Haploblepharus fuscus Smith, 1950 ZBK
(Fig. 7, Table 3)
Haploblepharus fuscus Smith, 1950 ZBK : 883.
Haploblepharus edwardsii : Günther , 1870: 401 (in part); Smith, 1949: 54 (in part).
Type Series and Locality. Holotype , RUSI 21 , adolescent male 530 mm TL (measured as 495 mm TL in the current study), collected off East London , Eastern Cape , South Africa (by J. L. B. Smith? ), approx. 33°00'S27°55'E , from the shore “in shallow water among rocks” using hook and line , in good condition, although there is a large gash present on the dorsal midline of the head above gill slits 3 to 5, and there are a number of patches where the skin has been damaged, particularly on the right trunk, that may be due to poor preservation (Fig. 7).
Paratype , missing presumed lost, possibly never catalogued, 680 mm TL male, collected from Knysna , Western Cape , South Africa (by J. L. B. Smith? ), approx. 34°03'S23°02.5'E , from the shore using hook and line .
Diagnosis. The largest member of the genus, H. fuscus ZBK has a relatively stocky body at all stages of maturity, particularly in mature individuals; abdomen width 11.0% TL for the holotype (mean 11.4% TL); snout rounded in the holotype, not coming to a point, becoming more broadly rounded in larger individuals, head width at the posterior margin of the orbit 12.0% TL in the holotype (mean 12.2% TL); head strongly depressed, head height at the posterior margin of the orbit 6.8% TL for the holotype (mean 6.1% TL); trunk depressed, trunk height 8.9% TL and trunk width 12.1% TL in the holotype (mean 8.7% TL and 12.2% TL, respectively); claspers of mature males stout, inner length 6.9 times the base in the holotype (mean inner length 4.3 times base). The holotype of H. fuscus ZBK has 84 rows of teeth in the upper jaw (mean 71.2) and 83 rows of teeth in the lower jaw (mean 74.7). The holotype of H. fuscus ZBK has a total of 134 (mean 134.8) vertebral centra. Haploblepharus fuscus ZBK always has a chocolate brown or dull brown background colouration, occasionally with indistinct saddles, and occasionally with white spots present, rarely with dark spots present.
Description. The morphometric and meristic data for H. fuscus ZBK are given in Table 3. Holotype, adolescent male 495 mm TL (mean of all specimens examined in Table 3, including the holotype, see Study material): H. fuscus ZBK is a stocky bodied Haploblepharus ZBK shark, although juveniles are slender, with a relatively broad head, head width at the pectoral origin 4.06 (3.91) times the preoral length; head length 1.24 (1.28) times distance from snout tip to first gill slit; height of first gill slit 1.86 (1.82) times the height of the fifth gill slit; eye length 5.67 (4.14) times longer than spiracle length; basimandibular cartilage found at the symphysis of the Meckels cartilage in the lower jaw; mouth length 1.33 (1.47) times the prenarial length; mouth width 8.5 (6.27) 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 (1.15) times the nasal flap width; head strongly depressed, head width at the posterior margin of the orbit 1.76 (2.0) times its height; head width 1.44 (1.60) times its height; trunk strongly depressed, trunk width 1.36 (1.40) times its height; abdomen depressed, abdomen width 1.29 (1.18) times its height; tail not depressed, tail width equal to its height; caudal peduncle strongly compressed, caudal peduncle width 0.70 (0.57) times its height; precaudal length 1.86 (1.79) times the distance from snout to first dorsal fin; dorsal fins rounded; height of first dorsal fin 0.96 (1.08) times that of the second dorsal fin; first dorsal fin length equal to the length of its anterior margin; second dorsal fin length 1.08 (1.04) times the length of its anterior margin; pectoral fin to pelvic fin space 1.14 (1.28) times the interdorsal space; pectoral and pelvic fins rounded; pectoral fin height 2.18 (1.98) times the height of the pelvic fin; pectoral fin length 0.91 (0.90) times the length of its anterior margin; pelvic fin length 1.35 (1.36) times the length of its anterior margin; claspers of mature males stout, clasper inner length 6.89 (4.33) times the base; anal fin to caudal fin space 1.48 (1.42) times the head height at the origin of the pectoral fin; length of anal fin base 1.19 (1.29) the length of the second dorsal fin base; anal fin length 1.37 (1.43) times the length of its anterior margin; distance from pectoral fin insertion to the midpoint of the first dorsal fin length 1.52 (1.53) times the caudal dorsal margin length. Vertebral counts: total 134 (128-141), 34 (34-40) monospondylous, 60 (52-62) precaudal diplospondylous and 40 (35-48) caudal diplospondylous vertebrae. Dental formula: upper jaw (left) 40 (23-43), (right) 44 (23-49); lower jaw (left) 43 (23-46), (right) 40 (25-46). Spiral valve turns: NA (9).
Size and sexual maturity. Haploblepharus fuscus ZBK is the largest Haploblepharus ZBK shark, both in terms of overall size and mass. In this study, males were found to be juvenile at 438 mm TL to 460 mm TL, adolescent at 495 mm TL to 543 mm TL, and mature at 550 mm TL to 649 mm TL. Females were found to be adolescent at 496 mm TL to 568 mm TL, and mature at 609 mm TL to 631 mm TL, no female juveniles were available for examination. There appears to be no sexual dimorphism in H. fuscus ZBK .
Colouration. Haploblepharus fuscus ZBK is the least patterned of the Haploblepharus ZBK sharks (Figs. 7 & 8). The holotype is a uniform chocolate brown, slightly paler ventrally, without saddles, spots, or any other markings(Fig. 7). In other H. fuscus ZBK individuals, the background dorsal colouration is most often uniform chocolate brown or dull grey brown with no markings, although sometimes individuals have small inconspicuous black spots (rarely) or inconspicuous white spots (Fig. 8A), never both; saddles variably present and inconspicuous when present (Fig. 8B), more conspicuous in smaller specimens, centre of saddle more orange than background with darker margins anteriorly and posteriorly, colour in the centre of the saddle dull, number of saddles and position on body highly variable, usually between 2 to 4, although never on the head; one individual had white spots and indistinct saddles. No markings on any fins. Ventral colouration uniform white, off white, cream or paler than the background dorsal colouration, pectoral and pelvic fin webs darker, anal fin with dorsal background colour.
Comparison with other species. Haploblepharus fuscus ZBK is the least patterned of the Haploblepharus ZBK sharks. The head is bluntly rounded as in H. pictus , and the body is depressed more so than in H. edwardsii and H. kistnasamyi . Claspers of mature males are equivalent in size to H. kistnasamyi and H. pictus , although longer and stouter than in H. edwardsii . Haploblepharus fuscus ZBK has the highest total tooth count of the Haploblepharus ZBK sharks. Haploblepharus fuscus ZBK is most similar to H. pictus in overall morphology, and distinguished from that species by having indistinct saddles and spots, when present, with spots and saddles rarely occurring together, more vertebrae and fewer teeth rows. The colouration of H. fuscus ZBK is distinct from H. edwardsii and H. kistnasamyi (except possibly for the juveniles of the latter species, which can be distinguished from H. fuscus ZBK in possessing obvious saddles; see Fig. 13), however, H. fuscus ZBK has also been confused with H. pictus .
Remarks. Smith (1950) made his original description of H. fuscus ZBK from two males, one 530 mm TL from East London, Eastern Cape (RUSI 21) and one 680 mm TL from Knysna, Western Cape. The only reference to the designation of a type specimen made by Smith was on the illustration of the 530 mm individual, to which he labelled “type”. This, in accordance to Articles 73.1.1 and 73.1.4 of the International Commission for Zoological Nomenclature (ICZN, 1999), establishes that individual (RUSI 21) as the holotype. In accordance with Recommendation 73D (ICZN, 1999), the remaining specimen is a paratype of H. fuscus ZBK , however, from examination of the SAIAB collection catalogue, this specimen apparently no longer exists, and there are no specimens of H. fuscus ZBK in that collection with the locality of Knysna, suggesting that the paratype was never catalogued. Bass et al. (1975) and Springer (1979) refer to the two specimens described by Smith, but make no mention of the status of the paratype.
The only material examined by Springer (1979) for H. fuscus ZBK was SAM 24545, which he illustrated and gave morphometric data for. In the present study, this specimen was determined to be H. pictus , therefore the morphometric proportions given by Springer for H. fuscus ZBK are actually referable to H. pictus .
The dentition and denticles of H. fuscus ZBK are described and illustrated by Smith (1950), Bass et al. (1975), and Compagno (1984b). Bass et al. (1975) found no evidence of sexual heterodonty in H. fuscus ZBK , and found the teeth to be quite different to other Haploblepharus ZBK in being longer and narrower.
The juveniles of this species are scarce (Bass et al., 1975; M. J. Smale, pers. comm.; current study), and there appears to be an unknown habitat that is used by H. fuscus ZBK for egg laying, and where juveniles spend that stage of their life history. Only two juvenile specimens could be found in the present study. RUSI 6079 is a juvenile male 460 mm TL, collected from Port Alfred, Eastern Cape (33°36’S 26°54’E), and SAM 32614 (Fig. 8B) is also a juvenile male 438 mm TL, collected from Langebaan Lagoon, Western Cape (Klein Oostervaal farm). Both of these juveniles are much larger than the juveniles of other members of the genus.
Haploblepharus fuscus ZBK is regularly caught by shore anglers, who often regard them as pests (Compagno, 1984b; pers. obs.). This species is also occasionally used in aquaria, however, there is no directed fishery for this species for the aquarium trade at present (pers. obs.).
The biology of this species is virtually unknown, which is somewhat disturbing given that this is an inshore shark and caught with relatively high frequency by shore anglers in the Eastern Cape (pers. obs.). The gathering of biological data for this species should be considered a priority given that it is an endemic with a restricted range, with a habitat preference that is in a zone that experiences significant fishing pressure.
Distribution. Haploblepharus fuscus ZBK is verified as occurring coastally, south of latitude 33°S in South Africa (Fig. 9). Haploblepharus fuscus ZBK is most commonly found in the Eastern Cape, from Storms River mouth to East London, however it is a rare visitor to the Western Cape and has been verified from a specimen collected in Langebaan Lagoon (SAM 32614), assuming that the locality given for this specimen is accurate. Haploblepharus fuscus ZBK almost certainly ranges north of East London, however, its northern limit on the east coast of South Africa is unknown. It apparently does not range as far north as Durban. Bass et al. (1975) record H. fuscus ZBK from southern kwaZulu-Natal, without specifics. Bass et al. (1975) and Bass (1986) also record a specimen from Bredasdorp, Western Cape (true locality probably Arniston, Stuisbaai or Cape Agulhas because Bredasdorp is more than 20 km inland), which they did not examine, however this specimen was examined by the author and was determined to be H. pictus (SAM 24545).
Etymology. Although Smith (1950) did not give the etymology in his original description, there is little doubt that the specific name comes from the Latin, fuscus, which means dusky, dark, or swarthy, and is in reference to the general drab brown colour of this species.
Common name. In an attempt to introduce species specific common names for this genus, this shark was given the name plain happy in Compagno & Human (2003). It is also known as the brown shyshark.
Study material. BAH 20020304.05 , male 340 mm TL, Hamburg , Eastern Cape , South Africa , 33°17.2'S27°28.9'E ; BAH 20020304.06 , male 580 mm TL, Hamburg , Eastern Cape , South Africa ; LJVC 820916 , gravid female 631 mm TL, Port Elizabeth , Eastern Cape , South Africa , 33°53'S25°39'E ; MJS 941004 , mature female 625 mm TL, Cape Recife , Eastern Cape , South Africa , 34°01.7'S25°42.1'E ; MJS 990217 , mature female 609 mm TL, Cape Recife , Eastern Cape , South Africa ; RUSI 21 holotype of Haploblepharus fuscus ZBK , see under Type Series and Locality for details; RUSI 3701 , mature male 605 mm TL, Cape Padrone , Eastern Cape , South Africa , 33°46'S26°28'E ; RUSI 6079, previously ORI 2470 , juvenile male 460 mm TL, Port Alfred , Eastern Cape , South Africa , 33°36'S26°54'E ; RUSI 6081, previously ORI 2787 , mature female 610 mm TL, Port Alfred , Eastern Cape , South Africa ; RUSI 6082, previously ORI 2471 , adolescent male 543 mm TL, Port Alfred , Eastern Cape , South Africa ; RUSI 7617 , adolescent female 568 mm TL, Cape Padrone , Eastern Cape , South Africa ; RUSI 10289 , mature male 551 mm TL, Algoa Bay , Eastern Cape , South Africa , 34°02'S25°42'E ; RUSI 12826 , mature male 602 mm TL, locality not recorded ; RUSI 13144 , 2 specimens, mature male 649 mm TL, Algoa Bay , Eastern Cape , South Africa ; RUSI 14005 , 2 specimens, one of which is referable to H. kistnasamyi , H. fuscus ZBK specimen is a mature male 637 mm TL, Cape Recife , Eastern Cape , South Africa ; RUSI 19993 , 3 specimens, 2 adolescent females 496 mm TL and 540 mm TL, mature male 550 mm TL, locality not recorded ; RUSI 25182 , mature male 577 mm TL, Paradise Beach , Eastern Cape , South Africa , 34°07'S24°52.5'E ; RUSI 25925 , gravid female 571 mm TL, Cape Recife , Eastern Cape , South Africa ; RUSI 41963 , mature male 559 mm TL, Fish River Lighthouse , Eastern Cape , South Africa , 33°29'S27°08'E ; SAM 32523 , 4 specimens, Storms River Mouth , Eastern Cape , South Africa , 34°01.3'S23°54.7'E ; SAM 32614 , immature male 438 mm TL, Langebaan Lagoon (Klein Ostervaal farm) , Western Cape , South Africa , 33°04.5'S18°02.5'E .
- Brett A. Human (2007): A taxonomic revision of the catshark genus Haploblepharus Garman 1913 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae). Zootaxa 1451, 1-40: 22-28, URL:http://www.zoobank.org/urn:lsid:zoobank.org:pub:1FEC783D-01D3-4458-9EE6-B499AF81A83F
H. fuscus ZBK -
BAH 20020304.05, male 340mm TL, Hamburg, Eastern Cape , South Africa , 33°17.2'S27°28.9'E ; BAH 20020304.06, male 580mm TL, Hamburg, Eastern Cape , South Africa ; LJVC 820916, gravid female 631mm TL, Port Elizabeth, Eastern Cape , South Africa , 33°53'S25°39'E ; LJVC 840308, adolescent female 586mm TL, Eastern Cape , South Africa (skeletonised) ; LJVC 850823, adult female 625mm TL, Eastern Cape , South Africa (skeletonised) ; LJVC 840323, 2 specimenes, adult male 640mm TL, Cape Recife, Eastern Cape , South Africa , adult female 730mm TL, Titsikama, Eastern Cape , South Africa (both skeletonised) ; MJS 941004, mature female 625mm TL, Cape Recife, Eastern Cape , South Africa , 34°01.7'S25°42.1'E ; MJS 990217, mature female 609mm TL, Cape Recife, Eastern Cape , South Africa ; RUSI 21 holotype of Haploblepharus fuscus Smith 1950 ZBK , adolescent male 495mm TL, East London, Eastern Cape , South Africa , approx. 33°00'S27°55'E ; RUSI 3701, mature male 605mm TL, Cape Padrone, Eastern Cape , South Africa , 33°46'S26°28'E ; RUSI 6079, previously ORI 2470, juvenile male 460mm TL, Port Alfred, Eastern Cape , South Africa , 33°36'S26°54'E ; RUSI 6081, previously ORI 2787, mature female 610mm TL, Port Alfred, Eastern Cape , South Africa ; RUSI 6082, previously ORI 2471, adolescent male 543mm TL, Port Alfred, Eastern Cape , South Africa ; RUSI 7617, adolescent female 568mm TL, Cape Padrone, Eastern Cape , South Africa ; RUSI 10289, mature male 551mm TL, Algoa Bay, Eastern Cape , South Africa , 34°02'S25°42'E ; RUSI 12826, mature male 602mm TL, locality not recorded ; RUSI 13144, 2 specimens, mature male 649mm TL, Algoa Bay, Eastern Cape , South Africa ; RUSI 14005, 2 specimens, one of which is referable to H. kistnasamyi , H. fuscus ZBK specimen is a mature male 637mm TL, Cape Recife, Eastern Cape , South Africa ; RUSI 19993, 3 specimens, 2 adolescent females 496mm TL and 540mm TL, mature male 550mm TL, locality not recorded ; RUSI 25182, mature male 577mm TL, Paradise Beach, Eastern Cape , South Africa , 34°07'S24°52.5'E ; RUSI 25925, gravid female 571mm TL, Cape Recife, Eastern Cape , South Africa ; RUSI 41963, mature male 559mm TL, Fish River Lighthouse, Eastern Cape , South Africa , 33°29'S27°08'E ; SAM 32523, 4 specimens, Storms River Mouth, Eastern Cape , South Africa , 34°01.3'S23°54.7'E ; SAM 32614, immature male 438mm TL, Langebaan Lagoon (Klein Ostervaal farm), Western Cape , South Africa , 33°04.5'S18°02.5'E .
- Brett A. Human, Leonard J. V. Compagno (2006): Description of Haploblepharus kistnasamyi, a new catshark (Chondrichthyes: Scyliorhinidae) from South Africa. Zootaxa 1318, 41-58: 56-56, URL:http://www.zoobank.org/urn:lsid:zoobank.org:pub:E9EF49D1-FC3C-4D6E-9A6D-A3C14B9E87A8
Habitat and Ecology
Very little of the life history is known for H. fuscus. Maximum size is reported at 73 cm total length (TL) (Compagno 1984). Males are juvenile at 43.8-46.0 cm TL, adolescent at 49.5-54.3 cm TL, and mature at 55.0-64.9 cm TL. Females are adolescent at 49.6-56.8 cm TL, and mature at 60.9-63.1 cm TL. The juveniles of this species are scarce and there appears to be an unknown habitat that is used by H. fuscus for egg laying, and where juveniles spend that stage of their life history (Human 2003, 2007). Other members of this genus produce a single eggcase per uterus which is assumed here for this species.
Depth range (m): 12.8016 - 18.288
Depth range (m): 12.8016 - 18.288
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.
Depth range (m): 12.8016 - 18.288
Depth range (m): 12.8016 - 18.288
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.
Life History and Behavior
IUCN Red List Assessment
Red List Category
Red List Criteria
- 2000Lower Risk/near threatened (LR/nt)
This species' inshore habitat is subject to heavy and increasing human utilizations, including extensive recreational diving and sport and commercial fishing along with coastal housing development, boating, commercial shipping, holiday-making, beach utilization and extensive pollution and habitat degradation of inshore environments.
Relevance to Humans and Ecosystems
The brown shyshark or plain happy (Haploblepharus fuscus) is a species of catshark, part of the family Scyliorhinidae. It is endemic to the shallow, coastal waters of South Africa from west of Cape Agulhas to KwaZulu-Natal. This benthic species is usually found over sandy or rocky bottoms. Measuring up to 73 cm (29 in) long, the brown shyshark is stoutly built, with a broad, flattened head and rounded snout. Unlike other shysharks, the brown shyshark has a plain brown color, though some individuals have faint "saddle" markings or light or dark spots. When threatened, this shark curls into a circle with its tail over its eyes, which is the origin of the name "shyshark". It feeds on bony fishes and lobsters, and is oviparous with females laying pairs of egg capsules. The International Union for Conservation of Nature (IUCN) has assessed this harmless species as Vulnerable. It is of no commercial or recreational interest, but its limited distribution makes its entire population vulnerable to increases in fishing pressure or habitat degradation.
Taxonomy and phylogeny
The brown shyshark was described by South African ichthyologist James Leonard Brierley Smith in a 1950 article for The Annals and Magazine of Natural History. He chose the specific epithet fuscus, which is Latin for "brown". The type specimen is a 57 cm (22 in) long adult male caught off East London, South Africa. A 2006 phylogenetic analysis based on three mitochondrial DNA genes found that the brown shyshark and the dark shyshark (H. pictus) are sister species. They are the more derived members of the genus relative to the basal puffadder shyshark (H. edwardsii).
A small species reaching a maximum known length of 73 cm (29 in), the dark shyshark has a stocky body and a short, broad head. The snout is blunt and dorsally flattened. The eyes are large and oval-shaped, with a rudimentary nictitating membrane (protective third eyelid) and a strong ridge underneath. The nostrils are very large, and are flanked by greatly expanded, triangular flaps of skin that reach the mouth. These nasal flaps cover a pair of deep grooves that connect the nasal excurrent (outflow) openings and the mouth. There are furrows at the corners of the mouth on both jaws. The teeth have a central cusp and a pair of smaller cusplets on the sides. The five pairs of gill slits are positioned on the upper sides of the body.
The first dorsal fin originates well behind the pelvic fin origins, and the second originates behind the anal fin origin. The pectoral fins are moderately large, and the dorsal, pelvic, and anal fins are of similar sizes. The caudal fin is short and broad, with a notch near the tip of the upper lobe and an indistinct lower lobe. The skin is thick and covered by well-calcified leaf-like dermal denticles. The coloration is a plain brown above and white below, though some individuals have a series of faint darker saddle-like markings or black or white spots.
Distribution and habitat
The brown shyshark has a restricted distribution along the coast of South Africa, from the Western and Eastern Cape Provinces to southern KwaZulu-Natal Province. It is usually found close to the bottom over sandy flats or rocky reefs, from the intertidal zone to a depth of 35 m (115 ft). However, it has been reported from as deep as 133 m (436 ft). The range of this species overlaps with the puffadder shyshark in the southeastern Cape region. There, the brown shyshark tends to favor shallow inshore habitats, while the puffadder shyshark inhabits deeper offshore waters.
Biology and ecology
The brown shyshark is a sedentary, bottom-dwelling species; one tag-recapture study found that recaptured sharks had moved no more than 8 km (5.0 mi) from their original tagging location. This shark is known to feed on bony fishes and lobsters. Like its relatives in the genus, it exhibits a curious response of curling into a ring with its tail covering its eyes when threatened, hence the name "shyshark". Reproduction is oviparous, with females depositing egg capsules (known as "mermaid's purses") two at a time. In captivity, the whelks Burnupena papyracea and B. lagenaria have been documented piercing the egg cases and extracting the yolk. Males reach sexual maturity at a length of 68–69 cm (27–27 in), and females at a length of 60–61 cm (24–24 in).
The brown shyshark is harmless to humans and not targeted by any commercial fisheries due to its small size, though it may be caught as bycatch. It is regarded as a minor pest species by recreational anglers and usually discarded or killed when hooked. The International Union for Conservation of Nature (IUCN) has assessed this species as Vulnerable. Although it is locally abundant, heavy fishing occurs throughout its small range and an increase in fishery activities or pollution could potentially affect the entire population. The brown shyshark adapts readily to captivity.
- Human, B. (2009). "Haploblepharus fuscus". IUCN Red List of Threatened Species. Version 2012.2. International Union for Conservation of Nature.
- Smith, J.L.B. (1950). "A new dog-fish from South Africa with notes on other Chondrichthyan fishes". Annals and Magazine of Natural History 3 (34): 878–887. doi:10.1080/00222935008654719.
- Human, B.A.; Owen, E.P.; Compagno, L.J.V.; Harley, E.H. (2006). "Testing morphologically based phylogenetic theories within the cartilaginous fishes with molecular data, with special reference to the catshark family (Chondrichthyes; Scyliorhinidae) and the interrelationships within them". Molecular Phylogenetics and Evolution 39 (2): 384–391. doi:10.1016/j.ympev.2005.09.009. PMID 16293425.
- Compagno, L.J.V. (1984). Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. Food and Agricultural Organization of the United Nations. p. 334. ISBN 92-5-101384-5.
- Compagno, L.J.V.; Dando, M.; Fowler, S. (2005). Sharks of the World. Princeton University Press. p. 235. ISBN 978-0-691-12072-0.
- Fowler, S.L., R.D. Cavanagh, M. Camhi, G.H. Burgess, G.M. Cailliet, S.V. Fordham, C.A. Simpfendorfer, and J.A. Musick (2005). Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes. International Union for Conservation of Nature and Natural Resources. pp. 265–266. ISBN 2-8317-0700-5.
- Kohler, N.E. and P.A. Turner (2001). "Shark tagging: a review of conventional methods and studies". Environmental Biology of Fishes 60: 191–223. doi:10.1023/A:1007679303082.
- Smith, C. and C. Griffiths (1997). "Shark and skate egg-cases cast up on two South African beaches and their rates of hatching success or causes of death". South African Journal of Zoology 32: 112–117.
- Froese, Rainer and Pauly, Daniel, eds. (2009). "Haploblepharus fuscus" in FishBase. August 2009 version.
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