Poroderma pantherinum (Smith in Mueller and Henle, 1838 )

Brett A. Human, 2006, A taxonomic revision of the catshark genus Poroderma Smith, 1837 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae)., Zootaxa 1229, pp. 1-32 : 13-24

Poroderma pantherinum (Smith in Mueller and Henle, 1838)

(Figure 3-7, Table 2)

Scyllium pantherinum Smith in Mueller 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 Mueller and Henle, 1838: 13 ( Fowler 1941: 38 attributes this name to Van Horst in Müller and Henle, although no reference to Van Horst is given in Müller and Henle).

Scyllium maeandrinum von Rapp in Mueller and Henle, 1838: 13 .

Scyllium variegatum Smith in Mueller and Henle, 1838: 14 ; Smith 1849: 11; Duméril 1853: 83; Bleeker 1878: 1, 8; Sauvage 1891: 511.

Scyllium pantherinum : Smith 1849: 13; Duméril 1853: 82; Bleeker 1878: 1, 8; Sauvage 1891: 511.

Scyllium africanum : Günther 1870: 405; Lampe 1914: 209.

Scyllium africanum var. pantherina : Günther 1870: 405; Lampe 1914: 212.

Scyllium africanum var. striata : Günther 1870: 405; Lampe 1914: 212.

Scyllium africanum var. variegata : Günther 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 Müller and Henle (1838 -1841) as Scyllium pantherinum ( Bigelow and Schroeder 1948; Compagno 1988).

Müller 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 Mueller and Henle (ex Smith) 1838 [and is] permanently invalid, preoccupied by S. pantherinum Mueller and Henle, 1838 ”. The author could not find any other reference to this ambiguity and disagrees with Eschmeyer (1998) on this point.

Günther (1870) and Gilchrist (1902) synonymised both Scyllium pantherinum and S. variegatum with P. africanum . Günther 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 ( Günther 1870). It seems that the Poroderma with stripes referred to by Günther 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’S 31°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’S 17°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 Müller 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. ANSP 53427 , holotype for Poroderma marleyi Fowler, 1934 , juvenile female 225 mm TL, “Natal coast, in 20 fathoms” , South Africa, collected by H.W. Bell Marley ; BAH 20000630.1 , juvenile female 420 mm TL, Black Sophie , Western Cape, 34°37'S 19°02'E ; BAH 20010000.7 , mature male 700 mm TL, De Monde, Struis Bay , Western Cape, approx. 34"42.8'S 20"07.3'E ; BAH 20010000.8 , adolescent female 641 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.9 , mature male 638 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.14 , mature female 589 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.15 , mature male 654 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.16 , adolescent male 672 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.17 , juvenile male 519 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.18 , mature male 666 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.19 , mature female 566 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.20 , adolescent female 566 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.21 , juvenile female 448 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.22 , mature female 577 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010000.23 , mature female 585 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010308.1 , mature female 584 mm TL, Klein Bay, Gans Bay , Western Cape, 34°35.1'S 19°20.7'E ; BAH 20010700.1 , mature male 685 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.2 , mature female 653 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.3 , mature female 618 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.4 , gravid female 640 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.5 , mature male 700 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.6 , gravid female 625 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.7 , mature male 654 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.8 , mature female 573 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.9 , mature male 692 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.10 , mature male 701 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.11 , mature male 680 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20010700.12 , mature male 685 mm TL, De Monde, Struis Bay , Western Cape ; BAH 20011119.1 , adolescent male 573 mm TL, Long Beach, Simonstown, False Bay , Western Cape, approx. 34"11.15'S 18"25.6'E ; BAH 20011119.2 , adolescent female 454 mm TL, Long Beach, Simonstown, False Bay , Western Cape ; BAH 20011119.3 , gravid female 620 mm TL, Long Beach, Simonstown, False Bay , Western Cape ; BAH 20011119.4 , juvenile male 390 mm TL, Long Beach, Simonstown, False Bay , Western Cape ; BAH 20011119.5 , mature female 552 mm TL, Millers Point, False Bay , Western Cape, approx. 34"14'S 18"28.6'E ; BAH 20011119.15 , juvenile female 353 mm TL, Long Beach, Simonstown, False Bay , Western Cape ; BAH 20020000.1 , adolescent female 471 mm TL, Western Cape ; BAH 20020000.2 , juvenile male 449 mm TL, Western Cape ; BAH 20020000.3 , adolescent female 477 mm TL, Western Cape ; BAH 20020000.4 , mature female 635 mm TL , BAH 20020000.5 , mature female 581 mm TL, Western Cape ; BAH 20020117.1 , adolescent female 588 mm TL, Granger Bay, Table Bay , Western Cape, approx. 33°54'S 18°24'E ; BAH 20020129.1 , juvenile female 485 mm TL, Millers Point, False Bay , Western Cape, approx. 34"14'S 18"28.6'E ; BAH 20020207.1 , juvenile male 424 mm TL, Millers Point, False Bay , Western Cape ; BAH 20020207.2 , mature female 614 mm TL, Millers Point, False Bay , Western Cape ; BAH 20020211.2 , mature male 610 mm TL, Granger Bay, Table Bay , Western Cape ; BAH 20020215.3 , mature male 607 mm TL, Buffels Bay, False Bay , Western Cape, approx. 34°19.3'S 18°28.1'E ; BAH 20020304.7 , mature female 565 mm TL, Hamburg , Eastern Cape, 33°17.2'S 27°28.9'E ; BAH 20020305.3 , mature female 650 mm TL, Mpekweni , Eastern Cape, 33°26.3'S 27°13.9'E ; BAH 20021000.1 , adolescent male 496 mm TL, Western Cape ; BAH 20030107.3 , mature male 675 mm TL, Jeffreys Bay , Eastern Cape, approx. 34"03.3'S 24"55.5'E ; BAH 20030107.4 , mature female 583 mm TL, Jeffreys Bay , Eastern Cape ; BAH 20030107.5 , adolescent female 560 mm TL, Jeffreys Bay , Eastern Cape ; BMNH 1845.7.3.130 , holotype for Scyllium variegatum Smith in Mueller and Henle, 1838 , mature female 612 mm TL, collected from Algoa Bay , Eastern Cape, although given as “Vom Cap” in the original description; dried mounted skin in fair condition, although the colour pattern has completely faded ; BMNH 1845.7.3.145, holotype for Scyllium pantherinum Smith in Mueller and Henle, 1838 , see under Type Series and Locality for details ; DAE 871903-1 , mature female 508 mm TL (recorded as 513 mm TL) ; LJVC 820816 , mature female 590 mm TL, West Beach, near Kleinemonde River , Eastern Cape, approx. 33°35'S 27°00'E ; LJVC 820927 , adolescent female 475 mm TL, Bushmans River, Kenton on Sea , Eastern Cape, approx. 33°41.7'S 26°39.7'E ; LJVC 910923 , 2 specimens, both juvenile males 328 mm and 408 mm TL, R.V. Africana Cruise 95, A12048 1046-044 , 34°07'S 22°26'E ; MJS uncat., 2 specimens, juvenile male 313 mm TL, adolescent female 507 mm TL, Algoa Bay , Eastern Cape, approx. 33"45'S 25"54'E ; MJS 851002 , juvenile female 94 mm TL, Eastern Cape ; MJS 860628 , mature female 620 mm TL, Cape Recife , Eastern Cape, approx. 34°01.7'S 25°42.1'E ; MJS 880627 , mature male 704 mm TL, Noordhoek , Eastern Cape, approx. 34°02'30"S 25°38'30"E ; MJS 940831 , juvenile female 257 mm TL, Algoa Bay , Eastern Cape ; MJS 941002 , mature male 664 mm TL, Algoa Bay , Eastern Cape ; MJS 941004 , mature female 630 mm TL, Cape Recife , Eastern Cape ; MJS 950201 , gravid female 655 mm TL, Algoa Bay , Eastern Cape ; MJS 950623 , 2 specimens, mature females 531 mm and 585 mm TL, near Struis Bay , Western Cape ; MJS 950913 -14, mature female 610 mm TL, Algoa Bay , Eastern Cape ; MJS 960423 , mature male 635 mm TL, Algoa Bay , Eastern Cape ; MJS 960522 , 3 specimens, mature females 601 mm and 650 mm TL, mature male 429 mm TL, near Struis Bay , Western Cape ; MJS 960525 , mature female 540 mm TL, Pringle Bay, False Bay , Western Cape, approx. 34°20.4'S 18°49.5'E ; MJS 970212 , juvenile female 223 mm TL, Algoa Bay , Eastern Cape ; MJS 981006 , mature male 720 mm TL, Bird Island, Algoa Bay , Eastern Cape, approx. 33°50.4'S 26°17.2'E ; MJS 020725 , juvenile female 165 mm TL, Algoa Bay , Eastern Cape ; RUSI 6002 , adolescent male 568 mm TL, “Natal” ; RUSI 10737 , mature female 507 mm TL, Umhlanga Rocks , kwaZulu-Natal, approx. 29°43'S 31°04'E ; RUSI 11960 , mature male 774 mm TL, Storms River Mouth , Eastern Cape, approx. 34°01'S 23°54'E ; RUSI 16728 , mature male 634 mm TL, Durban , kwaZulu-Natal, approx. 29°51'S 31°00'E ; RUSI 17326 , adolescent female 430 mm TL, Port St Johns , Eastern Cape, 31°35.8'S 29°32.9'E ; RUSI 25217 , mature female 627 mm TL, Oyster Bay, Cape St Francis , Eastern Cape, 34°10'S 24°39'E ; RUSI 25921 , mature male 733 mm TL, Cape Recife , Eastern Cape ; RUSI 25928 , mature female 545 mm TL, Shelly Beach, Port Alfred , Eastern Cape, 33°36'S 26°53'E ; RUSI 26441 , mature female 648 mm TL, Monteko Point , Eastern Cape, 31°34.9'S 29°37.6'E ; RUSI 26442 , adolescent female 479 mm TL, Monteko Point , Eastern Cape ; RUSI 26443 , mature female 572 mm TL, Monteko Point , Eastern Cape ; RUSI 27649 , 2 specimens, adolescent female 516 mm TL, mature female 640 mm TL, R.V. Africana Cruise 56, A6186 3063-002 , 35°04'18"S 20°21'24"E ; RUSI 48495 , 2 specimens, juvenile male 100 mm TL, juvenile female 113 mm TL, East London , Eastern Cape, approx. 33°02'S 27°54'E ; RUSI 53684 , mature female 523 mm TL, Port St Johns , Eastern Cape, 31°38'S 29°32'E ; SAM 22987 , 1 specimen, Gonubie, East London , Eastern Cape, 32°56.66'S 28°2.18'E ; SAM 24544 , 1 specimen ; SAM 28625 , juvenile female 356 mm TL, False Bay , Western Cape ; SAM 28626 , juvenile male 338 mm TL, False Bay , Western Cape ; SAM 28627 , juvenile male 289 mm TL, False Bay , Western Cape ; SAM 28628 , 1 specimen ; SAM 28629 , adolescent female 434 mm TL, False Bay , Western Cape ; SAM 28630 , adolescent male 469 mm TL, False Bay , Western Cape ; SAM 28631 , mature male 641 mm TL, False Bay , Western Cape ; SAM 28632 , juvenile male 389 mm TL, False Bay , Western Cape ; SAM 29305 , 1 specimen, Cape Peninsula , Western Cape, 34°21'4.5"S 18°29'8.5"E ; SAM 29944 , 1 specimen, Danger Point , Western Cape, 34°37.4'S 19°18.8'E ; SAM 33041 , mature male 610 mm TL, Pringle Bay, False Bay , Western Cape ; SAM 34378 , 1 specimen, Millers Point, False Bay , Western Cape ; SAM 35275 adolescent female 430 mm TL, Buffels Bay, False Bay , Western Cape .