Overview

Comprehensive Description

Biology

Occurs on the outer continental shelf and the upper slope. Found from shoreline to depths of 1,375 m (Ref. 26938). Feeds on small bottom fishes, crustaceans and bivalves. Ovoviviparous. Will bite when captured and can inflict nasty lacerations with its trap-like jaws and pointed teeth.
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Distribution

National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Western Atlantic: southern New England, USA to the Gulf of Mexico, Jamaica and Venezuela. Also in Columbia
  • North-West Atlantic Ocean species (NWARMS)
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Range Description

Found along the Atlantic coast of the United States from southern New England to Florida (Massachusetts, New York, Rhode Island, Delaware, Maryland, Virginia, North Carolina, South Carolina, south to the Florida Keys) and the Gulf of Mexico (Alabama, Mississippi, Louisiana, Texas and the Mexican coast). Reported but unconfirmed from Cuba, Nicaragua, Jamaica and Venezuela (Castro 1983, Compagno in prep. a).
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Western Atlantic: southern New England, USA to the Gulf of Mexico, Jamaica and Venezuela. Also in Columbia (Ref. 26938).
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Western Atlantic.
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Physical Description

Size

Maximum size: 1520 mm TL
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Max. size

152 cm TL (male/unsexed; (Ref. 247))
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Diagnostic Description

A ray-shaped shark. Head narrowing to distinct neck at base of pectoral fins (Ref. 26938).
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The main confusion among WNP squatinids lies in differentiating S. formosa and S. nebulosa from one another. The main character used in the past to differentiate these two species was the number of dermal lobes at the corners of the mouth (Chen 1963, Nakabo 2002, Compagno 2005a). Based upon our examination, this character does not appear to be robust in separating these species. Although all S. formosa individuals studied had one dermal lobe, S. nebulosa specimens had one or two lobes, invalidating this character as a method of separating the two species. Additionally, the number of dermal lobes was often difficult to determine since differentiation between lobes is often vague and these lobes are susceptible to damage during capture and or preservation of specimens. A combination of characters should be used to identify individuals of these two species, since the differences are subtle and a great deal of overlap exists among many characters. The best characters available, based on our findings, appear to be the shape of the caudal pelvic and dorsal fins, the ratio of standardized pelvic girdle distance with the standardized head length, and the shape and height of the upper lip arch. The upper lip arch appears to be of particular value as a field character. It is a feature that is not prone to damage and is easy to assess quickly in the field. More specimens are needed to assess the utility of this character but it appears to be useful for all WNP squatinids.

 

Based on the key characters presented herein, we provide the following key to the WNP squatinids:

 

Key to Western North Pacific Squatina Species

 

1. Pelvic fin tips do not extend to origin of first dorsal fin..............................................................................2.

 

- Pelvic fin tips extend to or surpass origin of first dorsal fin........................................................................3.

 

2. A prominent row of thorn-like denticles extending from mid-back to caudal peduncle; no distinct ocelli on posterior lobes of the pectoral fins................................................................................................ S. japonica

 

- No row of thorn-like denticles extending from the mid-back to the caudal peduncle, distinct paired ocelli on the posterior lobes of the pectoral fins......................................................................... S. tergocellatoides

 

3. Upper lip arch semi-circular in shape (>1.5% TL in height); dorsals are lobed with a curvilinear anterior margin; pelvic girdle distance 1.4 times or less head length; caudal fin is lobed, especially dorsally, with a curvilinear postventral caudal margin........................................................................................... S. formosa

 

- Upper lip arch is not semi-circular in shape (<1.5% TL in height); dorsals are not lobed (angular) without a curvilinear anterior margin (straight); pelvic girdle distance greater than 1.4 times head length; caudal fin is not lobed (angular), especially dorsally, without a curvilinear postventral caudal margin (straight) ... ..................................................................................................................................................... S. nebulosa

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Squatina Dumeril, 1806 is a monotypic genus within the family Squatinidae (Bonaparte, 1838). Although morphologically similar to batoids, squatinids are distinct from true batoids in that the squatinids have lateral gill openings, pectoral fin lobes that are free lateral to the gills, and possess a lower caudal fin lobe that is longer than the upper caudal lobe. Within the genus these sharks can be difficult to distinguish due to the lack of well defined characteristics. Adding to the confusion in the literature are the inadequate original descriptions of many species.

 

Sixteen valid species are recognized worldwide, with four reported to occur in the western North Pacific (WNP; Compagno et al., 2005). These four species include Squatina formosa Shen and Ting, 1972 , S. japonica Bleeker, 1858 , S. nebulosa Regan, 1906 , and S. tergocellatoides Chen, 1963 . Distinctions among these four WNP species hinge upon the nasal barbel shape, interorbital and interspiracle distances, ocellus patterns, number of dermal folds about the mouth, and the presence of midback thorns (Lindberg & Legeza, 1967; Shen & Ting, 1972; Nakabo, 2002). However, specific identification is hard to assign to individuals because many of these characters are difficult to distinguish, and many characters currently used are susceptible to damage during collection or from preservation. Additionally, inadequate original descriptions for some species and confusion within the subsequent literature have further obscured definitive characters among the WNP species. Because members of this genus are frequently targeted in fisheries in an area that has sparsely recorded catch information, and congeners are particularly sensitive to fishing pressure (Gaida, 1997; Stevens et al, 2000), it is imperative that adequate descriptions are available to identify individual species.

 

During two field expeditions to Taiwan, one of us (DAE) observed at least four species of Squatina that were frequently landed at fish markets around Taiwan. Attempts to identify the various species were often hampered by a lack of adequate fish keys and descriptive characters for the various Squatina species observed. Furthermore, we had the opportunity to examine the holotype and three paratypes of S. formosa as well as collect additional material.

 

Using this information, we provide a detailed redescription of three of the species, S. formosa , S. japonica , and S. nebulosa , with new material from Japanese and Taiwanese waters. In addition, we supply a list of important key characters for distinguishing all known WNP squatinids, especially to facilitate identification of the two most similar species, S. formosa and S. nebulosa .

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Ecology

Habitat

Habitat Type: Marine

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benthic
  • North-West Atlantic Ocean species (NWARMS)
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Habitat and Ecology

Habitat and Ecology
Little is known about this species. It is found seasonally in shallower water. Off the eastern United States it appears to move inshore in the spring and summer, and disappears, apparently into deeper water in other seasons.

Maturity is probably reached around 90 to 107 cm TL. Reproductive mode is aplacental viviparous with only the left ovary functional. Pupping occurs in June or July with pups measuring 28 to 30 cm TL at birth. Litters contain up to 25 pups. Birth usually occurs at depth (~20 to 30 m).

Eats small bottom fishes (flounders, skates and other bottom fishes), crustaceans and bivalves.

Life history parameters
Age at maturity (years): Unknown.
Size at maturity (total length): Female: 90 to 105 cm TL, median length 83.5 cm FL (Baremore and Carlson 2004); Male: 92 to 107 cm TL, median length 88.7 cm FL (Baremore and Carlson 2004).
Longevity (years): Unknown.
Maximum size (total length): 152 cm TL.
Size at birth: 28 to 30 cm TL.
Average reproductive age (years): Unknown.
Gestation time (months): Unknown.
Reproductive periodicity: Unknown.
Average annual fecundity or litter size: Up to 25/litter (Compagno in prep. a); 8 (+/- 1.82) pups (Baremore and Carlson 2004).
Annual rate of population increase: Unknown.
Natural mortality: Unknown.

Systems
  • Marine
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Environment

demersal; marine; depth range ? - 1390 m, usually ? - 128 m (Ref. 55306)
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Depth range based on 446 specimens in 1 taxon.
Water temperature and chemistry ranges based on 275 samples.

Environmental ranges
  Depth range (m): 6 - 1289
  Temperature range (°C): 3.798 - 24.310
  Nitrate (umol/L): 0.663 - 23.726
  Salinity (PPS): 32.865 - 36.472
  Oxygen (ml/l): 3.526 - 6.300
  Phosphate (umol/l): 0.124 - 1.339
  Silicate (umol/l): 1.657 - 16.452

Graphical representation

Depth range (m): 6 - 1289

Temperature range (°C): 3.798 - 24.310

Nitrate (umol/L): 0.663 - 23.726

Salinity (PPS): 32.865 - 36.472

Oxygen (ml/l): 3.526 - 6.300

Phosphate (umol/l): 0.124 - 1.339

Silicate (umol/l): 1.657 - 16.452
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Depth: 0 - 1390m.
Recorded at 1390 meters.

Habitat: demersal. Occurs on the outer continental shelf and the the upper slope. Feeds on small bottom fishes, crustaceans and bivalves. Ovoviviparous. Will bite when captured and can inflict nasty lacerations with its traplike jaws and pointed teeth. Ovoviviparous.
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Migration

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

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Trophic Strategy

Occurs on the outer continental shelf and the upper slope. Feeds on small bottom fishes, crustaceans and bivalves.
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Life History and Behavior

Life Cycle

Ovoviviparous, embryos feed solely on yolk (Ref. 50449).
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Squatina dumeril

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 12 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

ACCCTTTATTTAATTTTTGGTGCATGAGCAGGAATAGTAGGTACTGCCCTA---AGTATACTCATCCGAGCAGAATTAAGCCAGCCCGGAACACTCCTTGGGGAT---GATCAAATTTACAATGTAATTGTTACTGCCCACGCTTTCGTAATAATCTTTTTTATAGTTATACCAATCATAATCGGTGGATTTGGAAATTGATTAGTTCCATTAATA---ATTGGTGCACCAGATATAGCCTTTCCACGAATAAATAATATAAGCTTCTGACTTTTACCTCCTTCCCTATTATTATTACTTGCTTCAGCTGGAGTTGAAGCAGGGGCCGGCACTGGTTGAACAGTGTATCCCCCTCTTGCAGGAAATCTAGCTCACGCCGGAGCATCGGTAGATTTA---GCAATTTTTTCCCTACACTTAGCTGGTATCTCTTCAATTTTAGCCTCTATTAATTTCATTACAACTATTATTAACATAAAATCCCCAGCTACCTCCCAATATCAAACACCACTCTTTGTTTGATCAATTTTGGTAACTACTGTACTTCTCCTCCTTTCCCTCCCAGTTCTCGCAGCT---GCAATCACAATATTACTAACCGACCGTAATCTAAATACAACATTTTTTGACCCTGCAGGAGGGGGAGACCCAATCCTTTACCAACACTTA------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------TTC
-- end --

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Statistics of barcoding coverage: Squatina dumeril

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 12
Specimens with Barcodes: 19
Species With Barcodes: 1
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Conservation

Conservation Status

National NatureServe Conservation Status

United States

Rounded National Status Rank: NNR - Unranked

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NatureServe Conservation Status

Rounded Global Status Rank: GNR - Not Yet Ranked

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IUCN Red List Assessment


Red List Category
DD
Data Deficient

Red List Criteria

Version
3.1

Year Assessed
2006

Assessor/s
Heupel, M.R. & Carlson, J.K.

Reviewer/s
Cavanagh, R.D. & Kyne, P.M.

Contributor/s

Justification
Squatina dumeril is found in the western Atlantic in depths up to 1,290 m, but is seasonally recorded in shallower regions. This species is not known to be targeted or utilized, but is taken in trawl fisheries as accidental bycatch although no data is available. There is currently no information available on the life history of this species (beyond a limited description of its reproductive biology) and its full distribution is not defined, therefore it is listed as Data Deficient until more information is available. Studies are currently underway and this assessment should be reviewed in the near future.
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Population

Population
An apparently common but largely unknown species of temperate and subtropical waters of the Western North Atlantic continental shelf and slope. Typically found on or near the bottom from close inshore down to depths of 1,290 m, however, most records are from 40 to 250 m.

Population Trend
Unknown
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Threats

Major Threats
Accidental bycatch in trawl fisheries. In the Gulf of Mexico, Angel Sharks occur with relative frequency as bycatch in trawl fisheries targeting butterfish (Carlson, pers obs.) and it is possible they occur as bycatch in other fisheries. Note that S. dumeril is often found in deeper waters where shrimp trawling effort is low.

Utilisation
Not known to be utilised.
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Data deficient (DD)
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Management

Conservation Actions

Conservation Actions
In Atlantic US waters, angel sharks are currently listed as prohibited under the Federal Management for Atlantic tunas, swordfish and sharks, as a precautionary management measure.
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Relevance to Humans and Ecosystems

Benefits

Importance

fisheries: minor commercial; price category: medium; price reliability: very questionable: based on ex-vessel price for species in this family
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Wikipedia

Sand devil

The sand devil or Atlantic angel shark (Squatina dumeril) is a species of angel shark, family Squatinidae, native to the northwestern Atlantic Ocean. It occurs off the eastern United States, in the northern Gulf of Mexico, and possibly in parts of the Caribbean Sea. This bottom-dwelling shark is found in shallow inshore waters in summer and fall, and deep offshore waters in winter and spring. The sand devil's flattened body and enlarged pectoral and pelvic fins give it a ray-like appearance. There is a band of enlarged thorns running along the middle of its back. It is gray or brown in color, with scattered small dark spots. This species reaches 1.2–1.5 m (3.9–4.9 ft) in length.

The diet of the sand devil consists mainly of small teleost fishes and squid, which are captured via ambush attack. This species is viviparous, with the unborn young nourished by their yolk sacs. Females have a multi-year reproductive cycle and give birth to 4–25 pups in spring or early summer, following a 12-month gestation period. The sand devil is not aggressive, but may attempt to bite if harassed or captured. Though not valued economically, it is caught incidentally by commercial bottom trawl fisheries. The International Union for Conservation of Nature (IUCN) presently lacks sufficient information to assess the conservation status of this species.

Taxonomy and phylogeny[edit]

French naturalist Charles Alexandre Lesueur described the sand devil in an 1818 volume of Journal of the Academy of Natural Sciences of Philadelphia. Previously, it had been regarded as the same species as the European Squatina squatina. Lesueur based his account on a 1.2 m (3.9 ft) long adult male caught off the eastern United States, and named the species in honor of Auguste Duméril.[2]

Phylogenetic analysis, based on mitochondrial DNA, has concluded that the sand devil and the Pacific angel shark (S. californica) are sister species. The two of them in turn form a clade with other angel sharks found in the Americas. Molecular clock estimation has timed the divergence of S. dumeril and S. californica at c. 6.1 Ma, around the time that the Isthmus of Panama formed. It is likely that the rise of the Isthmus split the ancestral angel shark population, leading to their becoming separate species.[3]

Description[edit]

The sand devil is a ray-like shark with a color pattern of many small dark spots on a grayish or brownish background.

The sand devil has a flattened, moderately narrow body with greatly enlarged pectoral and pelvic fins. The skin folds along the sides of the head have smooth margins, without lobes. The large eyes are placed on top of the head and have prominent spiracles behind. The nostrils bear thin, pointed barbels with smooth or slightly fringed margins. The wide mouth is positioned terminally on the head. The jaws contain 10 upper and 9 lower tooth rows on each side, with toothless gaps at the middle. Each tooth has a broad base and a single pointed cusp with smooth edges. There are five pairs of gill slits located on the sides of the head.[4][5]

The pectoral fins are broad and angular with narrow rear tips; the front of the pectoral fin is separate from the head, forming a triangular lobe. The two dorsal fins are similar in size and shape, and are positioned far back on the body. The anal fin is absent. The lower lobe of the caudal fin is larger than the upper. The dermal denticles have rounded bases and three horizontal ridges. There is a distinctive band of larger thorns running along the midline of the back, from the nape to the caudal peduncle. The dorsal coloration varies from greenish or bluish gray to reddish brown, with a scattering of small, darker spots and sometimes irregular splotches; the underside is uniformly pale. This species reaches 1.3–1.5 m (4.3–4.9 ft) in length and at least 16 kg (35 lb) in weight.[4][5]

Distribution and habitat[edit]

The sand devil is found in the northwestern Atlantic Ocean from Massachusetts to the Florida Keys, where it is fairly common. It also occurs in the northern Gulf of Mexico, and there are additional unconfirmed records from Cuba, Jamaica, Nicaragua, and Venezuela. The southern extent of its range is uncertain due to confusion with the Gulf angel shark (S. heteroptera) and Mexican angel shark (S. mexicana). This bottom-dwelling shark is found over sandy or muddy bottoms on the continental shelf and slope.[4]

Off the eastern United States, the sand devil has been documented to migrate seasonally. In the summer, it moves towards the shore to depths of less than 35 m (115 ft), and many can be found in water only meters deep. During fall it can be found inshore, down to a depth of 90 m (300 ft). In winter and spring, it is found around the outer continental shelf at depths greater than 90 m (300 ft); individuals have been recorded as far as 140 km (87 mi) from land and 1,290 m (4,230 ft) deep.[4]

Biology and ecology[edit]

The Atlantic croaker is an important food source for the sand devil.

The sand devil is an ambush predator that spends much time buried in the bottom sediment. Its diet consists mainly of demersal teleost fishes such as croakers, goatfishes, and butterfishes. However, some types of demersal fishes such as jacks are rarely eaten, perhaps because they are more active and thus likely to escape attacks. Squid are a major secondary food source, particularly for smaller sharks. Crabs, shrimp, mantis shrimp, skates, and bivalves are infrequently eaten.[4][6] This species feeds both during the day and at night. It tends to select prey approximately 50–60% as long as its mouth is wide; this size is consistent with what is predicted from optimal foraging theory to yield the most efficient rate of energy return. The variety of prey taken is greatest in fall and least in winter, and smaller sharks have a more varied diet than larger ones.[6] In the northern Gulf of Mexico, the most important prey species are Atlantic croaker (Micropogonias undulatus), longspine porgy (Stenotomus caprinus), spot croaker (Leiostomus xanthurus), Gulf butterfish (Peprilus burti), red goatfish (Mullus auratus), dwarf goatfish (Upeneus parvus), and longfin inshore squid (Doryteuthis pealeii). The relative importance of each differs across seasons (for example, squid are more important in winter), perhaps due to seasonal variation in their availability.[6][7] A known parasite of the sand devil is the copepod Eudactylina spinula.[8]

Sand devil embryo with external yolk sac.

Like other angel sharks, the sand devil is viviparous with the developing embryos sustained by yolk. Mature females have one functional ovary, on the left, and two functional uteruses. Mating occurs in the spring; adult males have spines on the outer margins of their pectoral fins, which may help in gripping the female during copulation.[9] Females reproduce no more than once every two years, perhaps longer. Litter size varies from four to 25 pups, and does not appear related to the size of the female. The gestation period lasts roughly 12 months, and birthing occurs between February and June at depths of 20–30 m (66–98 ft).[4][9] The newborns measure 25–30 cm (9.8–11.8 in) in length. Males and females sexually mature at around 93 and 86 cm (37 and 34 in) long respectively; the fact that females mature at a smaller size than males is unusual among sharks.[9]

Human interactions[edit]

Though not normally aggressive towards humans, the sand devil can inflict serious wounds if provoked. Its common name refers to its habit of snapping vigorously at fishery workers when caught, and even out of the water it is capable of lunging upwards to bite.[5][4] This shark is caught incidentally in bottom trawls operated by commercial fisheries targeting other species. It is edible but seldom brought to market.[4] The International Union for Conservation of Nature (IUCN) presently lacks the data to list this species beyond Data Deficient.[1]

References[edit]

  1. ^ a b c Heupel, M.R.; Carlson, J.K. (2006). "Squatina dumeril". IUCN Red List of Threatened Species. Version 2012.2. International Union for Conservation of Nature. 
  2. ^ Lesueur, C.A. (1818). "Description of several new species of North American fishes". Journal of the Academy of Natural Sciences of Philadelphia 1 (2): 222–235, 359–368. 
  3. ^ Stelbrink, B.; von Rintelen, T.; Cliff, G.; Kriwet, J. (2010). "Molecular systematics and global phylogeography of angel sharks (genus Squatina)". Molecular Phylogenetics and Evolution 54 (2): 395–404. doi:10.1016/j.ympev.2009.07.029. PMID 19647086. 
  4. ^ a b c d e f g h Castro, J.H. (2011). The Sharks of North America. Oxford University Press. pp. 167–169. ISBN 978-0-19-539294-4. 
  5. ^ a b c Compagno, L.J.V. (1984). Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date. Food and Agricultural Organization of the United Nations. pp. 145–146. ISBN 92-5-101384-5. 
  6. ^ a b c Baremore, I.E.; Murie, D.J.; Carlson, J.K. (2010). "Seasonal and size-related differences in diet of the Atlantic angel shark Squatina dumeril in the northeastern Gulf of Mexico". Aquatic Biology 8 (2): 125–136. doi:10.3354/ab00214. 
  7. ^ Baremore, I.E.; Murie, D.J.; Carlson, J.K. (2008). "Prey selection by the Atlantic angel shark Squatina dumeril in the northeastern Gulf of Mexico". Bulletin of Marine Science 82 (3): 297–313. 
  8. ^ Pearse, A.S. (1950). "A new species of parasitic copepod from the angel shark". The Journal of Parasitology 36 (6): 515–516. doi:10.2307/3273282. 
  9. ^ a b c Baremore, I. E. (2010). "Reproductive aspects of the Atlantic angel shark Squatina dumeril". Journal of Fish Biology 76: 1682–1695. doi:10.1111/j.1095-8649.2010.02608.x. 
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