Blotchy Swellshark
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The blotchy swellshark or Japanese swellshark (''Cephaloscyllium umbratile'') is a common
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of catshark, belonging to the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Scyliorhinidae Scyliorhinidae is a Family (biology), family of sharks, one of a few families whose members share the common name catsharks, belonging to the Order (biology), order Carcharhiniformes, the ground sharks. Although they are generally known as cats ...
. The Blotchy swellshark is found at depths of in the northwestern
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...
, from
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
to
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
. 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 the Ancient Greek word (), meaning "the depths". ...
in nature 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 component, abiotic (non-living) processes such as deposition (geol ...
s. Reaching in length, this thick-bodied shark has a broad head, large mouth, and two unequally-sized
dorsal fin A dorsal fin is a fin on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...
s positioned far back past the
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hi ...
s. It can be identified by its dorsal coloration, consisting of seven brown "saddles" and extensive darker mottling on a light tan background. This species has often been confounded with the draughtsboard shark (''C. isabellum'') and the Sarawak pygmy swellshark (''C. sarawakensis'') in
scientific literature Scientific literature encompasses a vast body of academic papers that spans various disciplines within the natural and social sciences. It primarily consists of academic papers that present original empirical research and theoretical ...
. Voracious and opportunistic in feeding habits, the blotchy swellshark is known to consume numerous types of
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
es and
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s, including an unusually high diversity of
cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
es. Like other ''Cephaloscyllium'' species, it is capable of rapidly inflating its body as a defense against predators. This species is
oviparous Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...
, with females laying encapsulated eggs two at a time. There is no well-defined
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 ch ...
and reproduction occurs year-round. The eggs hatch after approximately one year. The blotchy swellshark is harmless and fares well in captivity. It is 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
commercial Commercial may refer to: * (adjective for) commerce, a system of voluntary exchange of products and services ** (adjective for) trade, the trading of something of economic value such as goods, services, information or money * a dose of advertising ...
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 zone, demersal trawling. Benthic tra ...
s, though its population does not seem to have suffered from fishing activity.


Taxonomy

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, 35,800 species of fish had been described as of March 2 ...
s
David Starr Jordan David Starr Jordan (January 19, 1851 – September 19, 1931) was the founding president of Stanford University, serving from 1891 to 1913. He was an ichthyologist during his research career. Prior to serving as president of Stanford Universi ...
and
Henry Weed Fowler Henry Weed Fowler (March 23, 1878 – June 21, 1965) was an American zoologist born in Holmesburg, Philadelphia, Holmesburg, Pennsylvania. He studied at Stanford University under David Starr Jordan. He joined the Academy of Natural Sciences in Ph ...
described the blotchy swellshark in a 1903 volume of ''Proceedings of the United States National Museum'', based on a long stuffed dry skin originally obtained from
Nagasaki , officially , is the capital and the largest Cities of Japan, city of Nagasaki Prefecture on the island of Kyushu in Japan. Founded by the Portuguese, the port of Portuguese_Nagasaki, Nagasaki became the sole Nanban trade, port used for tr ...
, Japan. They gave it the
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''umbratile'' (from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''umbratilis'', meaning "shaded") and assigned it to the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Cephaloscyllium''. The
taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
of the blotchy swellshark has a history of confusion. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
dried skin could not be located when shark expert
Stewart Springer Stewart Springer (5 June 190623 August 1991) was an American ichthyologist and herpetologist. He was an expert on shark behavior, classification (taxonomy), and population distribution. More than 35 species of sharks, skates, rays, and other cr ...
prepared his 1979 review of the catsharks, and in its absence he synonymized ''C. umbratile'' with '' C. isabellum'' on the basis of "inconclusive
morphometric Morphometrics (from Greek μορΦή ''morphe'', "shape, form", and -μετρία ''metria'', "measurement") or morphometry refers to the quantitative analysis of ''form'', a concept that encompasses size and shape. Morphometric analyses are co ...
differences". Some authors followed Springer's judgment while others, particularly in Japan, preferred to keep referring to ''C. umbratile''. The taxonomy of this species was further muddled by the application of the name ''C. umbratile'' to a similar but smaller species sharing part of its range. This second species, once referred to as "pseudo-''umbratile''" by
Leonard Compagno Leonard Joseph Victor Compagno (1943-2024) was 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 Agri ...
, has since been identified as '' C. sarawakensis''. Recently, the holotype was found again, and in 2008 ''Cephaloscyllium umbratile'' was re-described as distinct from ''C. isabellum'' by Jayna Schaaf-Da Silva and David Ebert.


Distribution and habitat

The blotchy swellshark is known to inhabit the northwestern
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...
from
Hokkaido is the list of islands of Japan by area, second-largest island of Japan and comprises the largest and northernmost prefectures of Japan, prefecture, making up its own list of regions of Japan, region. The Tsugaru Strait separates Hokkaidō fr ...
,
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
southward to
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
, including the
Yellow Sea The Yellow Sea, also known as the North Sea, is a marginal sea of the Western Pacific Ocean located between mainland China and the Korean Peninsula, and can be considered the northwestern part of the East China Sea. Names It is one of four ...
. Its range may extend as far as
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
. This abundant species is a bottom-dweller that inhabits 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 component, abiotic (non-living) processes such as deposition (geol ...
s on the
continental shelf A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an islan ...
, at depths of .


Description

The maximum reported length of the blotchy swellshark is . It has a firm, stout body with a soft, distensible abdomen, and a short, broad, flattened head. The snout is proportionately long and rounded, with large nostrils divided by short, triangular flaps of skin in front. The small, horizontally oval eyes are placed high on the head and 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. Most ...
s (protective third eyelids). A tiny spiracle lies closely behind each eye. Behind the spiracle are 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 c ...
s, which are short and become progressively smaller posteriorly. The capacious mouth forms a broad arch, and lacks furrows at the corners. The small teeth have a central cusp flanked by a smaller cusplet on both sides. There are around 59 tooth rows in the upper jaw and 62 tooth rows in the lower jaw. The
pectoral fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish aquatic locomotion, swim. Apart from the tail or caudal fin, fish fins have no direct connection with the vertebral column ...
s are moderately large and wide, with rounded tips. The
dorsal fin A dorsal fin is a fin on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...
s have rounded apexes and are placed well back on the body, the first originating behind the midpoints of the small
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hi ...
s. The first dorsal fin is about twice as high as the second. The
anal fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported o ...
is nearly as large as the first dorsal fin and placed slightly ahead of the second dorsal fin. The
caudal fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only ...
is large and broad, with the upper lobe longer than the lower and bearing a prominent ventral notch near the tip. The skin is thick and sparsely covered by large, well- calcified
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 scale (zoology), scales, which can also provide effective Underwater camouflage, camouflage through the us ...
s; each denticle has a diamond-shaped crown with three horizontal ridges. This shark is cream-colored with dark brownish to grayish mottling on the back and sides, and seven dark brown dorsal "saddles" on the body and tail. The mottling intensifies with age, while the saddles fade and may become obscured. Older sharks may also have a dark blotch on either side between the pectoral and pelvic fins. The underside is pale, with scant darker marks.


Biology and ecology

Like other members of its
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
, when threatened the blotchy swellshark is capable of rapidly inflating its
stomach The stomach is a muscular, hollow organ in the upper gastrointestinal tract of Human, humans and many other animals, including several invertebrates. The Ancient Greek name for the stomach is ''gaster'' which is used as ''gastric'' in medical t ...
with water or air. This allows the shark to wedge itself inside a rocky crevice, becoming extremely difficult to remove. This species is an opportunistic, highly voracious
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
; one recorded female long had 10 fish about long and 15 squid about long in her stomach. Predominantly
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
, this species is known to prey upon
hagfish Hagfish, of the Class (biology), class Myxini (also known as Hyperotreti) and Order (biology), order Myxiniformes , are eel-shaped Agnatha, jawless fish (occasionally called slime eels). Hagfish are the only known living Animal, animals that h ...
and at least 50 species of
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
es, including fast-swimming types that inhabit open water; significant prey species include the
mackerel Mackerel is a common name applied to a number of different species of pelagic fish, mostly from the family Scombridae. They are found in both temperate and tropical seas, mostly living along the coast or offshore in the oceanic environment. ...
''
Scomber japonicus The chub mackerel, Pacific mackerel, or Pacific chub mackerel (''Scomber japonicus'') is a species of fish in the tuna and mackerel family, Scombridae. This species of mackerel closely resembles the Atlantic chub mackerel. Characteristics The c ...
'', the
sardine Sardine and pilchard are common names for various species of small, oily forage fish in the herring suborder Clupeoidei. The term "sardine" was first used in English during the early 15th century; a somewhat dubious etymology says it com ...
'' Sardinops melanostictus'', the
filefish The filefish (Monacanthidae) are a diverse family of tropical to subtropical tetraodontiform marine fish, which are also known as foolfish, leatherjackets, or shingles. They live in the Atlantic, Pacific, and Indian Oceans. Filefish are closely ...
'' Thamnaconus modestus'', and the hakeling '' Physiculus japonicus''. Unusually for such a small shark, it also feeds on at least 10 species of
cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
es, including lantern sharks, catsharks (particularly the
cloudy catshark The cloudy catshark (''Scyliorhinus torazame'') is a common species of Scyliorhinidae, catshark, belonging to the family (biology), family Scyliorhinidae. It is a benthic fish, bottom-dweller that inhabits rocky reefs in the northwestern Pacific ...
, ''Scyliorhinus torazame'', and its eggs), the
electric ray The electric rays are a group of rays, flattened cartilaginous fish with enlarged pectoral fins, composing the order Torpediniformes . They are known for being capable of producing an electric discharge, ranging from 8 to 220 volts, depending ...
'' Narke japonica'', and skates (including their eggs). It also cannibalizes smaller members of its own species.
Cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
s, mostly the
squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...
'' Doryteuthis bleekeri'' and the
cuttlefish Cuttlefish, or cuttles, are Marine (ocean), marine Mollusca, molluscs of the order (biology), suborder Sepiina. They belong to the class (biology), class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique ...
'' Sepia'' spp., are also frequently taken, while
crab Crabs are decapod crustaceans of the infraorder Brachyura (meaning "short tailed" in Greek language, Greek), which typically have a very short projecting tail-like abdomen#Arthropoda, abdomen, usually hidden entirely under the Thorax (arthropo ...
s,
shrimp A shrimp (: shrimp (American English, US) or shrimps (British English, UK)) is a crustacean with an elongated body and a primarily Aquatic locomotion, swimming mode of locomotion – typically Decapods belonging to the Caridea or Dendrobranchi ...
, and
isopod Isopoda is an Order (biology), order of crustaceans. Members of this group are called isopods and include both Aquatic animal, aquatic species and Terrestrial animal, terrestrial species such as woodlice. All have rigid, segmented exoskeletons ...
s are occasionally consumed. The dietary composition of juveniles varies notably from place to place. The blotchy swellshark is
oviparous Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...
, and reproduction proceeds throughout the year with no obvious seasonal cycling. Adult females have a single functional
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
, on the right, and two functional
oviduct The oviduct in vertebrates is the passageway from an ovary. In human females, this is more usually known as the fallopian tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, or will dege ...
s. The species is thought to be relatively prolific, as the ovary contains numerous
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and special episodes of a series made specially for release in home video formats without prior showings on television or in theaters, though the ...
at various stages of development. Pairs of eggs are laid at a time, one per oviduct. Females have been documented producing eggs even after years without male contact, suggesting that they may be able to store
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
. The purse-shaped egg capsules are relatively large and thick, measuring around long and across. The capsule surface is smooth with lengthwise striations, and opaque cream in color with yellow margins. Long, coiled tendrils extend from the four corners of the capsule. When the
embryo An embryo ( ) is the initial stage of development for 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 sp ...
is long, the
external gills External gills are the gills of an animal, most typically an amphibian, that are exposed to the environment, rather than set inside the pharynx and covered by gill slits, as they are in most fishes. Instead, the respiratory organs are set on a fri ...
have been lost, the
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 scale (zoology), scales, which can also provide effective Underwater camouflage, camouflage through the us ...
s have begun to develop, and light brown saddles are present. The eggs take roughly one year to hatch; newly emerged sharks measure long. From a series of captive rearing experiments, Sho Tanaka reported that hatchling sharks grew in length by up to per day. Males and females attain
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans, it is related to both puberty and adulthood. ''Puberty'' is the biological process of sexual maturation, while ''adulthood'', the condition of being socially recognized ...
at the size of and respectively; the growth rate after maturity is very low. Known
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
s of this species include the
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
''Porrocaecum cephaloscyllii'', and the
leech Leeches are segmented parasitism, parasitic or Predation, predatory worms that comprise the Class (biology), subclass Hirudinea within the phylum Annelida. They are closely related to the Oligochaeta, oligochaetes, which include the earthwor ...
''Stibarobdella macrothela''.


Human interactions

Harmless to humans, the blotchy swellshark adapts readily to captivity and has reproduced in
public aquarium A public aquarium () or public water zoo is the aquatic counterpart of a zoo, which houses living aquatic animal and aquatic plant, plant specimens for public viewing. Most public aquariums feature tanks larger than those kept by home aquarists, ...
s. This species is caught incidentally by Japanese and Taiwanese
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 zone, demersal trawling. Benthic tra ...
ers and brought to market. Intensive
commercial fishing Commercial fishing is the activity of catching fish and other seafood for Commerce, commercial Profit (economics), profit, mostly from wild fisheries. It provides a large quantity of food to many countries around the world, but those who practice ...
within its range do not yet appear to have impacted its numbers. The
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the stat ...
(IUCN) has listed it as
Near Threatened A near-threatened species is a species which has been Conservation status, categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature (IUCN) as that may be vulnerable to Endangered species, endangerment in the ne ...
.


References

{{Taxonbar, from=Q1949737 Cephaloscyllium Taxa named by David Starr Jordan Taxa named by Henry Weed Fowler Fish described in 1903