Cephaloscyllium Laticeps By Bartholomew
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''Cephaloscyllium'' 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 ...
s, 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, commonly known as swellsharks because of their ability to inflate their bodies with water or air as a defense against
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. These sluggish, bottom-dwelling sharks are found widely in the
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and
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 ...
coastal waters of the
Indian Indian or Indians may refer to: Peoples South Asia * Indian people, people of Indian nationality, or people who have an Indian ancestor ** Non-resident Indian, a citizen of India who has temporarily emigrated to another country * South Asia ...
and
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...
s. They have stocky, spindle-shaped bodies and short, broad, and flattened heads. The mouth is capacious, containing many small teeth and lacking furrows at the corners. 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 placed far back on the body, with the first much larger than the second. Different species have various color patterns of saddles, blotches, reticulations, and/or spots. The largest members of the genus can grow over in length. Swellsharks prey on a variety of fishes 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, and 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 in pairs. They are harmless and have been deemed of having no commercial value.


Taxonomy

The genus ''Cephaloscyllium'' was proposed by American
ichthyologist Ichthyology is the branch of zoology devoted to the study of fish, including bony fish ( Osteichthyes), cartilaginous fish (Chondrichthyes), and jawless fish (Agnatha). According to FishBase, 33,400 species of fish had been described as of Octobe ...
Theodore Gill Theodore Nicholas Gill (March 21, 1837 – September 25, 1914) was an American ichthyologist, mammalogist, malacologist and librarian. Career Born and educated in New York City under private tutors, Gill early showed interest in natural histor ...
from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''kephale'' ("head") and ''skylion'' ("dogfish"), in an 1862 issue of ''Annals of the Lyceum of Natural History of New York''. However, most of Gill's contemporaries, particularly those in Europe, preferred to keep the swellsharks within the genus ''Scyllium'' (a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of ''
Scyliorhinus ''Scyliorhinus'' is a genus of catsharks in the family Scyliorhinidae. This genus is known in the fossil records from the Cretaceous period, late Albian age to the Pliocene epoch.Carrier, J. C.; Musick, J. A. & Heithaus, M. R. (2004)''Biology of ...
''). ''Cephaloscyllium'' did not gain wide acceptance until
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 ...
published "The Plagiostomia" in a 1913 volume of ''Memoirs of the Museum of Comparative Zoology'', in which he recognized three species: '' C. isabellum'', '' C. ventriosum'', and '' C. umbratile''. A long history of taxonomic confusion exists regarding the species within ''Cephaloscyllium'' owing to several factors, including variation in appearance (particularly between juveniles and adults), the existence of multiple undescribed species, a paucity of detailed scientific descriptions and
type material In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes t ...
, and the use of unreliable characters. Until recently, various authors had recognized five to eight species, along with at least five undescribed species in Australian waters and more in the western central Pacific and the Indian Ocean. In 2008, significant studies were made in resolving the taxonomy of ''Cephaloscyllium'', with a revision of the genus and an increase in the number of described species to 21.


Species

There are currently 17 recognized species in this genus: * ''
Cephaloscyllium albipinnum The whitefin swellshark (''Cephaloscyllium albipinnum'') is a little-known species of catshark, belonging to the family (biology), family Catshark, Scyliorhinidae, Endemism, endemic to southeastern Australia. It is found down, on the outer cont ...
''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Motomura & W. T. White, 2008
(whitefin swellshark) * '' Cephaloscyllium cooki''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Séret & W. T. White, 2008
(Cook's swellshark) * '' Cephaloscyllium fasciatum'' W. L. Y. Chan, 1966 (reticulated swellshark) * ''
Cephaloscyllium hiscosellum The Australian reticulate swellshark (''Cephaloscyllium hiscosellum'') is a little-known species of catshark in the family Scyliorhinidae. It is found off the coast of northwestern Australia at depths of . This shark has a stocky body and a sho ...
'' W. T. White & Ebert, 2008 (Australian reticulate swellshark) * '' Cephaloscyllium isabellum'' Bonnaterre, 1788 (draughtsboard shark) * '' Cephaloscyllium laticeps'' A. H. A. Duméril, 1853 (Australian swellshark) * '' Cephaloscyllium pictum''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Séret & W. T. White, 2008
(painted swellshark) * '' Cephaloscyllium sarawakensis'' Ka. Yano, A. Ahmad &
Gambang A gambang, properly called a gambang kayu ('wooden gambang') is a xylophone-like instrument used among people of Indonesia in gamelan and kulintang, with wooden bars as opposed to the metallic ones of the more typical metallophones in a gamelan. ...
, 2005
(Sarawak pygmy swellshark) * '' Cephaloscyllium signourum''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Séret & W. T. White, 2008
(flagtail swellshark) * '' Cephaloscyllium silasi''
Talwar The talwar (), also spelled ''talwaar'' and ''tulwar'', is a type of curved sword or sabre from the Indian subcontinent. Etymology and classification The word ''talwar'' originated from the Sanskrit word ''taravāri'' ( sa, तरवारि) ...
, 1974
(Indian swellshark) * '' Cephaloscyllium speccum''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Séret & W. T. White, 2008
(speckled swellshark) * '' Cephaloscyllium stevensi'' E. Clark & J. E. Randall, 2011 (Steven's swellshark) * '' Cephaloscyllium sufflans''
Regan The family name Regan, along with its cognates O'Regan, O Regan, Reagan, and O'Reagan, is an Anglicized form of the Irish surname Ó Riagáin or Ó Ríogáin, from Ua Riagáin. The meaning is likely to have originated in ancient Gaelic ''ri'' ...
, 1921
(balloon shark) * '' Cephaloscyllium umbratile'' D. S. Jordan & Fowler, 1903 (blotchy swellshark) * '' Cephaloscyllium variegatum''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Séret & W. T. White, 2008
(saddled swellshark) * '' Cephaloscyllium ventriosum''
Garman Garman is a surname or first name. Notable people with the name include: Sports * Ann Garman, All-American Girls Professional Baseball League player * Judi Garman (born 1954), American softball coach * Mike Garman (born 1949), American baseball pla ...
, 1880
(swellshark) * '' Cephaloscyllium zebrum''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Séret & W. T. White, 2008
(narrowbar swellshark) * ''Cephaloscyllium'' sp. 1 Not yet described (Philippine swellshark) * ''Cephaloscyllium'' sp. not yet described (tiger swellshark) * ''Cephaloscyllium'' sp. not yet described (New Guinea swellshark) * ''Cephaloscyllium'' sp. not yet described (dwarf balloon shark) * ''Cephaloscyllium'' sp. not yet described (dwarf oriental swellshark) * ''Cephaloscyllium'' sp. not yet described (New Caledonia swellshark) * ''Cephaloscyllium'' sp. not yet described (Red Sea swellshark)


Phylogeny and evolution

Morphological and
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies have shown the closest relative of ''Cephaloscyllium'' to be ''Scyliorhinus'', and these two genera and ''
Poroderma ''Poroderma'' is a genus of catsharks, and part of the family Scyliorhinidae. The color of these sharks are usually grey with dark stripes. They are found in South Africa and are nocturnal. They live preferably in shore areas, close to the bott ...
'' form the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Scyliorhininae, the most basal
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, ...
of the order
Carcharhiniformes Carcharhiniformes , the ground sharks, are the largest order of sharks, with over 270 species. They include a number of common types, such as catsharks, swellsharks, and the sandbar shark. Members of this order are characterized by the presen ...
. The oldest confirmed ''Cephaloscyllium''
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s come from
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
(23–5.3 Mya) deposits in
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
, though based on the rate of DNA divergence in sharks, the swellshark lineage likely dates back to the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
(145.5–65.5 Mya).


Distribution and habitat

''Cephaloscyllium'' is widely distributed in the Indian and Pacific Oceans, in
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 ...
and
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
waters, but not close to the
equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can als ...
. Swellshark diversity is greatest around
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and in the west-central Pacific, where multiple
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 ...
species are found. The most far-flung members of the genus are ''C. sufflans'' off southeastern Africa, ''C. silasi'' off southwestern
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
, ''C. umbratile'' in the northwestern Pacific, and ''C. ventriosum'' along the western coast of the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
. One
hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
states ''Cephaloscyllium'' originally evolved in Australia and
New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...
, and subsequently dispersed outwards in a series of colonization events, eventually reaching Africa and the Americas. ''Cephaloscyllium'' species, bottom-dwelling sharks, can be found 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 on upper continental and insular slopes.


Description

Swellsharks are stocky and spindle shaped, with the trunk tapering substantially to the
caudal peduncle 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 ...
. The head is short (comprising less than a fifth of the total length), broad, and flattened. The snout is very short and blunt, with the
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 preceded by laterally enlarged flaps of skin. The eyes are horizontally oval or slit-like and placed high on the head, with cat-like
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 and rudimentary nictitating eyelids. A broad ridge occurs beneath each eye, and a
spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...
behind. The mouth is very large and wide, containing numerous small, multicusped teeth; the upper teeth are exposed when the mouth is closed (except in ''C. silasi''). No furrows are found at the corners of the mouth. 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 generally large and broad, and 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 ...
s are small. 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 placed far back on the body: the first dorsal fin originates 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 ...
origins, while the second dorsal fin is positioned about opposite 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 ...
. The first dorsal and anal fins are much larger than the second dorsal fin. The tail is short; the
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 ...
is broad with a distinct lower lobe and a strong ventral notch near the tip of the upper 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 ...
dermal denticles. The coloration is typically grayish or brownish, with a variety of patterns that include saddles, blotches, reticulations, and/or spots. In several species, the juveniles differ substantially in coloration from the adults. ''Cephaloscyllium'' species generally fall into two size groups: dwarf species include ''C. fasciatum'' and ''C. silasi'', which are under long, and large species include ''C. umbratile'' and ''C. ventriosum'', which may exceed in length.


Biology and ecology

Members of the genus ''Cephaloscyllium'' are generally slow-moving, sedentary animals with an anguilliform (
eel Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
-like) mode of swimming. Of the species whose diets have been documented, they have been known to feed on a wide variety of benthic organisms, including other sharks and rays,
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,
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, 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. Swellsharks are perhaps best known for their ability to inflate themselves by rapidly swallowing water or air when threatened; this behavior may allow them to wedge themselves inside crevices, make themselves harder to swallow, and/or intimidate a would-be predator. 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 ...
, with females producing two eggs at a time, one in each
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 ...
. Individual eggs are enclosed in a flask-shaped capsules with tendrils at the corners that allow them to be attached to underwater structures.


Human interactions

Swellsharks are harmless to humans and generally of no economic value, but are susceptible to being caught 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 ...
in
artisanal An artisan (from french: artisan, it, artigiano) is a skilled craft worker who makes or creates material objects partly or entirely by hand. These objects may be functional or strictly decorative, for example furniture, decorative art ...
and
commercial fisheries Commercial fishing is the activity of catching fish and other seafood for commercial profit, mostly from wild fisheries. It provides a large quantity of food to many countries around the world, but those who practice it as an industry must often ...
. Several species (e.g. ''C. umbratile'' and ''C. ventriosum''), are known to be extremely hardy, capable of surviving out of water for extended periods and adapting readily to captivity.


References

{{Taxonbar, from=Q2510308 Shark genera Taxa named by Theodore Gill