1950)—Jensen’s skate

  • Amblyraja radiata (Donovan, 1808)—Thorny skate

  • Amblyraja reversa (Lloyd, 1906)—Reversed skate

  • Amblyraja robertsi (Hulley, 1970)—Bigmouth skate

  • Amblyraja taaf (Meisner, 1987)—Whiteleg skate

Genus Breviraja Bigelow and Schroeder, 1948—Lightnose skates. Found in the eastern and western Atlantic, mostly on continental slopes. Six described and possibly one undescribed species.

  • Breviraja claramaculata McEachran and Matheson, 1985—Bright spotted skate

  • Breviraja colesi Bigelow and Schroeder, 1948—Lightnose skate

  • Breviraja marklei McEachran and Miyake, 1987—Nove Scotia skate

  • Breviraja mouldi McEachran and Matheson, 1995—Blacknose skate

  • Breviraja nigriventralis McEachran and Matheson, 1985—Blackbelly skate

  • Breviraja spinosa Bigelow and Schroeder, 1950—Spiny skate

Genus Dactylobatus Bean and Weed, 1909—Skilletskates. Endemic to the western North Atlantic, primarily found on continental slopes. Two valid species.

  • Dactylobatus armatus Bean and Weed, 1909—Skillet skate

  • Dactylobatus clarki (Bigelow and Schroeder, 1958)—Hook skate

Genus Dipturus Rafinesque, 1810—Longnosed skates. A wide-ranging genus in cool temperate to tropical waters, found mostly on continental shelves and slopes. At least 31 species with 15 or more undescribed species; several species within this genus are questionably placed here.

  • Dipturus batis (Linnaeus, 1758)—Gray skate

  • Dipturus bullisi (Bigelow and Schroeder 1962)—Tortugas skate

  • Dipturus campbelli (Wallace, 1967)—Blackspot skate

  • Dipturus chilensis (Guichenot, 1848)—South American Yellownose skate

  • Dipturus crosnieri (Seret, 1989)—Madagascar skate

  • Dipturus diehli Soto and Mincarone, 2001—Thorny tail skate

  • Dipturus doutrei (Cadenat, 1960)—Javelin skate

  • Dipturus ecuadoriensis (Beebe and Tee-Van, 1941)—Ecuador skate

  • Dipturus garricki (Bigelow and Schroeder, 1958)—San Blas skate

  • Dipturus gigas (Ishiyama, 1958)—Giant skate

  • Dipturus gudgeri (Whitley, 1940)—Bight skate

  • Dipturus innominata (Garrick and Paul, 1974)—New Zealand smoothskate

  • Dipturus johannisdavisi (Alcock, 1899)—Travancore skate

  • Dipturus kwangtungensis (Chu, 1960)—Kwangtung skate

  • Dipturus laevis (Mitchell, 1817)—Barndoor skate

  • Dipturus lanceorostrata (Wallace, 1967)—Rattail skate

  • Dipturus leptocauda (Krefft and Stehmann, 1975)—Thintail skate

  • Dipturus linteus (Fries, 1838)—Sailray

  • Dipturus macrocauda (Ishiyama, 1955)—Japanese bigtail skate

  • Dipturus mennii Gomes and Parago, 200—South Brazilian skate

  • Dipturus nasutus (Banks in Müller and Henle 1841)—New Zealand longnose skate

  • Dipturus nidarosiensis Collett, 1880)—Norwegian skate

  • Dipturus olseni (Bigelow and Schroeder, 1951)—Spreadfin skate

  • Dipturus oregoni (Bigelow and Schroeder, 1958)—Hooktail skate

  • Dipturus oxyrinchus (Linnaeus, 1758)—Longnosed skate

  • Dipturus pullopunctata (Smith, 1964)—Slime skate

  • Dipturus springeri (Wallace, 1967)—Roughbelly skate

  • Dipturus stenorhynchus (Wallace, 1967)—Prownose skate

  • Dipturus teevani (Bigelow and Schroeder, 1951)—Florida skate

  • Dipturus tengu (Jordan and Fowler, 1903)—Japanese acutenose skate

  • Dipturus trachyderma Krefft and Stehmann, (1975)—Roughskin skate

Genus Fenestraja McEachran and Compagno, 1982—Pluto skates. A widely scattered group with representatives in the western North Atlantic, western Indian Ocean, and western Pacific, found mostly on continental and insular slopes. Eight species.

  • Fenestraja atripinna (Bigelow and Schroeder, 1950)—Blackfin skate

  • Fenestraja cubensis (Bigelow and Schroeder, 1950)—Cuban skate

  • Fenestraja ishiyamai (Bigelow and Schroeder, 1962)—Plain pygmy skate

  • Fenestraja maceachrani (Seret, 1989)—Madagascar skate

  • Fenestraja mamillidens (Alcock, 1889)—Prickly skate

  • Fenestraja plutonia (Garman, 1881)—Pluto skate

  • Fenestraja sibogae (Weber, 1913)—Siboga pygmy skate

  • Fenestraja sinusmexicanus (Bigelow and Schroeder, 1950)—Gulf of Mexico pygmy skate

Genus Gurgesiella de Buen, 1959—Finless pygmy skates. Western South Atlantic and eastern South Pacific, these skates occur on continental and insular slopes. Three described species.

  • Gurgesiella atlantica (Bigelow and Schroeder, 1962)—Atlantic skate

  • Gurgesiella dorsalifera McEachran and Compagno, 1980—Onefin skate

  • Gurgesiella furvescens (de Buen, 1959)—Dusky finless skate

Genus Leucoraja Malm, 1877—Rough skates. This genus is most diverse in the North Atlantic and Mediterranean Sea, but includes a few outlying endemic species off southern Africa and Australia, mostly found on continental shelves and slopes. Twelve described and three undescribed species.

  • Leucoraja circularis (Couch, 1838)—Sandy skate

  • Leucoraja compagnoi (Stehmann, 1995)—Tigertail skate

  • Leucoraja erinacea (Mitchell, 1825)—Little skate

  • Leucoraja fullonica (Linnaeus, 1758)—Shagreen skate

  • Leucoraja garmani (Whitley, 1939)—Rosette skate

  • Leucoraja lentiginosa (Bigelow and Schroeder, 1951)—Freckled skate

  • Leucoraja leucosticta (Stehmann, 1971)—Whitedappled skate

  • Leucoraja melitensis (Clark, 1926)—Maltese skate

  • Leucoraja naevus (Müller and Henle, 1841)—Cuckoo skate

  • Leucoraja ocellata (Mitchell, 1815)—Winter skate

  • Leucoraja wallacei (Hulley, 1970)—Yellowspot skate

  • Leucoraja yucatanensis (Bigelow and Schroeder, 1950)—Yucatan skate

Genus Malacoraja Stehmann, 1970—Soft skates. Restricted to the Atlantic Ocean, except for one undescribed species from the southwestern Indian Ocean, these are mostly deep-slope skates. Four described species and one possibly undescribed species.

  • Malacoraja kreffti (Stehmann, 1978)—Krefft’s skate

  • Malacoraja obscura de Carvalho, Gomes, and Gadig, 2005—Dusky skate

  • Malacoraja senta (Garman, 1885)—Smooth skate

  • Malacoraja spinacidermis (Barnard, 1923)—Roughskin skate

Genus Neoraja McEachran and Compagno, 1982—Pygmy skates. Western North Atlantic and eastern Atlantic, mostly found on continental slopes. Four species.

  • Neoraja africana Stehmann and Seret, 1983—African skate

  • Neoraja caerulea (Stehmann, 1976)—Blue skate

  • Neoraja carolinensis McEachran and Stehmann, 1984—Carolina skate

  • Neoraja stehmanni (Hulley, 1972)—African dwarf skate

Genus Okamejei Ishiyama, 1958—Spiny rasp skates. This genus is most diverse in the western Pacific, but with outlier species in the Indian Ocean, occurs on continental shelves and slopes. Fourteen described and at least two undescribed species.

  • Okamejei acutispina (Ishiyama, 1958)—Japanese sharpspine skate

  • Okamejei australis (Macleay, 1884)—Common skate

  • Okamejei boesemani (Ishihara, 1987)—Black sand skate

  • Okamejei cerva (Whitley, 1939)—Whitespotted skate

  • Okamejei heemstrai (McEachran and Fechhlem, 1982)—East African skate

  • Okamejei hollandi (Jordan and Richardson, 1909)—Yellowspot skate

  • Okamejei kenojei (Müller and Henle, 1841)—Ocellate spot skate

  • Okamejei koreana (Jeong and Nakabo, 1997)—Korean skate

  • Okamejei lemprieri (Richardson, 1845)—Thornback skate

  • Okamejei meerdervoorti (Bleeker, 1860)—Bigeye skate

  • Okamejei philipi (Lloyd, 1906)—Aden ringed skate

  • Okamejei pita (Fricke and Al-Hussar, 1995)—Pita skate

  • Okamejei powelli Alcock, 1898—Indian ringed skate

  • Okamejei schmidti Ishiyama, 1958—Browneye skate

Genus Raja Linnaeus, 1758—Ocellate skates. Primarily found in the eastern North Atlantic, Mediterranean and Black seas, but with some species ranging into the southwestern Indian Ocean, usually found on continental and insular shelves and slopes. Thirteen valid species, two dubious species, and at least one undescribed species.

  • Raja asterias Delaroche, 1809—Starry skate

  • Raja brachyura Lafont, 1873—Blonde skate

  • Raja clavata Linnaeus, 1758—Thornback skate

  • Raja herwigi Krefft, 1965—Cape Verde skate

  • Raja maderensis Lowe, 1838—Madera skate

  • Raja microocellata Montagu, 1818—Small-eyed skate

  • Raja miraletus Linnaeus, 1758—Twineyed skate

  • Raja montagui Fowler, 1910—Spotted skate

  • Raja polystigma Regan, 1923—Speckled skate

  • Raja radula Delaroche, 1809—Rough skate

  • Raja rondeleti Bougis, 1959—Rondelet’s skate

  • Raja straeleni Poll, 1951—False thornback skate or Biscuit skate

  • Raja undulata Lacepede, 1802—Undulate skate

Genus Rajella Stehmann, 1970—Gray skates. This genus is most diverse in the Atlantic, but with outliers in the eastern South Pacific, Indian Ocean, and Australia, mostly on continental and insular shelves and slopes. Fifteen valid species and at least four undescribed species, plus two dubious species.

  • Rajella annandalei (Weber, 1913)—Indonesian round skate

  • Rajella barnardi (Norman, 1935)—Bigthorn skate

  • Rajella bathyphila (Holt and Byrne, 1908)—Deepwater skate

  • Rajella bigelowi (Stehmann, 1978)—Bigelow’s or chocolate skate

  • Rajella caudispinosa (Von Bonde and Swart, 1923)—Munchkin skate

  • Rajella dissimilis (Hulley, 1970)—Ghost skate

  • Rajella eisenhardti Long and McCosker, 1999—Eisenhardt’s skate

  • Rajella fuliginea (Bigelow and Schroeder, 1954)—Sooty skate

  • Rajella fyllae (Lutken, 1887)—Round skate

  • Rajella kukujevi (Dolganov, 1985)—Mid-Atlantic skate

  • Rajella leopardus (von Bonde and Swart, 1923)—Leopard skate

  • Rajella nigerrima (de Buen, 1960)—Blackish skate

  • Rajella purpuriventralis (Bigelow and Schroeder, 1962)—Purplebelly skate

  • Rajella ravidula (Hulley, 1970)—Smoothback skate

  • Rajella sadowskii (Krefft and Stehmann, 1974)—Brazilian skate

Genus Rostroraja Hulley, 1972—Spearnose skates. A single species ranging from the eastern Atlantic to the southwestern Indian Ocean, mostly found on continental shelves and uppermost slopes.

  • Rostroraja alba (Lacepede, 1803)—Spearnose skate

Species nominally retained in Raja representing undescribed genus-group taxa

Genus “A” for North Pacific Raja Assemblage McEachran and Dunn, 1998. Six species.

  • Raja binoculata Girard, 1854—Big skate

  • Raja cortezensis McEachran and Miyake, 1988—Sea of Cortez skate

  • Raja inornata Jordan and Gilbert, 1881—California skate

  • Raja pulchra Liu, 1932–Mottled skate

  • Raja rhina Jordan and Gilbert, 1880—Longnose skate

  • Raja stellulata Jordan and Gilbert, 1880—Starry skate

Genus “B” for Amphi-American Raja Assemblage McEachran and Dunn, 1998. Seven species.

  • Raja ackleyi Garman, 1881—Ocellate skate

  • Raja bahamensis Bigelow and Schroeder, 1965—Bahama skate

  • Raja cervigoni Bigelow and Schroeder, 1964—Venezuela skate

  • Raja eglanteria Bosc, 1802—Clearnose skate

  • Raja equatorialis Jordan and Bollman, 1890—Equatorial skate

  • Raja texana Chandler, 1921—Roundel skate

  • Raja velezi Chirichigno, 1973—Rasptail skate

Genus “C” for western South Pacific species with provisional generic and familiar placement. Two described and at least three undescribed species.

  • Raja polyommata Ogilby, 1910—Argus skate

  • Raja whitleyi Iredale, 1938—Melbourne skate

Family Anacanthobatidae von Bonde and Swart, 1924—Legskates

McEachran and Dunn (1998) reduced this group to a sister taxon within the subfamily Rajinae. A small group, only two genera, these skates have distinctive leglike anterior pelvic fin lobes, and includes 18 valid species plus at least five undescribed species.

Genus Anacanthobatis von Bonde and Swart, 1924—Smooth legskates. A wide ranging, but sporadically distributed group found in the western North and Central Atlantic, Indian Ocean, and the western Central Pacific, on continental and insular slopes. Ten described and at least five undescribed species.

  • Anacanthobatis americanus Bigelow and Schroeder 1962—American legskate

  • Anacanthobatis borneensis Chan, 1965—Borneo skate

  • Anacanthobatis donghaiensis (Deng, Xiong and Zhan, 1983)—East China legskate

  • Anacanthobatis folirostris (Bigelow and Schroeder, 1951)—Leafnose legskate

  • Anacanthobatis longirostris Bigelow and Schroeder 1962—Longnose legskate

  • Anacanthobatis marmoratus (von Bonde and Swart, 1924)—Spotted legskate

  • Anacanthobatis melanosomus (Chan, 1965)—Blackbellied legskate

  • Anacanthobatis nanhaiensis (Meng and Li, 1981)—South China legskate

  • Anacanthobatis ori (Wallace, 1967)—Black legskate

  • Anacanthobatis stenosomus (Li and Hu, 1982)—Narrow legskate

Genus Cruriraja Bigelow and Schroeder 1948—Legskates. Western North and Central Atlantic, eastern South Atlantic, and southwestern Indian Ocean, found on continental and insular shelves and slopes. Eight valid species.

  • Cruriraja andamanica (Lloyd, 1909)—Andaman legskate

  • Cruriraja atlantis Bigelow and Schroeder, 1948—Atlantic skate

  • Cruriraja cadenati Bigelow and Schroeder 1962—Broadfoot legskate

  • Cruriraja durbanesis (von Bonde and Swart, 1924)—Smoothnose legskate

  • Cruriraja ‘parcomaculata’ Smith, 1964 (not von Bonde and Swart, 1924)—Roughnose skate

  • Cruriraja poeyi Bigelow and Schroeder, 1948—Cuban legskate

  • Cruriraja rugosa Bigelow and Schroeder 1958—Rough legskate

  • Cruriraja triangularis Smith, 1964 (=C. parcomaculata von Bonde and Swart, 1924)—Triangular skate

Rights and permissions

Reprints and permissions

About this article

Cite this article

Ebert, D.A., Compagno, L.J.V. Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei). Environ Biol Fish 80, 111–124 (2007). https://doi.org/10.1007/s10641-007-9247-0

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10641-007-9247-0

Keywords

Navigation