Squaliolus Laticaudus
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The spined pygmy shark (''Squaliolus laticaudus'') 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 squaliform shark in 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 ...
Dalatiidae The Dalatiidae are the family of kitefin sharks of the order Squaliformes (the term "kitefin shark" also refers specifically to the species ''Dalatias licha''). Members of this family are small, under long, and are found worldwide. They have cig ...
found widely in all oceans. Growing no larger than roughly , it is one of the smallest sharks alive, with this record beaten by the
dwarf lanternshark The dwarf lanternshark (''Etmopterus perryi'') is a species of dogfish shark in the family Etmopteridae and is the smallest shark in the world, reaching a maximum known length of . It is known to be present only on the upper continental slopes of ...
. This shark has a slender, cigar-shaped body with a sizable conical snout, a long but low second
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 ...
, and an almost symmetrical
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 ...
. Its
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 ...
''S. aliae'' and it are the only sharks with a spine on the first dorsal fin and not the second. Spined pygmy sharks are dark brown to black, with numerous
bioluminescent Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorganisms including ...
organs called
photophore A photophore is a glandular organ that appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors, ...
s on their ventral surface. The shark is believed to use these photophores to match ambient light conditions, which break up its silhouette and help the shark to avoid being seen by predators below. Usually inhabiting nutrient-rich waters over upper continental and insular slopes, the spined pygmy shark feeds on small
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
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
. Like its prey, it is a diel vertical migrator, spending the day at close to deep and moving towards a depth of at night. Reproduction is presumably
aplacental viviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
, with females giving birth to litters of up to four pups. This diminutive shark has no economic value. 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 ...
has assessed this species as of
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 faces little threat from
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 ...
and has a wide distribution.


Taxonomy and phylogeny

The spined pygmy shark was one of many new species discovered during the course of the 1907–1910 Philippine Expedition of the U.S. Fish Commission Steamer ''Albatross''. It was described 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 ...
s
Hugh McCormick Smith Hugh McCormick Smith, also H. M. Smith (November 21, 1865 – September 28, 1941) was an American ichthyologist and administrator in the United States Bureau of Fisheries. Biography Smith was born in Washington, D.C. In 1888, he received a Docto ...
and
Lewis Radcliffe Lewis Radcliffe (1880–1950) was a naturalist, malacologist, and ichthyologist. He was Deputy Commissioner of the United States Bureau of Fisheries until 1932 and was the assistant naturalist under Hugh McCormick Smith for the 1907-1910 Philippin ...
in a 1912 paper for the
scientific journal In academic publishing, a scientific journal is a periodical publication intended to further the progress of science, usually by reporting new research. Content Articles in scientific journals are mostly written by active scientists such as s ...
''Proceedings of the United States National Museum'', based on two specimens collected in
Batangas Bay Batangas Bay is a semi-enclosed body of water situated in the province of Batangas in Luzon island in the Philippines. The bay is an extension of the Verde Island Passage and is separated from the Balayan Bay to the west by the Calumpan Peninsula ...
, south of
Luzon Luzon (; ) is the largest and most populous island in the Philippines. Located in the northern portion of the Philippines archipelago, it is the economic and political center of the nation, being home to the country's capital city, Manila, as ...
in the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. One of these, a 15-cm-long adult male, was designated the
type specimen In biology, a type is a particular wiktionary:en:specimen, 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 a ...
. Smith and Radcliffe coined the new
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 ...
''Squaliolus'' for this shark, and 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 ...
''laticaudus'', 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 ...
''latus'' meaning "broad" or "wide", and ''cauda'' meaning "tail". The spined pygmy shark may also be referred to as the dwarf shark or the bigeye dwarf shark. Based on similarities in their
claspers 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 ...
(male
intromittent organ An intromittent organ is any external organ of a male organism that is specialized to deliver sperm during copulation. Intromittent organs are found most often in terrestrial species, as most non-mammalian aquatic species fertilize their eggs e ...
s), the closest relative of the spined pygmy shark and the related ''S. aliae'' is thought to be the pygmy shark (''Euprotomicrus bispinatus'').


Distribution and habitat

The spined pygmy shark has a wide distribution around the world. In the
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
, it occurs off
Bermuda ) , anthem = "God Save the King" , song_type = National song , song = " Hail to Bermuda" , image_map = , map_caption = , image_map2 = , mapsize2 = , map_caption2 = , subdivision_type = Sovereign state , subdivision_name = , e ...
, the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
,
Suriname Suriname (; srn, Sranankondre or ), officially the Republic of Suriname ( nl, Republiek Suriname , srn, Ripolik fu Sranan), is a country on the northeastern Atlantic coast of South America. It is bordered by the Atlantic Ocean to the north ...
, southern
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and northern
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
in the west, and off northern
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
,
Madeira ) , anthem = ( en, "Anthem of the Autonomous Region of Madeira") , song_type = Regional anthem , image_map=EU-Portugal_with_Madeira_circled.svg , map_alt=Location of Madeira , map_caption=Location of Madeira , subdivision_type=Sovereign st ...
,
Cape Verde , national_anthem = () , official_languages = Portuguese , national_languages = Cape Verdean Creole , capital = Praia , coordinates = , largest_city = capital , demonym ...
, and the
Azores ) , motto =( en, "Rather die free than subjected in peace") , anthem= ( en, "Anthem of the Azores") , image_map=Locator_map_of_Azores_in_EU.svg , map_alt=Location of the Azores within the European Union , map_caption=Location of the Azores wi ...
in the east. In the
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or ~19.8% of the water on Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia to the east. To the south it is bounded by th ...
, this species has only been recorded off
Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constituti ...
. In the
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 ...
, it is found off southern
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
,
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the nort ...
, and the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. The spined pygmy shark is found at depths of and seldom approaches the surface, unlike the related pygmy shark and
cookiecutter shark The cookiecutter shark (''Isistius brasiliensis''), also called the cigar shark, is a species of small squaliform shark in the family Dalatiidae. This shark occurs in warm, oceanic waters worldwide, particularly near islands, and has been reco ...
(''Isistius brasiliensis''). This shark prefers areas of high biological productivity over upper continental and insular slopes. It may also be found over outer shelves, but avoids central
ocean basin In hydrology, an oceanic basin (or ocean basin) is anywhere on Earth that is covered by seawater. Geologically, ocean basins are large  geologic basins that are below sea level. Most commonly the ocean is divided into basins foll ...
s. The range of this species does not overlap that of the pygmy shark, which has a similar ecology, and is also largely separate from that of the cookiecutter shark.


Description

One of the world's smallest sharks, the spined pygmy shark attains a maximum recorded length of for males and for females. This species has an elongated, spindle-shaped body with a long, bulbous, moderately pointed snout. The eyes are large, with the upper rim of the orbit almost straight. Each nostril is preceded by a short flap of skin. The mouth has thin, smooth lips and contains 22–31 tooth rows in the upper jaw and 16–21 tooth rows in the lower jaw. The upper teeth are narrow and smooth-edged with single upright cusps. The bases of the lower teeth are broad and interlocked to form a continuous cutting surface, with each tooth bearing a single upright, smooth-edged, knife-like cusp. The openings of 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 minute and uniform in size. The two species of ''Squaliolus'' are the only sharks with a spine on the first
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 ...
but not the second. The spine is
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, being typically exposed in males and enclosed by skin in females. The first dorsal fin is tiny and originates over the trailing margin of 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. The second dorsal fin is low, with a base twice as long as that of the first, and originates over the anterior half 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. The pectoral fins are short and triangular, with the rear margin slightly curved. The pelvic fins are long and low, and there is no
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
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 ...
is slender and laterally expanded into weak keels. 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 and paddle-like, with the upper and lower lobes of similar size and shape, and a deep notch in the trailing margin of the upper lobe. 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 scales, which can also provide effective camouflage through the use of reflection and colouration, as we ...
s are flat and blocky, not elevated on stalks or bearing marginal teeth. The coloration is dark brown to black, with light fin margins. The underside is densely carpeted by light-emitting
photophore A photophore is a glandular organ that appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors, ...
s, which extend to the tip of the snout and around the eyes and nostrils, and thin to almost non-existent on the back. This species has on average only 60
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e, the fewest of any shark.


Biology and ecology

The diet of the spined pygmy shark consists mainly 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 (including the dragonfish ''
Idiacanthus ''Idiacanthus'' is a genus of barbeled dragonfishes, the larvae of which are noted for exhibiting the Stylophthalmine trait. Species There are currently three recognized species in this genus: * '' Idiacanthus antrostomus'' C. H. Gilbert, 1890 ...
'', the
lanternfish Lanternfishes (or myctophids, from the Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family Myctophidae. One of two families in the order Myctophiformes, the Myctophidae are represente ...
''
Diaphus ''Diaphus'' is a genus of lanternfish Lanternfishes (or myctophids, from the Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family Myctophidae. One of two families in the order Myct ...
'', and the
bristlemouth The Gonostomatidae are a family of mesopelagic marine fish, commonly named bristlemouths, lightfishes, or anglemouths. It is a relatively small family, containing only eight known genera and 32 species. However, bristlemouths make up for their ...
''
Gonostoma ''Gonostoma'' is a genus of bristlemouths. Species There are three extant species recognized in this genus: * '' Gonostoma atlanticum'' Norman, 1930 (Atlantic fangjaw) * '' Gonostoma denudatum'' Rafinesque, 1810 * '' Gonostoma elongatum'' Günt ...
'') and
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
(including members of the genera ''
Chiroteuthis ''Chiroteuthis'' is a genus of chiroteuthid squid, comprising two subgenera. The hectocotylus is absent from all members of the genus; instead, a penis extending from the mantle opening is utilised. The genus is characterised by enlarged, lidde ...
'' and ''
Histioteuthis ''Histioteuthis'' is a genus of squid in the family Histioteuthidae. It goes by the common name cock-eyed squid, because in all species the right eye is normal-sized, round, blue and sunken; whereas the left eye is at least twice the diameter of ...
''). Catch records suggest that the spined pygmy shark follows its prey on their
diel vertical migration Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word ''diel'' comes from the Latin ''dies'' day, and means a 24-h ...
s, spending the day close to a depth of and ascending towards a depth of at night. The ventral photophores of the spined pygmy shark have been theorized to function in
counter-illumination Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength. Marine animals of ...
, a form of camouflage in which the shark disguises its silhouette from would-be
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 by matching the ambient light welling down from above. There is no evidence that this shark swallows its shed teeth like the pygmy and cookiecutter sharks. The spined pygmy shark is
aplacental viviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
like the rest of its family, with 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 ...
s being sustained by a
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 ...
until birth. Adult females have two functional
ovaries 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. T ...
that may each contain up to 12 mature eggs. However, the actual litter size is much smaller; a pregnant female caught off southern Brazil in 1999 contained four near-term pups. The young are born at long. Males mature sexually at a length of , and females at a length of . The spined pygmy shark was widely considered to be the smallest living shark species until the discovery of the
dwarf lanternshark The dwarf lanternshark (''Etmopterus perryi'') is a species of dogfish shark in the family Etmopteridae and is the smallest shark in the world, reaching a maximum known length of . It is known to be present only on the upper continental slopes of ...
(''Etmopterus perryi''), though the
pygmy ribbontail catshark The pygmy ribbontail catshark (''Eridacnis radcliffei'') is a species of finback catshark, family Proscylliidae, distributed patchily in the western Indo-Pacific from Tanzania to the Philippines. It occurs around the edges of continental and ins ...
(''Eridacnis radcliffei'') is also known to mature at a size comparable to these two species. Whether one of these sharks is definitively smaller than the others cannot yet be stated with certainty, because of the difficulties involved in assessing reproductive maturity in sharks.Martin, R.A. (July 28, 1999
What is the Smallest Species of Shark?
ReefQuest Centre for Shark Research. Retrieved on September 27, 2009.


Human interactions

Spined pygmy sharks have no commercial value; they sometimes appear in the
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 ...
of
trawl Trawling is a method of fishing that involves pulling a fishing net through the water behind one or more boats. The net used for trawling is called a trawl. This principle requires netting bags which are towed through water to catch different speci ...
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, both ...
, but are generally too small to be captured. In light of its wide distribution and the absence of substantial threats from human activity, 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 this species as of
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 ...
.


References

{{Taxonbar, from=Q952103 Squaliolus Fish described in 1912