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Scyliorhinus ugoi Soares, Gadig & Gomes, 2015 Paratypes: MZUSP 110448; one adult male; 480 mm TL (cited as 465 mm TL in Soares et al., 2015); Alagoas, North-eastern Brazil, 9ºS, 34º50’W; depth not listed; collector not listed; 07 Nov 1997; MZUSP 110449; one adult male; 437 mm TL (cited as 445 mm TL in Soares et al., 2015); Rio Grande do Norte, North-eastern Brazil,; 6º14’S, 34º51’W); depth not listed; collector not listed; 30 Nov 1998.

The complete mitochondrial DNA sequence of the Nursehound Scyliorhinus stellaris has been determined for the first time and compared with congeneric species. The mitogenome sequence was 16,684 bp in length. The mitogenome is composed of 13 PCGs, 2 rRNAs, 22 transfer RNA genes and non-coding regions. The gene order of the newly sequenced mitogenome is analogous to the organization described in other vertebrate genomes. The typical conservative blocks in the control region were indicated. The phylogenetic analysis revealed a monophyletic origin of the Scyliorhininae subfamily, and within it, two subclades were identified. A significant divergence of Scyliorhinus spp. together with Poroderna patherinum in relation to the group of Cephaloscyllium spp. was observed, except for Scyliorhinus torazame, more related to this last cited clade. A hypothesis of a divergent evolution consequent to a selective pressure in different geographic areas, which lead to a global latitudinal diversity gradient, has been suggested to explain this phylogenetic reconstruction. However, convergent evolution on mitochondrial genes could also involve different species in some areas of the world.

The genus Scyliorhinus is part of the family Scyliorhinidae, the most diverse family of sharks and of the subfamily Scyliorhininae along with Cephaloscyllium and Poroderma. This study reviews the phylogenetic relationships of species of Scyliorhinus in the subfamily Scyliorhininae. Specimens of all Scyliorhinus species were examined as well as specimens of four of the 18 species of Cephaloscyllium, two species of Poroderma, representatives of almost all other catshark (scyliorhinid) genera and one proscylliid (Proscyllium habereri). A detailed morphological study, including external and internal morphology, morphometry and meristic data, was performed. From this study, a total of 84 morphological characters were compiled into a data matrix. Parsimony analysis was employed to generate hypotheses of phylogenetic relationships using the TNT 1.1. Proscyllium habereri was used to root the cladogram. The phylogenetic analysis, based on implied weighting (k = 3; 300 replications and 100 trees saved per replication), resulted in three equally most parsimonious cladograms with 233 steps, with a CI of 0.37 and an RI of 0.69. The monophyly of the subfamily Scyliorhininae is supported as well as of the genus Scyliorhinus, which is proposed to be the sister group of Cephaloscyllium. The phylogenetic relationships amongst Scyliorhinus species are presented for the first time.

Lantern sharks are small shark species that can be seen at depths between 70 and 2000 meters. Due to their luminescent characteristics, they have been called “Lantern shark”. In total eleven specimens of the velvet belly lantern shark, Etmopterus spinax (Linnaeus, 1758), were caught in the deep seas of Northern Cyprus by using a bottom trawl. Sampling was carried out using 13 trawling operations. The collected samples were placed in 4% formalin and stored at the Museum of the Systematic, Faculty of Fisheries, Mersin University, (catalogue number: MEUFC-18-11-082). As a sampling area, the depths between 274 and 641 m were selected. Other cartilaginous fish caught during sampling except E. spinax were Galeus melastomus (1 individual), Squalus acanthias (4 individuals), Scyliorhinus canicula (85 individuals). E. spinax made up 10.89% of all cartilaginous fishes which were caught. Species identification for all fishes caught is made with the help of morphological features.

Scyliorhinus hachijoensis sp. nov. (Table 1 and Figs. 1���9) New English name: Cinder cloudy catshark; new Japanese name: Fukami-torazame. Scyliorhinus sp.: Hagiwara, 1993: 1, 9, Tables 1 ���3, Fig. 3 (keeping and reproduction) Holotype. NSMT-P135960, adult male, 370 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 200���300 m depth, 24 October 2018). Paratypes. MSM-19-292, female, 331 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300���400 m depth, 12 July 2018); MSM-19-293, female, 354 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300��� 400 m depth, 12 July 2018); MSM-19-294, female, 326 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300���400 m depth, 12 July 2018); MSM-19-295, female, 354 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300���400 m depth, 12 July 2018); MSM-19-296, female, 368 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300���400 m depth, 12 July 2018); MSM-19-297, female, 382 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300���400 m depth, 12 July 2018); MSM-19-298, female, 366 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 200���300 m depth, 18 August 2018); MSM-19-299, female, 350 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 200���300 m depth, 18 August 2018); MSM- 19-300, female, 356 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 200���300 m depth, 18 August 2018); MSM-19-301, female, 368 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 200���300 m depth, 18 August 2018); NSMT-P 135961, female, 322 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 400���500 m depth, 12 July 2018); NSMT-P 135962, female, 348 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 500 m depth, 11 July 2018); NSMT-P 135963, male, 384 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); NSMT-P 135964, female, 364 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); NSMT-P 135965, male, 419 mm TL (off the east coast of Torishima Island, Tokyo, Japan, 600���650 m depth, 27 June 2018); NSMT-P 136476, female, 386 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300 m depth, 15 November 2018); NSMT-P 136477, female, 342 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); NSMT-P 136478, male, 346 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); SPMN-PI 45542, female, 370 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300 m depth, 15 November 2018); SPMN-PI 45543, female, 364 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300 m depth, 15 November 2018); SPMN-PI 45544, female, 345 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300 m depth, 15 November 2018); SPMN-PI 45545, female, 372 mm TL (off the east coast of Hachijojima Island, Tokyo, Japan, 300 m depth, 15 November 2018); SPMN-PI 45546, female, 301 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); SPMN-PI 45547, female, 355 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); SPMN-PI 45548, female, 305 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); SPMN-PI 45549, male, 294 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); SPMN-PI 45550, male, 335 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018); SPMN-PI 45551, female, 356 mm TL (off the east coast of Mikurajima Island, Tokyo, Japan, 100���200 m depth, 24 October 2018). Other materials. HUMZ113575, female, 353 mm TL (off Torishima Island, Tokyo, Japan); HUMZ113576, male, 372 mm TL (off Torishima Island, Tokyo, Japan). Diagnosis. A species of Scyliorhinus distinguished by its anterior nasal flaps not reaching the upper lip (vs. flaps reaching upper lip, and sometimes covering it, in S. canicula, S. cervigoni, S. comoroensis, S. duhamelii, S. garmani and S. stellaris); nasoral grooves absent and posterior nasal flaps situated posterior to excurrent apertures (vs. nasoral grooves prexents and posterior nasal flaps laterally situated in S. canicula and S. duhamelii); mouth length less than half of mouth width (vs. mouth length more than or equal half of mouth width except in S. torazame and S. ugoi); anal fin height more than caudal peduncle height (vs. less than caudal peduncle height in S.boa, S. duhamelii, S. torazame and S. torrei), and greater than or equal to half of mouth width (vs. less than half of mouth width in S. boa, S. capensis, S. duhamelii, S. haeckelii, S. hesperius, S.meadi, S. torazame, S. torrei and S. ugoi); saddles darker than the background color (vs. inconspicuous or absent in S. boa, S. cabofriensis, S. cervigoni, S. duhamelii, S. garmani and S. torrei, and dark lines in S. retifer); body grayish brown to dark brown with well-defined light spots and small dark spots (vs. spots absent in S. retifer, yellow to golden spots in S. capensis, light spots absent in S. cervigoni, S. garmani, S. meadi and S. retifer, and dark spots absent in S. capensis, S. comoroensis, S. hesperius, S. meadi, S. torazame and S. torrei); light spots spiracle-sized or larger (vs. predominantly smaller than spiracles in S. boa, S. cabofriensis, S. canicula, S. duhamelii, S. stellaris and S. ugoi); dark spots smaller than spiracles (vs. predominantly larger than spiracles in S. cervigoni, S. duhamelii, S. garmani, S. haeckelii and S. stellaris); number of monospondylous vertebrae 34���36 (vs. counts higher except in S. duhamelii, S. torazame and S. torrei); clasper with hooks (vs. absent in all other species except S. torazame); accessory terminal cartilage present (vs. absent in S. cabofriensis, S. cervigoni, S. comoroensis, S. duhamelii, S. haeckelii, S. stellaris, S. torrei and S. ugoi); egg case surface with irregularities (vs. smooth in all other species). Description. Morphometric measurements are given in Table 1. Body slender and tapering to caudal fin (Fig. 1), precaudal length 77.3 %TL (72.0���76.4 %TL in palatypes). Prepectoral length 0.4 times prepelvic length. Pectoral���pelvic space 1.5 times pelvic���anal space (1.7���2.5 times in female, 1.4���1.9 times in male). Interdorsal space 1.6 times dorsal���caudal space. Trunk shorter than tail; snout���vent length 44.1 %TL (42.0���44.6 %TL). First dorsal origin above insertion of pelvic fin, second dorsal origin forward to anal insertion. No interdorsal, postdorsal, or postanal ridges; lateral crest on caudal peduncle absent. Head (Fig. 2) moderately broad and depressed. Head length (HL) 18.9 %TL (17.2���19.6 %TL) and 1.4 times head width. Snout short, prenasal length 2.3 %TL (2.1���2.6 %TL) and 0.6 times preoral length. Preoral length 3.8 %TL (3.2���4.4 %TL), 0.4 times mouth width and 0.9 times preorbital length. Preorbital length 4.1 %TL (4.1���5.1 %TL), 0.5 times interorbital space. Eyes (Fig. 2A) large and slitlike, eye length 4.1 %TL (3.3���4.4 %TL), 0.2 times head length, with lower edges medial to horizontal head rim in dorsal view; subocular ridge strong. Nictitating lower eyelid of rudimentary type, with shallow subocular pocket and secondary lower eyelid free from upper eyelid. Eye length 5.3 times and interorbital distance 9.6 times spiracle length. Spiracles (Fig. 2A) close behind but clearly separated from eyes, positioned dorsolaterally on head and somewhat lower than level of eye notch. ......Continued on the next page ......Continued on the next page First and second gill openings about equal in height, with last three tapering down to fifth (Fig. 2A, B). First gill opening height 1.6 times that of fifth. All gill openings slightly concave and not elevated on dorsolateral surface of head. Gill filaments not visible externally. Nostrils (Fig. 2C) with broad incurrent apertures, without nasoral grooves or nasal barbels and with small and oval excurrent apertures. Anterior nasal flaps large, triangular, and covering posterior nasal flap and excurrent aperture, extending just anterior to mouth, close to but not touching upper lip. Mesonarial ridge distinct but not exceeding posterior border of anterior nasal flap. Posterior nasal flaps small and rectangular, situated on posterior border of excurrent aperture. Internarial distance 0.3 times interorbital distance. Mouth (Fig. 2A, C) arched, moderately wide, and short. Mouth length 2.9 %TL (2.5���4.4 %TL), 0.3 times mouth width and 0.2 times head length; mouth width 0.7 times head width. Upper labial furrows absent. Lower labial furrows short and narrow, length 0.1 times mouth width. Tooth counts 39/36 (36���49/ 32���43 in paratypes); teeth morphologically similar in both jaws. Monognathic heterodonty gradual and well developed; anterolateral teeth abruptly larger than symphysial ones and smaller distally; posterior teeth with smaller principal cusps than anterolateral teeth (Fig. 3). Sexual heterodonty weak. Symphysial teeth with principal cusp and one weak cusplet on each side. Anterolateral teeth with more developed principal cusp than symphysial teeth and two to four cusplets; most outside cusplets small and undeveloped when with three or four cusplets. Posterior teeth have two or three cusplets and tilted to the outside; principal cusp weakly developed. Pectoral fins (Figs. 1, 4A, B) large and rounded-triangular, not falcate, with broadly convex anterior margins and almost straight posterior and inner margins. Pectoral fin inner corner obtuse angle (96�� in holotype, 90�����102�� in paratypes). Pectoral origin under third gill opening, with base length 0.8 times mouth width. Pectoral anterior margin 2.4 times base length and 1.5 times posterior margin. Pelvic fins (Figs. 1, 4A, B) broadly triangular, with rounded outer corner. Pelvic anterior margin 0.9 times base length. In males, pelvic inner margins fused together forming a pelvic apron, elongated posteriorly and covering claspers. In females, the pelvic fin apex located obviously posterior to middle of fin, and angle of pelvic fin inner corner 63�����75�� in paratypes. Claspers (Fig. 5A) moderately long and cylindrical, extending beyond free rear tips of pelvic fins in adult specimens. Clasper inner length 14.1 %TL (14.1���17.2 %TL in mature males of paratypes), 1.4 times (1.4���1.8 times in mature males of paratypes) pelvic anterior margin and 1.6 times (1.4���1.7 times) clasper outer length. Most of clasper surface (except part of medial surface of clasper shaft and part of exorhipidion, rhipidion, and terminal dermal cover) covered by dermal denticles with anteriorly directed crowns. Clasper hooks present, forming row from behind apopyle to posterior tip of exorhipidion, running along medial border of exorhipidion (Fig. 5A, B). Rhipidion well developed, partly covered medially by prominent exorhipidion and anteriorly by cover rhipidion. Rhipidion insertion point located in anterior portion of dorsal terminal 2 cartilage and extending to end of glans. Cover rhipidion expanded medially, reaching exorhipidion with nearly straight border at the shaft and meandering border at glans. Both cover rhipidion and exorhipidion cover clasper groove. Envelope, pseudopera, and pseudosiphon absent. Terminal dermal cover smooth, and contacting exorhipidion and rhipidion. Clasper skeleton relatively simple (Fig. 5C). Ventral and dorsal terminal cartilage forms spoon-shaped structure; dorsal terminal begins anteriorly but ends together with ventral cartilage. Accessory terminal cartilage quadrangle situated posteriorly and inside ventral marginal cartilage. Dorsal terminal 2 cartilage beginning posteriorly and ventrally to dorsal marginal cartilage, elongated and rod-like, laterally positioned on dorsal terminal cartilage and extending throughout its length. Ventral terminal 2 cartilage elongated and oval, laterally and posteriorly positioned on ventral terminal cartilage and extending to half its length. First dorsal fin (Fig. 1) sub-rectangular and triangular, not falcate, with nearly straight anterior margin, round apex, and angular free rear tip. First dorsal base 0.8 times the interdorsal space. First dorsal anterior margin length 1.5 times its base, and first dorsal height 0.9 times its base. Second dorsal fin (Fig. 1) triangular, not falcate and smaller than first dorsal fin, with nearly straight anterior margin, straight posterior margin, rounded apex, and sharper free rear tip than first dorsal fin. Second dorsal fin base 1.4 times its height and 0.9 times dorsal���caudal distance. Second dorsal anterior margin 1.3 times its base. Second dorsal fin base 0.8 times first dorsal fin base, second dorsal fin height 0.6 times first dorsal fin height. Anal fin (Fig. 1) triangular, slightly high, apically narrow, and not falcate. Anal fin anterior margin nearly straight, apex rounded, free rear tip acutely pointed, and posterior margin straight. Anal fin base 0.8 times interdorsal space and 1.3 times dorsal���caudal distance. Anal fin anterior margin 1.5 times its posterior margin and 1.0 times its base. Anal fin height 0.5 times its base and 1.4 times caudal peduncle height. Anal fin larger than second dorsal fin, base 1.4 times second dorsal fin base, and anal fin height 1.0 times second dorsal fin height. Caudal fin (Fig. 1) narrow-lobed and asymmetrical, with developed terminal lobe. All caudal fin margins nearly straight, all tips rounded. Dorsal caudal margin without lateral undulations and crest of denticles. Dorsal caudal margin 0.3 times precaudal length and 2.1 times preventral caudal margin. Subterminal margin 0.6 times terminal margin. Total vertebral counts 112 (109���115 in paratypes), monospondylous vertebral counts 34 (34���36), and precaudal vertebral counts 77 (71���78). Intestinal valve type conicospiral, with 8 turns (7���8 turns in paratypes). Dermal denticles (Fig. 6) with flat, elongated, teardrop-shaped crowns. Crown with a strong medial ridge extending its entire length onto a long principal cusp. Dermal denticles above pectoral fin present three ridges, with medial ridge less prominent than in denticles of other regions. Denticles below dorsal fins are longer and present a prominent medial ridge, extending to distal tip of cusplet. In all regions, lateral ridges do not extend beyond intersection between principal cusp and cusplets, and are less prominent. Lateral cusps of denticles well developed above pectoral fin, but not well developed below dorsal fins. Color pattern (Figs. 1, 4A, B) with dark brown saddles on a brown background. Light spots present on dorsolateral surfaces and fins, spiracle-sized or larger. Dark spots present on lateral body and fins, smaller than spiracles and light spots. Outlines of dark spots not distinct. Light spots on saddles slightly smaller than on body, with spots largest on fins and smallest on saddles. Saddles darker than background, with pectoral saddles extending diagonally forward. Ventral region white to cream, turning dark anterior of the nostrils. Distribution. This species was recorded from the waters around the Izu Islands, Japan (Fig. 7). All specimens were captured by longline fishing for Splendid alfonsino, at depths of ca. 100���200 m around Mikurajima Island, ca. 200���400 m around Hachijojima Island, and ca. 500���600 m around Torishima Island. Biological data. Males ranged in size from 294 to 416 mm TL (n = 6) and females from 301 to 386 mm TL (n = 22). Size at first maturity is not precisely defined, but males had well-developed claspers at 346 mm TL and females had egg cases at 342 mm. Reproduction is by single oviparity, one egg case for each oviduct. Egg cases are amber in color, with tendrils and surface irregularities that resemble wrinkles (Fig. 8). Maximum case length without tendrils (ML) was 44.6���56.25 mm, and maximum case width was 19.45���22.5 mm (33.4���42.1 %ML) (n = 7). Stomachs were found to contain small crustaceans, small bony fishes, and polychaets. Genetic data. A total of 2164 base pairs (bp) from three mitochondrial DNA regions were examined: 487 bp for the 16S region, 664 bp for COI, and 1013 bp for Cytb. The monophyly of S. hachijoensis was strongly supported by molecular phylogenetic analysis (Fig. 9). The genetic distances between S. hachijoensis and S. torazame were 0.3702 (16S), 0.0295 (COI) and 0.0403 (Cytb), respectively. Etymology. The species name ��� hachijoensis ��� refers to the species��� main collection area, Hachijojima Island. The English name is derived from ���Cinderella���, because the dark spots on the body surface are similar to black ashes ���cinder���. The Japanese name ���Fukami��� means ���deep sea���. Remarks. Clasper hooks are a unique character observed in S. torazame (Soares & de Carvalho 2019, 2020), and the same character was also observed in S. hachijoensis. Both male and female S. hachijoensis can be easily distinguished from eleven other Scyliorhinus species by color pattern: S. cervigoni Maurin & Bonnet, 1970, S. garmani (Fowler, 1934), S. meadi Springer, 1966 and S. retifer (Garman, 1881) have no light spots, S. capensis (M��ller & Henle, 1838) has light yellow to golden spots, and S. torrei Howell-Rivero, 1936 has beige to cream spots. Scyliorhinus canicula, S. cabofriensis Soares, Gomes & de Carvalho, 2016, S. cervigoni, S. garmani, S. haeckelii (Miranda Ribeiro, 1907) and S. stellaris (Linnaeus, 1758) have conspicuous dark spots, and S. boa (Goobe & Bean, 1896) has various dark spots (Springer 1979; Ebert et al. 2013a; Soares & de Carvalho 2019). Scyliorhinus comoroensis Compagno, 1988 and S. duhamelii (Garman, 1913) (as well as S. canicula, S. cervigoni, S. garmani and S. stellaris, which are also listed above) can be distinguished from S. hachijoensis because the nasal flaps reach or cover the upper lip (Springer 1979; Ebert et al. 2013a; Soares & de Carvalho 2019). Scyliorhinus hesperius Springer, 1966 and S. ugoi Soares, Gomes & Gadig, 2015 are very similar to S. hachijoensis in color pattern, but differ in the number of monospondylous vertebrae and size of maturity. The number of monospondylous vertebrae is 39���42 in S. hesperius and 38���39 in S. ugoi. Adult males mature at least 420 mm TL in S. hesperius, and by at least 450 mm TL and 470 mm TL in male and female specimens of S. ugoi, respectively. By contrast, the number of monospondylous ve, Published as part of Ito, Nanami, Fujii, Miho, Nohara, Kenji & Tanaka, Sho, 2022, Scyliorhinus hachijoensis, a new species of catshark from the Izu Islands, Japan (Carcharhiniformes: Scyliorhinidae), pp. 331-349 in Zootaxa 5092 (3) on pages 333-345, DOI: 10.11646/zootaxa.5092.3.5, http://zenodo.org/record/5881301, {"references":["Hagiwara, S. (1993) Keeping and Reproduction of Chondrichthyans in Captivity at Shimoda Floating Aquarium. Report of Japanese Society for Elasmobranch Studies, 30, 1 - 18.","Soares, K. D. A. & de Carvalho, M. R. (2019) The catshark genus Scyliorhinus (Chondrichthyes: Carcharhiniformes: Scyliorhinidae): taxonomy, morphology and distribution. Zootaxa, 4601 (1), 1 - 147. https: // doi. org / 10.11646 / zootaxa. 4601.1.1","Springer, S. (1979) A revision of the catsharks, family Scyliorhinidae. NOAA technical report NMFS circular, 422, 1 - 97. https: // doi. org / 10.5962 / bhl. title. 63029","Ebert, D. A., Fowler, S. & Compagno, L. (2013 a) Sharks of the World: A Fully Illustrated Guide. Wild Nature Press, Plymouth, 528 pp.","Compagno, L. J. V. (1988) Sharks of the Order Carcharhiniformes. Princeton University Press, Princeton, New Jersey, 542 pp.","Soares, K. D. A., Gadig, O. F. B. & Gomes, U. L. (2015) Scyliorhinus ugoi, a new species of catshark from Brazil (Chondrichthyes: Carcharhiniformes: Scyliorhinidae). Zootaxa, 3937 (2), 347 - 361. https: // doi. org / 10.11646 / zootaxa. 3937.2.6","Carpenter, K. E. (1998) FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Vol. 2. Cephalopods, crustaceans, holothurians and sharks. FAO, Rome, 710 pp. [pp. 687 - 1396]"]}

Poroderma pantherinum (Smith, in Müller & Henle, 1838c) Leopard Catshark Scyllium pantherinum Smith, in M̹ller & Henle, 1838c: 13. Syntypes: BMNH 1845.7.3.145 (stuffed); RMNH uncatalogued (1, lost); UTZI uncatalogued (1, lost). According to Fricke et al. (2020) appeared first as name only in Smith, 1838: 85. Local synonymy: Poroderma pantherinum: Smith, 1837: 85 (name only); Smith, in M̹ller & Henle, 1838c: 13; Garman, 1913: 70; Fowler, 1925b: 189; Fowler, 1941: 37; Smith 1949a: 53, fig. 33; Smith, 1965: 53, fig. 33; Bass et al., 1975a: 30, fig. 18; Springer, 1979: 114; Compagno, 1984b: 349, fig.; Bass, 1986: 95, fig. 11.15, pl. 3; Compagno, 1988a: 117; Compagno et al., 1989: 48, pl.; Compagno, 1999: 119; Heemstra & Heemstra, 2004: 67; Compagno et al., 2005: 243, fig., pl. 39; Human, 2006a: 6, fig. 3; Ebert et al., 2013 a: 370, fig., pl. 50; Mann, 2013: 187; NPOA, 2013: 47; da Silva et al., 2015: 248; Ebert & van Hees, 2015: 147; Weigmann, 2016: 878. Poroderma submaculatum Smith, 1837: 85 (name only). Poroderma variegatum Smith, 1837: 85 (name only). Scyllium leopardinum M̹ller & Henle, 1838c: 13 (Fowler, 1941: 38 attributes this name to Van Horst in M̹ller & Henle, although no reference to Van Horst is given in M̹ller & Henle). Scyllium maeandrinum von Rapp in M̹ller & Henle, 1838c: 13. Scyllium variegatum Smith in M̹ller & Henle, 1838c: 14; Smith, 1849: 11, fig. 3, pl. 25; Duméril 1853: 83; Bleeker 1878: 1, 8; Sauvage 1891: 511. Scyllium pantherinum: Smith in M̹ller & Henle, 1838c: 13; Smith 1849: 11, fig. 2, pl. 25; Gray, 1851: 31; Bleeker, 1860: 57; Duméril, 1865: 322; Gilchrist, 1902: 164. Scyllium africanum: G̹nther, 1870: 405; Lampe, 1914: 209. Scyllium africanum var. pantherina: G̹nther, 1870: 405; Lampe, 1914: 212. Scyllium africanum var. striata: G̹nther, 1870: 405; Lampe, 1914: 212. Scyllium africanum var. variegata: G̹nther, 1870: 405; Lampe, 1914: 212. Scyliorhinus pantherinus: Regan, 1908b: 456. Scyllium africanum var. punctata Lampe, 1914: 212. Scylliorhinus pantherinus: Smith, 1849: 11, fig. 2, pl. 25; Thompson, 1914: 138; Gilchrist & Thompson, 1916: 270; Gilchrist, 1922a: 4; Barnard, 1925: 40, fig. 5, pl. 2; Barnard, 1927: 1013. Scyliorhinus regani: Fowler, 1925b: 188, fig. 1; Fowler, 1926: 32, fig. 3, not Holohalaelurus regani (Gilchrist, 1922b). Poroderma marleyi Fowler, 1934a: 234 (original description); Fowler, 1941: 38; Bigelow & Schroeder, 1948b: 197; Smith, 1949a: 53, pl. 2; Springer & Garrick, 1964: 86; Smith, 1965: 53, pl. 2; Bass et al., 1975a: 29, fig. 17; Springer, 1979: 114; Compagno, 1984b: 348, fig.; Bass, 1986: 94, fig. 11.14; Compagno, 1988a: 118; Human, 2006a: fig. 5. Scyliorhinus leopardus: Fowler, 1935: 361. Conoporoderma pantherinum: Bigelow & Schroeder, 1948b: 197. South Africa voucher material: SAIAB 5910, SAIAB 6000, SAIAB 6001, SAIAB 6002, SAIAB 10273, SAIAB 10372, SAIAB 10737, SAIAB 11960, SAIAB 12016, SAIAB 13130, SAIAB 16728, SAIAB 17326, SAIAB 17783, SAIAB 18179, SAIAB 19992, SAIAB 25181, SAIAB 25214, SAIAB 25215, SAIAB 25216, SAIAB 25217, SAIAB 25336, SAIAB 25921, SAIAB 25921, SAIAB 25928, SAIAB 26283, SAIAB 26284, SAIAB 26285, SAIAB 26286, SAIAB 26287, SAIAB 26288, SAIAB 26289, SAIAB 26441, SAIAB 26442, SAIAB 26443, SAIAB 27206, SAIAB 27207, SAIAB 27649, SAIAB 34577, SAIAB 37057, SAIAB 39990, SAIAB 48495, SAIAB 53684, SAIAB 53685, SAIAB 189034. South African distribution: Endemic. Saldanha Bay (WC) to Durban (KZN), but most common on the south and southeast coasts (Human, 2006a). Remarks: Fowler (1934a) described a new Poroderma species (P. marleyi) based on an extreme color morph from off Durban. However, examination of additional color morphs throughout its range reveals this species to be a junior synonym of P. pantherinum (Human, 2006a). Furthermore, it appears that “marleyi” color morph occurs at the extreme ends of its range, both in the Saldaha (WC) and KZN areas (D.A. Ebert, unpubl. data) Conservation status: LC (2020). Genus Scyliorhinus Blainville, 1816 Scyliorhinus (subgenus of Squalus) Blainville, 1816: 121. Type species: Scyliorhinus canicula (Linnaeus, 1758). Type by subsequent designation., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on pages 53-54, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["M ʾ ller, J. & Henle, F. G. J. (1838 c) s. n. In: Systematische Beschreibung der Plagiostomen. Plagiostomen, Berlin, pp. 1 - 28.","Fricke, R., Eschmeyer, W. N. & Van der Laan, R. (Eds.) (2020) Eschmeyer's catalog of fishes: genera, species, and references. Available from: http: // researcharchive. calacademy. org / research / ichthyology / catalog / fishcatmain. asp (accessed 30 June 2020)","Smith, A. (1838) (On the necessity for a revision of the groups included in the Linnean genus Squalus). Proceedings of the Zoological Society of London, 5 (57), 85 - 86.","Smith, A. (1837) (On the necessity for a revision of the groups included in the Linnean genus Squalus). Annals of Natural History, 1 (1), 72 - 74.","Garman, S. (1913) The Plagiostomia (sharks, skates, and rays). Memoirs of the Museum of Comparative Zoology, Harvard College, 36, 1 - 515.","Fowler, H. W. (1925 b) Fishes from Natal, Zululand and Portuguese East Africa. Proceedings of the Academy of Natural Sciences, Philadelphia, 1925, 77, 187 - 268.","Fowler, H. W. (1941) The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostarophysi obtained by the U. S. Bureau of Fisheries streamer \" Albatross \" in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bulletin United States National Museum, 100, 1 - 879.","Smith, J. L. B. (1949 a) The Sea Fishes of Southern Africa. South Africa Central News Agency Ltd., 550 pp.","Smith, J. L. B. (1965) The Sea Fishes of Southern Africa. 5 th Edition. Central News Agency Ltd., 580 pp.","Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1975 a) Sharks of the east coast of southern Africa. II. The families Scyliorhinidae and Pseudotriakidae. Investigational Report. Oceanographic Research Institute, Durban, 37, 1 - 64.","Springer, S. (1979) A revision of the catsharks, family Scyliorhinidae. NOAA (National Oceanic and Atmospheric Administration) Technical Report NMFS (National Marine Fisheries Service) Circular, 422, 1 - 152. https: // doi. org / 10.5962 / bhl. title. 63029","Compagno, L. J. V. (1984 b) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 2. Carcharhiniformes. FAO, Rome, pp. 251 - 655.","Bass, A. J. (1986) Families Chlamydoselachidae, Heterodontidae, Orectolobidae, Rhinicodontidae, Scyliorhinidae, Pseudotriakidae, Sphyrnidae, Lamnidae, Cetorhinidae, Alopiidae, Pseudocarchariidae, Squatinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 47 - 48 + 64 - 66 + 87 - 102 + 103 + 107.","Compagno, L. J. V. (1988 a) Sharks of the Order Carcharhiniformes. Princeton University Press, Princeton, New Jersey, 486 pp.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Heemstra, P. C. & Heemstra, E. (2004) Coastal Fishes of Southern Africa. National Inquiry Service Centre and South African Institute for Aquatic Biodiversity, Grahamstown, 488 pp.","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Human, B. A. (2006 a) A taxonomic revision of the catshark genus Poroderma Smith, 1837 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae). Zootaxa, 1229 (1), 1 - 32. https: // doi. org / 10.11646 / zootaxa. 1229.1.1","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Mann, B. Q. (2013) Southern African marine linefish species profiles. Oceanographic Research Institute, Special Publication, 9, 1 - 343.","NPOA. (2013) National Plan of Action for the Conservation and Management of Sharks (NPOA-Sharks). Department of Agriculture, Forestry and Fisheries (DAFF), Rogge Bay, Cape Town, 63 pp.","da Silva C., Booth, A. J., Dudley, S. F. J., Kerwath, S. E., Lamberth, S. J., Leslie, R. W., McCord, M. E., Sauer, W. H. H. & Zweig, T. (2015) A description and updated overview of the status and management of South Africa's chondrichthyan fisheries. South African Journal of Marine Science, 37, 233 - 248. https: // doi. org / 10.2989 / 1814232 X. 2015.1044471","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Smith, A. (1849) Pisces. In: Illustrations of the Zoology of South Africa. Vol. 4. Smith, Elder, and Co., London, 77 pp.","Dumeril, A. H. A. (1853) Monographie de la tribu des Scylliens ou Roussettes (poissons plagiostomes) comprenant deux especes nouvelles. Revue et Magasin de Zoologie, Series 2, 5, 8 - 25 + 73 - 87 + 119 - 130.","Gray, J. E. (1851) List of the Specimens of Fish in the Collection of the British Museum. Part I. Chondropterygii. British Museum (Natural History), London, 160 pp.","Bleeker, P. (1860 a) Elfde bijdrage tot de kennis der vischfauna van Amboina. Acta Societatis Regiae Scientiarum Indo-Neerlandicae, 8 (5), 1 - 14. https: // doi. org / 10.5962 / bhl. title. 144153","Dumeril, A. H. A. (1865) Histoire naturelle des poissons ou ichthyologie generale. Tome Premier. I. Elasmobranches. Plagiostomes et Holocephales ou Chimeres. Vol. 1. Librarie Encylopedique de Roret, Paris, 720 pp. https: // doi. org / 10.5962 / bhl. title. 2111","Gilchrist, J. D. F. (1902) Catalogue of fishes recorded from South Africa. Cape of Good Hope, Department of Agriculture, Marine Investigations in South Africa, 1, 97 - 179.","Lampe, M. 1914. Die fische der Deutschen S ʾ dpolar-Expedition 1901 - 1903. 3. Die hochsee- und k ʾ stenfische. Deutsch S ̡ dpolar-Expedition, 15 (2), 203 - 256. https: // doi. org / 10.1071 / 9780643109148","Regan, C. T. (1908 b) A synopsis of the sharks of the Family Scyliorhinidae. Annals and Magazine of Natural History, Series 8, 1, 453 - 465. https: // doi. org / 10.1080 / 00222930808692434","Thompson, W. W. (1914) Catalogue of the fishes of the Cape Province (Pt 1). Marine Biological Report, Cape Town, 2 (8), 132 - 167.","Gilchrist, J. D. F. & Thompson, W. W. (1916) A catalogue of the sea fishes recorded from Natal. Part I. Annals of the Durban Museum, 1 (3), 255 - 290.","Gilchrist, J. D. F. (1922 a) Fisheries and Marine Biological Survey. Report no. 2 for the Year 1921. Annexure A. List of Fishes, etc., procured; Annexure B., List of stations of SS \" Pickle \"; Annexure C., Journal of SS \" Pickle \". Report of the Fisheries and Marine Biological Survey, Union of South Africa, 2, 1 - 85.","Barnard, K. H. (1925) A monograph of the marine fishes of South Africa. Part I (Amphioxus, Cyclostomata, Elasmobranchii, and Teleostei-Isospondyli to Heterosomata). Annals of the South African Museum, 21, 1 - 418.","Barnard, K. H. (1927) A monograph of the marine fishes of South Africa. Part II (Teleostei - Discocephali to end. Appendix). Annals of the South African Museum 21, 419 - 1065.","Fowler, H. W. (1926) Descriptions of three new fishes from the Natal coast. Annals of the Natal Government Museum, 5 (3), 399 - 402.","Gilchrist, J. D. F. (1922 b) Deep-sea fishes procured by the S. A. \" Pickle \" (Part I.). Special Report (3), Report of the Fisheries and Marine Biological Survey, Union of South Africa, 2, 41 - 79.","Fowler, H. W. (1934 a) Descriptions of new fishes obtained 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Proceedings of the Academy of Natural Sciences of Philadelphia, 85 (for 1933), 233 - 367.","Bigelow, H. B. & Schroeder, W. C. (1948 b) Sharks. Fishes of the Western North Atlantic Part 1. Memoirs of the Sears Foundation for Marine Research, Series 1, 1, 56 - 576.","Garrick, J. A. F. & Springer, S. (1964) Isistius plutodus, a new squaloid shark from the Gulf of Mexico. Copeia, 1964 (4), 678 - 682. https: // doi. org / 10.2307 / 1441443","Fowler, H. W. (1935) South African fishes received from Mr. H. W. Bell-Marley in 1935. Proceedings of the Academy of Natural Sciences of Philadelphia, 87, 361 - 408.","Blainville, H. de (1816) Prodrome d'une nouvelle distribution systematique du regne animal. Bulletin de la Societe Philomatique, Paris, 8, 105 - 112, 121 - 124.","Linnaeus, C. (1758) Systema Naturae, Ed. X. (Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Impensis Direct. Laurentii Salvii, Holmiae, 824 pp. https: // doi. org / 10.5962 / bhl. title. 542"]}

Scyliorhinus capensis (Müller & Henle, 1838c) Yellowspotted Catshark Scyllium capense M̹ller & Henle, 1838c: 11. Lectotype: BMNH 1845.7.3.141 (lectotype designated by Soares & de Carvalho, 2019). Type locality: Cape of Good Hope, Western Cape Province, South Africa. Local synonymy: Scyllium capense: Smith, 1837: 85 (name only); M̹ller & Henle, 1838c: 11; Gray, 1851: 31; Bleeker, 1860b: 57; Duméril, 1865: 320; G̹nther, 1870: 404; Gilchrist, 1902: 165 (listed, “ Cape Seas”). Scyliorhinus capensis: Regan, 1908b: 458; Fowler, 1941: 35; Smith, 1949a: 54, fig. 38, pl. 2; Smith, 1965: 54, fig. 38, pl. 2; Bass et al., 1975a: 32, fig. 19; Springer, 1979: 132, fig. 84; Compagno, 1984b: 359, fig.; Bass, 1986: 95, figs. 11, 16, pl. 3; Compagno, 1988a: 122; Compagno, 1988b: 606, figs. 2, 6b, 7b, 8c–d (compared to S. comoroensis); Compagno et al., 1989: 46, pl.; Compagno et al., 1991: 83; Ebert et al., 1996: 236; Compagno, 1999: 119; Compagno et al., 2005: 248, fig., pl. 41; Ebert et al., 2006: 1053; Ebert, 2013: 186, fig. 250; Ebert et al., 2013 a: 379, fig., pl. 51; Ebert & Mostarda, 2013: 66, fig.; Ebert, 2015: 153, fig. 167; Ebert & Mostarda, 2015: 48, fig.; NPOA, 2013: 47; da Silva et al., 2015: 248; Ebert & van Hees, 2015: 147; Weigmann, 2016: 879 (listed); Soares & de Carvalho, 2019: 32. Catulus capensis: Garman, 1913: 74. Scylliorhinus capensis: Thompson, 1914: 137; Gilchrist, 1921: 71 (listed, Cape of Good Hope); Gilchrist, 1922b: 45; von Bonde, 1923: 5; Barnard, 1925: 40, fig. 8b; von Bonde, 1934: 15; Barnard, 1947: 16, fig. 1, pl. 3. Scyliorhinus (Scyliorhinus) capensis: Norman, 1935: 36. Haploblepharus capensis: White, 1937: 121. South Africa voucher material: Lectotype: BMNH 1845.7.3.141. Paralectotypes: BMNH 1845.7.3.144; BMNH 1953.5.10.2. Soares & de Carvalho (2019) lists 81 additional institutional specimens in appendix. South African distribution: Most common in the west from the Orange River (NC) to Cape Agulhas (WC), but does extend at least to Waterloo Bay (EC) (Soares & de Carvalho, 2019) where it is uncommon, and with a record of one specimen from KZN (Bass et al., 1975a; Compagno et al., 1989). Remarks: Scyliorhinus capensis is a near endemic to South Africa, although there is at least one record of this species from southwest of L̹deritz, southern Namibia (Compagno et al., 1991). Conservation status: NT (2020)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 54, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["M ʾ ller, J. & Henle, F. G. J. (1838 c) s. n. In: Systematische Beschreibung der Plagiostomen. Plagiostomen, Berlin, pp. 1 - 28.","Soares, K. D. A. & Carvalho, M. R. de (2019) The catshark genus Scyliorhinus (Chondrichthyes: Carcharhiniformes: Scyliorhinidae): taxonomy, morphology and distribution. Zootaxa, 4601 (1), 1 - 147. https: // doi. org / 10.11646 / zootaxa. 4601.1.1","Smith, A. (1837) (On the necessity for a revision of the groups included in the Linnean genus Squalus). Annals of Natural History, 1 (1), 72 - 74.","Gray, J. E. (1851) List of the Specimens of Fish in the Collection of the British Museum. Part I. Chondropterygii. British Museum (Natural History), London, 160 pp.","Bleeker, P. (1860 b) Zesde bijdrage tot de kennis der vischfauna van Japan. Acta Societatis Regiae Scientiarum Indo-Neerlandicae, 8 (1), 1 - 104. https: // doi. org / 10.5962 / bhl. title. 144153","Dumeril, A. H. A. (1865) Histoire naturelle des poissons ou ichthyologie generale. Tome Premier. I. Elasmobranches. Plagiostomes et Holocephales ou Chimeres. Vol. 1. Librarie Encylopedique de Roret, Paris, 720 pp. https: // doi. org / 10.5962 / bhl. title. 2111","Gilchrist, J. D. F. (1902) Catalogue of fishes recorded from South Africa. Cape of Good Hope, Department of Agriculture, Marine Investigations in South Africa, 1, 97 - 179.","Regan, C. T. (1908 b) A synopsis of the sharks of the Family Scyliorhinidae. Annals and Magazine of Natural History, Series 8, 1, 453 - 465. https: // doi. org / 10.1080 / 00222930808692434","Fowler, H. W. (1941) The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostarophysi obtained by the U. S. Bureau of Fisheries streamer \" Albatross \" in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bulletin United States National Museum, 100, 1 - 879.","Smith, J. L. B. (1949 a) The Sea Fishes of Southern Africa. South Africa Central News Agency Ltd., 550 pp.","Smith, J. L. B. (1965) The Sea Fishes of Southern Africa. 5 th Edition. Central News Agency Ltd., 580 pp.","Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1975 a) Sharks of the east coast of southern Africa. II. The families Scyliorhinidae and Pseudotriakidae. Investigational Report. Oceanographic Research Institute, Durban, 37, 1 - 64.","Springer, S. (1979) A revision of the catsharks, family Scyliorhinidae. NOAA (National Oceanic and Atmospheric Administration) Technical Report NMFS (National Marine Fisheries Service) Circular, 422, 1 - 152. https: // doi. org / 10.5962 / bhl. title. 63029","Compagno, L. J. V. (1984 b) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 2. Carcharhiniformes. FAO, Rome, pp. 251 - 655.","Bass, A. J. (1986) Families Chlamydoselachidae, Heterodontidae, Orectolobidae, Rhinicodontidae, Scyliorhinidae, Pseudotriakidae, Sphyrnidae, Lamnidae, Cetorhinidae, Alopiidae, Pseudocarchariidae, Squatinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 47 - 48 + 64 - 66 + 87 - 102 + 103 + 107.","Compagno, L. J. V. (1988 a) Sharks of the Order Carcharhiniformes. Princeton University Press, Princeton, New Jersey, 486 pp.","Compagno, L. J. V. (1988 b) Scyliorhinus comoroensis sp. n., a new catshark from the Comoro Islands, western Indian Ocean (Carcharhiniformes, Scyliorhinidae). Bulletin du Museum National d'Histoire Naturelle, (Serie 4), Section A: Zoologie Biologie et Ecologie Animales, 10 (3), 603 - 625.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V., Ebert, D. A. & Cowley, P. D. (1991) Distribution of offshore demersal cartilaginous fishes (class Chondrichthyes) of the west coast of southern Africa, with notes on their systematics. South African Journal of Marine Science, 11, 43 - 139. https: // doi. org / 10.2989 / 025776191784287664","Ebert, D. A., Cowley, P. D. & Compagno, L. J. V. (1996) A preliminary investigation of the feeding ecology of catsharks (Scyliorhinidae) off the west coast of southern Africa. South African Journal of Marine Science, 17, 233 - 240. https: // doi. org / 10.2989 / 025776196784158563","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Ebert, D. A., Compagno, L. J. V. & Cowley, P. D. (2006) Reproductive biology of catsharks (Chondrichthyes: Scyliorhinidae) from off the west coast of southern Africa. ICES Journal of Marine Science, 63, 1053 - 1065. https: // doi. org / 10.1016 / j. icesjms. 2006.04.016","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. & Mostarda, E. (2013) Identification Guide to the Deep-sea Cartilaginous Fishes of the Indian Ocean. FishFinder Programme, FAO, Rome, 76 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & Mostarda, E. (2015) Identification Guide to the Deep-sea Cartilaginous Fishes of the Southeastern Atlantic Ocean. FishFinder Programme, FAO, Rome, 70 pp.","NPOA. (2013) National Plan of Action for the Conservation and Management of Sharks (NPOA-Sharks). Department of Agriculture, Forestry and Fisheries (DAFF), Rogge Bay, Cape Town, 63 pp.","da Silva C., Booth, A. J., Dudley, S. F. J., Kerwath, S. E., Lamberth, S. J., Leslie, R. W., McCord, M. E., Sauer, W. H. H. & Zweig, T. (2015) A description and updated overview of the status and management of South Africa's chondrichthyan fisheries. South African Journal of Marine Science, 37, 233 - 248. https: // doi. org / 10.2989 / 1814232 X. 2015.1044471","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Garman, S. (1913) The Plagiostomia (sharks, skates, and rays). Memoirs of the Museum of Comparative Zoology, Harvard College, 36, 1 - 515.","Thompson, W. W. (1914) Catalogue of the fishes of the Cape Province (Pt 1). Marine Biological Report, Cape Town, 2 (8), 132 - 167.","Gilchrist, J. D. F. (1921) Fisheries and Marine Biological Survey. Report no. 1 for the Year 1920. Annexure A. List of Fishes, etc., procured; Annexure B., List of stations of SS \" Pickle \"; Annexure C., Journal of SS \" Pickle \". Report of the Fisheries and Marine Biological Survey, Union of South Africa, 1, I - Ill.","Gilchrist, J. D. F. (1922 b) Deep-sea fishes procured by the S. A. \" Pickle \" (Part I.). Special Report (3), Report of the Fisheries and Marine Biological Survey, Union of South Africa, 2, 41 - 79.","von Bonde, C. (1923) Shallow-water fishes procured by the S. S. \" Pickle \". Special Report 1 in Report on the Fisheries and Marine Biological Survey, Union of South Africa, 3, 1 - 40.","Barnard, K. H. (1925) A monograph of the marine fishes of South Africa. Part I (Amphioxus, Cyclostomata, Elasmobranchii, and Teleostei-Isospondyli to Heterosomata). Annals of the South African Museum, 21, 1 - 418.","von Bonde, C. (1934) Shark fishing as an industry. Union of South Africa, Department of Commerce and Industry, Fisheries and Marine Biological Survey, Investigational Report, Series 2, 1 - 19.","Barnard, K. H. (1947) A Pictorial Guide to South African Fishes. Marine and Freshwater. Maskew Miller Ltd, Cape Town, 226 pp.","Norman, J. R. (1935) Coast fishes, Part. I. The South Atlantic. ' Discovery' Reports, 12, 1 - 58.","White, E. G. (1937) Interrelationships of the elasmobranchs with a key to the order Galea. Bulletin of the American Museum of Natural History, 74 (2), 25 - 138."]}

Scyliorhinus canicula (Linnaeus 1758) ���Small-spotted catshark Taxonomy. First record from Lebanon as Scyllium caninula, Cuv. by Gruvel (1931: 73). The species was never recorded afterwards. Distribution. Western Baltic Sea, North Sea, western Mediterranean Sea, northeastern Atlantic: Shetland Islands and southern Norway south to Senegal. Conservation. IUCN: Global (LC: 1 December 2008); Med. (LC: 25 March 2016). Capture and threats: FIT, FIB, unknown. Occurrence: Unknown. Remarks. Gruvel (1931: 74) mentioned that the species, along with Scyliorhinus stellaris (Linnaeus 1758), were abundant and hated by fishermen., Published as part of Bariche, Michel & Fricke, Ronald, 2020, The marine ichthyofauna of Lebanon: an annotated checklist, history, biogeography, and conservation status, pp. 1-157 in Zootaxa 4775 (1) on page 12, DOI: 10.11646/zootaxa.4775.1.1, http://zenodo.org/record/3983887, {"references":["Gruvel, A. (1931) Les Etats de Syrie. Richesses marines et fluviales. Exploitation actuelle-Avenir. Societe d'Editions Geographiques, Maritimes et Coloniale, 453 pp."]}

Scyliorhinus stellaris (Linnaeus 1758) ���Nursehound Taxonomy. First record from Lebanon as Scyllium catulus, Cuv. by Gruvel (1931: 73). The species was never recorded afterwards. Distribution. North Sea, Mediterranean Sea, eastern North Atlantic: Shetland Islands and southern Norway to Morocco. Conservation. IUCN: Global (NT: 14 February 2006); Med. (NT: 25 March 2016). Capture and threats: FIT, FIB, unknown. Occurrence: Unknown. Remarks. Gruvel (1931: 74) mentioned that the species, along with Scyliorhinus canicula (Linnaeus 1758) were abundant and hated by fishermen., Published as part of Bariche, Michel & Fricke, Ronald, 2020, The marine ichthyofauna of Lebanon: an annotated checklist, history, biogeography, and conservation status, pp. 1-157 in Zootaxa 4775 (1) on page 12, DOI: 10.11646/zootaxa.4775.1.1, http://zenodo.org/record/3983887, {"references":["Gruvel, A. (1931) Les Etats de Syrie. Richesses marines et fluviales. Exploitation actuelle-Avenir. Societe d'Editions Geographiques, Maritimes et Coloniale, 453 pp."]}

Intersexual differences in morphometrics were investigated in five species of the catshark genus Scyliorhinus. ANCOVA was used to test 59 measurements, considering capture location and total length as covariates. In all examined species, pelvic-anal distances and pelvic-fin inner margin lengths were greater in males than in females, representing a clear pattern for the genus.

The catshark genus Scyliorhinus belongs to the family Scyliorhinidae, the most diverse family of sharks, and currently presents 16 valid species according to most modern accounts. The long history of taxonomic rearrangements and inaccurate descriptions of many species have contributed to misidentification of specimens and lack of information on the distributional range and diagnostic characters of its species. Species of Scyliorhinus are reviewed and redescribed here, with detailed descriptions on external morphology, neurocranium, claspers, dermal denticles, and tooth morphology provided for the first time for all species. Sixteen species are recognized as valid: Scyliorhinus boa (Goode & Bean, 1896), S. cabofriensis Soares, Gomes & de Carvalho, 2016, S. canicula (Linnaeus, 1758), S. capensis (Müller & Henle, 1838), S. cervigoni Maurin & Bonnet, 1970, S. comoroensis Compagno, 1988, S. duhamelii (Garman, 1913), S. garmani (Fowler, 1934), S. haeckelii (Miranda Ribeiro, 1907), S. hesperius Springer, 1966, S. meadi Springer, 1966, S. retifer (Garman, 1881), S. stellaris (Linnaeus, 1758), S. torazame (Tanaka, 1908), S. torrei Howell-Rivero, 1936, and S. ugoi Soares, Gomes & Gadig, 2015. The main taxonomic decisions herein taken include the resurrection of S. duhamelii (previously a junior synonym of S. canicula) and the synonimization of S. tokubee with S. torazame. Information on geographic distribution was updated for most species, especially for those with wide ranges (S. canicula, S. haeckelii, S. retifer, and S. stellaris).

Scyliorhinus ugoi Soares, Gomes & Gadig, 2015 (Figs. 10C, 11) Common names: cação-gato negrinho (Brazil), dark freckled catshark (United States). Scyliohinus sp.: Gomes et al., 2010: 85–86, fig. 110 (catalogue, Rio de Janeiro, Brazil). Scyliorhinus hesperius: Gadig & Gomes, 2003: 22 (catalogue, Brazil). Scyliorhinus ugoi Soares, Gomes & Gadig, 2015: 348 –361, figs. 1-9 (original description, type locality: Bahia, Brazil); Weigmann, 2016: 44 (listed); Rincón et al., 2017: 94 –95, fig. 4j (catalogue, Brazil). Holotype. MNRJ 42619, female, 496 mm TL (southern Bahia, Brazil). Paratypes. MZUSP 110448, male, 465 mm TL (Alagoas, northeastern Brazil, 9°S 34°50’W); MZUSP 110449, male, 445 mm TL (Rio Grande do Norte, northeastern Brazil, 6°14’S 34°51’W); UERJ 1725, male, 530 mm TL (somewhere between southern Bahia and northern Rio de Janeiro; neurocranium, jaws and claspers); UERJ 2179, male, 415 mm TL (southern Bahia, Brazil) . Additional material examined. 21 specimens (see Appendix). Diagnosis. Scyliorhinus ugoi differs from all congeners by presenting a color pattern composed of saddles with antero- and posteromedial projections (vs. projections absent in other species); dark spots spiracle-sized to greater than the spiracles (vs. dark spots absent in S. capensis, S. comoroensis, S. hesperius, S. meadi, S. torazame, and S. torrei; reticulate pattern in S. retifer; spots predominantly smaller in S. boa and S. cabofriensis), spots restricted to saddles and not bordering them (vs. spots over entire body in S. cabofriensis; bordering saddles in S. boa); adult males at least 445 mm TL and adult females at least 500 mm TL (vs. adult sizes smaller than 420 mm TL in S. boa, S. cabofriensis, S. haeckelii, S. torazame, and S. torrei; sizes greater than 500 mm TL in S. capensis, S. cervigoni, S. meadi, and S. stellaris). The following combination of characters, although less conspicuous, also helps distinguish this species: lunate spots and double points absent (vs. present in S. cabofriensis and S. haeckelii); anterior nasal flaps not reaching the upper lip (vs. flaps reaching the lip, sometimes covering it in S. canicula, S. cervigoni, S. comoroensis, S. duhamelii, S. garmani, and S. stellaris); mesonarial ridge not exceeding the posterior border of anterior nasal flaps (vs. exceeding it in S. stellaris); pelvic apron extending to 2/3 the length of pelvic inner margins (vs. extending through almost entire length of pelvic margins in S. canicula, S. capensis, S. duhamelii, S. torazame, and S. torrei); clasper with cover rhipidion covered by dermal denticles (vs. absent in S. boa, S. cervigoni, S. hesperius, and S. retifer); terminal dermal cover smooth (vs. rough in S. canicula and S. capensis); terminal 3 cartilage absent (vs. present in S. boa, S. canicula, S. capensis, S. retifer, and S. torazame); dorsal terminal 2 cartilage reduced and subtriangular (vs. elongated in S. boa, S. canicula, S. comoroensis, S. duhamelii, S. retifer, S. stellaris, S. torazame, and S. torrei); counts of monospondylous vertebrae 38–39 (vs. 46 in S. capensis; 40–45 in S. cervigoni; 48 in S. garmani; 46–48 in S. meadi; 43–47 in S. stellaris; 30–35 in S. torrei). Etymology. The specific name ‘ugoi’ was dedicated to Ugo de Luna Gomes, son of the elasmobranch researcher Ulisses Leite Gomes. Remarks. Soares et al. (2015) described the clasper of S. ugoi with an accessory dorsal marginal cartilage (RD2), which would support the rhipidion (p. 352, 353, 356, figs. 5, 6). In this study, this structure is reidentified as a dorsal terminal 2 cartilage (Fig. 8C), following Jungersen (1899) and Compagno (1988a). Some specimens captured in Pernambuco and Rio Grande do Norte, Brazil (UERJ 2178, UFPB 5294 and other uncatalogued specimens), present a color pattern distinct with grayish background color, dark spots smaller than the spiracles throughout the body and light spots less frequent inside the saddles. These specimens are tentatively classified as S. ugoi and we highlight the need to obtain more specimens for a more detailed study of these populations. Specimens USNM 221611 and USNM 221652, USNM 221561 and, UF 77857, previously identified as Scyliorhinus boa, S. hesperius and S. retifer, respectively, present a color pattern similar to S. ugoi with antero- and posteromedial projections in the saddles, light and dark spots similar in size or smaller than the spiracle, scattered all over the body. These specimens are tentatively identified here as S. ugoi, which in turn extends the geographic range of this species from northern Rio de Janeiro, Brazil to Barbados, Caribbean Sea (Fig. 9). No data on the conservation status of this species was found.

Scyliorhinus haeckelii (Miranda Ribeiro, 1907) (Figs. 10B, 11) Common names: freckled catshark, polkadot catshark (United States), ca����o-pinto (Brazil). Catulus haeckelii Miranda Ribeiro, 1907: 163 ���165, fig. 8 (original description, type locality: Rio de Janeiro, Brazil). Catulus boa: Garman, 1913: 17 (only the part referring to the holotype of S. haeckelii). Scyliorhinus boa: Bigelow & Schroeder, 1948: 204 ���207, fig. 32 (only the part referring to the illustration of the holotype of S. haeckelii and remarks about it); Springer, 1966: 602 (only the part referring to the holotype of S. haeckelii). Scyliorhinus retifer besnardi Springer & Sadowsky, 1970: 94 ���97, fig. 2 (original description, type locality: Rio Grande do Sul, Brazil); Cadenat & Blache, 1981: 183 ���184, fig. 123c (catalogue, western coast of Africa); Menni, Ringuelet & Aramburu, 1984: 97 ���98 (catalogue, Argentina and Uruguay) [synonymy follows Soares et al., 2016: 513 ���523]. Scyliorhinus retifer haeckelii Springer & Sadowsky, 1970: 92 ���93 (taxonomic review, Western Central Atlantic). Scyliorhinus retifer: Regan, 1908: 457 (only the part referring to the holotype of S. haeckelii); Figueiredo, 1977: 13, fig. 16 (catalogue, Brazil). Scyliorhinus besnardi: Springer, 1979: 126 ���128, figs. 79, 80 (taxonomic review); Compagno, 1984: 357 (FAO catalogue); Compagno et al., 2005: 246, pl. 42 (compilation); Ebert et al., 2013a: 374, 377, pl. 52 (compilation). Scyliorhinus haeckelii: Springer, 1979, pp. 135���137, fig. 86 (taxonomic review); Compagno, 1984: 362 ���363 (FAO catalogue); Gomes & Tom��s, 1991: 193 ���200 (sexual dimorphism); Gomes & de Carvalho, 1995: 232 ���236, fig. 2 (egg capsules); Compagno, 1999: 480 (listed); Soto, 2001: 68 ���69 (catalogue, Brazil); Gadig & Gomes, 2003: 22 (catalogue, Brazil); Bernardes et al., 2005: 59 (catalogue); Compagno et al., 2005: 250, pl. 42 (compilation); Ebert et al., 2013a: 374, 381, pl. 52 (compilation); Soares et al., 2016: 513 ���523, figs. 9���23, 25, 26 (taxonomic review, Western Central Atlantic); Weigmann, 2016: 43 (listed); Rinc��n et al., 2017: 94 ���95 (catalogue, Brazil). Scyliorhinus haeckelii / besnardi group Gomes et al., 2010: 84 ���85, fig. 109 (catalogue, Rio de Janeiro, Brazil); Soares et al., 2015: 1 (compared to S. ugoi, new species described). Holotype. MNRJ 494, male, 316 mm TL (Ilha Rasa, Rio de Janeiro, Brazil). Additional material examined. 127 specimens (see Appendix). Diagnosis. Scyliorhinus haeckelii differs from all congeners by presenting a color pattern composed of dark spots predominantly greater than the spiracles (vs. dark spots absent in S. capensis, S. comoroensis, S. hesperius, S. meadi, S. torazame, and S. torrei; reticulate pattern in S. retifer; spots predominantly smaller in S. boa and S. cabofriensis); anterior nasal flaps not reaching the upper lip (vs. flaps reaching the upper lip in S. canicula, S. cervigoni, S. comoroensis, S. duhamelii, S. garmani, and S. stellaris); interdorsal distance greater than the anal base (vs. smaller or equal to anal base in S. canicula, S. capensis, S. cervigoni, S. comoroensis, S. duhamelii, S. garmani, S. stellaris, and S. torazame); dorsal terminal 2 cartilage reduced and subtriangular (vs. elongated in S. boa, S. canicula, S. comoroensis, S. duhamelii, S. retifer, S. stellaris, S. torazame, and S. torrei); groove on the distal portion of the ventral terminal cartilage rudimentary or absent (vs. groove well developed in S. cabofriensis). The following combination of characters, although less conspicuous, also helps to distinguish this species: dark spots restricted to saddles and not bordering them (vs. spots scattered throughout the body in S. cabofriensis and S. duhamelii; bordering saddles in S. boa); spots double, lunate and with clear centers present on dorsolateral surfaces below the lateral line (vs. absent in S. boa, S. cervigoni, S. garmani, and S. ugoi); saddles without antero- and posteromedial projections (vs. projections present in S. ugoi); mesonarial ridge not exceeding the posterior border of the anterior flaps (vs. exceeding in S. stellaris); pelvic apron extending to 2/3 of length of pelvic inner margins (vs. extending for almost the entire length in S. canicula, S. capensis, S. duhamelii, S. torazame, and S. torrei); clasper with cover rhipidion covered by dermal denticles (vs. denticles absent in S. boa, S. cervigoni and S. retifer); clasper terminal dermal cover smooth (vs. rough in S. canicula and S. capensis); terminal 3 cartilage absent (vs. present in S. boa, S. canicula, S. capensis, S. retifer and, S. torazame); counts of monospondylous vertebrae 36���40 (vs. 44���46 in S. capensis; 40���45 in S. cervigoni; 48 in S. garmani; 46���48 in S. meadi; 43���47 in S. stellaris; 30���35 in S. torrei); adult males at least from 350 mm TL and adult females from 390 mm TL (vs. adult males greater than 450 mm TL in S. capensis, S. cervigoni, S. meadi, and S. stellaris; 269 mm TL and 294 mm TL, respectively, in S. torrei). Remarks. Soares et al. (2016) described the clasper of S. haeckelii with an accessory dorsal marginal cartilage (RD2), which would support the rhipidion (p. 519���520, figs. 14, 16). In this study, this structure is reidentified as a terminal dorsal 2 cartilage (Fig. 8b), following Jungersen (1899) and Compagno (1988a). Additionally, specimen UERJ 2230.2, previously identified as S. cabofriensis, is reidentified as S. haeckelii by presenting a color pattern more similar to the latter. Springer (1979) included specimen USNM 188061, captured just north of the mouth of the Amazonas River, in his list of examined material of S. haeckelii. This specimen is a 143 mm TL juvenile male that presents a color pattern composed of saddles slightly darker than the background, dark spots more frequent in the border of saddles and light spots present from the pectoral saddles (same color pattern presented by the specimen illustrated in Oliveira et al., 2015: 31). Despite the difficulty in identifying juvenile specimens, due to ontogenetic variation in color pattern, clasper morphology and morphometrics, the presence of dark spots bordering the saddles is understood here as a diagnostic character for S. boa. This specimen is therefore reidentified here as S. boa. With the reidentification of specimens captured from Colombia to northern Brazil, the geographic distribution of S. haeckelii is restricted here to southwestern Atlantic, from southern Bahia, Brazil, to northern Argentina (Fig. 11). Conservation status ���Data Deficient��� (Rincon 2004)., Published as part of Soares, Karla D. A. & De, Marcelo R., 2019, The catshark genus Scyliorhinus (Chondrichthyes: Carcharhiniformes: Scyliorhinidae): taxonomy, morphology and distribution, pp. 1-147 in Zootaxa 4601 (1) on pages 68-69, DOI: 10.11646/zootaxa.4601.1.1, http://zenodo.org/record/2669727, {"references":["Miranda Ribeiro, A. M. (1907) Fauna brasiliense. Peixes. II (Desmobranchios). Archivos Museu Nacional, 14, 2 - 20.","Garman, S. (1913) The Plagiostomia (Sharks, skates and rays). Memoirs of the Museum of Comparative Zoology at Harvard College, 36, 1 - 515.","Bigelow, H. B. & Schroeder W. C. (1948) Fishes of the Western North Atlantic. Part I. Lancelets, Cyclostomes and Sharks. Yale University, New Haven, 576 pp.","Springer, S. (1966) A review of Western Atlantic cat sharks, Scyliorhinidae, with descriptions of a new genus and five new species. Fishery Bulletin U. S. Fish and Wildlife Service, 65 (3), 581 - 624.","Springer, S. & Sadowsky, V. (1970) Subspecies of the western Atlantic catshark, Scyliorhinus retifer. Proceedings of the Biological Society of Washington, 83 (7), 83 - 98.","Cadenat, J. & Blache, J. (1981) Requins de Mediterranee et d'Atlantique (plus particulierement de la Cote Occidentale d'Afrique). Fauna Tropicale, 21, 1 - 330.","Menni, R. C., Ringuelet, R. A. & Aramburu, R. H. (1984) Peces marinos de la Argentina y Uruguay. Editorial Hemisferio Sur S. A, Buenos, Aires, 359 pp.","Soares, K. D. A, Gomes, U. L. & de Carvalho, M. R. (2016) Taxonomic review of catsharks of the Scyliorhinus haeckelii group, with the description of a new species (Chondrichthyes: Carcharhiniformes: Scyliorhinidae). Zootaxa, 4066 (5), 501 - 534. https: // doi. org / 10.11646 / zootaxa. 4066.5.1","Regan, C. T. (1908) A synopsis of the sharks of the family Scyliorhinidae. The Annals and Magazine of Natural History, 8 (1), 453 - 465. https: // doi. org / 10.1080 / 00222930808692434","Figueiredo, J. L. (1977) Manual de Peixes Marinhos do Sudeste do Brasil. I. Introducao. Cacoes, Raias e Quimeras. Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, 104 pp. https: // doi. org / 10.5962 / bhl. title. 109986","Springer, S. (1979) A revision of the catsharks, Family Scyliorhinidae. NOAA technical report NMFS CIRC, 422, 1 - 97.","Compagno, L. J. V. (1984) FAO species catalogue. Fol. 4. Sharks of the world: An annotated and illustrated catalogue of shark species known to date. Part 2. Carcharhiniformes. In: FAO Fisheries Synopsis No. 125. Fol. 4. Part 2. FAO, Rome, pp. 251 - 655.","Compagno, L. J. V., Dando, M. & Fowler, S. (2005) Sharks of the World. Princeton University Press, Princeton, 368 pp.","Gomes, U. L. & Tomas, A. R. G. (1991) Dimorfismo sexual secundario no cacao Scyliorhinus haeckelli Ribeiro, 1907 (Elasmobranchii, Scyliorhinidae). Anais da Academia Brasileira de Ciencias, 63 (2), 193 - 200.","Gomes, U. L. & de Carvalho, M. R. (1995) Egg Capsules of Schroederichthys tenuis and Scyliorhinus haeckelii (Chondrichthyes, Scyliorhinidae). Copeia, 1995 (1), 232 - 236. https: // doi. org / 10.2307 / 1446823","Compagno, L. J. V. (1999) Checklist of living elasmobranchs. In: Hamlett, W. C. (Ed.), Sharks, skates, and rays: the biology of elasmobranch fishes. Johns Hopkins University Press, Maryland, pp. 471 - 498.","Soto, J. M. R. (2001) Annotated systematic checklist and bibliography of the coastal and oceanic fauna of Brazil. I. Sharks. Mare Magnum, 1 (1), 51 - 120.","Gadig, O. B. F. & Gomes, U. L. (2003) Classe Chondrichthyes. In: Menezes, N. A., Buckup, P. A., Figueiredo, J. L. & Moura, R. L. (Eds.), Catalogo das especies de peixes marinhos do Brasil. Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, pp. 21 - 22.","Bernardes, R. A., de Figueiredo, J. L., Rodrigues, A. R., Fischer, L. G., Vooren, C. M., Haimovici, M. & Rrossi-Wongtschowski, C. L. D. B. (2005) Peixes de Zona Economica Exclusiva da Regiao Sudeste-Sul do Brasil: Levantamento com armadilhas, pargueiras e rede de arrasto de fundo. Editora da Universidade de Sao Paulo, Sao Paulo, 295 pp.","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88 (3), 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Rincon, G., Mazzoleni, R. C., Palmeira, A. R. O. & Lessa, R. (2017) Deep-Water Sharks, Rays, and Chimaeras of Brazil. In: Rodrigues Filho, L. F. S. (Ed.), Chondrichthyes-Multidisciplinary Approach Edition. Chapter 5. Available from: https: // www. intechopen. com / books / chondrichthyes-multidisciplinary-approach / deep-water-sharks-rays-and-chimaeras-of-brazil (accessed 30 Mars 2018) https: // doi. org / 10.5772 / intechopen. 69471","Gomes, U. L., Signori, C. N., Gadig, O. B. F. & Santos, H. R. S. (2010) Guia para Identificacao de Tubaroes e Raias do Rio de Janeiro. Technical Books, Rio de Janeiro, 234 pp.","Soares, K. D. A, Gadig, O. B. F. & Gomes, U. L. (2015) Scyliorhinus ugoi, a new species of catshark from Brazil (Chondrichthyes: Carcharhiniformes: Scyliorhinidae). Zootaxa, 3937 (2), 347 - 361. https: // doi. org / 10.11646 / zootaxa. 3937.2.6","Jungersen, H. F. E. (1899) On the apendices genitales in the greenland shark Somniosus microcephalus (Bl. Schn.) and another selachians. Danish Ingolf Expedition. Fol. II. Bianco luno, Copenhagen, 88 pp.","Oliveira, J. E. L., Nobrega, M. F., Garcia, J., Sampaio, C., Di Dario, F., Fischer, L. G. & Mincarone, M. M. (2015) Biodiversidade Marinha da Bacia Potiguar / RN: Peixes do Talude Continental. Museu Nacional, Rio de Janeiro, 218 pp.","Rincon, G. (2004) Scyliorhinus haeckelii. The IUCN Red List of Threatened Species, 2004, e. T 44589 A 10909893."]}

Scyliorhinus stellaris (Linnaeus, 1758) (Figs. 70–76, Tabs. 3, 9, 16) Common names: Large spotted dogfish, Larger spotted dogfsh, Greater spotted dogfsh, Nurse hound, Bull huss, Spotted dogfsh (United Kingdom); Le squale rochier, Chat rochier, Rousette a grandes taches, Panthêre de mer, Cat rouquiera, Pinto reussou, Vache (France); Pantherhai, Hundshai, Grossgefleckter Katzenhai (Germany); Grote gevlekte hondshaai (Netherlands); Bounce (Belgium); Bruxa, Cacåo, Carraca, Cascarra, Gata, Bata roxa, Pata roxa denisa (Portugal); Alitán, Gat, Gatet, Gato, Gaton, Muxina, Pintarroja (Spain). Squalus stellaris Linnaeus, 1758:235 (original description, type locality: Mediterranean Sea); Houttuin, 1764: 508 –509 (compilation); Linnaeus, 1766: 399 (compilation); Berkenhout, 1769: 36 (catalogue, Great Britain and Ireland); Heppe, 1787: 212 (compilation); Bonnaterre, 1788: 7 (compilation); Duperrey, 1830: 83 (catalogue, expedition ‘La Coquille’); Lacépède, 1830: 384–387 (compilation). Scyllium stellaris: Risso, 1826:116 (catalogue, Southern Europe); Jenyns, 1835: 496 –497 (catalogue, United Kingdom). Catulus saxatilis: Valmont, 1769: 51 (listed). Scyllium catulus: Cuvier, 1817: 124 (compilation); Cuvier, 1829: 386 (compilation); Cuvier et al., 1834: 586 (brief account, classification); Partington, 1837: 651 (compilation); Parnell, 1838: 407 –409 (catalogue, North Sea); Müller & Henle, 1838 –41: 9–11 (brief description, systematics of Elasmobranchii); Gosse, 1851: 310 (compilation); Hamilton, 1854: 300 - 301 (compilation); Duméril, 1865: 316 –318 (compilation); Bocage & Capello, 1866: 11 (catalogue, Portugal); Moreau, 1881: 280 –284, fig. 36 (catalogue, France); Day, 1884: 312–314, fig. 2, pl. 159 (catalogue, Great Britain and Ireland); Gordon, 1902: 135 (catalogue, Great Britain); Coward, 1910: 161 (catalogue, Cheschire and Liverpool Bay); Leigh- Sharpe, 1920: 247–251, fig.1 (clasper description) [synonymy follows Springer 1973:19]. Scyllium stellare: Fleming, 1828:165 (catalogue, United Kingdom); Schinz, 1840: 461 (catalogue, Europe); Thompson, 1856: 247 (listed); Günther, 1870: 403 (catalogue, British Museum); Doderlein, 1880: 22–24 (catalogue, Mediterranean Sea); Balfour, 1881: 656 –671 (anatomy and development of paired fins); Faber, 1883: 181 –182 (catalogue, Adriatic Gulf); Jungersen, 1899: 34 –36, pl. II, figs. 16-18. VI, figs. 65, 66 (clasper description); Danois, 1913: 15, figs. 4, 8 (compilation). Scylliorhinus stellaris: Blainville, 1830:71, pl. 17 (listed, classification). Catulus stellaris: Hoffman & Jordan, 1892: 233 (catalogue, Greece); Garman, 1913: 75 –76 (brief account). Scyliorhinus stellaris: Regan, 1908: 457 (brief account); Ford, 1921: 492 –493 (diet); Bigelow & Schroeder, 1948: 203 (catalogue, comparison with species of Northwestern Atlantic); Tortonese, 1956: 130, figs. 66–69 (catalogue, Italy); Wheeler, 1969: 44, fig. 15 (catalogue, British Islands); Springer, 1973: 19 (listed, Northeastern Atlantic and Mediterranean Sea); Springer, 1979: 143 –144, figs. 3, 94 (taxonomic review); Cadenat & Blache, 1981: 178 –181, figs. 121b, 122 (except part referring to specimens of S. cervigoni); Compagno 1984; 366–367 (FAO catalogue); Compagno, 1999: 480 (listed); Soldo et al., 2000: 355 –356 (tooth morphology); Compagno et al. 2005: 252, pl. 41 (compilation); Serena, 2005: 38, pl. IV, 31 (FAO catalogue, Mediterranean Sea); Capapé et al., 2006: 29 –36 (reproductive biology); Fricke et al., 2007: 13 (listed); George, 2009 (listed, North Sea): 35; Ebert et al. 2013a: 372, 383, pl. 51 (compilation); Ebert & Stehmann, 2013: 211 –2012, figs. 242, 243 (egg capsules); Sabata & Clò, 2013: 178 –179 (reproductive biology); Bilecenoglu et al., 2014: 904 (catalogue, Turkey); Gordon et al., 2016: 272 –273, fig. 8c (egg capsules); Weigmann, 2016: 44 (listed). Haploblepharus stellaris: White, 1937: 121 (listed, systematics). Scyllium (Betascyllium) catulus: Leigh-Sharpe, 1924: 325 (clasper description, classification). Scyllium (Betascyllium) stellare: Leigh-Sharpe, 1924: 326 (clasper description, classification). Neotype. NHMUK 1976.7.30.10, male, 476 mm TL (Banyuls, Pyrenees, France) [designated herein]. Additional material examined. 98 specimens (see Appendix). Diagnosis. Scyliorhinus stellaris differs from all congeners by presenting a mesonarial ridge extending posteriorly beyond the posterior border of anterior nasal flap (vs. not exceeding it in all other species); pelvic fins trapezoidal (vs. subtriangular in other species, except S. garmani and S. torazame); color pattern composed of dark spots with well-defined borders and predominantly greater than spiracles (vs. dark spots absent in S. capensis, S. comoroensis, S. hesperius, S. meadi, S. torazame, and S. torrei; reticulated pattern in S. retifer; dark spots predominantly smaller than spiracle in S. boa and S. cabofriensis; spots scattered, diffuse in S. duhamelii). The following combination of characters, although less conspicuous, also helps distinguish this species: spots scattered along the dorsolateral surface (vs. spots predominantly restricted to saddles in S. boa, S. cervigoni and S. haeckelii); anterior nasal flaps reaching the upper lip (vs. not exceeding in others, except in S. cervigoni, S. comoroensis, S. duhamelii, and S. garmani); distance between anterior nasal flaps two times smaller than width of anterior nasal flap (vs. 6–7.5 times in S. canicula; 3.5–5 times smaller in S. duhamelii); nasoral grooves absent and posterior nasal flaps situated on posterior border of excurrent apertures (vs. grooves present and flaps laterally situated in S. canicula and S. duhamelii); interdorsal distance 0.6–1.0 times the anal base (vs. greater than the anal base in S. boa, S. cabofriensis, S. haeckelii, S. hesperius, S. meadi, S. retifer, S. torrei, and S. ugoi); mandibular canal of lateral line system with 6 or 7 pores (vs. 4–5 in S. duhamelii; 3–4 in S. hesperius); oral canal of lateral line system with 7–10 pores (vs. 5–6 in S. hesperius; 10–12 in S. duhamelii; 9–13 in S. torrei); commissural teeth with two cusplets (vs. one in S. cervigoni, S. torazame and S. torrei; three or more in S. boa, S. canicula and S. hesperius); upper tooth rows 41–51 (vs. 33–42 in S. torrei); lower tooth rows 34–50 (vs. 48–85 in S. capensis); pelvic apron extending to 2/ 3 length of pelvic inner margins (vs. extending through almost pelvic entire length in S. canicula, S. capensis, S. duhamelii, S. torazame, and S. torrei); clasper with terminal dermal cover smooth (vs. rough in S. canicula and S. capensis); cover rhipidion covered by dermal denticles (vs. denticles absent in S. boa, S. cervigoni, S. hesperius, and S. retifer); terminal 3 cartilage absent (vs. present in S. boa, S. canicula, S. capensis, S. retifer, and S. torazame); dorsal terminal 2 cartilage elongated and corresponding to 1/4 of dorsal terminal cartilage (vs. reduced and subtriangular in S. cabofriensis, S. capensis, S. cervigoni, S. haeckelii, and, S. ugoi; 1/3 of dorsal terminal cartilage in S. boa and S. comoroensis; cartilages same length in S. torazame); counts of monospondylous vertebrae 43–47 (vs. lower counts in than S. boa, S. cabofriensis, S. canicula, S. comoroensis, S. duhamelii, S. haeckelii, S. hesperius, S. torazame, and S. torrei; 48 in S. garmani); adult males between 700–740 mm TL and adult females between 770–790 mm TL (vs. adult sizes smaller in other species, except S. capensis, S. cervigoni and S. meadi). Description. Morphometric and meristic data are given in Table 16, and neurocranial measurements in Table 9. Body robust, tapering considerably posterior to cloaca (Figs. 70, 71). Prepectoral length 0.5 times the prepelvic length. Trunk shorter than tail; snout-vent length 0.8–0.9 (0.8) times vent-caudal length. Pectoral-pelvic space 1.9–3.1 (2.1) times the pelvic-anal space. Interdorsal space 1.8–2 (1.8) times the dorsal-caudal space (Tab. 16). No interdorsal, postdorsal or postanal ridges; lateral crest on caudal peduncle absent. ......continued on the next page Head broad and depressed; head length 1.4–1.6 (1.8) times head width (Figs. 70, 71). Snout relatively short, preoral length 0.5–0.7 (0.6) times mouth width and 0.7–0.9 (0.7) times smaller than preorbital length. Prenasal length 0.4–0.6 (0.5) times internarial space; preorbital length 0.8–0.9 (0.8) times interorbital space. Eye large and slitlike, eye length 2.1–4.5 (2.8) times its height and 0.2 times smaller than head length (Figs. 70, 71). Eye dorsolateral on head, with lower edge medial to horizontal head rim in dorsal view; subocular ridge strong. Nictitating lower eyelid of rudimentary type, with shallow subocular pouch and secondary lower eyelid free from upper eyelid. Spiracle close behind but well separated from eyes, dorsolaterally on head and somewhat lower than level of eye notch. Spiracle diameter goes 2.3–4.5 (3.4) times in eye length and 5.1–8.8 (7.5) times in interorbital distance. First two gill openings about equally wide; first one twice as long as fifth. All gill openings slightly concave and not elevated on dorsolateral surface of head; gill filaments not visible externally. Nostril with broad incurrent aperture, without nasoral groove or nasal barbel, and small and oval excurrent aperture. Anterior nasal flap large, triangular, covering posterior nasal flap and excurrent aperture, and touching the upper lip (Figs. 72 A–B). Mesonarial ridge distinct and extending posteriorly beyond the posterior border of the anterior nasal flap. Posterior nasal flap rectangular, situated on the posterior border of the excurrent aperture. Mesonarial superior and inferior flaps conical and corresponding to 1/3 of anterior nasal flap. Internarial space 0.7–0.8 (0.7) times the interorbital space. Mouth arched, moderately wide and short, its length goes 1.4–1.9 (1.6) times in mouth width (Figs. 72 A–B). Lower labial furrow short and narrow, 2.8–4 (3.9) times smaller than mouth width. Dorsal labial cartilage 1.3 times the ventral cartilage; anterior tip of dorsal labial cartilage reaching the orbital process of the palatoquadrate. Tongue flat and rounded, light-colored, with oral papillae hardly detectable. Monognathic heterodonty gradual well developed; anterior teeth abruptly larger than the parasymphysial ones and lateral teeth smaller distally, with smaller and thicker principal cusps (Fig. 73). Sexual heterodonty pronounced with females presenting shorter principal cusp and more developed cusplets than males. Tooth counts 20–26 16–25/16–25 0– 1 16–24. Parasymphysial teeth with a principal cusp flanked by one cusplet on each side; cusplets half the height and the width of the principal cusp. Protuberances on medial portion of the crown base and striae restricted to the crown base. Anterior teeth larger than the parasymphysial and principal cusp less stout. Anterior teeth with two to four cusplets; marginal cusplets poorly developed and 1/3 the height of proximal cusplets in upper teeth and poorly developed in lower teeth. Proximal cusplets 1/3 the height and the width of the principal cusp in teeth in both jaws. Protuberances on the crown base and striae extending to one half the height of crown. Lateral teeth with three cusplets; principal cusp slightly oblique, two cusplets at the mesial edge and one at the distal edge. Mesial proximal and distal cusplets one halt to 2/3 the height of the principal cusp; mesial marginal cusplet half the height of proximal one. Protuberances on the crown base and striae running from base toward the apex of the principal cusp. Commissural teeth with two cusplets; principal cusp stronger and laterally situated. Cusplets 2/3 the height and the width of the principal cusp. Protuberances on the crown base and striae throughout the crown. Ectodermal pits present in lateral and commissural teeth, restricted to the crown base. Lateral trunk denticles with flat, elongated teardrop-shaped crowns, 1.6–1.8 times as long as wide (Tab. 3); anterior part covered with ectodermal pits. Dermal denticles above the pectoral fin presenting five ridges, median ridge less prominent than in denticles of other regions and not extending to the intersection between principal cusp and cusplets. Denticles below dorsal fins longer and with prominent median and lateral ridges, extending to the distal tip of cusplets (Fig. 74). Pectoral base 0.7–0.8 (0.8) times mouth width (Fig. 72D). Pectoral anterior margin 2.1–2.2 (2.3) times its base and 1.4–1.7 (1.5) times the posterior margin. Pectoral fin skeleton aplesodic with radials mostly divided into three segments. Propterygium and mesopterygium trapezoidal; the former smaller than the latter. Propterygium with one proximal segment; mesopterygium with 3–4 proximal segments fused proximally. Metapterygium with 8–9 segments. Metapterygial axis rectangular and corresponding to 1/4 of metapterygium. Pelvic fin subrectangular in females and trapezoid in males (Figs. 72 E–F); pelvic anterior margin 1–1.3 (1.1) times the posterior margins and 0.9–1 time the pelvic base. Pelvic inner margins of males fused by 2/3 of their extension; claspers of juveniles evident without lifting the pelvic apron. Clasper short and cylindrical (Fig. 72F), sometimes extending beyond free rear tips of pelvic fins; clasper inner length 1.4–1.7 (1.3) times the pelvic anterior margin, 1.7–3 (2.4) times the clasper outer length and 3.7–4.8 (4.4) times the clasper base. Most of clasper surface except ventrolateral and dorsomedial surface of glans, rhipidion, and terminal dermal cover, covered by dermal denticles with anteriorly directed crowns (Fig. 75A). Clasper hooks absent. Rhipidion well-developed, partly covered medially by a prominent exorhipidion and anteriorly by the cover rhipidion; insertion of rhipidion at posterior portion of dorsal terminal 2 cartilage and extending to the end of glans. Cover rhipidion expanded medially reaching the exorhipidion, and sometimes covered by this anteriorly; both cover rhipidion and exorhipidion covering the clasper groove. Envelope absent; pseudosiphon distinct and robust. Terminal dermal cover extending for 1/3 of the ventral terminal cartilage, covering the posterior edges of exorhipidion and cover rhipidion. Clasper skeleton relatively simple (Fig. 75B). Ventral terminal cartilage beginning anteriorly, but ending together with the dorsal terminal. Terminal 3 cartilage absent. Dorsal terminal 2 cartilage elongated and rod-like, medially positioned on the dorsal terminal cartilage; this cartilage supports the rhipidion and corresponds to 1/4 the length of dorsal terminal cartilage. Ventral terminal 2 cartilage slender, above the ventral terminal cartilage, and corresponding to 2/3 of this; ventral terminal 2 cartilage beginning at the same level that dorsal terminal 2. First dorsal fin subrectangular or triangular, with nearly straight anterior margin, rounded apex and angular free rear tip (Figs. 70, 71). First dorsal fin origin slightly anterior to the pelvic fin insertion. First dorsal fin insertion opposite to the anterior 1/3 of pelvic-anal distance. Anterior margin 1.5 (1.3) times first dorsal fin base; first dorsal fin height 0.8 times its base. Second dorsal fin triangular and smaller than the first (Figs. 70, 71). Second dorsal fin origin slightly posterior to the anal midbase and insertion opposite to the posterior end of the anal fin. Anterior margin 1.3 times base of second dorsal fin; second dorsal base 1.6–1.8 (1.6) times its height and 0.4–0.6 (0.6) times the dorsal-caudal distance. First dorsal fin 1.3–1.4 (1.4) times larger than the second dorsal fin. Anal fin rounded, apically narrow and not falcate (Figs. 70, 71); anal fin base 1.6–1.8 (1.6) times the second dorsal fin base. Anal fin anterior margin nearly straight, apex rounded, free rear tip acutely pointed, and inner margin nearly straight. Anal fin base 1.4–2.1 (1.4) times the interdorsal distance and 2–2.2 (2.2) times the dorsalcaudal distance. Anal fin anterior margin 1.8–2.1 (1.8) times the posterior margin; anal fin height 0.3–0.4 (0.4) times its base. Caudal fin narrow-lobed and asymmetrical (Figs. 70, 71). Dorsal caudal lobe 1.2–1.4 (1.5) times larger than preventral lobe; subterminal caudal margin 0.7–0.8 (0.7) times the terminal margin. Caudal crest of enlarged denticles absent on caudal fin margins. Neurocranium broad and somewhat flattened, corresponding to 8.9–9.5% TL. Rostrum length 1.4–1.7 times the distance between lateral rostral cartilages. Nasal capsule wider than long, oval-shaped and expanded laterally; width 1.1–1.2 times its length. Basal plate flat with narrow borders, its width 2–2.1 times smaller than nasobasal length. Orbital region 2.5–2.6 times smaller than nasobasal length. Otic capsule short, its length 4.4–4.6 times smaller than nasobasal length and width 2.7–2.8 times otic capsule length. Width across postorbital processes 1.1–1.2 times the preorbital processes width (Tab. 9). Coloration in alcohol. Body beige with dark brown spots greater than spiracles and distributed along the dorsolateral surface; spots smaller in dorsal regions than in lateral regions (Figs. 70, 71). Lunate and clear center spots present. Longitudinal dorsal stripe extending from level of spiracle to caudal peduncle in some specimens; seven to eight saddles present or absent. Subsaddles between saddles anterior to the first dorsal fin. Juveniles with numerous dark and small spots, usually with light spots, and sometimes saddles distinct in relation to the background. Belly and ventral surface of paired fins with or without dark spots. Distribution. This species is distributed along the Balearic and Tyrrhenian Seas, in the continental shelves of Italy, Tunisia, France and Spain, with two records in the Adriatic Sea close to Venice, Italy, and Rovinj, Croatia. Also found along the Northeastern Atlantic in the continental shelves of Portugal, France and the United Kingdom, as well as one record for the North Sea (55°N, 3°E). No records exist from northwestern Africa (Fig. 76). Biological data. Adult males between 700–740 mm TL; largest male examined 956 mm. Adult females between 770–790 mm TL; largest female examined 971 mm TL. Maximum size recorded 1900 mm (Fischer et al. 1987); sizes larger in the northern part of its distribution (up to 1500 mm TL) than in the Mediterranean Sea (750 mm TL) (Tortonese 1956). Egg deposition occurs, Published as part of Soares, Karla D. 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