Haploblepharus Pictus
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The dark shyshark or pretty happy (''Haploblepharus pictus'') is a
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
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 ...
, belonging to 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,
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
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
waters off 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 ...
and western
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 ...
. It is
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
in nature and inhabits shallow,
inshore A shore or a shoreline is the fringe of land at the edge of a large body of water, such as an ocean, sea, or lake. In physical oceanography, a shore is the wider fringe that is geologically modified by the action of the body of water past a ...
waters and favors rocky
reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock out ...
s and
kelp forest Kelp forests are underwater areas with a high density of kelp, which covers a large part of the world's coastlines. Smaller areas of anchored kelp are called kelp beds. They are recognized as one of the most productive and dynamic ecosystems on Ea ...
s. Growing to long, this small, stocky shark has a wide, flattened head with a rounded snout and a large flap of skin extending from before the nostrils to the mouth. Its dorsal coloration is extremely variable and may feature black-edged orange to blackish saddles and/or white spots on a light brown to nearly black background. When threatened, the dark shyshark curls into a ring with its tail covering its eyes, hence the name "shyshark". It preys mainly on small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
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
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s. Reproduction is
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 ...
and proceeds throughout the year. Females lay two egg cases at a time, which hatch after 6–10 months. This harmless species is of little commercial significance due to its small size. It is frequently caught by recreational fishers and persecuted as a pest. 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 listed the dark shyshark under
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
, as it is common and does not appear to be substantially threatened by fishing 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 ...
.


Taxonomy and phylogeny

German physicians and biologists
Johannes Peter Müller Johannes Peter Müller (14 July 1801 – 28 April 1858) was a German physiologist, comparative anatomy, comparative anatomist, ichthyology, ichthyologist, and herpetology, herpetologist, known not only for his discoveries but also for his ability ...
and
Friedrich Gustav Jakob Henle Friedrich Gustav Jakob Henle (; 9 July 1809 – 13 May 1885) was a German physician, pathologist, and anatomist. He is credited with the discovery of the loop of Henle in the kidney. His essay, "On Miasma and Contagia," was an early argument for ...
originally described the dark shyshark in their 1838–41 ''Systematische Beschreibung der Plagiostomen'', based on five specimens caught off the
Cape of Good Hope The Cape of Good Hope ( af, Kaap die Goeie Hoop ) ;''Kaap'' in isolation: pt, Cabo da Boa Esperança is a rocky headland on the Atlantic coast of the Cape Peninsula in South Africa. A common misconception is that the Cape of Good Hope is t ...
and deposited in the
Rijksmuseum van Natuurlijke Historie The Rijksmuseum van Natuurlijke Historie (National Museum of Natural History) was a museum on the Rapenburg in Leiden, the Netherlands. It was founded in 1820 by Royal Decree from a merger of several existing collections. This happened on the initi ...
in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration wit ...
,
The Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
. Because of the shark's ornate coloration, they gave it the
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''pictum'' from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for "painted". Originally placed in the now-obsolete genus ''Scyllium'', subsequent authors moved this species to the genus ''
Haploblepharus ''Haploblepharus'' is a genus of catshark, and part of the family Scyliorhinidae, containing four species of shysharks. Their common name comes from a distinctive defensive behavior in which the shark curls into a circle and covers its eyes with ...
'', coined by American
zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal kingdom, including the anatomy, structure, embryology, evolution, Biological clas ...
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. The dark shyshark was often regarded to be the same as the puffadder shyshark (''H. edwardsii'') until 1975, with the publication of A.J. Bass, Jeanette D'Aubrey, and Nat Kistnasamy's review of southern African sharks. It continues to be confused for the other three shyshark species because of its extremely variable coloration. The
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 ...
"pretty happy" ("happy" refers to the genus name ''Haploblepharus'') was recently introduced to the public as an easily remembered alternative to the colloquial names "shyshark" and "doughnut", which can apply to several species and have confounded research efforts. Brett Human's 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, found that the dark shyshark and the brown shyshark (''H. fuscus'') are
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
, and that the two make up the more
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
within the genus.


Description

Reaching no more than long, the dark shyshark differs from other shysharks in that it is slender-bodied when juvenile and stocky-bodied when adult. Its head is short, wide, and flattened, with a bluntly rounded snout and very large nostrils. The anterior rims of the nostrils bear greatly enlarged lobes of skin, which are fused into a single flap that reaches the mouth; the flap conceals the nasal excurrent openings and a pair of grooves that run between them and the mouth. The large, horizontally oval eyes are equipped with 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 ...
s (protective third eyelids), and have strong ridges beneath. The mouth is short but wide and bears furrows at the corners extending onto both jaws. There are 45–83 upper and 47–75 lower tooth rows; each tooth has a long central cusp flanked by a pair of smaller 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 fairly high on 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 of nearly equal size and placed far back on the body, with the first originating over the latter third of 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 ...
bases and the second over the latter half of 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 ...
base. The pectoral and pelvic fins are broad and rounded; the males have stout
clasper In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's c ...
s. The pelvic and anal fins are about as large as the dorsal fins. 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 about a fifth of the total body length and has a strong notch near the tip of the upper lobe and an indistinct lower lobe. 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 ...
arrowhead-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 of the dark shyshark is highly variable, and particular individuals can resemble any of the other shyshark species. The background color ranges from light brown to reddish to grayish to almost black above, transitioning abruptly to white or cream below, sometimes with dark blotches beneath the paired fins. There may be 6–8 orange, brown, or blackish variably shaped saddles along the back and tail, edged more or less obviously in black. There may also be white spots within or within and between the saddles.


Distribution and habitat

The range of the dark shyshark is restricted to the coastal waters of southern Africa, from north of
Lüderitz Lüderitz is a town in the ǁKaras Region of southern Namibia. It lies on one of the least hospitable coasts in Africa. It is a port developed around Robert Harbour and Shark Island. The town is known for its colonial architecture, includi ...
in 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 ...
to east of the
Storms River Storms River ( af, Stormsrivier) is a river in the Eastern Cape province of South Africa. The river mouth is located in the Tsitsikamma National Park. The 5 day Otter Trail starts at the Storms River mouth. See also * Tsitsikamma National Pa ...
mouth in the
Eastern Cape Province The Eastern Cape is one of the provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha. The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in 1994 ...
of
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 ...
. It is abundant, particularly west of
Cape Agulhas Cape Agulhas (; pt, Cabo das Agulhas , "Cape of the Needles") is a rocky headland in Western Cape, South Africa. It is the geographic southern tip of the African continent and the beginning of the dividing line between the Atlantic and Indian ...
. This species is a bottom-dweller found close
inshore A shore or a shoreline is the fringe of land at the edge of a large body of water, such as an ocean, sea, or lake. In physical oceanography, a shore is the wider fringe that is geologically modified by the action of the body of water past a ...
, from the
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species o ...
to a depth of . It prefers rocky reefs and
kelp forest Kelp forests are underwater areas with a high density of kelp, which covers a large part of the world's coastlines. Smaller areas of anchored kelp are called kelp beds. They are recognized as one of the most productive and dynamic ecosystems on Ea ...
s, but also presumably traverses sandy flats in between patches of more suitable
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
. As this shark is not known to travel long distances, there is likely subpopulation differentiation across parts of its range.


Biology and ecology

The dark shyshark is a
generalist A generalist is a person with a wide array of knowledge on a variety of subjects, useful or not. It may also refer to: Occupations * a physician who provides general health care, as opposed to a medical specialist; see also: ** General pract ...
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 ...
whose main food sources are, in descending order of importance, small
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
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
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s. Larger sharks consume proportionately more crustaceans.
Polychaete worm Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are mad ...
s and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s are also taken on occasion, and
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
may be swallowed incidentally. This species is preyed upon by the
broadnose sevengill shark The broadnose sevengill shark (''Notorynchus cepedianus'') is the only extant member of the genus ''Notorynchus'', in the family Hexanchidae. It is recognizable because of its seven gill slits, while most shark species have five gill slits, with ...
(''Notorynchus cepedianus''), and potentially also other large 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, it adopts a characteristic defense posture in which it curls into a ring with its tail over its eyes; this behavior likely makes the shark harder to swallow and is the origin of the common names "shyshark" and "doughnut". In captivity, the
whelks Whelk (also known as scungilli) is a common name applied to various kinds of sea snail. Although a number of whelks are relatively large and are in the family Buccinidae (the true whelks), the word ''whelk'' is also applied to some other mari ...
'' Burnupena papyracea'' and '' B. lagenaria'' have been documented piercing the egg cases of this species and extracting the
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example bec ...
. A known
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
of the dark shyshark is the trypanosome ''Trypanosoma haploblephari'', which infests the
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the c ...
. Like other members of its genus, the dark shyshark is
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 ...
; adult females have a single functional
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
and two functional
oviduct The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
s. There appears to be no 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 reproduction occurs year-round. Females produce mature eggs two at a time, one per oviduct. The eggs are enclosed in purse-shaped capsules measuring long and across; each capsule is plain amber to dark brown in color and bears thin, coiled tendrils at the four corners. In one observation of an egg that hatched after 104 days, the developing
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
had external gill filaments until it was 50 days old, and completely absorbed its
yolk sac The yolk sac is a membranous sac attached to an embryo, formed by cells of the hypoblast layer of the bilaminar embryonic disc. This is alternatively called the umbilical vesicle by the Terminologia Embryologica (TE), though ''yolk sac'' is far ...
shortly before hatching. Eggs in nature typically hatch in 6–10 months, with the newly hatched shark measuring long. Both sexes grow at approximately the same rate, reaching
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
at around 15 years of age. Mature males and females range from and long respectively. The
maximum lifespan Maximum life span (or, for humans, maximum reported age at death) is a measure of the maximum amount of time one or more members of a population have been observed to survive between birth and death. The term can also denote an estimate of the maxi ...
is 25 years.


Human interactions

The dark shyshark poses no danger to humans and is too small to be of commercial importance. Many are hooked by sport fishers casting from the shore, who regard the shark as a pest and often kill it. It may also be caught by subsistence fishers and in
lobster trap A lobster trap or lobster pot is a portable trap that traps lobsters or crayfish and is used in lobster fishing. In Scotland (chiefly in the north), the word creel is used to refer to a device used to catch lobsters and other crustaceans. A l ...
s and
bottom trawl Bottom trawling is trawling (towing a trawl, which is a fishing net) along the seafloor. It is also referred to as "dragging". The scientific community divides bottom trawling into benthic trawling and demersal trawling. Benthic trawling is towing ...
s, though not in substantial quantities. This shark occasionally finds its way into the
aquarium trade Fishkeeping is a popular hobby, practiced by aquarists, concerned with keeping fish in a home aquarium or garden pond. There is also a piscicultural fishkeeping industry, serving as a branch of agriculture. Origins of fishkeeping Fish have ...
, though there is no direct fishing for this purpose. As the dark shyshark remains common and does not seem heavily threatened by human activity, it has been assessed as
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
by 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). Its small range does raise concerns that an increase in regional 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 impact the entire population.


References

{{Taxonbar, from=Q660343 Haploblepharus Fish described in 1838