The puffadder shyshark (Haploblepharus edwardsii), also known as the Happy Eddie, is a species of catshark, belonging to the family Scyliorhinidae, endemic to the temperate waters off the coast of South Africa. This common shark is found on or near the bottom in sandy or rocky habitats, from the intertidal zone to a depth of . Typically reaching in length, the puffadder shyshark has a slender, flattened body and head. It is strikingly patterned with a series of dark-edged, bright orange "saddles" and numerous small white spots over its back. The Natal shyshark (H. kistnasamyi), formally described in 2006, was once considered to be an alternate form of the puffadder shyshark.
When threatened, the puffadder shyshark (and other members of its genus) curls into a circle with its tail covering its eyes, giving rise to the local names "shyshark" and "doughnut". It is a predator that feeds mainly on crustaceans, polychaete worms, and small bony fishes. This shark is oviparous and females deposit egg capsules singly or in pairs onto underwater structures. Harmless to humans, the puffadder shyshark is usually discarded by commercial and recreational fishers alike for its small size. It has been assessed as endangered by the International Union for Conservation of Nature (IUCN), as its entire population is located within a limited area and could be affected by a local increase in fishing pressure or habitat degradation.
Taxonomy and phylogeny
The first known reference to the puffadder shyshark in literature was by prominent English naturalist George Edwards in 1760, by the name Catulus major vulgaris, of three individuals caught off the Cape of Good Hope that have since been lost. In 1817, French zoologist Georges Cuvier described this species as "Scyllium D'Edwards", based on Edwards' account, though he was not considered to be proposing a true scientific name. In 1832, German zoologist Friedrich Siegmund Voigt translated Cuvier's description under the name Scyllium edwardsii, thus receiving attribution for the species. However, in 2001 M.J.P. van Oijen discovered that Swiss naturalist Heinrich Rudolf Schinz had provided an earlier translation of Cuvier's text with the proper scientific name in 1822, and subsequently the International Commission on Zoological Nomenclature (ICZN) rendered a decision that this species is properly attributed to Schinz. In 1913, American zoologist Samuel Garman created the new genus Haploblepharus for this and other shyshark species.
Two forms of puffadder shyshark were once recognized: "Cape" and "Natal", which differed in appearance and habitat preferences. In 2006, the "Natal" form was described as a new species, the Natal shyshark. A 2006 phylogenetic analysis, based on three mitochondrial DNA genes, found that the puffadder shyshark is the most basal member of its family, with a sister relationship to the clade containing the dark shyshark (H. pictus) and the brown shyshark (H. fuscus). The Natal shyshark was not included in the study, though it is very close morphologically to this species. The common name "puffadder shyshark" refers to the puff adder (Bitis arietans), a widely distributed African viper with similar coloration. "Happy Eddie" (from the scientific name Haploblepharus edwardsii) is used by academics for this shark, and was recently introduced to the public as an easily remembered alternative to the ambiguous vernaculars "shyshark" and "doughnut", which can apply to several different species and have confounded research efforts.
Description
The puffadder shyshark is more slender than other shysharks, with a short, broad, dorsally flattened head and a narrowly rounded snout. The large, oval-shaped eyes have cat-like slit pupils, a simple nictitating membrane (a protective third eyelid), and a prominent ridge underneath. The nostrils are very large, with a pair of greatly expanded, triangular flaps of skin in front that are fused together and reach the mouth. There is a deep groove connecting the excurrent (outflow) opening of each nostril to the mouth, obscured by the nasal flaps. The mouth is short with furrows at the corners on both jaws. There are 26–30 tooth rows in the upper jaw and 27–33 tooth rows in the lower jaw. Tooth shape is sexually dimorphic: those of males are longer and three-pointed, while those of females are shorter and five-pointed. Unusually, the two halves of the lower jaw are connected by a special cartilage, which allows a more even distribution of teeth and may increase bite strength.