Author: Linnaeus, 1758

Raja (Raja) clavata Linnaeus, 1758
Diagnosis: upper surfaces always wholly prickly, even in early juveniles; underside wholly prickly in large females, only snout and margins of disc in young and large males. Orbital thorns separate, a regular row of 30-50 thorns from nape to first dorsal (number reduced in large males), 0-2 thorns between separate dorsal fins. Additional large 'buckler' thorns with swollen bases scattered on upper surface of disc in adults, also on underside (mainly in large females). Thorn pattern extremely variable. Colour: upper surface very variable, all shades of brown, variegated with dark and light spots and blotches, often marbled or producing patterns like eye-spots; plain coloured specimens recorded; tail usually with light and dark crossbars; underside white, margins of disc often grey. Size: to about 90 cm TL.

Habitat: benthic, from very shallow waters to about 300 m; common in most areas and important commercially in north-western Europe, also locally in Mediterranean. Food: all kinds of bottom animals, preference for crustaceans. Reproduction: oviparous; about 150 egg-cases laid per year, in spring (north-western Europe) or winter and spring (Mediterranean), 60-90 by 49-69 mm (excluding horns); development of embryos about 5 months (usually in inshore waters).

Distribution: Atlantic coasts from Madeira and Morocco northward to Iceland and Norway (south of Arctic Circle), as well as North Sea and western Baltic (rare), also whole Mediterranean and western part of Black Sea. Elsewhere, southward to South Africa and south-western Indian Ocean.

Complementary iconography. Clark, 1930, in Faune ichthyol. Atl. N., fiche 43 Muus & Dahlström, 1965: 53, fig. | Bini, 1967: 157-158, fig.
Eggs, embryonic and young stages. Williamson, 1913: 2, fig. 8 (egg-capsule) | Clark, 1922: 593-601, fig. 1 (egg-capsule), fig. 2-4 (young).

Kingdom Animalia
Phylum Chordata
Subphylum Vertebrata
Superclass Gnathostomata
Class Chondrichthyes
Superorder Hypotremata
Family Rajidae
Genus Raja
Species Raja clavata

Status in World Register of Marine Species

Accepted name: Raja clavata Linnaeus, 1758

Scientific synonyms and common names

Raja (Raja) clavata Linnaeus, 1758

Synonyms:

Raja (Raja) clavata Linnaeus, 1758
Raja clavata Linnaeus, 1758, Syst. Nat., ed. X: 232 ('Habitat in Oceano Europaeo'). No type material.
Raja rubus Schneider, 1801, Syst. Ichthyol.: 366-367 ('Habitat in Oceano Europaeo'). No type material.
Raja rubus: Donovan, 1802: 2 p., pl. 20 Risso, 1810: 7 Blainville, 1820 1830: 21-22, pl. 3c, fig. 1.
Raja aspera Risso, 1810, Ichthyol. Nice: 5-6 (Mediterranean). Holotype: MNHN no. B 843.
Raja pontica Pallas, 1811, Zoogr. rosso-asiat., 3: 58-60, pl. 8A-B ('In littore Tauriae'). No type material.
Dasybatis clavata: Bonaparte, 1840: fasc. 29, punt. 154, 1 pl., fig. 4.
Raja capensis Müller & Henle, 1841, System. Beschr. Plag.: 151 ('Vom Cap'). Syntype: MNHN no. 1333; 3 additional syntypes: RMNH no. D 2492, D 2493, D 2507.
Raja clavata: Müller & Henle, 1841: 135-137 Duméril, 1865: 528-531, pl. 12 Günther, 1870: 456-457 Moreau, 1881: 391-394, fig. 69 Möbius & Heincke, 1883: 154-156, fig. Smitt, 1895: 1104-1107, pl. 47 Jenkins, 1925: 338-339, pl. 134 Clark, 1926: 2428, pl. 8-10 Lozano Rey, 1928: 541-550, fig. 172, pl. 10 Nobre, 1935: 480-481 Ehrenbaum, 1936: 297-298, pl. 22 Fowler, 1936: 110-111, fig. 43 Poll, 1947: 88-90, fig. 53-56 Dieuzeide et al., 1953: 79-82, fig. Roland, 1953: 141-159, fig. 1-5 Albuquerque, 1954-1956: 160-162, fig. 97, 97A Tortonese, 1956: 244-247, fig. 135-136 Soljan, 1963: 100, fig. Svetovidov, 1964: 33-37, fig. 8 Aloncle, 1966: 42-43, fig. 1-4 Banarescu, 1969: 92-94, fig 49 Wheeler, 1969: 100-101, fig. Stehmann, 1970: 149, pl. 2, 10, 18.
Raja rizacanthus Regan, 1906, Ann. Natal Mus., 1:1-6 (Coast of Natal). Holotype: BMNH no. 1905.6.8.13.
Betaraia clavata: Leigh-Sharpe, 1924: 568 Capapé, 1976 a: 907, 6 fig.

Common names:

Nagelroche [De]
thornback ray [En]
raya de clavos [Es]
raie bouclée [Fr]
razza chiodata [It]
raia lenga [Pr]
stekelrog [Ne]

*Nobre, A. 1935. Fauna marinha de Portugal. 1. Vertebrados (Mamiferos, Reptis e Peixes), Porto.: Ixxxiv+1-21 (Mamiferos); 1-5 (Reptis); 1-574 (Peixes), 77 pl., 64 phot.

Albuquerque, R. M. 1954-1956. Peixes de Portugal e ilhas adjacentes. Chavas para a sua determinação. Port. Acta biol., ser. B,5:xvi+1167 p., 445 fig.

Aloncle, H. 1966. A propos d'un caractère anatomique intéressant dans la détermination des Rajidae. Bull Inst. Pêch. marit. Maroc, (14): 42-50, fig. 1-14.

Banarescu, P. 1969. Cyclostomata si Chondrichthyes. Fauna Repub. pop. rom., Bucuresti, 12 (1): 106 p., 50 fig.

Bini, G. 1967-72. Atlante dei pesci delle Coste italiane. Mondo Sommerso, Milano, 9 vol: I, 1967, Leptocardi, Ciclostomi, Selaci, 206 p., 66 fig+64 col. fig. II, 1971, Osteitti (Acipenseriformi, Clupeiformi, Mictofiformi, Anguilliformi), 300 p., 73 col. fig. III, 1970, Notacantiformi . . . Zeiformi, 229 p., 34 fig+63 col. fig. IV, 1968, Perciformi (Mugiloidei, Percoidei), 163 p., 34 fig+49 col. fig. V, 1968, Perciformi (Percoidei), 175 p., 22 fig.+56 col. fig. VI, 1968, Perciformi (Trichiuroidei . . . Blennioidei), 177 p., 48 fig+57 col. fig. VII,1969, Perciformi (Ofidioidei . . . Dactilopteroidei), 196 p., 57 fig.+59 col. fig. VIII,1968, Pleuronettiformi, Echeniformi, Gobioesociformi, Tetraodontiformi, Lofiformi, 164 p., 34+63 fig. IX, 1972, Introduzione. Parte generale. Aggiornamenti. Indici. 176 p.

Blainville, H. M. Ducrotay de. 1825. Vertébrés. Classe V. Poissons. Sous classe Ire. Poissons cartilagineux. In: P. Vieillot et al., 1820-30, Faune francaise ou histoire naturelle générale et particulière des animaux qui se trouvent en France, livr. 13-14, 96 p., 22 pl. (num. I to 68 with lacks).

Bloch, M. E.; Schneider, J. G. 1801. M. E. Blochii Systema Ichthyologiae iconibus cx illustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit J. G. Schneider, Saxo. Berolini: Ix+584 p., 110 pl.

Bonaparte, C. L. 1832-41, Iconografia della Fauna italica per le quattro classi degli Animali Vertebrati, tomo III, Pesci, Roma, without pagination, 75 puntate (in 30 fasc.), 78 pl.

Capapé, C. 1976a. Contribution a la biologie des Rajidae des cates tunisiennes. 111. Raja clavata Linne, 1758, repartition geographique et bathymetrique, sexualite, reproduction et fecondite. Bull Mus. Nat. Hist. Nat., (3) 393, Zool. 27S: 907-922, 6 fig.

Clark, R. S.1922. Rays and Skates (Raiae). No. 1. Egg-Capsules and Young. J. mar. biol. Ass. U.K (n.s.), 12 (4): 577-643, fig 1-20.

Clark, R. S.1926. Rays and Skates, a revision of the european species. Fisheries, Scotland, Scient. Invest., (1) :1-66, fig. 1-42, pl. 1-36.

Clark, R. S.1929-32. Selachii, Raiidae. In: L. Joubin, ed., Faune ichthyol. Atl. N., CIEM, Copenhagen: fiches 34bis-57.

Dieuzeide, R.; Novella, M. (collab. J. Roland), 1953-55. Catalogue des Poissons des Côtes algériennes. Bull. Stn Aquic. Pêch. Castiglione, I (n.s.), (4), 1952 [1953]: 1-135, 73 fig. n. num.; II (n.s.), (5), 1953 [1954] :1-258, 135 fig. n. num.; III (n.s.), (6), 1954 [1955] :1-384, 202 fig. n. num.

Donovan, E. 1802-08. The Natural History of British Fishes including scientific and general descriptions of the most interesting species, and an extensive selcction of accurately finished coloured plates, London. 1, 1802: [1-70], pl. I-XXIV; 2, 1903: [71-164], pl. XXV-XLVIII; 3, 1804: [165-300], pl. XLIX-LXXII; 4, 1806: [301-406], pl. LXXIII-XCVI; 5, 1808: [407-516], pl. XCVIICXX.

Duméril, A. H. A.1865. Histoire naturelle des poissons ou ichthyologie générale, Paris, 2 vol., 1: 720 p., Atlas: 14 pl.

Ehrenbaum, E. 1936. Naturgeschichte und wirtschaftliche Bedeutung der Seefische Nordeuropas, in Handbuch der Seefischerei N. Eur., 2: x+337 p., 276 fig., 2 tabl., 26 pl.

Fowler, H. W.1936. The Marine Fishes of West Africa, based on the collection of the American Museum Congo Expedition 1909-15. Bull. am. Mus. nat. Hist., 70 (1), Jan. 21, 1936: vii+606 p., fig. 1-275; (2), Nov. 18,1936: 607-1493, fig. 276-567.

Günther, A.1870. Catalogue of the fishes in the British Museum. 8. Catalogue of the Physostomi containing the families Gymnotidae, Symbranchidae, Muraenidae, Pegasidae and of the Lophobranchii, Plectognathi, Dipnoi, Ganoidei, Chondropterygii, Cyclostomata, Leptocardii in the collection of the British Museum. London, xxv+549 p.

Jenkins, J. T. 1925. The fishes of the British Isles both fresh water and salt. London: vii+ 376 p., 143 pl.

Leigh-Sharpe, W. H. 1924. The comparative morphology of the secondary sexual characters of elasmobranch fishes. The claspers, clasper siphons, and claspr glands. VII. J. Morph., 39 (2): 567-577, fig. 1-15.

Linnaeus, C. 1758. Systema Naturae, ed. X, vol. 1, 824 p. Nantes & Pisces: 230-338. (Reprint, 1956, London.)

Lozano y Rey, L.1928. Fauna Ibérica. Pesces (Generalidades, Ciclóstomos y Elasmobranquios). Mus. Nac. Ciencias Nat., Madrid, 1:1-692,197 fig., 20 pl.

Möbius, K.; Heincke, F. 1883. Die Fische der Ostsee in Bericht der Comission zur wissenschaftlichen Untersuchung der Deutschen Meere in Kiel. Berlin :1-206, fig., 1 map.

Moreau, E.1881-91. Histoire naturelle des poissons de la France, Paris, I, 1881: I-VII+ 1-480, fig. 1-82; II, 1881:1-572, fig. 83-145; III, 1881:1-697, fig. 146-220; Suppl., 1891: 1-144, fig. 221-227.

Müller, J.; Henle, J.1841. Systematische Beschreibung der Plagiostomen. Berlin, xxii+204 p., 60 pl.

Muus, B. J.; Dahlström, P. 1965. Meeresfische in Farben, 244 p., 173 numb. and many unnumb. fig.

Pallas, P. S.1811 [probably 1814]. Zoographia rosso-asiatica, sistens omnium animalium in extenso Imperio Rossico et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones anatomen atque icones plurimorum, 3 vol. Petropoli, 3, 428 p. (pl. [1], XIII, XIV, XV, XXI, XX (sic)) (89 pl. are mentioned in the text but are unpublished. Perhaps published in Icones ad Zoographiam Rosso-Asiaticam, 1834-427.). (2nd ed., 1831, 3 vol. and Atlas, Petropoli.)

Poll, M.1947. Poissons marins in Faune de Belgique. Bruxelles, 452 p., 267 fig., 2 pl., 2 maps n. num. (inset).

Regan, C. T.1906b. Descriptions of new or little known fishes from the coast of Natal. Ann. Natal Mus., 1: 1-6.

Risso, A.1810. Ichthyologie de Nice, ou histoire naturelle des poissons du département des Alpes Maritimes. Paris, xxxvi+388 p., 11 pl. (Reprint, 1966, Asher, Amsterdam.)

Roland, J. 1953. Diagnoses de quelques Raies des Côtes algériennes. Bull. Stn Aquic. Pêch. Castiglione (n.s.), 1952 (4) :139-274, fig. 1-32.

Smitt, F. A. 1893-95. (ed.) A history of Scandinavian fishes, Stockholm and Paris, Atlas, I, 1893, pl. i-xxvii.

Soljan, T. 1963. Fishes of the Adriatic (Ribe Jadrana). Fauna et flora adriatica 1 (revised and enlarged for the English edition), 428 p., many unnumbered fig.

Stehmann, M. 1970. Vergleichend morphologische und anatomische Untersuchungen zur Neuordnung der Systematik der nordostatlantischen Rajidae (Chondrichthyes, Batoidei). Arch. FischWiss., 21(2): 73-163,1 fig., pl. 1-27.

Svetovidov, A. N. 1964. Rîbî Chernogo Morya. [The fishes of the Black Sea]. Opred Faune SSSR, 86 :1-552, fig. 1-191 (in Russian).

Tortonese, E. 1956b. Leptocardia, Ciclostomata, Selachii. Fauna Ital., 2: 334 p., 163 fig.

Wheeler, A. 1969. The Fishes of the British Isles and North-West Europe. Macmillan, London, Melbourne and Toronto: i-xvii+1-163, 5+177 fig., +392 fig. (princ. sp.)+92 n. num. fig., 16 pl., maps.

Williamson, H. C. 1913. On the eggs of certain skates (Raia). Scient. Invest. Fishery Bd Scotl., 1912 [1913], (1): 1-6, pl. 1-5.

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