Squatina Squatina
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''Squatina squatina'', the angelshark or monkfish, is a species of
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
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 ...
Squatinidae (known generally also as
angel shark The angelsharks are a group of sharks in the genus ''Squatina'' of the family Squatinidae. They commonly inhabit sandy seabeds close to in depth. Many species are now classified as critically endangered by the International Union for Conservat ...
s), that were once widespread in the coastal waters of the northeastern
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 ...
. Well-adapted for
camouflaging Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
itself on the sea floor, the angelshark has a flattened form with enlarged
pectoral Pectoral may refer to: * The chest region and anything relating to it. * Pectoral cross, a cross worn on the chest * a decorative, usually jeweled version of a gorget * Pectoral (Ancient Egypt), a type of jewelry worn in ancient Egypt * Pectoralis ...
and
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, giving it a superficial resemblance to a ray. This species can be identified by its broad and stout body, conical
barbels In fish anatomy and turtle anatomy, a barbel is a slender, whiskerlike sensory organ near the mouth. Fish that have barbels include the catfish, the carp, the goatfish, the hagfish, the sturgeon, the zebrafish, the black dragonfish and some ...
, thornless back (in larger individuals), and grayish or brownish dorsal coloration with a pattern of numerous small light and dark markings (that is more vivid in juveniles). It measures up to long. Like other members of its family, the angelshark is a
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey us ...
that buries itself in sediment and waits for passing prey, mostly
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 ...
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, but also
skate Skate or Skates may refer to: Fish *Skate (fish), several genera of fish belonging to the family Rajidae * Pygmy skates, several genera of fish belonging to the family Gurgesiellidae * Smooth skates or leg skates, several genera of fish belongin ...
s 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. An
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 ...
species, females bear litters of seven to 25 pups every other year. The angelshark normally poses little danger to humans, though if provoked, it is quick to bite. Since the mid-20th century, intense
commercial fishing 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 ...
across the angelshark's range has decimated its population via
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 ...
– it is now locally extinct or nearly so across most of its northern range, and the prospects of the remaining fragmented subpopulations are made more precarious by its slow rate of reproduction. As a result, the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
has assessed this species as Critically Endangered.


Taxonomy and phylogeny

The angelshark was originally described by the Swedish natural historian
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, known as the "father of
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
", in the 1758 tenth edition of ''
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nomen ...
'' as ''Squalus squatina''. He did not designate a
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 ...
. The word ''squatina'' is the name for
skate Skate or Skates may refer to: Fish *Skate (fish), several genera of fish belonging to the family Rajidae * Pygmy skates, several genera of fish belonging to the family Gurgesiellidae * Smooth skates or leg skates, several genera of fish belongin ...
in
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 ...
; it was made the genus name for all angel sharks by the French zoologist André Duméril in 1806. Other
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 ...
s used for this species include angel, angel fiddle fish, angel puffy fish, angel ray, angelfish, escat jueu, fiddle fish, monk, and monkfish. Stelbrink and colleagues (2010) conducted a
phylogenetic 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 ...
study based on
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 ...
, and found that the
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 ...
of the angelshark is the sawback angelshark (''S. aculeata''). The two species formed a
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, ...
with a number of Asian angelshark species.


Description

One of the largest members of its family, female angelsharks can attain a length of and males ; the maximum reported weight is . This species shares in common with other angelsharks a flattened body and large, wing-like
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 whose anterior lobes are not fused to the head. The head and body are very broad and stocky, with small eyes positioned dorsally and followed by a pair of larger
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 ...
s. A pair of unadorned
barbel Barbel may refer to: *Barbel (anatomy), a whisker-like organ near the mouth found in some fish (notably catfish, loaches and cyprinids) and turtles *Barbel (fish), a common name for certain species of fish **''Barbus barbus'', a species of cyprinid ...
s occurs in front of the nares, as well as a smooth or weakly fringed flap. Folds of skin with a single triangular lobe are present on the sides of the head. The teeth are small, sharp, and of similar shape in both jaws. The pectoral and pelvic fins are wide with rounded tips; 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 positioned on the muscular tail behind the pelvic fins. 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 ...
is absent, and 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 ...
has a larger lower lobe than upper. 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 small, narrow, and pointed, and cover the entire upper and most of the lower body surface. There are patches of small spines on the snout and over the eyes. Small individuals have a row of thorns down the middle of the back. The coloration is gray to reddish or greenish brown above, with many small black and white spots, and white below. Juveniles are more ornately patterned than adults, with pale lines and darker blotches. The dorsal fins have a darker leading margin and lighter trailing margin. Some individuals have a white spot on the back of the "neck".


Distribution and habitat

The angelshark occurs in 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 of the northeastern Atlantic, from southern
Norway Norway, officially the Kingdom of Norway, is a Nordic country in Northern Europe, the mainland territory of which comprises the western and northernmost portion of the Scandinavian Peninsula. The remote Arctic island of Jan Mayen and t ...
and
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...
to the
Western Sahara Western Sahara ( '; ; ) is a disputed territory on the northwest coast and in the Maghreb region of North and West Africa. About 20% of the territory is controlled by the self-proclaimed Sahrawi Arab Democratic Republic (SADR), while the r ...
and the
Canary Islands The Canary Islands (; es, Canarias, ), also known informally as the Canaries, are a Spanish autonomous community and archipelago in the Atlantic Ocean, in Macaronesia. At their closest point to the African mainland, they are west of Morocc ...
, including around the
British Isles The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isles, ...
and in the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
. According to the
IUCN 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 ...
, it is possible that it has been
extirpated Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
from the
North Sea The North Sea lies between Great Britain, Norway, Denmark, Germany, the Netherlands and Belgium. An epeiric sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian S ...
. It remains extant around the Canary Islands, Algeria, Tunisia, Libya, Israel, Turkey, northern Cyprus, eastern Greece (the
Aegean Sea The Aegean Sea ; tr, Ege Denizi (Greek language, Greek: Αιγαίο Πέλαγος: "Egéo Pélagos", Turkish language, Turkish: "Ege Denizi" or "Adalar Denizi") is an elongated embayment of the Mediterranean Sea between Europe and Asia. It ...
), the Adriatic Sea of eastern Italy, Sicily, Malta, Corsica, Ireland and western Britain/Wales . Its modern presence in parts of the Mediterranean is unknown, such as around Madeira, the Azores, Morocco, Egypt, continental Spain and France, Crete, Syria, Sardinia, western Greece and western Italy. This
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 ...
shark inhabits 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 island ...
, preferring soft substrates such as mud or sand, and can be found from near the coast to a depth of . It sometimes enters
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
environments. Northern angelshark subpopulations migrate northward in summer and southward in winter.


Biology and ecology

During daytime, the angelshark usually lies motionless on the sea floor, buried under a layer of sediment with only its eyes showing. At night, it becomes more active, and may sometimes be seen swimming above the bottom. Aggregations numbering up to a hundred have been observed off
Gran Canaria Gran Canaria (, ; ), also Grand Canary Island, is the third-largest and second-most-populous island of the Canary Islands, an archipelago off the Atlantic coast of Northwest Africa which is part of Spain. the island had a population of that co ...
in the summer.Murch, A
Common Angel Shark Information and Pictures
Elasmodiver.com. Retrieved on July 8, 2009.
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 ...
s of this species include the
tapeworm Eucestoda, commonly referred to as tapeworms, is the larger of the two subclasses of flatworms in the class Cestoda (the other subclass is Cestodaria). Larvae have six posterior hooks on the scolex (head), in contrast to the ten-hooked Cestodar ...
s ''Grillotia smaris-gora'', ''G. angeli'', and '' Christianella minuta'', the
fluke Fluke may refer to: Biology * Fluke (fish), a species of marine flatfish * Fluke (tail), the lobes of the tail of a cetacean, such as dolphins or whales, ichthyosaurs, mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ...
''Pseudocotyle squatinae'', the
monogenea Monogeneans are a group of ectoparasitic flatworms commonly found on the skin, gills, or fins of fish. They have a direct lifecycle and do not require an intermediate host. Adults are hermaphrodites, meaning they have both male and female reprod ...
n ''Leptocotyle minor'', and the
isopod Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, an ...
''Aega rosacea''. The angelshark is an ambush predator that feeds mainly on bottom-dwelling bony fishes, especially
flatfish A flatfish is a member of the Ray-finned fish, ray-finned demersal fish order (biology), order Pleuronectiformes, also called the Heterosomata, sometimes classified as a suborder of Perciformes. In many species, both eyes lie on one side of the ...
es, though it also preys on skates and invertebrates. Prey reported taken include the
hake The term hake refers to fish in the: * Family Merlucciidae of northern and southern oceans * Family Phycidae (sometimes considered the subfamily Phycinae in the family Gadidae) of the northern oceans Hake Hake is in the same taxonomic order (Gad ...
''
Merluccius merluccius ''Merluccius merluccius'', the European hake, is a merluccid hake of the genus '' Merluccius''. Other vernacular names include Cornish salmon and herring hake. It is a predatory species which was often netted alongside one of its favoured prey, t ...
'', the
bream Bream ( ) are species of freshwater and marine fish belonging to a variety of genera including ''Abramis'' (e.g., ''A. brama'', the common bream), ''Acanthopagrus'', '' Argyrops'', ''Blicca'', '' Brama'', ''Chilotilapia'', '' Etelis'', ''Lepo ...
''
Pagellus erythrinus The common pandora (''Pagellus erythrinus'') is a fish of the sea bream family, Sparidae. It is a popular food fish in Mediterranean countries, with delicate white flesh. It has a slim, oval fusiform body, with a smallish mouth and scales cover ...
'', grunts in the genus ''
Pomadasys ''Pomadasys'' is a genus of grunts native to the waters of the eastern Atlantic Ocean and through the Indian Ocean to the Pacific coast of the Americas. The name of this genus is a compound of ''poma'' meaning "lid" or "covering" and ''dasys'' m ...
'', the flatfishes ''
Bothus ''Bothus'' is a genus of flatfish in the family Bothidae (lefteye flounders) from the Pacific, Indian and Atlantic Oceans. Some species in this genus have spots consisting of blue rings. Species There are currently 16 recognized species in thi ...
'' spp., '' Citharus linguatula'', and ''
Solea solea The common sole, Dover sole, or black sole (''Solea solea'') is a species of flatfish in the family Soleidae. It is one of the largest fish in the ''Solea'' genus. It lives on the sandy or muddy seabed of the northern Atlantic and the Mediterra ...
'', the
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 ...
''
Loligo vulgaris The European squid or common squid (''Loligo vulgaris'') is a large squid belonging to the family Loliginidae. It occurs abundantly in coastal waters from the North Sea to at least the west coast of Africa. This species lives from sea level to d ...
'', the
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...
es ''
Sepia officinalis #REDIRECT Common cuttlefish #REDIRECT Common cuttlefish #REDIRECT Common cuttlefish {{redirect category shell, {{R from other capitalisation{{R from move ... {{redirect category shell, {{R from other capitalisation{{R from move ...
{{redire ...
'' and '' Sepiola'' spp., and the
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
s '' Medorippe lanata'', '' Geryon trispinosus'', ''
Dromia personata ''Dromia personata,'' also known as the sponge crab or sleepy crab, is a species of crab found in the North Sea, the Mediterranean Sea, and connecting parts of the northeastern Atlantic Ocean. Like most other epibenthic crustaceans, the biomass ...
'', '' Goneplax rhomboides'', '' Liocarcinus corrugatus'', and '' Atelecyclus rotundatus''. The stomachs of some examined specimens have also contained
seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families (Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the orde ...
or birds (in one case an entire
cormorant Phalacrocoracidae is a family of approximately 40 species of aquatic birds commonly known as cormorants and shags. Several different classifications of the family have been proposed, but in 2021 the IOC adopted a consensus taxonomy of seven ge ...
). Individual sharks select sites that offer the best ambush opportunities, and if successful, may remain there for several days. Angelsharks are aplacental viviparous, meaning the young hatch inside the mother's
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
and are nourished 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. 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 ...
, with the right ovary containing more
oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...
s and the right uterus correspondingly containing more embryos; this functional asymmetry is not present in other angel shark species. Unlike most sharks, in which
vitellogenesis Vitellogenesis is the process of yolk protein formation in the oocytes of non mammalian vertebrates during sexual maturation. The term ''vitellogenesis'' comes from the Latin ''vitellus'' ("egg yolk"). Yolk proteins, such as Lipovitellin and Phosv ...
(yolk formation) occurs concurrently with pregnancy, in the angelshark, the onset of vitellogenesis is delayed until halfway through the
gestation period In mammals, pregnancy is the period of reproduction during which a female carries one or more live offspring from implantation in the uterus through gestation. It begins when a fertilized zygote implants in the female's uterus, and ends once it ...
. The mature ova measure across and are not enclosed in a capsule. The reproductive cycle has been estimated at 2 years with
ovulation Ovulation is the release of eggs from the ovaries. In women, this event occurs when the ovarian follicles rupture and release the secondary oocyte ovarian cells. After ovulation, during the luteal phase, the egg will be available to be fertilized ...
taking place in spring, though this periodicity is ill-defined. The litter size ranges from seven to 25 and is correlated with the size of the mother; the young are gestated for 8–10 months. Parturition occurs from December to February in the Mediterranean and in July off England, with the newborns measuring long. Males and females mature at lengths of and , respectively.


Human interactions

The angelshark is generally not aggressive towards humans, though it can deliver a severe bite if disturbed. When approached underwater, the angelshark usually remains still or swims away, though one circling a diver with its mouth open is recorded. Fishery workers, in particular, should treat it with caution; in the 1776 edition of ''British Zoology'', Thomas Pennant wrote that it is "extremely fierce and dangerous to be approached. We know of an instance of a fisherman, whose leg was terribly torn by a large one of this species, which lay within his nets in shallow water, and which he went to lay hold of incautiously." Humans have used the angelshark for thousands of years.
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
authors, such as Diphilus and Mnesitheus, described its meat as "light" and "easily digestible", and
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic '' ...
noted in his ''
Naturalis Historia The ''Natural History'' ( la, Naturalis historia) is a work by Pliny the Elder. The largest single work to have survived from the Roman Empire to the modern day, the ''Natural History'' compiles information gleaned from other ancient authors. ...
'' (77–79 AD) that its rough skin was valued by craftsmen for polishing
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin th ...
and
ivory Ivory is a hard, white material from the tusks (traditionally from elephants) and teeth of animals, that consists mainly of dentine, one of the physical structures of teeth and tusks. The chemical structure of the teeth and tusks of mammals is ...
.
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
recorded elements of its natural history, including that it bore live young, and correctly recognized that it was a shark despite its resemblance to rays and skates. The use of this species for food has continued into modern times; it is sold fresh or dried and salted, often under the name "monkfish" (which also refers to the
goosefish Goosefishes are anglerfishes in the family Lophiidae found in the Arctic, Atlantic, Indian, and Pacific Oceans, where they live on sandy and muddy bottoms of the continental shelf and continental slope, to depths of more than . Like most other an ...
es of the genus ''Lophius''). The angelshark may also be a source for
shark liver oil Shark liver oil is an oil obtained from the livers of sharks. It has been used for centuries as a folk remedy to promote the healing of wounds and as a remedy for respiratory tract and digestive system problems.
and
fishmeal Fish meal is a commercial product made from whole wild-caught fish, bycatch and fish by-products to feed farm animals, e.g., pigs, poultry, and farmed fish.R. D. Miles and F. A. Chapman.FA122: The Benefits of Fish Meal in Aquaculture DietsFisherie ...
.


Conservation status

Sources from the 19th and early 20th centuries indicate that the angelshark was once abundant all around the coasts of
Western Europe Western Europe is the western region of Europe. The region's countries and territories vary depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the ancient Mediterranean ...
. Yarrell (1836), Day (1880–04), and Garstang (1903) all noted that the angelshark was common around the British Isles, and Rey (1928) recorded that this species was common around the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
and in the Mediterranean. However, from the latter half of the 20th century onwards, the angelshark has come under intense pressure from commercial fisheries operating across much of its range. Due to its benthic, near-shore habits, individuals of all ages are susceptible to incidental capture by
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,
trammel net A fishing net is a net used for fishing. Nets are devices made from fibers woven in a grid-like structure. Some fishing nets are also called fish traps, for example fyke nets. Fishing nets are usually meshes formed by knotting a relatively thin ...
s, and bottom longlines; the low reproductive rate of this shark limits its capacity to withstand population depletion. This has also led to habitat loss caused by the development of coastal areas for commercialism and tourism. Angelshark numbers have declined precipitously across most of its range; it is now believed to be extinct in the North Sea and most of the northern Mediterranean, and has become extremely rare elsewhere. During the comprehensive Mediterranean International Trawl Survey program from 1995 to 1999, only two angelsharks were captured from 9,905 trawls. Similarly, another survey by the Italian National Project (National Group for Demersal Resource Evaluation) around the same period caught angelsharks in only 38 of 9,281 trawls. Fishery data compiled by the Working Group for Elasmobranch Fishes (WGEF) show that no angelsharks have been landed in the Northeast Atlantic since 1998. Fewer than a dozen angelsharks are thought to remain in Irish waters. Healthy subpopulations of angelsharks are thought to still persist in areas off
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
and around the Canary Islands, though a more thorough assessment is urgently needed. As a result of these steep population declines and the ongoing threat from
demersal The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer of ...
fisheries, the
IUCN 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 the angelshark as Critically Endangered. An assessment of the angelshark population by the IUCN showed a decrease in population of over 90%. The assessment also showed that there was no signs of recovery of the population. It was listed on Annex III of the 1976
Barcelona Convention The Convention for the Protection of the Marine Environment and the Coastal Region of the Mediterranean, originally the Convention for Protection of the Mediterranean Sea against Pollution,pollution Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, the ...
in the Mediterranean Sea. In 2012 it was moved to Annex II, making it illegal to catch and keep in countries bordering the Mediterranean Sea (if caught, it must be released). This species is protected within three marine reserves in the
Balearic Islands The Balearic Islands ( es, Islas Baleares ; or ca, Illes Balears ) are an archipelago in the Balearic Sea, near the eastern coast of the Iberian Peninsula. The archipelago is an autonomous community and a province of Spain; its capital is ...
, although it has not been reported from this area since the mid-1990s. In 2008, the angelshark also received full legal protection from human activities in the waters off England and
Wales Wales ( cy, Cymru ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by England to the Wales–England border, east, the Irish Sea to the north and west, the Celtic Sea to the south west and the ...
from the coast to a distance of , under the UK
Wildlife and Countryside Act The Wildlife and Countryside Act 1981 is an Act of Parliament in the United Kingdom implemented to comply with European Council Directive 79/409/EEC on the conservation of wild birds. In short, the act gives protection to native species (especia ...
. Since 2010, it has been illegal to keep angelsharks caught in waters of the European Union (if caught, it must be released). The
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
and
Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to th ...
have pushed, unsuccessfully, for this species to be listed on the
Convention for the Protection of the Marine Environment of the North-East Atlantic Convention may refer to: * Convention (norm), a custom or tradition, a standard of presentation or conduct ** Treaty, an agreement in international law * Convention (meeting), meeting of a (usually large) group of individuals and/or companies in a ...
Priority List of Threatened and Endangered Species. A captive breeding program has been initiated at Deep Sea World,
North Queensferry North Queensferry is a village in Fife, Scotland, situated on the Firth of Forth where the Forth Bridge, the Forth Road Bridge, and the Queensferry Crossing all meet the Fife coast, some from the centre of Edinburgh. It is the southernmost sett ...
, with the first live pups born in 2011. In 2019, a population of angelsharks was discovered off the coast of
Wales Wales ( cy, Cymru ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by England to the Wales–England border, east, the Irish Sea to the north and west, the Celtic Sea to the south west and the ...
, indicating that the species had begun a potential return to the region.


References


External links


"''Squatina squatina'', Angelshark" at FishBase"Species description of ''Squatina squatina''" at Shark-References.com
* {{Good article Squatinidae Fish of the East Atlantic Fish of the Mediterranean Sea Fauna of the Canary Islands Marine fauna of North Africa Marine fish of Europe Critically endangered fish Critically endangered fauna of Africa Critically endangered biota of Europe Fish described in 1758 Taxa named by Carl Linnaeus