imported>OAbot: Open access bot: doi updated in citation with #oabot.

2025-06-19T01:04:23Z

Open access bot: doi updated in citation with #oabot.

New page

{{Short description|Species of shark}}
{{Good article}}
{{Speciesbox
| image = Squatina dumeril nefsc.jpg
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 18 November 2021">{{cite iucn |author=Baremore, I. |date=2019 |title=''Squatina dumeril'' |volume=2019 |page=e.T197087A22515424 |doi=10.2305/IUCN.UK.2019-1.RLTS.T197087A22515424.en |access-date=18 November 2021}}</ref>
| genus = Squatina
| species = dumeril
| authority = [[Charles Alexandre Lesueur|Lesueur]], 1818
| range_map = Squatina dumeril distmap.png
| range_map_caption = Range of the sand devil<ref name="iucn status 18 November 2021" />
}}

The '''sand devil''' or '''Atlantic angel shark''' (''Squatina dumeril'') is a [[species]] of [[angelshark]], [[family (biology)|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 [[benthic fish|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 fin|pectoral]] and [[pelvic fin]]s give it a [[Batoidea|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 {{convert|1.2|-|1.5|m|ft|abbr=on}} in length.

The diet of the sand devil consists mainly of small [[teleost]] fishes and [[squid]], which are captured via [[ambush predator|ambush attack]]. This species is [[viviparous]], with the unborn young nourished by their [[yolk sac]]s. 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 [[bycatch|caught incidentally]] by [[commercial fishing|commercial]] [[bottom trawl]] [[fisheries]].

==Taxonomy and phylogeny==
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 an adult male {{convert|1.2|m|ft|abbr=on}} long caught off the eastern United States, and named the species in honor of [[André Marie Constant Duméril]].<ref name="lesueur"/>

[[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 [[speciation|divergence]] of ''S. dumeril'' and ''S. californica'' at ''c.'' 6.1 [[Ma (unit)|Ma]], around the time that the [[Isthmus of Panama]] formed. The rise of the Isthmus likely split the ancestral angel shark population, leading to their becoming separate species.<ref name="stelbrink et al"/>

==Description==
The sand devil has a flattened, moderately narrow body with greatly enlarged [[pectoral fin|pectoral]] and [[pelvic fin]]s. 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 [[Spiracle (vertebrates)|spiracle]]s behind. The nostrils bear thin, pointed [[barbel (anatomy)|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 slit]]s located on the sides of the head.<ref name="castro"/><ref name="compagno"/>

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 fin]]s 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 denticle]]s 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 {{convert|1.3|-|1.5|m|ft|abbr=on}} in length and at least {{convert|16|kg|lb|abbr=on}} in weight.<ref name="castro"/><ref name="compagno"/>

<gallery mode="packed">
Squatina dumeril SI.jpg|The sand devil is a ray-like shark with a color pattern of many small dark spots on a gray-brown background.
Squatina dumeril SI2.jpg|Underside
Squatina dumeril jaws.jpg|Jaws
Squatina dumeril jaws2.jpg|Jaws
Squatina dumeril central teeth2.jpg|Central teeth
Squatina dumeril lower teeth.jpg|Lower teeth
</gallery>

==Distribution and habitat==
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 [[disparate angelshark]] (''S. heteroptera'') and [[Mexican angelshark]] (''S. mexicana''). Because they inhabit deep-waters and because of their low commercial value the effort put into data collection is low, as with many species under these circumstances. Although their appearances are fragmented they still are classified under large broader spaces due to the lack of research done. As a result, ranges of deep-water shark species are often considered continuous across broad expanses despite records of occurrence, in many cases, being spatially fragmented.<ref>{{Cite journal|last=Driggers|first=Williams B..|date=2018|title=Distribution of angel sharks (Squatinidae) in United States waters of the western North Atlantic Ocean|url=https://spo.nmfs.noaa.gov/sites/default/files/pdf-content/fish-bull/driggers_0.pdf|journal=Fishery Bulletin|volume=116|issue=3–4 |pages=337–347|doi=10.7755/FB.116.3-4.11 |doi-broken-date=1 November 2024 |s2cid=54807345 }}</ref> This [[benthic fish|bottom-dwelling]] shark is found over sandy or muddy bottoms on the [[continental shelf]] and [[continental slope|slope]].<ref name="castro"/>

Off the eastern [[United States]], the sand devil has been documented to [[fish migration|migrate]] seasonally. In the summer, it moves towards the shore to depths of less than {{convert|35|m|ft|abbr=on}}, and many can be found in water only meters deep. During fall it can be found [[inshore]], down to a depth of {{convert|90|m|ft|abbr=on}}. In winter and spring, it is found around the outer continental shelf at depths greater than {{convert|90|m|ft|abbr=on}}; individuals have been recorded as far as {{convert|140|km|mi|abbr=on}} from land and {{convert|1290|m|ft|abbr=on}} deep.<ref name="castro"/>

==Biology and ecology==
[[Image:Atlantic Croaker.jpg|thumb|left|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 fish|demersal]] [[teleost]] fishes such as [[Sciaenidae|croakers]], [[goatfish]]es, and [[Stromateidae|butterfish]]es. However, some types of demersal fishes such as [[Carangidae|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. [[Crab]]s, [[shrimp]], [[mantis shrimp]], [[Rajidae|skates]], and [[bivalve]]s are infrequently eaten.<ref name="castro"/><ref name="baremore et al 2010"/> 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.<ref name="baremore et al 2010"/> 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.<ref name="baremore et al 2010"/><ref name="baremore et al 2008"/> A known [[parasite]] of the sand devil is the [[copepod]] ''[[Eudactylina spinula]]''.<ref name="pearse"/>

[[Image:Sand devil embryo.jpg|thumb|150px|Sand devil embryo with external yolk sac.]]
Like other angel sharks, the sand devil is [[viviparous]] with the developing [[embryo]]s sustained by [[yolk]]. Mature females have one functional [[ovary]], on the left, and two functional [[uterus]]es. [[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 (zoology)|copulation]].<ref name="baremore"/> 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 {{convert|20|-|30|m|ft|abbr=on}}.<ref name="castro"/><ref name="baremore"/> The newborns measure {{convert|25|-|30|cm|in|abbr=on}} in length. Males and females [[sexually mature]] at around {{convert|93|and|86|cm|in|abbr=on}} long respectively; the fact that females mature at a smaller size than males is unusual among sharks.<ref name="baremore"/>

==Human interactions==
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.<ref name="castro"/><ref name="compagno"/> This shark is [[bycatch|caught incidentally]] in [[bottom trawl]]s operated by [[commercial fishing|commercial fisheries]] targeting other species. It is edible but seldom brought to market.<ref name="castro"/> The [[International Union for Conservation of Nature]] (IUCN) presently rates this species as [[least concern]].<ref name="iucn status 18 November 2021" />

==References==
{{Reflist|colwidth=30em|refs=

<ref name="baremore">{{cite journal |author=Baremore, I. E. |year=2010 |title=Reproductive aspects of the Atlantic angel shark ''Squatina dumeril'' |journal=Journal of Fish Biology |volume=76 |issue=7 |pages=1682–1695 |doi=10.1111/j.1095-8649.2010.02608.x|pmid=20557624 |bibcode=2010JFBio..76.1682B }}</ref>

<ref name="baremore et al 2010">{{cite journal |title=Seasonal and size-related differences in diet of the Atlantic angel shark ''Squatina dumeril'' in the northeastern Gulf of Mexico |author1=Baremore, I.E. |author2=Murie, D.J. |author3=Carlson, J.K. |journal=Aquatic Biology |volume=8 |issue=2 |pages=125–136 |doi=10.3354/ab00214 |year=2010|doi-access=free }}</ref>

<ref name="baremore et al 2008">{{cite journal |title=Prey selection by the Atlantic angel shark Squatina dumeril in the northeastern Gulf of Mexico |author1=Baremore, I.E. |author2=Murie, D.J. |author3=Carlson, J.K. |journal=Bulletin of Marine Science |volume=82 |issue=3 |pages=297–313 |year=2008}}</ref>

<ref name="castro">{{cite book |author=Castro, J.H. |title=The Sharks of North America |publisher=Oxford University Press |year=2011 |isbn=978-0-19-539294-4 |pages=167–169}}</ref>

<ref name="compagno">{{cite book |author=Compagno, L.J.V. |year=1984 |title=Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date |publisher=Food and Agricultural Organization of the United Nations |isbn=92-5-101384-5 |pages=145–146}}</ref>

<ref name="lesueur">{{cite journal |author=Lesueur, C.A. |year=1818 |title=Description of several new species of North American fishes |journal=Journal of the Academy of Natural Sciences of Philadelphia |volume=1 |issue=2 |pages=222–235, 359–368}}</ref>

<ref name="pearse">{{cite journal |title=A new species of parasitic copepod from the angel shark |author=Pearse, A.S. |journal=The Journal of Parasitology |volume=36 |issue=6 |year=1950 |pages=515–516 |doi=10.2307/3273282|jstor=3273282 |doi-access=free }}</ref>

<ref name="stelbrink et al">{{cite journal |doi=10.1016/j.ympev.2009.07.029 |author1=Stelbrink, B. |author2=von Rintelen, T. |author3=Cliff, G. |author4=Kriwet, J. |title=Molecular systematics and global phylogeography of angel sharks (genus ''Squatina'') |journal=Molecular Phylogenetics and Evolution |volume=54 |issue=2 |year=2010 |pages=395–404 |pmid=19647086|bibcode=2010MolPE..54..395S }}</ref>

}}

==External links==
{{Commons category|Squatina dumeril}}
*[http://www.fishbase.us/summary/Squatina-dumeril.html ''Squatina dumeril'', Sand devil] at [http://www.fishbase.org/search.php FishBase]
*[http://www.iucnredlist.org/details/60248/0 ''Squatina dumeril'' (Atlantic Angel Shark, Sand Devil)] at [http://www.iucnredlist.org/ IUCN Red List]
*[http://www.flmnh.ufl.edu/fish/Gallery/Descript/AtlanticAngel/AtlanticAngel.html Biological Profiles: Atlantic angel shark] at [http://www.flmnh.ufl.edu/fish/ Florida Museum of Natural History Ichthyology Department]

{{Squatiniformes}}

{{Taxonbar|from=Q756873}}

[[Category:Squatinidae|sand devil]]
[[Category:Fauna of the Southeastern United States]]
[[Category:Fish of the Caribbean]]
[[Category:Fish of the Dominican Republic]]
[[Category:Fish described in 1818|sand devil]]

Sand devil - Revision history

imported>OAbot: Open access bot: doi updated in citation with #oabot.