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Haploblepharus pictus ( Mueller & Henle, 1838)

(Fig. 5, Table 2)

Scyllium pictum Mueller & Henle, 1838 ZBK : 4.

Haploblepharus edwardsii : Günther , 1870: 401 (in part); Regan, 1908: 463 (in part); Garman, 1913: 102 (in part); Thompson, 1914: 137 (in part); Gilchrist & Thompson, 1916: 283 (in part); Fowler, 1941: 64 (in part); von Bonde, 1945a: 1 (in part); von Bonde, 1945b: 220 (in part); Smith, 1949: 54 (in part); Smith, 1950: 879 (in part).

Type series and locality. Probable syntypes (see Remarks below), five specimens in four lots. RMNH 4161 , two specimens, mature male 522 mm TL, and mature female 490 mm TL, both are dried specimens. The female appears to be in poor condition and the male in very poor condition. RMNH 4162 , immature female 135 mm TL, in reasonable condition. RMNH 4163 , immature male 199 mm TL, in reasonable condition. RMNH 4164 , immature female 143 mm TL, in good condition.

The entire syntypic series collected “Vom Cap der guten Hoffnung” (from Cape of Good Hope) , South Africa , 1826-1834 (Leg. & don.: H.B. van Horstok ).

Diagnosis. The author was not able to personally measure the putative type series (see remarks below), and given the highly contorted condition of those specimens, the measurements obtainable from those specimens would likely be inaccurate. The following is based on the mean measurements of H. pictus specimens examined by the author (see study material). Haploblepharus pictus is distinguished from other Haploblepharus ZBK sharks in having a slender body in juvenile individuals but becoming stockier in mature individuals, abdomen width 11.2% TL; snout broadly rounded, not coming to a point; head width at the posterior margin of the orbit 13.0% TL; head strongly depressed, head height at the posterior margin of the orbit 6.7% TL; trunk somewhat depressed, trunk height 10.2% TL and trunk width 12.3% TL; claspers of mature males stout, inner length 6.2 times the base. Haploblepharus pictus specimens have 45-83 rows of teeth in the upper jaw (mean 72.0) and 47-75 rows of teeth in the lower jaw (mean 62.8). The number of vertebral centra in H. pictus specimens ranges from 93-133 (mean 124.5) total. Haploblepharus pictus has a highly variable colour pattern that can superficially resemble H. edwardsii or H. fuscus ZBK in some individuals, although H. pictus more commonly has 6-8 distinct, dark saddles.

Description. Morphometric and meristic data are given in Table 2. The following is based on the mean measurements of H. pictus specimens examined by the author (see Study material): H. pictus is a slender to moderately stocky bodied Haploblepharus ZBK shark with a relatively broad head, head width at the pectoral origin3.63 times the preoral length; head length 1.26 times distance from snout tip to first gill slit; height of first gill slit 1.76 times the height of the fifth gill slit; eye length 4.29 times longer than spiracle length; basimandibular cartilage found at the symphysis of the Meckels cartilage in the lower jaw; mouth length 1.23 times the prenarial length; mouth width 6.6 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 1.19 times the nasal flap width; head very strongly depressed, head width at the posterior margin of the orbit 1.95 times its height; head width 1.48 times its height; trunk depressed, trunk width 1.21 times its height; abdomen not depressed, abdomen width 0.99 times its height; tail not depressed, tail width 1.04 times its height; caudal peduncle strongly compressed, caudal peduncle width 0.61 times its height; precaudal length 1.76 times the distance from snout to first dorsal fin; dorsal fins rounded; height of first dorsal fin 1.04 times that of the second dorsal fin; first dorsal fin length equal to the length of its anterior margin; second dorsal fin length equal to the length of its anterior margin; pectoral fin to pelvic fin space 1.33 times the interdorsal space; pectoral and pelvic fins rounded; pectoral fin height 1.8 times the height of the pelvic fin; pectoral fin length 0.94 times the length of its anterior margin; pelvic fin length 1.46 times the length of its anterior margin; claspers of mature males stout, clasper inner length 6.17 times the base; anal fin to caudal fin space 1.1 times the head height at the origin of the pectoral fin; length of anal fin base 1.21 the length of the second dorsal fin base; anal fin length 1.4 times the length of its anterior margin; distance from pectoral fin insertion to the midpoint of the first dorsal fin length 1.51 times the caudal dorsal margin length. Vertebral counts: total 124.5 (93-133), 35 (30-38) monospondylous, 52.2 (35-57) precaudal diplospondylous and 37.7 (28-45) caudal diplospondylous vertebrae. Dental formula: upper jaw (left) 35.6 (23-40), (right) 36.4 (22-43); lower jaw (left) 31.6 (23-37), (right) 31.2 (24-38).

Size and sexual maturity. In this study, H. pictus males were found to be embryonic at 103 mm TL, juvenile at 165 mm TL to 369 mm TL, adolescent at 282 mm TL to 407 mm TL, and mature at 400 mm TL to 570 mm TL. Females were found to be juvenile at 110 mm TL to 306 mm TL, adolescent at 430 mm TL to 533 mm TL, and mature at 359 mm TL to 597 mm TL. There is no sexual dimorphism apparent in this species.

Colouration. Haploblepharus pictus has a background dorsal colouration pale to very dark brown, almost black, pale grey brown to dark grey brown, red brown, or grey; white spots variably present (Figs. 5A, C-E) or absent (Fig. 5F), occurring on the saddles when spots present, between and on the saddles, but never just between the saddles; number of saddles, when present, 6 to 8, variably placed; centre of saddles range from orange, brown, dark brown to almost black, margins variably present, when margins present may be indistinct; background colouration extending to dorsal surface of pectoral and pelvic fins, including spots when present; saddles not extending to dorsal surface of pectoral and pelvic fins. Ventral colouration abruptly uniform white (Fig. 5B), cream or red brown (latter probably a preservation artefact); ventral surfaces of pectoral and pelvic fins, as well as anal fin, same colouration as the dorsal background colouration, occasionally dark spots and blotches present.

Comparison with other species. Haploblepharus pictus has a highly variable colour pattern and is easily misidentified for other members of the genus when using colour pattern alone. Haploblepharus pictus is a relatively large member of the genus, second to H. fuscus ZBK in overall size, and is more robust than H. edwardsii or H. kistnasamyi . The head is more bluntly rounded than in H. edwardsii and H. kistnasamyi , and about equally so with H. fuscus ZBK . Head and body is depressed more so than in H. kistnasamyi and H. edwardsii , but less so than H. fuscus ZBK . Claspers of mature males are equivalent in size to H. fuscus ZBK and H. kistnasamyi , although longer and stouter than in H. edwardsii . Haploblepharus pictus has a lower total tooth count than H. fuscus ZBK , but higher than either H. edwardsii or H. kistnasamyi . Haploblepharus pictus is most similar to H. fuscus ZBK in overall morphology, and is distinguished from that species by having distinct saddles and spots, less vertebrae, and more tooth rows. The variable colour pattern of H. pictus can superficially resemble all other Haploblepharus ZBK species, particularly H. edwardsii (Fig. 5D).

Remarks. Müller & Henle (1838-1841) refer to specimens in the Museum in Leyden (now the Rijksmuuseum van Natuurlijke Historie, Leiden, RMNH) and in the British Museum (BMNH), but give no further details of the type material. RMNH 4161 and RMNH 4164 bear the name Scyllium edwardsii , while RMNH 4162 and RMNH 4163, have two names written on the label, Scyllium edwardsii and Scyllium pictum ZBK . All four lots were in the museum during the time of Müller and Henles’ visit to the museum (M. van Oijen, pers. comm.). Photographs of these specimens (supplied by M. van Oijen) were examined by the author. The specimens are in poor condition, and although the colour patterning on most of the specimens are considerably faded, saddles are visible on all specimens, and all specimens appear to be H. pictus . Although there is no label identifying them as such, the RMNH specimens are probably the syntypes for H. pictus . Of the scyliorhinid specimens in the BMNH that have been assigned a species name, none are referable to H. pictus . As there are many lots in the BMNH under the name Scyllium ZBK sp., it was not practical to examine all of these specimens remotely, and the status of types for H. pictus in the BMNH remains unresolved.

Eschmeyer (1998) refers to two possible syntypes in the Naturhistorisches Museum, Vienna, Lot NMW 78529, represented by an immature female (NMW 78529:1 approx. 200 mm TL) and an immature male (NMW 78529:2, approx. 110 mm TL). These specimens were purchased in 1842 from Ludwig Parreyss by the Naturhistorisches Museum, and may have been acquired from the museums in Leiden or London, however there are no labels accompanying the specimens indicating from where the specimens were procured, or whether they were used as type material. If these specimens did come from either the Leiden or London museums, then the specimens were probably available at the time Müller and Henle visited those museums. The only label accompanying the specimens currently, is a label dated at the time Steindachner was at the museum, which was some time after the specimens were deposited there (H. Wellendorf, pers. comm.). Photos taken of these specimens (supplied by H. Wellendorf) were examined by the author and identified as H. pictus , however, NMW 78529 probably does not represent type material. Springer (1979) made no reference to type material for H. pictus , and Compagno (1984b, 1988) did not locate type material for H. pictus .

Since the description of H. pictus by Müller & Henle (1838-1841), many authors have chosen to synonymise it with H. edwardsii (e.g., Fowler, 1941; Smith, 1950), and this was due, in the most part, to the loss of the syntypes of H. edwardsii . Bass et al. (1975) revived H. pictus , which has been followed by subsequent authors. Bass et al. (1975) report that Squalus pictus ZBK described by Lichtenstein in 1823 is probably not H. pictus and may or may not be referable to H. edwardsii . There is no type material for this species and the description is insufficient to unambiguously identify this species.

Haploblepharus pictus is regularly misidentified, mostly for H. edwardsii (Fig. 5D), but also for H. fuscus ZBK (Fig. 5F). This misidentification has led to an easterly range extension in its distribution (Fig. 6, and see below). The problems with misidentification may also underestimate the abundance of this species and overestimate the abundance of H. edwardsii . It is hoped that the current revision will aid in the identification of these species in the field and allow for a more accurate representation of distribution and abundance for both H. pictus and H. edwardsii .

The morphometric proportions given here agree with those given by Bass et al. (1975) and Springer (1979). The dentition and denticles have been described and illustrated by Bass et al. (1975) and Compagno (1984b). Bass et al. (1975) found that there was strong sexual heterodonty in H. pictus . Vertebral counts given here agree with Bass et al. (1975) and Springer (1979).

Haploblepharus pictus has a highly variable colour pattern (Fig. 5), some of these stunning, with no identifiable forms, as opposed to Poroderma pantherinum for example, which also has a highly variable colour pattern, but with identifiable forms (Compagno et al., 1989; Human, 2006a). Saddles are rarely absent, and some specimens resembling H. edwardsii has led to the misidentification of H. pictus as that species.

Yeld & Smit (2006) have identified a blood parasite in this species (along with H. edwardsii ) that constitutes the first record of a blood parasite from a South African chondrichthyan, and H. pictus is the type host for Trypanosoma haploblephari Yeld & Smit 2006 (Mastigophora: Trypanosomatidae).

Distribution. Figure 6 shows the distribution of H. pictus as verified by this study. Haploblepharus pictus occurs north of Lüderitz , Namibia, extending south into South Africa to Cape of Good Hope, then east to into the Indian Ocean, east of the Storms River mouth, Eastern Cape.

The present revision of Haploblepharus ZBK has led to a major range extension for H. pictus , which was previously thought to occur only as far east as Cape Agulhas (Bass et al., 1975; Compagno, 1988; Compagno et al., 1989). The previous distribution of H. pictus was due to misidentification of this species with H. edwardsii and H. fuscus ZBK , the former in particular, east of Cape Agulhas.

Haploblepharus pictus seems to be more abundant west of Cape Agulhas (which makes it somewhat surprising that it was not recorded by Clark (1997)), although it appears not to be uncommon east of Cape Agulhas. Dr. P. D. Cowley (SAIAB) has shown the author photographs of a number of live Haploblepharus ZBK specimens from Titsikama National Park, Eastern Cape, with colour patterns that resemble H. edwardsii , however, these specimens proved to be H. pictus .

Etymology. Not provided by Müller & Henle in their original description, however, the Latin word, pictus, means painted or coloured and is a clear reference to the variegated colour pattern of this species.

Common name. The common name, pretty happy, was introduced in Compagno & Human (2003). It is otherwise known as the dark shyshark, or skaamoog in Afrikaans.

Study material. BAH 20000617.08 , male 560 mm TL, Millers Point, False Bay , Western Cape , South Africa , 34°14'S18°28.6'S ; BAH 20000617.09 , male 700 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20001007.03 , male 545 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20010308.04 , 1 specimen, Klein Bay, Gans Bay , Western Cape , South Africa , 34°35.1'S19°20.7'E ; BAH 20010405.06 , male 595 mm TL, Granger Bay, Table Bay , Western Cape , South Africa , 33°54'S18°24'E ; BAH 20010405.07 , male 489 mm TL, Granger Bay, Table Bay , Western Cape , South Africa ; BAH 20010405.08 , male 580 mm TL, Granger Bay, Table Bay , Western Cape , South Africa ; BAH 20010405.09 , male 641 mm TL, Granger Bay, Table Bay , Western Cape , South Africa ; BAH 20010405.10 , female 420 mm TL, Granger Bay, Table Bay , Western Cape , South Africa ; BAH 20010405.11 , male 580 mm TL, Granger Bay, Table Bay , Western Cape , South Africa ; BAH 20011119.17 , 1 specimen, Long Beach, False Bay , Western Cape , South Africa , 34°11.15'S18°25.6'E ; BAH 20021014.01 , gravid female 433 mm PCL, Froggy Pond, False Bay , Western Cape , South Africa , 34°12.26'S18°27.56'E ; BAH 20021014.02 , adolescent female 493 mm TL, Froggy Pond, False Bay , Western Cape , South Africa ; BAH 20021014.04 , mature male 559 mm TL, Froggy Pond, False Bay , Western Cape , South Africa ; BAH 20021014.05 , mature male 540 mm TL, Froggy Pond, False Bay , Western Cape , South Africa ; BAH 20021014.06 , mature female 506 mm TL, Froggy Pond, False Bay , Western Cape , South Africa ; BAH 20021015.03 , adolescent female 496 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20021015.04 , mature male 514 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20021015.05 , mature male 570 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20021015.06 , mature male 484 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20021017.01 , mature male 517 mm TL, Froggy Pond, False Bay , Western Cape , South Africa ; BAH 20021020.01 , mature male 460 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BAH 20021020.02 , adolescent female 455 mm TL, Millers Point, False Bay , Western Cape , South Africa ; BGR.1.H , deposited at SAM , embryonic male 103 mm TL, Berg River Mouth, St Helena Bay , Western Cape , South Africa , 32°46'S18°09'E ; MJS 960526 , adolescent female 430 mm TL, Rocky Bank, False Bay , Western Cape , South Africa , 34°24.4'S18°35.5'E ; MJS 970124 , gravid female 517 mm TL, Slangkop , Western Cape , South Africa , 34°10.97'S18°18.46'E ; MJS 970128 , 3 specimens, mature male 554 mm TL, two mature females 547 mm TL and 597 mm TL, Die Kelders, Gans Bay , Western Cape , South Africa , 34°30.76'S19°20.24'E ; NMW 78529 , 2 specimens, immature male ~110 mm TL, immature female ~200 mm TL, “Cap” , South Africa ; RMNH 4161, probable syntypes, see under Type Series and Locality for details; RMNH 4162, probable syntype, see under Type Series and Locality for details; RMNH 4163, probable syntype, see under Type Series and Locality for details; RMNH 4164, probable syntype, see under Type Series and Locality for details; RUSI 4115 , juvenile male 165 mm TL, Woodstock Power Station (site no longer in existence, reclaimed land), Cape Town , Western Cape , South Africa , 33°55'S18°26'E ; RUSI 4117 , adolescent female 483 mm TL, Robben Island, Table Bay , Western Cape , South Africa , 33°48'S18°22'E ; RUSI 6083, previously ORI 6083 , mature male 565 mm TL, Cape Point , Western Cape , South Africa , 34°21'4.5"S18°29'8.5"E ; RUSI 6084, previously ORI 2874 , adolescent female 533 mm TL, Cape Point , Western Cape , South Africa ; RUSI 6160, previously ORI 2932 , juvenile male 199 mm TL, Woodstock (presently reclaimed land), Cape Town , Western Cape , South Africa , 33°55'S18°26'E ; RUSI 39993 , 3 specimens, two juvenile males 320 mm TL and 369 mm TL, mature male 537 mm TL, Platboom, Cape Point , Western Cape , South Africa , 34°20.2'S18°26.65'E ; RUSI 44494 , mature male 432 mm TL, Berg River Mouth, St Helena Bay , Western Cape , South Africa ; RUSI 48499 , gravid female 359 mm TL, Hottentot Bay , Namibia , 26°07'S14°57'E ; RUSI 48500 , mature male, Lüderitz , Namibia , 26°39'S15°09'E ; SAM 3225 , mature male, Table Bay, Cape Town , Western Cape , South Africa , 33°53.85'S18°26.1'E ; SAM 10142 , adolescent female, Saldanha Bay , Western Cape , South Africa , 32°58.5'S17°51'E ; SAM 21941 , adolescent male 282 mm TL, Simonstown Naval Dockyard, False Bay , Western Cape , South Africa , 34°11.1'S18°26.3'E ; SAM 22996 , 3 specimens, Saldanha Bay , Western Cape , South Africa ; SAM 23096 , juvenile male 335 mm TL, Salt River Power station filter screens, Table Bay , Western Cape , South Africa , 33°55'S18°26'E ; SAM 23296 , 2 specimens, juvenile male 218mm TL, juvenile female 242 mm TL, North of Orange River Mouth , Namibia (no coordinates) ; SAM 23307 , juvenile male 135 mm TL, 70 miles (112 km) North of Orange River Mouth , Namibia , approx. 28°02'S15°45'E ; SAM 23578 , embryonic male 83 mm TL, Schaapen Island, Langebaan Lagoon , Western Cape , South Africa , 33°05.6'S18°01.5’E ; SAM 23598 , 4 specimens, 2 juvenile males 128 mm TL and 276 mm TL, 2 juvenile females, 110 mm TL and 128 mm TL, Hout Bay , Western Cape , South Africa , 34°03'S18°21'E ; SAM 23811 , 1 specimen, South of Cape Columbine , Western Cape , South Africa , 32°47.5'S17°46.8'E ; SAM 24345 , 5 specimens, 2 juvenile males 235 mm TL and 268 mm TL, adolescent male 407 mm TL, mature male 516 mm TL, adolescent female 411 mm TL, Lüderitz , Namibia ; SAM 24545 , 1 specimen, Bredasdorp (erroneous, true locality probably Arniston, Struis Bay or Cape Agulhas) , Western Cape , South Africa ; SAM 26385 , juvenile female 306 mm TL, Lamberts Bay , Western Cape , South Africa , 32°04.5'S18°18.5'E ; SAM 29303 , 13 specimens, Cape Peninsula , Western Cape , South Africa , 34°21'4.5"S18°29'8.5"E ; SAM 29337 , mature female, Mowe Point , Namibia (no coordinates) ; SAM 32526 , mature female, Cape Point , Western Cape , South Africa ; SAM 32556 , mature female, Cape Point , Western Cape , South Africa ; SAM 32617 , mature female 553 mm TL, Titsikama , Eastern Cape , South Africa , 34°08.1'S24°26.4'E .

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