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Haploblepharus edwardsii (Schinz, 1822)
(Fig. 3, Table 1)
Squalus edwartsii Schinz, 1822 ZBK : 214 (in van Oijen, 2001).
Scyllium edwardsii : Cuvier, 1817: 124 (in van Oijen, 2001); Müller & Henle, 1838: 4; Duméril , 1853: 79; Günther , 1870: 401; Gilchrist, 1902: 164; Lampe, 1914: 213; Bass et al., 1975: 17.
Scyllium edwardsii : Voigt, 1832: 504 (in van Oijen, 2001); Eschmeyer, 1998, CD-ROM.
Catulus edwardii ZBK : Smith, 1837: 85.
Scyliorhinus edwardsii : Regan, 1908: 463.
Scylliorhinus edwardsi : Thompson, 1914: 137.
Scylliorhinus edwardsii : Gilchrist & Thompson, 1916: 283; Gilchrist, 1922: 4; Barnard, 1925: 41.
Type series and locality. Neotype , designated herein, SAM 36079 (previously BAH 20021015.1) , mature male 391mm TL, collected from Millers Point, False Bay , Western Cape , South Africa , 34°14’S18°28.6’E , collected by Brett Human on 15th October 2002 , in excellent condition (Fig. 3).
Diagnosis. Haploblepharus edwardsii is distinguished from its congeners by having a slender body at all stages of maturity, abdomen width 12.1% TL for the neotype (mean 11.1% TL); neotype with snout acutely rounded, often coming to a point in other specimens, head width at the posterior margin of the orbit 11.9% TL in the neotype (mean 11.6% TL); head strongly depressed, head height at the posterior margin of the orbit 6.6% TL for the neotype (mean 5.9% TL); trunk depressed, trunk height 9.1% TL and trunk width 12.8% TL in the neotype (mean 8.5% TL and 11.7% TL, respectively); claspers of mature males slender, inner length 8.2 times the base in the neotype (mean inner length 6.2 times base). The neotype of H. edwardsii has 58 rows of teeth in the upper jaw (mean 58.3) and 46 rows of teeth in the lower jaw (mean 52.4). The neotype of H. edwardsii has a total of 121 (mean 128.9) vertebral centra. Haploblepharus edwardsii always has eight to, occasionally, ten distinct saddles that have orange centres with dark borders.
Description. Morphometric and meristic data are given in Table 1. Neotype, mature male 391mm TL (mean of all specimens examined in Table 1, including the neotype, see Study material): H. edwardsii is a slender bodied Haploblepharus ZBK shark with a relatively narrow head, head width at the pectoral origin 3.67 (3.08) times the preoral length; head length 1.22 (1.27) times distance from snout tip to first gill slit; height of first gill slit 1.44 (1.82) times the height of the fifth gill slit; eye length 4.0 (5.0) times longer than spiracle length; basimandibular cartilage found at the symphysis of the Meckels cartilage in the lower jaw; mouth length 1.0 (1.0) times the prenarial length; mouth width 5.8 (6.46) times the upper labial furrow length; labial cartilages present; nasal lobes fused into a nasal flap that covers the excurrent apertures and extends to the mouth; interorbital width 1.18 (1.25) times the nasal flap width; head strongly depressed, head width at the posterior margin of the orbit 1.80 (1.97) times its height; head width 1.45 (1.58) times its height; trunk strongly depressed, trunk width 1.41 (1.38) times its height; abdomen depressed, abdomen width 1.25 (1.21) times its height; tail slightly depressed, tail width 1.17 (1.04) times its height; caudal peduncle strongly compressed, caudal peduncle width 0.51 (0.63) times its height; precaudal length 1.71 (1.74) times the distance from snout to first dorsal fin; dorsal fins rounded; height of first dorsal fin 0.87 (1.06) times that of the second dorsal fin; first dorsal fin length 1.05 (0.96) times the length of its anterior margin; second dorsal fin length 0.97 (0.98) times the length of its anterior margin; pectoral fin to pelvic fin space 1.31 (1.33) times the interdorsal space; pectoral and pelvic fins rounded; pectoral fin height 1.86 (1.77) times the height of the pelvic fin; pectoral fin length 0.91 (0.91) times the length of its anterior margin; pelvic fin length 1.40 (1.46) times the length of its anterior margin; claspers of mature males slender, clasper inner length 8.2 (6.2) times the base; anal fin to caudal fin space 1.35 (1.54) times the head height at the origin of the pectoral fin; length of anal fin base 1.36 (1.39) the length of the second dorsal fin base; anal fin length 1.53 (1.49) times the length of its anterior margin; distance from pectoral fin insertion to the midpoint of the first dorsal fin length 1.7 (1.67) times the caudal dorsal margin length. Vertebral counts: total 121 (128.9), 33 (33-40) monospondylous, 54 (53-61) precaudal diplospondylous and 34 (33-42) caudal diplospondylous vertebrae. Dental formula: upper jaw (left) 28 (26-35), (right) 30 (25-33); lower jaw (left) 23 (23-30), (right) 23 (23-30). Spiral valve turns: NA (7).
Size and sexual maturity. In this study, H. edwardsii males were found to be juvenile at 305mm TL, adolescent at 462mm TL, and mature at 365mm TL to 555mm TL. Females were found to be mature at 365mm TL to 554mm TL and no juvenile or adolescent females were available for examination in this study. There is no sexual dimorphism apparent in this species. There is a large size range in the mature individuals for both sexes. Haploblepharus edwardsii apparently grows to a smaller total length west of Cape Agulhas (about 482 mm TL for males and about 477 mm TL for females, from the data in the current study) and to a much larger maximum size east of this point (see maximum sizes above). This size difference may be due to the deeper habit of H. edwardsii east of Cape Agulhas (see Distribution for Haploblepharus ZBK above).
Colouration. A stunningly patterned shark (Fig. 3); H. edwardsii has a background dorsal colouration pale brown, pale grey brown to dark grey brown; blanketed with many small white spots; with 8, 9, or occasionally10 prominent saddles, 2 or 3 (or 4) before the first dorsal origin, one on the first dorsal fin, one between the dorsal fins, one on the second dorsal fin, one on the caudal peduncle and two on the caudal fin; centre of saddles are brilliant orange, orange brown or dull brown (latter probably a preservation artefact), with dark brown to almost black margins anteriorly and posteriorly; background colouration extending to dorsal surface of pectoral and pelvic fins, although white spots usually larger there than on the body; saddles not extending to dorsal surface of pectoral and pelvic fins; some individuals with no markings on any fins (probably preservation artefact). Ventral colouration uniform white or off white to dull brown (latter probably a preservation artefact); ventral surfaces of pectoral and pelvic fins, as well as anal fin, with same colouration as the dorsal background, particularly in darker specimens, otherwise no distinct patterning on pectoral or pelvic fins; anal fin variably with or without patterning, fin webs slightly darker, the lack of patterning is a probable preservation artefact.
Comparison with other species. Haploblepharus edwardsii has a consistent colour pattern between individuals, always possessing bright orange saddles on a paler background. The head is more narrowly pointed than in H. fuscus ZBK and H. pictus , about the same in H. kistnasamyi . Head and body more depressed in H. edwardsii than in H. kistnasamyi , but less so than H. fuscus ZBK and H. pictus . Claspers of mature males are more slender than in other members of the genus. Haploblepharus edwardsii has a dental formula about equivalent to H. kistnasamyi , but has fewer teeth than either H. fuscus ZBK or H. pictus . Haploblepharus edwardsii is most similar to H. kistnasamyi in overall morphology and colouration, but is separated from that species by having a more depressed body and a colour pattern that is distinct from that species.
Remarks. Edwards (1764) described and figured three individuals referable to H. edwardsii , from the Cape of Good Hope, South Africa (van Oijen, 2001). These specimens, apparently lost (Smith, 1950; van Oijen, 2001), represent the syntypic series for H. edwardsii , and not RMNH 4161-4164, as stated by Eschmeyer(1998), which may represent the type series for Scyllium pictum ZBK (van Oijen, 2001).
According to van Oijen (2001), Cuvier described this species in 1817 from a reproduction of the illustrations and translation of the text of Edwards (1764) by Houttuyn (1776). However, Cuvier did so under the name “sq. d’Edwards”, which is a vernacular name, therefore is not available as a species name, as was also concluded by Compagno (1984b). Voigt (1832) translated Cuviers’ text and formalised the name, and all subsequent authors have attributed the name to Voigt. However, Schinz had already translated Cuviers’ text and formalised the species name in 1822, a document that had been overlooked until the study of van Oijen (2001), and since Schinz pre-dates Voigt, the name edwardsii must be attributed to Schinz (ICZN, 2003).
Schinz erroneously spelt Cuviers’ new species and described it under the name Squalus edwartsii ZBK (van Oijen, 2001). The name edwartsii has been placed on the Official Index of Rejected and Invalid Specific Names in Zoology, with edwardsii placed on the Official List of Specific Names in Zoology (Opinion 2056, ICZN, 2003).
This species is commonly misidentified with H. pictus and many registered specimens examined in this study that were identified as H. edwardsii were actually H. pictus . This may have a number of consequences, which includes erroneous abundance estimates for H. edwardsii (which may be overestimated) and H. pictus , as well as confounding other records of this species, such as locality and catch records, and biological and life history observations. For example, under his account of H. edwardsii , Smith (1949) illustrated H. fuscus ZBK (which he described the following year) and gave the distribution as ranging to Port Nolloth, which is almost certainly referable to H. pictus . Furthermore, Günther (1870), also under the name H. edwardsii , included H. pictus as a synonym of this, however gave a colour description that is referable to H. fuscus ZBK .
A contributing factor to the confusion between H. edwardsii and H. pictus may be the illustrations that have been used for H. edwardsii . Although the original illustrations by Edwards have not been examined by the author, the earliest illustration examined by the author was that by Müller & Henle (1838-1841), which may be the first illustration of a mature specimen and is one of the most accurate in terms of the colour for H. edwardsii , as was also noted by Bass et al. (1975). Bass (1986) illustrated his description of H. edwardsii with a line drawing of H. kistnasamyi , and although the illustrations within the colour plates appear to be based on H. edwardsii , the colours are not accurate. Given the wide usage of this book, these illustrations have probably been a major contributor to the misidentification problem. Recent illustrations that accurately depict the colouration of H. edwardsii are provided in Heemstra & Heemstra (2004: 66, although the body shape appears to be based on H. kistnasamyi ) and Compagno et al. (2005: pl. 40).
Another contributing factor to the misidentification of H. edwardsii with H. pictus is the position of the first dorsal fin relative to the pelvic fin ( Müller & Henle, 1838-1841; Bass et al., 1975), which has been used as a diagnostic character in species identification keys. The position of the origin of the first dorsal fin in relation to the insertion of the pelvic fin is variable in both species (Bass et al., 1975; present study) and the relative position of these fins is not a reliable character for separating these two species.
The morphometric proportions given here agree with Bass et al. (1975), although their proportions are confounded by the presence of H. kistnasamyi specimens in their measurements. Bass (1973) noted that only a few measurements examined by him displayed ontogenetic allometry, and most measurements were isometric.
The denticles of H. edwardsii have been described and illustrated by Smith (1950), Bass et al. (1975), and Compagno (1984b). A number of studies have found strong sexual heterodonty in this species (Bass et al., 1975; M. Marks, pers. comm.). Vertebral counts of H. edwardsii are slightly higher in Bass et al. (1975) than those given here (mean total count 128.9, n = 12), and is probably due to the inclusion of smaller specimens here. The counts given here agree with that obtained by Springer & Garrick (1964).
Haploblepharus edwardsii are known to be predated upon, and played with, by cape fur seals ( Arctocephalus pusillus pusillus ), and that kelp gulls ( Larus dominicanis vetula ) opportunistically scavenge H. edwardsii from the fur seals (Martin, 2004). Haploblepharus edwardsii are commonly caught by recreational anglers from the shore and are also commonly sighted by SCUBA divers (Bass et al., 1975; Compagno, 1984b; Compagno & Smale, 1989; pers. obs.). This species also forms part of the bycatch of demersal trawlers (Compagno, 1984b; Compagno & Smale, 1989), and are caught for the aquarium trade (pers. obs.). Haploblepharus edwardsii , along with H. pictus , have recently been reported as hosting the first blood parasite described in chondrichthyan fishes from South African waters (Yeld & Smit, 2006).
Distribution. Haploblepharus edwardsii has been verified in the current study as occurring from Langebaan Lagoon, Western Cape in the southeast Atlantic, southeast to Cape Agulhas and into the Indian Ocean as far east as the western shore of Algoa Bay (Fig. 4). Previously, it was thought that H. edwardsii ranged as far north as Durban (Bass et al., 1975; Compagno, 1984b, 1988; Compagno et al., 1989), however, these distributions included H. kistnasamyi and almost certainly included misidentifications of H. fuscus ZBK and H. pictus . Records of “Cape” H. edwardsii from East London (Bass et al., 1975) could either be this species, or juveniles of H. kistnasamyi .
Martin (2004) reported a westward range extension into False Bay, Western Cape, for H. edwardsii . However, the occurrence of H. edwardsii in False Bay has been previously reported by Müller & Henle (1838- 1841), Günther (1870), Gilchrist (1902), Lampe (1914), Thompson (1914), Barnard (1925), Bass et al. (1975), Springer (1979), Bass (1986), Compagno (1988), Smith & Griffiths (1997), Branch et al. (1999), and van Oijen (2001).
Smith (1950) reports this species from as far north in the southeast Atlantic as Port Nolloth, however, his description of H. edwardsii includes H. pictus , which he synonymised with this species, therefore, this record is most probably referable to H. pictus . Compagno (1988) examined a specimen of H. edwardsii AMNH 40988 that was labelled as coming from Congo, and as Compagno reasoned, this is probably erroneous.
Etymology. Named after G. Edwards, who was the first to illustrate this species in 1764 (van Oijen, 2001).
Common name. In an attempt to introduce species specific common names for this genus that will be easily remembered by the general public, this shark was given the name happy Eddie (Compagno & Human, 2003), which is the name referred to this shark by academics familiar with it. It is also known as the puffadder shyshark, and in Afrikaans as a skaamoog (shyshark).
Study material. BAH 19991222.02 , gravid female 540 mm TL, Buffels Bay, False Bay , Western Cape , South Africa , 34°19.3'S18°28.1'E ; BAH 20000304.01 , mature female 415 mm TL, Rooi Els, False Bay , Western Cape , South Africa , 34°18'S18°48.8'E ; BAH 20000930.03 , mature male 414 mm TL, Buffels Bay, False Bay , Western Cape , South Africa ; BAH 20011119.16 , gravid female 364 mm TL, Longbeach, False Bay , Western Cape , South Africa , 34°11.15'S18°25.6'E ; BAH 20020417.01 , mature female 477 mm TL, R.V. Africana Cruise 167 , A20810 007-2231, 95 m ; BAH 20021015.02 , mature female 365 mm TL, Millers Point, False Bay , Western Cape , South Africa , 34°14'S18°28.6'S ; BAH 20021017.02 , mature male 370 mm TL, Froggy Pond, False Bay , Western Cape , South Africa , 34°12.26'S18°27.56'E ; BAH 20021017.03 , mature male 390 mm TL, Froggy Pond, False Bay , Western Cape , South Africa ; RUSI 6071, previously ORI 2475 , adolescent male 462 mm TL, Algoa Bay , Eastern Cape , South Africa , approx. 34°02'S25°42'E ; RUSI 6073, previously ORI 2473 , mature male 536 mm TL, Algoa Bay , Eastern Cape , South Africa ; RUSI 13145 , mature female 522 mm TL, Plettenberg Bay , Western Cape , South Africa , approx. 34°04'S23°23'E ; RUSI 13146 , gravid female 554 mm TL, Algoa Bay , Eastern Cape , South Africa ; SAM 10141 , immature male 305 mm TL, False Bay , South Africa ; SAM 12980 , 3 specimens, Kalk Bay, False Bay , Western Cape , South Africa , 34°7.6'S18°27'E ; SAM 17758 , 2 specimens ; SAM 22021 , mature male 371 mm PCL, Simonstown Lighthouse, False Bay , Western Cape , South Africa , 34°11.1'S18°26.3'E ; SAM 23199 , 1 specimen, Cape Peninsula, False Bay , Western Cape , South Africa , 34°21'4.5"S18°29'8.5"E ; SAM 28636 , 24 specimens, those measured in the current study -mature male 365 mm TL, mature male 367 mm TL, mature male 406 mm TL, mature male 430 mm TL, mature male 472 mm TL, mature male 472 mm TL, mature male 482 mm TL, mature female 408 mm TL, gravid female 434 mm TL, R.V. Africana Cruise 120 , A47043 2220 (erroneous station number, R. Leslie, pers. comm.) ; SAM 29307 , 6 specimens, Cape Peninsula, False Bay , Western Cape , South Africa ; SAM 32524 , 1 specimen, R.V. Africana Cruise 82 , A10145 047-3415 , 35°00'S22°25'E ; SAM 32525 , mature male 555 mm TL, Storms River Mouth , Eastern Cape , South Africa , 34°01.3'S23°54.7'E ; SAM 32555 , 1 specimen, Storms River Mouth , Eastern Cape , South Africa ; SAM 32616 , 1 specimen, Langebaan Lagoon (Klein Ostervaal farm) , Western Cape , South Africa , 33°04.5'S18°02.5'E ; SAM 33203 , 1 specimen, R.V. Africana Cruise 111 , A14791 053-2160 , 34°55'48"S21°35'24" ; SAM 34640 , 1 specimen, Bloubergstrand, Cape Town , Western Cape , South Africa , 33°48.5'E18°27'E ; SAM 34647 , mature female 462 mm TL, R.V. Africana Cruise 111 , A14732 001-1002 , 34°54.9'S20°02'E ; SAM 36079, designated neotype of H. edwardsii , see under Type Series and Locality for details.