Haploblepharus
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''Haploblepharus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
catshark Catsharks are ground sharks of the family Scyliorhinidae. They are the largest family of sharks with around 160 species placed in 17 genera. Although they are generally known as catsharks, some species can also be called dogfish due to previous n ...
, and part of the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Scyliorhinidae, containing four
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of shysharks. Their
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
comes from a distinctive defensive behavior in which the shark curls into a circle and covers its eyes with its tail. The genus is
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to
southern Africa Southern Africa is the southernmost subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number of ...
, inhabiting shallow coastal waters. All four species are small, stout-bodied sharks with broad, flattened heads and rounded snouts. They are characterized by very large
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
s with enlarged, triangular flaps of skin that reach the mouth, and deep grooves between the nostrils and the mouth. Shysharks are bottom-dwelling
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s of
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
es and
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s. They are
oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and ...
, with the females laying egg capsules. These harmless sharks are of no
commercial Commercial may refer to: * a dose of advertising conveyed through media (such as - for example - radio or television) ** Radio advertisement ** Television advertisement * (adjective for:) commerce, a system of voluntary exchange of products and s ...
or
recreational Recreation is an activity of leisure, leisure being discretionary time. The "need to do something for recreation" is an essential element of human biology and psychology. Recreational activities are often done for enjoyment, amusement, or pleasure ...
interest, though their highly limited distributions in heavily fished South African waters are of potential conservation concern.


Taxonomy and phylogeny

The genus ''Haploblepharus'' was created by American zoologist
Samuel Garman Samuel Walton Garman (June 5, 1843 – September 30, 1927), or "Garmann" as he sometimes styled himself, was a naturalist/zoologist from Pennsylvania. He became noted as an ichthyologist and herpetologist. Biography Garman was born in Indian ...
in 1913, in the 36th volume of ''Memoirs of the Museum of Comparative Zoology, at Harvard College'', to contain the puffadder shyshark, then known as ''Squalus edwardsii''. The name is derived from the Greek ''haplóos'' meaning "single", and ''blepharos'' meaning "eyelid".Bester, C
Biological Profiles: Puffadder Shyshark
Florida Museum of Natural History Ichthyology Department. Retrieved on August 31, 2009.
In 1988,
Leonard Compagno Leonard Joseph Victor Compagno is an international authority on shark taxonomy and the author of many scientific papers and books on the subject, best known of which is his 1984 catalogue of shark species produced for the Food and Agriculture Organi ...
placed ''Haploblepharus'' with ''
Halaelurus ''Halaelurus'' is a genus of catsharks in the family Scyliorhinidae. Species * '' Halaelurus boesemani'' S. Springer & D'Aubrey, 1972 (speckled catshark) * '' Halaelurus buergeri'' ( J. P. Müller & Henle, 1838) (blackspotted catshark) * '' H ...
'' and ''
Holohalaelurus ''Holohalaelurus'' is a genus of catshark in the family Scyliorhinidae, commonly known as Izak catsharks or hallelujah sharks. Member species of this genus are distributed in the western Indian Ocean off the coasts of various Southern African a ...
'' in the tribe Halaelurini of the family Scyliorhinidae, based on morphological characters. This interpretation was corroborated by a 2006
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
based on three
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s. However, that study also found that within this group ''Haploblepharus'' is most closely related to the genus ''Halaelurus'', which is inconsistent with the presence of three autapomorphic traits in ''Haploblepharus'' that are not shared by either ''Halaelurus'' or ''Holohalaelurus''. Within the genus, the puffadder shyshark is the most basal species.


Species

* '' Haploblepharus edwardsii'' ( Schinz, 1822) (puffadder shyshark) * '' Haploblepharus fuscus''
J. L. B. Smith James Leonard Brierley Smith (26 September 1897 – 8 January 1968) was a South African ichthyologist, organic chemist, and university professor. He was the first to identify a taxidermied fish as a coelacanth, at the time thought to be long ex ...
, 1950
(brown shyshark) * '' Haploblepharus kistnasamyi''
Human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
& Compagno, 2006
(Natal shyshark) * '' Haploblepharus pictus'' ( J. P. Müller &
Henle Henle can refer to: * Friedrich Gustav Jakob Henle, a German physician, pathologist and anatomist (1809–1885) ** Loop of Henle in the kidney, named after Henle *Fritz Henle, a photographer, known as "Mr. Rollei" for his use of the 2.25" square for ...
, 1838)
(dark shyshark)


Distribution and habitat

All four shyshark species are
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to the southern tip of Africa; three of them are found only off
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
, while the range of the dark shyshark extends to southern
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
. They are bottom-dwelling in nature and usually found in shallow, coastal waters over sandy or rocky bottoms.


Description

Different shyshark species are very similar to one another in appearance but can be reliably differentiated using morphological measurements. However, in the field the only way to readily tell them apart is by their different color patterns, and even this may be problematic as individuals of the same species can vary considerably in coloration. All four species are small, seldom exceeding in length. A shyshark has a stout, spindle-shaped body and a short head comprising less than one-fifth of the total length. The head is broad and dorsally flattened, with a rounded snout. The large, oval eyes have cat-like slit
pupil The pupil is a black hole located in the center of the iris of the eye that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company. It appears black ...
s, a rudimentary
nictitating membrane The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
(protective third
eyelid An eyelid is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. The human eyel ...
), and a prominent ridge running underneath. A distinguishing trait of this genus are the large
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
s partially covered by greatly expanded, triangular flaps of skin that overlap the mouth and cover a pair of deep grooves between the nasal excurrent (outflow) openings and the mouth. The mouth is short and curved, with furrows at the corners extending onto both jaws. The teeth have a central cusp and smaller lateral cusplets. The five pairs of
gill slit Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of cartilaginous fish such as sharks and rays, as well as deep-branching vertebrates such as lampreys. In con ...
s are located on the upper side of the body. The two
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through conv ...
s are set far back on the body, the first originating behind the
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods. Structure and function Structure In actinopterygians, the pelvic fin consists of two en ...
midbases and the second originating behind the
anal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
midbase. The
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
s are moderately large; the dorsal, pelvic, and anal fins are of similar size. The broad
caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
comprises a fifth of the total length; the upper lobe has a deep ventral notch near the tip and the lower lobe is virtually absent. The skin is thick and covered by well-
calcified Calcification is the accumulation of calcium salts in a body tissue. It normally occurs in the formation of bone, but calcium can be deposited abnormally in soft tissue,Miller, J. D. Cardiovascular calcification: Orbicular origins. ''Nature Mat ...
, leaf-shaped
dermal denticle A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as we ...
s. The coloration is brown above and white below; the brown shyshark is usually plain while the other species have varying patterns of darker "saddles" and white spots.


Biology and ecology

Shysharks feed on a variety of small benthic
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
es and
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s. In turn, they are preyed upon by larger fishes and
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s. When threatened, these sharks adopt a curious posture in which they curl their bodies into a ring and cover their eyes with their tails, which is the origin of the name "shyshark". This behavior likely serves to make the shark more difficult for a
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
to swallow. Shysharks are
oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and ...
with females producing egg capsules (known as "
mermaid's purse An egg case or egg capsule, often colloquially called a mermaid's purse, is the casing that surrounds the eggs of oviparous sharks, skates and chimaeras. Egg cases typically contain one embryo, except for big skate and mottled skate egg cases ...
s") two at a time. Reproduction has been investigated in the puffadder and dark shysharks; neither species appears to have a distinct
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
, and their eggs take around three and a half weeks to hatch.Dainty, A.M. (2002). Biology and ecology of four catshark species in the southwestern Cape, South Africa. M.Sc. thesis, University of Cape Town.


Human interactions

Due to their small size, shysharks pose no danger to humans and are not sought by either
commercial Commercial may refer to: * a dose of advertising conveyed through media (such as - for example - radio or television) ** Radio advertisement ** Television advertisement * (adjective for:) commerce, a system of voluntary exchange of products and s ...
or recreational fishers. They are taken as
bycatch Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
and hooked by shore anglers, who regard them (and other catsharks) as minor pests. Although shysharks are not uncommon, their extremely limited distribution within heavily fished waters raises concerns that a local increase in fishing pressure or
habitat degradation Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
could affect entire species populations. As a result, the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN) has assessed two shyshark species as
Near Threatened A near-threatened species is a species which has been categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature as that may be vulnerable to endangerment in the near future, but it does not currently qualify fo ...
.


References

{{Taxonbar, from=Q2093983 Shark genera * Taxa named by Samuel Garman