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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"]}

Poroderma pantherinum (Smith in M��ller and Henle, 1838) (Figure 3-7, Table 2) Scyllium pantherinum Smith in M��ller and Henle, 1838: 13. Poroderma pantherinum Smith, 1837: 85 (name only). Poroderma submaculatum Smith, 1837: 85 (name only). Poroderma variegatum Smith, 1837: 85 (name only). Scyllium leopardinum M��ller and Henle, 1838: 13 (Fowler 1941: 38 attributes this name to Van Horst in M��ller and Henle, although no reference to Van Horst is given in M��ller and Henle). Scyllium maeandrinum von Rapp in M��ller and Henle, 1838: 13. Scyllium variegatum Smith in M��ller and Henle, 1838: 14; Smith 1849: 11; Dum��ril 1853: 83; Bleeker 1878: 1, 8; Sauvage 1891: 511. Scyllium pantherinum: Smith 1849: 13; Dum��ril 1853: 82; Bleeker 1878: 1, 8; Sauvage 1891: 511. Scyllium africanum: G��nther 1870: 405; Lampe 1914: 209. Scyllium africanum var. pantherina: G��nther 1870: 405; Lampe 1914: 212. Scyllium africanum var. striata: G��nther 1870: 405; Lampe 1914: 212. Scyllium africanum var. variegata: G��nther 1870: 405; Lampe 1914: 212. Scyliorhinus pantherinus: Regan 1908: 456. Scyllium africanum var. punctata Lampe 1914: 212. Scylliorhinus pantherinus: Thompson 1914: 138; Gilchrist and Thompson 1916: 270; Gilchrist 1922: 4; Barnard 1925: 40; Barnard 1927: 1013. Scyliorhinus regani: Fowler 1925: 188; Fowler 1926: 32, not Holohalaelurus regani (Gilchrist, 1922). Poroderma marleyi Fowler, 1934: 234; Fowler 1941: 38; Bigelow and Schroeder 1948: 197; Smith 1949: 53; Springer and Garrick 1964: 86; Bass et al. 1975: 29; Springer 1979: 114; Buxton et al. 1984: 186; Compagno 1984b: 348; Wallace et al. 1984: 159; Compagno 1986: 7; Bass 1986: 94. Scyliorhinus leopardus: Fowler 1935: 361. Conoporoderma pantherinum: Bigelow and Schroeder 1948: 197. Type Series and Locality. Holotype, BMNH 1845.7.3.145, mature female 664 mm TL, collected from Algoa Bay, Eastern Cape, South Africa, although given as ���Vom Cap der gutten Hoffnung��� (from the Cape of Good Hope) in the original description, dried mounted skin in fair condition, the colour pattern has faded considerably, but is still discernable on the dorsal surface (Fig. 3A-B). Diagnosis. P. pantherinum is distinguished from P. africanum by having a narrower, longer and more acutely pointed snout than in P. africanum; head width 12.9% TL for the holotype (mean 12.9% TL). The upper labial furrow is longer and thinner than in P. africanum; upper labial furrow length 1.5 (mean 1.0) times its width. The nasal barbels of P. pantherinum are more elongate than P. africanum, extending over the upper lip and overhanging the mouth in P. pantherinum; nasal barbel length 1.1 (mean 0.9) times nostril width. Although the colour patterning of P. pantherinum is highly variable, it never forms the bold anteroposterior striping observed in P. africanum. Description. Morphometric and meristic data are given in Table 2. Holotype, mature female 664 mm TL (mean of all specimens examined in Table 2, including the holotype, see Study material): although a robust shark, P. pantherinum is the smaller of the Poroderma sharks. Precaudal length 1.25 (1.25) times the preanal length; snout narrow and pointed; distance from snout to 2nd dorsal fin origin 1.2 (1.25) times the distance from snout to 1st dorsal fin origin; mouth width 2.1 (1.7) times the preoral length; prenarial length 1.8 (1.1) times the mouth length; nasal barbels elongated and extending over the upper lip; 2nd dorsal base length 1.2 (1.0) times dorsal caudal space; distance from snout to pectoral fin origin 1.4 (1.1) times the caudal dorsal margin length; pelvic fin base length 1.2 (1.1) times the pectoral fin base length; height of pectoral fin 2.3 (1.85) times that of the pelvic fin; distance between the pelvic and anal fin 1.3 (1.1) times the length of the anal fin base. Vertebral counts: NA (32-46) monospondylous, NA (29-42) precaudal diplospondylous, 31 (27-45) caudal diplospondylous vertebrae. Dental formula: upper jaw (left) 23 (20-30), (right) 20 (18-27); lower jaw (left) 20 (13-25), (right) 18 (14-26). Spiral valve turns: NA (8-13). Size and Sexual Maturity. In the current study P. pantherinum was found to grow to at least 774 mm TL and weigh at least 2.4 kg, making it a far less robust shark than its congener. Compagno (1986) lists the SA angling record for this species as 3.2 kg, which may be outdated now. Males are juvenile at 100 mm TL to 519 mm TL, adolescent at 469 mm TL to 672 mm TL, and mature at 607 mm TL to 774 mm TL. Females are juvenile at 94 mm TL to 485 mm TL, adolescent at 430 mm TL to 641 mm TL, and mature at 507 mm TL to 666 mm TL. There appears to be some sexual dimorphism in this species, with males growing slightly larger than the females, and maturing at a larger size than the females. There also appears to be a geographical component to the maximum size obtained, as larger specimens were consistently recorded from the Eastern Cape, compared to the Western Cape (Bass et al. 1975; current study). Colouration of the Holotype. Colouration partly faded on the holotype (Fig. 3A-B), background dorsal colouration pale grey, grey, to brown (latter is preservation artefact); many small brown spots and reticulations, not forming large whorls, and small rosettes; most clearly seen along dorsal midline with patterning fading rapidly ventrolaterally; rosettes becoming more common proceeding posteriorly along the dorsal midline; patterning undiscernible posterior to the insertion of the 1st dorsal fin; some patterning visible on the bases of the pectoral fins, similar to dorsal patterning; no patterning evident on any other fins. Ventral background colouration brown (preservation artefact). According to the illustration by Smith (1849), this specimen originally had many rosettes on the dorsolateral surface; extending posteriorly to the dorsal origin of the caudal fin; short broken lines along the lateral surface from the origin of the 1st dorsal fin to the posterior of the caudal peduncle; all fins with many small spots. Colouration of non-type specimens. Range of colouration of P. pantherinum observed from non-type specimens (Figs. 4-6), dorsal background colouration very pale grey to charcoal grey, to glossy jet black; with combinations of large black spots (Figs. 5 & 6A), small black spots (Fig. 6B), large black blotches, whole or broken rosettes (Fig. 6C- D), with spots sometimes fused into short thin broken lines to long thin lines running for most of the length of the body anteroposteriorly (Fig. 6E), also laterally across the dorsal midline, particularly just posterior to the orbits when present; patterning usually present on the dorsal surface of the pectoral and pelvic fins, as well as the dorsal, anal and caudal fins; or patterning absent in black specimens (Fig. 6F). Ventral background colouration white, very pale grey to charcoal grey, to black; sometimes strongly demarcated from the dorsal background colouration; patterning often found ventrally, but not along the ventral midline; patterning often present on the ventral surface of the pectoral and pelvic fins. The colour pattern of P. pantherinum exhibits an extraordinary amount of variation, and colouration and patterning of this species is best understood by referring to Figs. 3-6 and the discussion below. The greatly variable colour pattern of P. pantherinum has been noted by most authors and is summed by Barnard (1925: 40) who stated ������a series of examples can be obtained showing an almost complete transition from the plain striped pattern of africanus [= P. africanum] to the purely ocellate one of pantherinus [= P. pantherinum]���. Compagno et al. (1989) classified the different colour patterns of P. pantherinum into three forms and gave illustrations for them - typical (although there is no true typical form), marleyi (Figs. 5 and 6A) and salt and pepper (Fig. 6B). To this, one could add a melanistic form, which include sharks that have an exceptionally dark dorsal surface, approaching jet black with a strong sheen, with a variable number of stripes running anteroposteriorly along the entire dorsal surface, although stripes may be absent, with a variable number of spots either present or absent. Rosettes have not been observed in the melanistic form by the author. The author has observed some individuals of the latter form, which lack any type of patterning (Fig. 6F). A photograph taken by the author of a specimen (BAH 20020211.2) of the melanistic form was figured by Dainty (2002). The typical form can be extended to include individuals with spots fused into irregular stripes that generally run anteroposteriorly, but occasionally one or two stripes may be seen running transverse to the body axis between, and just posterior to, the orbits (Fig. 6E), as described by Garman (1913), and illustrated by Springer (1979). Fowler (1941: 38) described P. pantherinum ���with many thick-set white spots���. In observing more than approximately two hundred P. pantherinum sharks (including museum specimens and live specimens in their natural habitat), the author has not seen an individual with white spots, and the white spots of Fowler may refer to the rosettes commonly observed in this species. Comparison with other species. P. pantherinum is a less stockier shark than P. africanum at all maturity stages for both sexes, and grows to a smaller total length. P. pantherinum never has the bold stripes seen on P. africanum. Remarks. Smith (1837) presented pantherinum, submaculatum and variegatum, as well as africanum (= Squalus africanus Gmelin, 1789) by name only, as species for his new genus Poroderma, and failed to characterise both the genus and the species. P. pantherinum was first characterised by M��ller and Henle (1838-1841) as Scyllium pantherinum (Bigelow and Schroeder 1948; Compagno 1988). M��ller and Henle (1838-1841) also described S. variegatum, however they did not adopt the taxon submaculatum and did not provide a synonymy for it. The type of S. variegatum, BMNH 1845.7.3.130, is in a similar condition to the type of S. pantherinum, except that the colour pattern has completely faded (Fig. 4A-B). Smith (1849) separated S. variegatum from S. pantherinum purely on the basis of patterning, and the stripes (created from the fusion of spots) observed on S. variegatum was used as the primary basis for the separation of the two species. The illustrations by Smith (1849) for S. variegatum and S. pantherinum appear to be based on the type material for them, and the data on the labels accompanying the holotypes confirm this. The date of the publication means that the type specimens would have been available for the illustration, and the locality given by Smith, at least for S. variegatum, agrees with the type locality for that specimen. Presently, the colour pattern has completely faded on the S. variegatum type, however the colour pattern is still visible on the S. pantherinum type and agrees with the illustration of Smith. Eschmeyer (1998) reports that the illustration of S. variegatum is indeed that of the holotype for that species, however, the illustration of S. pantherinum is ���apparently independent of S. pantherinum M��ller and Henle (ex Smith) 1838 [and is] permanently invalid, preoccupied by S. pantherinum M��ller and Henle, 1838 ���. The author could not find any other reference to this ambiguity and disagrees with Eschmeyer (1998) on this point. G��nther (1870) and Gilchrist (1902) synonymised both Scyllium pantherinum and S. variegatum with P. africanum. G��nther justified his synonymy of these species with P. africanum because he found the nasal barbels to extend to the upper lip in Poroderma with stripes (G��nther 1870). It seems that the Poroderma with stripes referred to by G��nther are actually specimens of P. pantherinum with that patterning. The taxon P. marleyi Fowler, 1934 has been a somewhat enigmatic taxonomic problem for South African ichthyologists. Only a few specimens identified as P. marleyi exist in collections and the species was considered rare. The author was unable to personally examine the P. marleyi holotype (ANSP 53427; see Study material for details), however, examination of photographs taken of the type for this study show it to be in good condition, although somewhat distorted (Fig. 5). Fowler erroneously illustrated (with different illustrations), and described P. marleyi under the name Scyliorhinus regani von Bonde [= Holohalaelurus regani (Gilchrist, 1922)], in Fowler (1925) and Fowler (1926), which he acknowledged in a later publication (Fowler 1934). It should be noted that although the date of the proceedings that Fowler published his description of Conoporoderma and P. marleyi is shown as 1933, the publication date is 20th January 1934, hence is the date that should be referred to for these taxa, as has been done by Compagno (1984b) and Eschmeyer (1998). Proportional measurements, given by Fowler (1934), of the holotype of P. marleyi are indistinguishable from those of P. pantherinum examined here, the only difference being the low tooth count of the P. marleyi holotype in the upper jaw, which seems very low (22 teeth total compared to an average of 45.1 (n = 43) for P. pantherinum in this study) and may be erroneous. Springer and Garrick (1964) provided a vertebral count of the holotype with 104 vertebrae in total, which also agrees with the vertebral count for P. pantherinum obtained in this study. Bass et al. (1975) listed P. marleyi as valid based on a single specimen (ORI 934). This specimen is currently in the SAIAB collection (RUSI 6002) and was examined by the author. There are no morphometric or meristic data that separate this animal from P. pantherinum. The basis then for the separation of P. marleyi from P. pantherinum is the large black spots that are considered a distinguishing feature of P. marleyi. Given the enormous variability of the colour pattern of P. pantherinum, the patterning of P. marleyi amounts to an extreme colour pattern variant of P. pantherinum. The holotype of P. marleyi is only 225 mm TL and must be considered a juvenile of P. pantherinum, with the implication that the marleyi colour pattern is paedomorphic (see below). The proportional dimensions and vertebral counts given here for P. pantherinum are closely matched by those given by Bass et al. (1975). Vertebral counts given by Springer (1979) and Compagno (1988) also agree with the current study (mean total vertebral count 116.4, n = 52). Bass et al. (1975) describe and illustrate the teeth of this species and noted that sexual heterodonty occurs in this species. P. pantherinum also exhibits ontogenetic heterodonty, between adolescent and mature specimens of males only (M. Marks pers. comm.). Compagno (1988) described strong gradient heterodonty for all species of Poroderma, and also noted weak sexual heterodonty. Tooth counts provided here (mean upper total 45.1, lower total 42, n = 42) agree with those given by Fowler (1925) and Compagno (1988). The denticles of P. pantherinum have been described by Bass et al. (1975) and Compagno (1988), and illustrated by Bass et al. (1975). The spiral valve counts given by Compagno (1988), agree with counts obtained during this study (mean 10.1, min. 8, max. 13, n = 45). There is a general trend in the colour pattern observed and the geographic locality of the specimen (Bass et al. 1975; pers. obs.). The melanistic form appears to be exclusive to False Bay. The salt and pepper form and the marleyi form appear to be exclusive to the Eastern Cape and kwaZulu-Natal, while the typical form and intermediates are encountered throughout the range of P. pantherinum. There is also an ontogenetic component apparent to the patterning in this animal (Bass et al. 1975; current study). The smaller individuals generally tend to have complete rosettes and/or large solid spots, and as the animal grows these large spots diffuse into rosettes, broken rosettes or scattered spots, where the patterning then tends to converge towards one of the colour forms described above, otherwise the spots can fuse into longitudinal stripes with varying degrees of spotting and rosettes. It appears that the marleyi colour form is a type of paedomorphosis, in that the juvenile colouration is retained throughout the life of the animal. Furthermore, hatching studies undertaken for this species by Dr M. J. Smale of the Port Elizabeth Museum, show that P. pantherinum invariably hatches with a marleyi pattern (Fig. 7) in animals collected from the Port Elizabeth area (principally Algoa Bay), which apparently diffuses as the animals grow (M. J. Smale pers. comm.) and corroborates the pattern trend observed in the current study. Distribution. P. pantherinum is endemic to South Africa, predominantly on the south and southeast coasts, but also ranging into the warm sub-tropical waters of kwaZulu-Natal, at least to Durban (Fig. 8). Fowler (1925) reported a specimen from off the Tugela River (29��13.5���S31��28���E) and P. pantherinum is unlikely to occur north of this, however, it may range further north of Cape Town on the west coast, but probably not as far north as P. africanum might. Compagno et al. (1989) report this shark from Saldanha Bay (32��58.5���S17��51���E). Gilchrist (1922) described this species as uncommon in Table Bay, although it was recorded from there (in Granger Bay), in this study. Like P. africanum, P. pantherinum prefers shallow inshore waters with a rocky substratum (Wallace et al. 1984; Compagno et al. 1989; current study). The greatest verified depth for this species from specimens examined in this study was 49 m, although it almost certainly occurs deeper than this. Smith (1949) reports this species as occurring to 150 fathoms (~274 m), and Fowler (1925) reports a specimen caught in a Cape trawl at 140 fathoms (~256 m). Lampe (1914) refers to a specimen (70 cm long) under the name Scyllium africanum var. punctata in the Berliner Zoologischen Museum (now the Humboldt Museum) recorded from Mauritius. There are three specimens with the name P. africanum var. striata in that collection, two specimens in ZMB 18973, and one specimen in ZMB 19145, however, the locality given for both lots is Simonstown (False Bay, Western Cape). Bleeker (1878) (which Bass et al. 1975 incorrectly refer to as Bleeker (1879)) also lists Scyllium pantherinum and S. variegatum, by name only, as occurring in Mauritius. A specimen exists in the Zoological Museum, Hamburg, recorded from Mauritius that may be referable to this species (see account for P. africanum). It is highly unlikely that Poroderma occurs in tropical waters, and even less likely that Poroderma occurs in Mauritius. Sauvage (1891) listed Scyllium pantherinum and S. variegatum, by name only, as occurring in Madagascar along with S. africanum. It is unlikely that this species occurs east of the Mozambique Channel and the locality records for the specimens examined by Sauvage (if any in the case of P. pantherinum specimens) had erroneous data. Etymology. Not provided by either Smith (1837) or M��ller and Henle (1838-1841), but likely to be derived from the Latin word panthera, meaning cat, and is likely to be a reference to the colour pattern of the holotype, consisting of panther like spots, rather than a reference to the placement of this species within the catsharks. Common name. P. pantherinum is most commonly known as the leopard catshark. Study material. A, Published as part of Brett A. Human, 2006, A taxonomic revision of the catshark genus Poroderma Smith, 1837 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae)., pp. 1-32 in Zootaxa 1229 on pages 13-24

Haematophagous larvae of a gnathiid isopod were collected from the gills, nares and buccal cavity of a single leopard catshark Poroderma pantherinum (Smith, 1838) at Jeffreys Bay and five puffadder shysharks Haploblepharus edwardsii (Voight, 1832) and one blackspotted electric ray Torpedo fuscomaculata Peters, 1855, at the De Hoop Nature Reserve on the South African south coast. Larvae were kept in fresh seawater until their moult into adult stages. The morphology of the adult males did not conform to that of any known species and they are therefore described as Gnathia pantherina sp. n. The descriptions of the adult male, female and praniza larva are based on light and scanning electron microscopy observations. Characteristic features of this species include the large size of all the final life-cycle stages, the deeply divided mediofrontal process of the male, the morphology of the pylopods and maxillipedes of the female, and the number of teeth on the mandibles (eight) and maxillules (seven) of the praniza larvae.

This paper examines the alpha level taxonomy of the genus Poroderma Smith, 1837 (Chondrichthyes: Scyliorhinidae), which is endemic to South Africa. A novel species identification key is presented for Poroderma, and a comprehensive review of the taxonomy (including type material and synonyms) and distribution of all taxa is presented. Conoporoderma Fowler, 1934 is shown to be a junior synonym of Poroderma Smith, 1837, P. marleyi Fowler, 1934 is shown to be a junior synonym of P. pantherinum, and a new colour form of P. pantherinum is described. This review is the first to offer a comprehensive description of the colour variation exhibited by P. pantherinum. The genus Poroderma contains the contemporary taxa P. africanum (Gmelin, 1789) and P. pantherinum (Smith in Muller and Henle, 1838).