View source for Biology:Puffadder shyshark

{{Short description|Species of shark}}
{{Speciesbox
| image = haploblepharus_edwardsii_Puffadder_shyshark.jpg
| image_alt = A small shark with a broad, flattened head, covered with small white spots and orange-brown saddle-like markings, resting on a gravel sea bed
| status = EN
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 19 November 2021">{{cite iucn |author=Pollom, R. |author2=Da Silva, C. |author3=Gledhill, K. |author4=Leslie, R. |author5=McCord, M.E. |author6=Winker, H. |date=2020 |title=''Haploblepharus edwardsii'' |volume=2020 |page=e.T39345A124403633 |doi=10.2305/IUCN.UK.2020-2.RLTS.T39345A124403633.en |access-date=19 November 2021}}</ref>
| genus = Haploblepharus
| species = edwardsii
| authority = (Schinz, 1822)
| range_map = Haploblepharus edwardsii distmap.png
| range_map_caption = Range of the puffadder shyshark
| range_map_alt = Partial world map with a blue outline along the coast of South Africa
| synonyms = ''Scyllium edwardsii'' <small>Schinz, 1822</small>
}}

The '''puffadder shyshark''' ('''''Haploblepharus edwardsii'''''), also known as the '''Happy Eddie''', is a [[Biology:Species|species]] of [[Biology:Catshark|catshark]], belonging to the family [[Biology:Catshark|Scyliorhinidae]], [[Biology:Endemism|endemic]] to the temperate waters off the coast of {{wipe|South Africa}}. This common shark is found on or near the bottom in sandy or rocky [[Earth:Habitat|habitat]]s, from the [[Earth:Intertidal zone|intertidal zone]] to a depth of {{convert|130|m|ft|abbr=on}}. Typically reaching {{convert|60|cm|in|abbr=on}} 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 [[Biology:Natal shyshark|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 [[Biology:Crustacean|crustacean]]s, 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 [[Organization:International Union for Conservation of Nature|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==
[[File:The Greater Cat-Fish.jpg|thumb|upright=0.8|left|George Edwards' etching of the puffadder shyshark from ''Gleanings of natural history'' (1760).]]
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'',<ref>{{ cite book | last=Edwards | first=George |  year=1760 | title=Gleanings of Natural History, exhibiting figures of quadrupeds, birds, insects, plants &c... | language=English, French | volume=Part 2 | location=London | publisher=Printed for the author, at the College of Physicians | url=https://www.biodiversitylibrary.org/page/50289590 | pages=169–170, Plate 289 }}</ref> of three individuals caught off the Cape of Good Hope that have since been lost. In 1817, French zoologist [[Biography:Georges Cuvier|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 [[Organization:International Commission on Zoological Nomenclature|International Commission on Zoological Nomenclature]] (ICZN) rendered a decision that this species is properly attributed to Schinz.<ref>{{ cite book | last=Schinz | first=Heinrich Rudolf | author-link=Heinrich Rudolf Schinz | date=1822 | title=Das Thierreich eingetheilt nach dem Bau der Thiere als Grundlage ihrer Naturgeschichte und der vergleichenden Anatomie | volume=2 | language=German | location=Stuttgart und Tübingen | publisher=in der J.G. Cotta'schen Buchhandlung | page=214, Footnote | url=https://www.biodiversitylibrary.org/page/51548805 }}</ref><ref>{{ cite journal | author=International Commission on Zoological Nomenclature |  year=2003 | title=Opinion 2056 (Case 3186). ''Squalus edwardsii'' (currently ''Haploblepharus edwardsii''; Chondrichthyes, Carcharhiniformes): attributed to Schinz, 1822 and ''edwardsii'' conserved as the correct original spelling of the specific name | journal=Bulletin of Zoological Nomenclature | volume=60 | issue=3 | page=250 | url=https://www.biodiversitylibrary.org/page/34358000 }}</ref><ref name="human">{{cite journal |author=Human, B.A. |title=A taxonomic revision of the catshark genus ''Haploblepharus'' Garman 1913 (Chondrichthyes: Carcharhiniformes: Scyliorhinidae) |journal=Zootaxa |volume=1451 |pages=1–40 |year=2007 | doi=10.11646/zootaxa.1451.1.1 }}</ref> In 1913, American zoologist Samuel Garman created the new genus ''Haploblepharus'' for this and other shyshark species.<ref name="compagno"/>

Two [[Biology:Form (zoology)|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 [[Biology:Natal shyshark|Natal shyshark]].<ref>{{cite journal |author1=Human, B.A.  |author2=Compagno, L.J.V.  |name-list-style=amp |title=Description of ''Haploblepharus kistnasamyi'', a new catshark (Chondrichthyes: Scyliorhinidae) from South Africa |journal=Zootaxa |volume=1318 |pages=41–58|year=2006|doi=10.11646/zootaxa.1318.1.2 }}</ref> A 2006 phylogenetic analysis, based on three [[Biology:Mitochondrial DNA|mitochondrial DNA]] [[Biology:Gene|gene]]s, found that the puffadder shyshark is the most [[Biology:Basal (phylogenetics)|basal]] member of its family, with a sister relationship to the [[Biology:Clade|clade]] containing the [[Biology:Dark shyshark|dark shyshark]] (''H. pictus'') and the [[Biology:Brown shyshark|brown shyshark]] (''H. fuscus''). The Natal shyshark was not included in the study, though it is very close [[Biology:Morphology|morphologically]] to this species.<ref>{{cite journal |author=Human, B.A. |author2=E.P. Owen |author3=L.J.V. Compagno |author4=E.H. Harley |name-list-style=amp |title=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 |journal=Molecular Phylogenetics and Evolution |volume=39 |issue=2 |date=May 2006 |pages=384–391 |doi=10.1016/j.ympev.2005.09.009 |pmid=16293425}}</ref> The [[Biology:Common name|common name]] "puffadder shyshark" refers to the [[Biology:Bitis arietans|puff adder]] (''Bitis arietans''), a widely distributed African [[Biology:Viperidae|viper]] with similar coloration.<ref name="ferrari and ferrari"/> "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.<ref name="human"/>

==Description==
[[File:Haploblepharus edwardsii, Cape Town.jpg|thumb|left|alt=Two small sharks lying on white gravel|Puffadder shysharks have a distinctive color pattern of orange "saddles" and small white spots.]]
The puffadder shyshark is more slender than other shysharks, with a short, broad, dorsally flattened head and a narrowly rounded snout.<ref name="human"/> The large, oval-shaped eyes have cat-like slit [[Biology:Pupil|pupil]]s, a simple [[Biology:Nictitating membrane|nictitating membrane]] (a protective third [[Biology:Eyelid|eyelid]]), and a prominent ridge underneath. The [[Medicine:Nostril|nostril]]s 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.<ref name="compagno"/> 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.<ref name="bester">Bester, C. [http://www.flmnh.ufl.edu/fish/Gallery/Descript/PuffadderShyshark/PuffadderShyshark.html Biological Profiles: Puffadder Shyshark] {{Webarchive|url=https://web.archive.org/web/20120524153533/http://www.flmnh.ufl.edu/fish/Gallery/Descript/PuffadderShyshark/PuffadderShyshark.html |date=2012-05-24 }}. Florida Museum of Natural History Ichthyology Department. Retrieved on August 31, 2009.</ref> Unusually, the two halves of the lower jaw are connected by a special [[Biology:Cartilage|cartilage]], which allows a more even distribution of teeth and may increase bite strength.<ref name="van der elst">{{cite book |title=A Guide to the Common Sea Fishes of Southern Africa |author=Van der Elst, R. |edition=third |publisher=Struik |year=1993 |isbn=978-1-86825-394-4 |page=71}}</ref>

The five pairs of [[Biology:Gill slit|gill slit]]s are positioned somewhat on the upper surface of the body. The dorsal, [[Biology:Pelvic fin|pelvic]], and anal fins are all of similar size. The dorsal fins are located far back on the body, the first originating behind the pelvic fin origins and the second behind the anal fin origin. The pectoral fins are broad and of moderate size. The short, broad caudal fin comprises about one-fifth of the body length and has a deep ventral notch near the tip of the upper lobe and a barely developed lower lobe. The skin is thick and covered by well-[[Biology:Calcification|calcified]], leaf-shaped dermal denticles.<ref name="compagno"/> The dorsal coloration consists of a light to dark brown background with a series of 8–10 striking yellowish to reddish brown "saddles" with darker margins, all covered by a profusion of small white spots. The underside is white. This species attains a length of {{convert|60|cm|in|abbr=on}}, with a maximum record of {{convert|69|cm|in|abbr=on}}.<ref name="compagno et al"/> Sharks found west of Cape Agulhas are smaller than those found east, reaching only {{convert|48|cm|in|abbr=on}} long.<ref name="human"/>

==Distribution and habitat==
The range of the puffadder shyshark is limited to the [[Earth:Continental shelf|continental shelf]] along the coast of {{wipe|South Africa}}, from Langebaan Lagoon in Western Cape Province to the western shore of Algoa Bay. Previous records of it being found as far north as Durban are now thought to be misidentifications of other species.<ref name="human"/> This bottom-dwelling shark is most common over sandy or rocky bottoms. It is found in progressively deeper water towards the northeastern portion of its range, from {{convert|0|-|15|m|ft|abbr=on}} off [[Place:Cape Town|Cape Town]] to {{convert|40|-|130|m|ft|abbr=on}} off KwaZulu-Natal; this distribution pattern may reflect this shark's preference for cooler waters.<ref name="compagno">{{cite book |author=Compagno, L.J.V. |year=1984 |title=Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date |place=Rome |publisher=Food and Agricultural Organization |isbn=978-92-5-101384-7 |pages=332–333}}</ref>

==Biology and ecology==
[[File:Haploblepharus edwardsii false bay.jpg|thumb|The puffadder shyshark is a slow-moving, benthic species.]]
Quite common within its small range, the sluggish and reclusive puffadder shyshark is often seen lying still on the sea floor.<ref name="ferrari and ferrari">{{cite book |author1=Ferrari, A.  |author2=A. Ferrari  |name-list-style=amp |title=Sharks |url=https://archive.org/details/sharks0000ferr  |url-access=registration  |publisher=Firefly Books |year=2002 |isbn=978-1552096291 |page=[https://archive.org/details/sharks0000ferr/page/131 131]}}</ref><ref name="heemstra and heemstra">{{cite book |title=Coastal Fishes of Southern Africa |author1=Heemstra, E.  |author2=P. Heemstra  |name-list-style=amp |publisher=NISC and SAIAB |year=2004 |isbn=978-1-920033-01-9 |page=67}}</ref> It is gregarious and several individuals may rest together.<ref name="compagno et al"/> A generalist predator with grasping [[Biology:Dentition|dentition]], the puffadder shyshark is known to take a variety of small [[Earth:Benthic zone|benthic]] prey: [[Biology:Crustacean|crustacean]]s (including [[Biology:Crab|crab]]s, shrimp, crayfish, [[Biology:Mantis shrimp|mantis shrimp]], and [[Biology:Hermit crab|hermit crab]]s), [[Biology:Annelida|annelid worms]] (including [[Biology:Polychaete|polychaete]]s), bony fishes (including [[Biology:Anchovy|anchovies]], [[Biology:Trachurus|jack mackerels]], and [[Biology:Goby|gobies]]), [[Biology:Cephalopod|cephalopod]]s (including [[Biology:Squid|squid]]), and fish offal.<ref name="bester"/> Overall, the most important component of this shark's diet is crustaceans, followed by polychaetes and then fishes. Males seem to prefer polychaetes, while females prefer crustaceans.<ref name="compagno et al"/><ref name="dainty">Dainty, A.M. (2002). Biology and ecology of four catshark species in the southwestern Cape, South Africa. M.Sc. thesis, University of Cape Town.</ref> It has been observed attacking a common octopus (''Octopus vulgaris'') by tearing off an arm with a twisting motion.<ref>{{cite journal |title=Diets of common suprabenthic reef fish in False Bay, South Africa |author1=Lechanteur, Y.A.R.G.  |author2=C.L. Griffiths  |name-list-style=amp |journal=African Zoology |volume=38 |issue=2 |pages=213–227 |date=October 2003}}</ref>

The puffadder shyshark is preyed upon by larger fishes, such as the [[Biology:Broadnose sevengill shark|broadnose sevengill shark]] (''Notorynchus cepedianus'').<ref>{{cite journal |author=Ebert, D.A. |title=Diet of the seven gill shark Notorynchus cepedianus in the temperate coastal waters of southern Africa |journal=South African Journal of Marine Science |volume=11 |issue=1 |date=December 1991 |pages=565–572 |doi=10.2989/025776191784287547|doi-access=free }}</ref> The Cape fur seal (''Arctocephalus pusillus pusillus'') has been documented capturing and playing with puffadder shysharks, tossing them into the air or gnawing on them. The shark is often injured or killed during these encounters; the seal may eat torn-off pieces of flesh, but seldom consumes the entire shark. On occasion, [[Biology:Kelp gull|black-backed kelp gull]]s (''Larus dominicanus vetula'') take advantage of this behavior and steal the sharks from the seals.<ref>{{cite journal |author=Martin, R.A. |title=Natural mortality of puffadder shysharks due to Cape fur seals and black-backed kelp gulls at Seal Island, South Africa |journal=Journal of Fish Biology |year=2004 |volume=64|issue=3|pages=711–716 |doi=10.1111/j.1095-8649.2004.00339.x}}</ref> When threatened or disturbed, the puffadder shyshark adopts a characteristic posture in which it curls into a ring and covers its eyes with its tail; this reaction is the basis for the common names "shyshark" and "doughnut", and is likely meant to make the shark harder for a predator to swallow.<ref name="human"/><ref name="compagno et al"/>

The eggs of the puffadder shyshark are fed upon by the [[Biology:Buccinidae|whelks]] ''[[Biology:Burnupena papyracea|Burnupena papyracea]]'' and ''[[Biology:Burnupena lagenaria|B. lagenaria]]'', at least in captivity.<ref>{{cite journal |author1=Smith, C.  |author2=C. Griffiths  |name-list-style=amp |year=1997 |title=Shark and skate egg-cases cast up on two South African beaches and their rates of hatching success or causes of death |journal=South African Journal of Zoology |volume=32 |issue=4  |pages=112–117|doi=10.1080/02541858.1997.11448441  |doi-access=free}}</ref> Known [[Biology:Parasite|parasite]]s of this species include the [[Biology:Trypanosoma|trypanosome]] ''Trypanosoma haploblephari'', which infests the [[Biology:Blood|blood]],<ref>{{cite journal |author1=Yeld, E.M.  |author2=N.J. Smit  |name-list-style=amp |title=A new species of ''Trypanosoma'' (Kinetoplastida: Trypanosomatidae) infecting catsharks from South Africa |journal=Journal of the Marine Biological Association of the United Kingdom |year=2006 |volume=86 |issue=4 |pages=829–833 |doi=10.1017/S0025315406013750|s2cid=86498165 }}</ref> the [[Biology:Nematode|nematode]] ''Proleptus obtusus'', which infests the [[Chemistry:Intestine|intestine]],<ref>{{cite journal |author=Moravec, F. |author2=J.G. Van As |author3=I. Dykova |name-list-style=amp |title=''Proleptus obtusus'' Dujardin, 1845 (Nematoda: Physalopteridae) from the puffadder shyshark ''Haploblepharus edwardsii'' (Scyliorhinidae) from off South Africa |journal=Systematic Parasitology |volume=53 |issue=3 |date=November 2002 |pages=169–173 |doi=10.1023/A:1021130825469 |pmid=12510161|s2cid=22873948 }}</ref> and the [[Biology:Copepod|copepod]]s ''Charopinus dalmanni'' and ''Perissopus oblongatus'', which infest the skin.<ref>{{cite journal |author=Dippenaar, S.M. |title=Reported siphonostomatoid copepods parasitic on marine fishes of southern Africa |journal=Crustaceana |volume=77 |issue=11 |pages=1281–1328 |doi=10.1163/1568540043165985 |year=2004}}</ref> Another parasite is the praniza [[Biology:Larva|larva]]l stage of the isopod ''Gnathia pantherina'', which infests the [[Medicine:Nostril|nares]], mouth, and gills. The deep-penetrating mouthparts of these larvae significantly damage local tissue, causing bleeding and [[Medicine:Inflammation|inflammation]].<ref>{{cite journal |author=Hayes, P.M. |author2=N.J. Smit |author3=A.J. Davies |name-list-style=amp |title=Pathology associated with parasitic juvenile gnathiids feeding on the puffadder shyshark, ''Haploblepharus edwardsii'' (Voight) |journal=Journal of Fish Diseases |volume=30 |issue=1 |pages=55–58 |pmid=17241405 |year=2007 |doi=10.1111/j.1365-2761.2007.00777.x}}</ref>

===Life history===
[[File:Puffadder shyshark egg case.jpg|thumb|alt=Close-up of a rounded egg case with brown and cream bands, with coral in the background|The egg case of the puffadder shyshark has a different color pattern than those of other shysharks.]]
The puffadder shyshark is oviparous; there is no distinct breeding season and reproduction occurs year-round.<ref name="dainty"/> Females deposit [[Biology:Egg case (Chondrichthyes)|egg capsules]] one or two at a time, attaching them to vertical structures such as sea fans.<ref name="compagno et al">{{cite book |author=Compagno, L.J.V. |author2=M. Dando |author3=S. Fowler |name-list-style=amp |title=Sharks of the World |publisher=Princeton University Press |year=2005 |isbn=978-0-691-12072-0 |pages=234–235}}</ref> The thin-walled egg cases are brown with distinctive pale transverse bands; and have a slightly furry texture and long adhesive tendrils at the corners. They are smaller than those of other shyshark species, measuring {{convert|3.5|-|5|cm|in|abbr=on}} long and {{convert|1.5|-|3|cm|in|abbr=on}} across.<ref name="van der elst"/><ref>{{cite journal |title=Shark and skate egg-cases cast up on two South African beaches and their rates of hatching success, or causes of death |author1=Smith, C.  |author2=C. Griffiths  |name-list-style=amp |journal=South African Journal of Zoology |volume=32 |issue=4 |pages=112–117 |date=October 1997|doi=10.1080/02541858.1997.11448441  |doi-access=free }}</ref> The young shark hatches after three months, and measures around {{convert|9|cm|in|abbr=on}} long.<ref name="iucn status 19 November 2021" /> The length at [[Biology:Sexual maturity|maturation]] for both sexes has been reported as anywhere from {{convert|35|to|55|cm|in|abbr=on}} by various sources; this high degree of variation may reflect regional differences as sharks from deeper waters in the eastern part of its range seem to mature at a larger size than those from the west.<ref name="human"/> The age at maturation is estimated to be around 7 years, and the maximum lifespan is at least 22 years.<ref name="dainty"/>

==Human interactions==
Harmless to humans, the puffadder shyshark can be easily caught by hand.<ref name="fowler et al">{{cite book |author=Fowler, S.L. |author2=R.D. Cavanagh |author3=M. Camhi |author4=G.H. Burgess |author5=G.M. Cailliet |author6=S.V. Fordham |author7=C.A. Simpfendorfer |author8=J.A. Musick |name-list-style=amp |title=Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes |publisher=International Union for Conservation of Nature and Natural Resources |year=2005 |isbn=978-2-8317-0700-6 |page=265}}</ref> Not targeted by commercial fisheries because of its small size, it is taken incidentally and discarded by bottom trawlers operating between Mossel Bay and East London, and by fishing boats in False Bay. Many are hooked by recreational anglers casting from the shore, who also generally discard or kill them as minor pests.<ref name="van der elst"/> Some local exploitation of this species does occur for [[Biology:Lobster|lobster]] bait and the [[Biology:Fishkeeping|aquarium trade]]. The [[Organization:International Union for Conservation of Nature|International Union for Conservation of Nature]] (IUCN) has assessed the puffadder shyshark as endangered. The small range of this shark lies entirely within a heavily fished region, and any increase in fishing activities or habitat degradation could potentially impact the entire population.<ref name="fowler et al"/>

==References==
{{Reflist|colwidth=30em}}

==External links==

{{Taxonbar|from=Q612558}}

[[Category:Haploblepharus|puffadder shyshark]]

{{Sourceattribution|Puffadder shyshark}}