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{{short description|Suborder of fishes}}
{{otheruses}}
{{About|the fish|other uses|Stingray (disambiguation)}}
{{Cleanup|date=October 2007}}
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{{Taxobox
{{Automatic taxobox
| color = pink
| name =Stingrays
| name = Stingrays
| fossil_range = {{fossil range|Hauterivian|recent|[[Early Cretaceous]] to recent<ref>{{cite journal |last1=Marmi |first1=Josep |last2=Vila # |first2=Bernat |last3=Oms |first3=Oriol |last4=Galobart |first4=Àngel |last5=Cappetta |first5=Henri |title=Oldest records of stingray spines (Chondrichthyes, Myliobatiformes) |journal=Journal of Vertebrate Paleontology |date=18 May 2010 |volume=30 |issue=3 |pages=970–974 |doi=10.1080/02724631003758011 |bibcode=2010JVPal..30..970M }}</ref>}}
| image = Stingray2.jpg
| image = SStringray.jpg
| image_width = 250px
| image_caption = [[Bluespotted stingray]], ''Taeniura lymma''
| image_caption = [[Southern stingray]] (''Hypanus americanus'')
| regnum = [[Animal]]ia
| taxon = Myliobatoidei
| authority = [[Leonard Joseph Victor Compagno|Compagno]], 1973
| phylum = [[Chordata]]
| subdivision_ranks = Families
| classis = [[Chondrichthyes]]
| subdivision = * '''Hexatrygonoidea'''
| subclassis = [[Elasmobranchii]]
**[[Hexatrygonidae]]
| ordo = [[Rajiformes]]
| familia = '''Dasyatidae'''
* '''Urolophoidea'''
**[[Plesiobatidae]]
| subdivision_ranks = Genera
**[[Urolophidae]]
| subdivision =
* '''Dasyatoidea'''
''[[Dasyatis]]''<br/>
**[[Urotrygonidae]]
''[[Himantura]]''<br/>Urogymnus]]''<br/>
**[[Dasyatidae]]
See text for species.
**[[Potamotrygonidae]]
**[[Gymnuridae]]
**{{extinct}}[[Dasyomyliobatidae]]
**[[Myliobatidae]]
**{{extinct}}[[Rhombodontidae]]
}}
}}
Dasyatids are common in tropical coastal waters throughout the world, and there are [[fresh water]] species in [[Asia]] (their large [[pectoral wings]] (commonly mistaken as "fins"). Their [[stinger]] is a razor-sharp, barbed, or serrated cartilaginous spine which grows from the ray's whip-like tail (like a fingernail), and can grow as long as 37 cm (about 14.6 inches). On the underside of the spine are two grooves containing [[venom (poison)|venom]]-secreting glandular tissue. The entire spine is covered with a thin layer of skin called the integumentary sheath, in which venom is concentrated.<ref>{{cite journal | author = Meyer P | title = Stingray injuries | journal = Wilderness Environ Med | volume = 8 | issue = 1 | pages = 24-8 | year = 1997 | id = PMID 11990133}}</ref> This gives them their common name of '''stingrays''' (a [[Portmanteau]] of "Stinger" and "Ray"), but the name can also be used to refer to any poisonous ray.


'''Stingrays''' are a group of sea [[Batoidea|rays]], a type of [[cartilaginous fish]]. They are classified in the suborder '''Myliobatoidei''' of the order [[Myliobatiformes]] and consist of eight families: [[Hexatrygonidae]] (sixgill stingray), [[Plesiobatidae]] (deepwater stingray), [[Urolophidae]] (stingarees), [[Urotrygonidae]] (round rays), [[Dasyatidae]] (whiptail stingrays), [[Potamotrygonidae]] (river stingrays), [[Gymnuridae]] (butterfly rays) and [[Myliobatidae]] (eagle rays).<ref name="nelson">{{cite book |title=Fishes of the World | vauthors = Nelson JS |edition=fourth |publisher=John Wiley |year=2006 |isbn=978-0-471-25031-9 |pages=76–82}}</ref><ref>{{cite book | vauthors = Helfman GS, Collette BB, Facey DE |title=The Diversity of Fishes |publisher=Blackwell Science |year=1997 |isbn=978-0-86542-256-8 |page=180}}</ref>
Some adult rays may be no larger than a human palm, while other species, like the [[short-tail stingray]], may have a body of six feet in diameter, and an overall length, including their tail, of fourteen feet.
There are about 220 known stingray species organized into 29 genera.


Stingrays are common in coastal [[tropical]] and [[subtropical]] marine waters throughout the world. Some species, such as the [[thorntail stingray]] (''Dasyatis thetidis''), are found in warmer [[temperate]] oceans and others, such as the [[deepwater stingray]] (''Plesiobatis daviesi''), are found in the [[deep ocean]]. The [[Potamotrygonidae|river stingrays]] and a number of [[whiptail stingray]]s (such as the [[Niger stingray]] (''Fontitrygon garouaensis'')) are restricted to [[fresh water]]. Most myliobatoids are [[demersal]] (inhabiting the next-to-lowest zone in the [[water column]]), but some, such as the [[pelagic stingray]] and the [[eagle ray]]s, are [[pelagic]].<ref>{{cite web | url = http://www.flmnh.ufl.edu/fish/gallery/descript/pelagicstingray/pelagicstingray.html | title = Pelagic Stingray | vauthors = Bester C, Mollett HF, Bourdon J | publisher = [[Florida Museum of Natural History]], Ichthyology department | date = 2017-05-09 | access-date = 2009-09-29 | archive-date = 2016-01-15 | archive-url = https://web.archive.org/web/20160115085500/http://www.flmnh.ufl.edu/fish/Gallery/Descript/PelagicStingray/PelagicStingray.html | url-status = live }}</ref>
Stingrays may also be called the '''whip-tailed rays''' though this usage is much less common.


Stingray species are [[List of threatened rays|progressively becoming threatened]] or vulnerable to [[extinction]], particularly as the consequence of [[unregulated fishing]].<ref>[http://www.cites.org/common/com/ac/25/E25i-06.pdf The Future of Sharks: A Review of Action and Inaction] {{Webarchive|url=https://web.archive.org/web/20130512102129/http://www.cites.org/common/com/ac/25/E25i-06.pdf |date=2013-05-12 }} CITES AC25 Inf. 6, 2011.</ref> As of 2013, 45 species have been listed as [[Vulnerable species|vulnerable]] or [[endangered species|endangered]] by the [[International Union for Conservation of Nature|IUCN]]. The status of some other species is poorly known, leading to their being listed as [[data deficient]].{{fact|date=November 2024}}
A group or collection of stingrays is commonly referred to as a "fever" of stingrays.


==Feeding habits==
== Evolution ==
[[File:Heliobatis radians Green River Formation (cropped).jpg|thumb|[[Early Eocene]] fossil stingray ''[[Heliobatis radians]]''|266x266px]]
[[Image:Freshwater ray.jpg|thumb|left|Underside of [[freshwater ray]] showing mouth]]
Stingrays diverged from their closest relatives, the [[Panray|panrays]], during the [[Late Jurassic]] period, and diversified over the course of the [[Cretaceous]] into the different extant families today. The earliest stingrays appear to have been benthic, with the ancestors of the eagle rays becoming pelagic during the early [[Late Cretaceous]].<ref name=":0">{{cite journal |last1=Marramà |first1=Giuseppe |last2=Carnevale |first2=Giorgio |last3=Giusberti |first3=Luca |last4=Naylor |first4=Gavin J. P. |last5=Kriwet |first5=Jürgen |title=A bizarre Eocene dasyatoid batomorph (Elasmobranchii, Myliobatiformes) from the Bolca Lagerstätte (Italy) reveals a new, extinct body plan for stingrays |journal=Scientific Reports |date=1 October 2019 |volume=9 |issue=1 |page=14087 |doi=10.1038/s41598-019-50544-y |pmid=31575915 |pmc=6773687 |bibcode=2019NatSR...914087M }}</ref><ref name="wiley" />
Stingrays are flat so as to hide on the depths of the sea. They ruffle up the sand and hide beneath it. Since their eyes are on top of their body and their mouths on the bottom, stingrays cannot see their prey. Instead, they use the sense of smell and electro-receptors, similar to those of the [[shark]]. They feed primarily on [[mollusk]]s and [[crustacean]]s and occasionally on small fish. Their mouths contain powerful, shell-crushing teeth. Rays settle on the bottom while feeding, sometimes leaving only their eyes and tail visible.


===Fossils===
==Stinging mechanism==
[[File:Lessiniabatis fossil.png|left|thumb|The bizarre ''[[Lessiniabatis]]'' of Early Eocene Italy]]
[[Image:Himantura granulata 2.jpg|thumb|right|250px|A stingray in the [[Maldives]].]]
[[Permineralization|Permineralized]] stingray teeth have been found in [[sedimentary]] deposits around the world as far back as the [[Early Cretaceous]]. The oldest known stingray taxon is "''Dasyatis''" ''speetonensis'' from the [[Hauterivian]] of [[England]], whose teeth most closely resemble that of the extant [[sixgill stingray]] (''Hexatrygon''). Although stingray [[teeth]] are rare on [[seafloor|sea bottoms]] compared to the similar [[shark teeth]], [[scuba diver]]s searching for the latter do encounter the teeth of stingrays.<ref name=":0" /><ref>{{Cite web |title=Heliobatis radians Stingray Fossil from Green River |url=http://www.fossilmall.com/Science/Sites/GreenRiver/WF14/heliobatis.htm |access-date=2023-02-14 |website=www.fossilmall.com}}</ref>
[[Image:Stringray sting.jpg|thumb|right|250px|A stingray's stinger]]
[[Image:DSC01207.JPG|thumb|right|250px|3 yellow-spotted stingrays at the [[Dallas World Aquarium]].]]
[[Image:Sting ray - melbounre aquarium.jpg|thumb|right|250px|Stingrays form a large part of the underwater display at the [[Melbourne Aquarium]].]]
Dasyatids generally do not attack aggressively or even actively defend themselves. When threatened, their primary reaction is to swim away. However, when they are attacked by predators or stepped on, the barbed stinger in their [[tail]] is whipped up. This attack is normally ineffective against their main [[predator]], [[shark]]s.<ref>{{cite web | url = http://www.caribbeanmag.com/search/articles/Cayman_Islands/Stingray_city___diving___snorkeling_grand_cayman.html | title = Stingray City - About Stingrays | publisher = Caribbean Magazine}}</ref> Humans are usually stung in the [[foot]] region (depending on the size of the stingray); it is also possible, although less likely, to be stung by brushing against the stinger. The stinger often breaks off in the wound, which is non-fatal to the stingray, and will be regrown. Contact with the stinger causes local [[Physical trauma|trauma]] (from the cut itself), pain and swelling from the venom, and possible later infection from bacteria on parts of the stinger left in the wound. Immediate injuries to humans include, but are not limited to: [[venom|poisoning]], punctures, severed [[artery|arteries]] and possibly death. Fatal stings are extremely rare.


Full-body stingray fossils are very rare but are known from certain [[lagerstätte]] that preserve soft-bodied animals. The extinct ''[[Cyclobatis]]'' of the [[Cretaceous]] of [[Lebanon]] is thought to be a [[Skate (fish)|skate]] that had [[Convergent evolution|convergently evolved]] a highly stingray-like body plan, although its exact taxonomic placement is still uncertain.<ref>{{cite journal |last1=Marramà |first1=Giuseppe |last2=Schultz |first2=Ortwin |last3=Kriwet |first3=Jürgen |title=A new Miocene skate from the Central Paratethys (Upper Austria): the first unambiguous skeletal record for the Rajiformes (Chondrichthyes: Batomorphii) |journal=Journal of Systematic Palaeontology |date=3 June 2019 |volume=17 |issue=11 |pages=937–960 |doi=10.1080/14772019.2018.1486336 |pmc=6510527 |pmid=31156351 |bibcode=2019JSPal..17..937M }}</ref> True stingray fossils become more common in the Eocene, with the extinct freshwater stingrays ''[[Heliobatis]]'' and ''[[Asterotrygon]]'' known from the [[Green River Formation]].<ref>{{cite journal |last1=De Carvalho |first1=Marcelo R. |last2=Maisey |first2=John G. |last3=Grande |first3=Lance |title=Freshwater Stingrays of the Green River Formation of Wyoming (Early Eocene), with the Description of a New Genus and Species and an Analysis of ITS Phylogenetic Relationships (Chondrichthyes: Myliobatiformes) |journal=Bulletin of the American Museum of Natural History |date=June 2004 |volume=284 |pages=1–136 |doi=10.1206/0003-0090(2004)284<0001:FSOTGR>2.0.CO;2 }}</ref> A diversity of stingray fossils is known from the Eocene [[Monte Bolca]] formation from [[Italy]], including the early [[Urolophidae|stingaree]] ''[[Arechia]]'', as well as ''[[Dasyomyliobatis]]'', which is thought to represent a [[Transitional fossil|transitional form]] between stingrays and [[Eagle ray|eagle rays]], and the highly unusual ''[[Lessiniabatis]]'', which had an extremely short and slender tail with no sting.<ref name=":0" /><ref name="wiley">{{Cite journal |last1=Marramà |first1=G. |last2=Villalobos-Segura |first2=E. |last3=Zorzin |first3=R. |last4=Kriwet |first4=J. |last5=Carnevale |first5=G. |year=2023 |title=The evolutionary origin of the durophagous pelagic stingray ecomorph |journal=Palaeontology |volume=66 |issue=4 |at=e12669 |bibcode=2023Palgy..6612669M |doi=10.1111/pala.12669 |pmc=7614867 |pmid=37533696}}</ref>


==Anatomy==
Treatment for stings includes application of near-scalding water, which helps ease pain by denaturing the complex venom protein, and [[antibiotic]]s. Immediate injection of [[local anesthetic]] in and around the wound is very helpful, as is the use of adjunct opiates such as intramuscular pethidine. Local anesthetic brings almost instant relief for several hours. Other possible pain remedies include [[papain]] ([[papaya]] extract, contained in unseasoned powdered meat tenderizer), which may break down the protein of the toxins, although this may be more appropriate for [[jellyfish]] and similar stings. One odd but usual method of pain relief is to urinate on the wound. The actual urine and sometimes vinegar are not benefitial themselves, but the warmth of the urine provides relief. Pain normally lasts up to 48 hours, but is most severe in the first 30&ndash;60 minutes and may be accompanied by [[nausea]], [[Fatigue (physical)|fatigue]], [[headache]]s, [[fever]] and chills. All stingray injuries should be medically assessed; the wound needs to be thoroughly cleaned, and [[surgical]] exploration is often required to remove any barb fragments remaining in the wound. Following cleaning, a [[radiograph]] is helpful to confirm removal of all the fragments.<ref>{{cite journal | author = Flint D, Sugrue W | title = Stingray injuries: a lesson in debridement | journal = N Z Med J | volume = 112 | issue = 1086 | pages = 137-8 | year = 1999 | id = PMID 10340692 }}</ref> However, not all remnants are radio-opaque; [[Medical ultrasonography|ultrasound]] imaging is useful in difficult cases.
[[File:Dasyatis say njsm (annotated).jpg|thumb|400px|{{center|dorsal (topside) ← &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; &nbsp; → ventral (underside)<br />External anatomy of a male [[bluntnose stingray]] (''Hypanus say'')}}]]
{{multiple image
| align = right
| caption_align = center
| direction = horizontal
| width1 = 170
| image1 = Stingray teeth and jaws.jpg
| alt1 =
| caption1 = Stingray jaw and teeth.<br />The teeth are modified [[placoid scale]]s.
| width2 = 226
| image2 = Taeniura lymma by Marek Jakubowski.jpg
| alt2 =
| caption2 = Like in other rays, the [[bluespotted ribbontail ray]] (''Taeniura lymma'') breathes though [[Spiracle (vertebrates)|spiracle]]s just behind the eyes when it hunts in [[seafloor sediment]].
}}


=== Jaw and teeth ===
== Reproduction ==
The mouth of the stingray is located on the [[Anatomical term of location#Dorsal and ventral|ventral]] side of the vertebrate. Stingrays exhibit hyostylic jaw suspension, which means that the mandibular arch is only suspended by an articulation with the [[hyomandibula]]. This type of suspensions allows for the upper jaw to have high mobility and protrude outward.<ref>{{cite book|url=https://books.google.com/books?id=7qXMBQAAQBAJ&q=biology+of+sharks+and+their+relatives+2nd+edition+jaws&pg=PA176|title=Biology of Sharks and Their Relatives|edition=Second|last1=Carrier|first1=Jeffrey C.|last2=Musick|first2=John A.|last3=Heithaus|first3=Michael R.|name-list-style=vanc|date=2012-04-09|publisher=CRC Press|isbn=9781439839263|access-date=2020-11-21|archive-date=2022-01-10|archive-url=https://web.archive.org/web/20220110002705/https://books.google.com/books?id=7qXMBQAAQBAJ&q=biology+of+sharks+and+their+relatives+2nd+edition+jaws&pg=PA176|url-status=live}}</ref> The teeth are modified [[placoid scale]]s that are regularly shed and replaced.<ref>{{Cite book|url=https://books.google.com/books?id=E0Oy1i-8vfsC&q=teeth+that+are+modified+placoid+scales&pg=PA26|title=Biology Of Fishes|last=Khanna|first=D. R.|date=2004|publisher=Discovery Publishing House|isbn=9788171419081|access-date=2020-11-21|archive-date=2022-01-10|archive-url=https://web.archive.org/web/20220110002707/https://books.google.com/books?id=E0Oy1i-8vfsC&q=teeth+that+are+modified+placoid+scales&pg=PA26|url-status=live}}</ref> In general, the teeth have a root implanted within the connective tissue and a visible portion of the tooth, is large and flat, allowing them to crush the bodies of hard shelled prey.<ref name="Morphology does not predict">{{cite journal |last1=Kolmann |first1=M. A. |last2=Crofts |first2=S. B. |last3=Dean |first3=M. N. |last4=Summers |first4=A. P. |last5=Lovejoy |first5=N. R. |title=Morphology does not predict performance: jaw curvature and prey crushing in durophagous stingrays |journal=Journal of Experimental Biology |date=13 November 2015 |volume=218 |issue=24 |pages=3941–3949 |doi=10.1242/jeb.127340 |pmid=26567348|doi-access=free }}</ref> Male stingrays display [[sexual dimorphism]] by developing [[Cusp (anatomy)|cusps]], or pointed ends, to some of their teeth. During mating season, some stingray species fully change their tooth morphology which then returns to baseline during non-mating seasons.<ref>{{Cite journal|last1=Kajiura|first1=null|last2=Tricas|first2=null|date=1996|title=Seasonal dynamics of dental sexual dimorphism in the Atlantic stingray Dasyatis sabina|journal=The Journal of Experimental Biology|volume=199|issue=Pt 10|pages=2297–2306|doi=10.1242/jeb.199.10.2297|pmid=9320215|doi-access=free}}</ref>
Mating season occurs in the winter. When a male is courting a female, he will follow her closely, biting at her pectoral disc. During mating, the male will go on top of the female (his belly on her back) and put one of his claspers into her vent.[http://www.elasmo-research.org/education/topics/lh_stingray_city.htm]


=== Spiracles ===
Most rays are [[viviparous]], bearing live young in "litters" of five to ten. The female holds the embryos in the womb without a placenta. Instead, the embryos absorb nutrients from a yolk sac, and after the sac is depleted the mother provides uterine milk. [http://www.flmnh.ufl.edu/fish/Gallery/Descript/AtlanticStingray/AtlanticStingray.html]
[[Spiracle (vertebrates)|Spiracle]]s are small openings that allow some fish and amphibians to breathe. Stingray spiracles are openings just behind its eyes. The respiratory system of stingrays is complicated by having two separate ways to take in water to use the oxygen. Most of the time stingrays take in water using their mouth and then send the water through the gills for [[gas exchange]]. This is efficient, but the mouth cannot be used when hunting because the stingrays bury themselves in the ocean sediment and wait for prey to swim by.<ref>{{Cite web|url = http://bioweb.uwlax.edu/bio203/s2009/long_nico/Adaptation.htm|title = Stingray|website = bioweb.uwlax.edu|access-date = 2018-05-12|archive-date = 2018-07-23|archive-url = https://web.archive.org/web/20180723120346/http://bioweb.uwlax.edu/bio203/s2009/long_nico/Adaptation.htm|url-status = live}}</ref> So the stingray switches to using its spiracles. With the spiracles, they can draw water free from sediment directly into their gills for gas exchange.<ref name=":7">{{Cite book|title=Vertebrates: Comparative Anatomy, Function, Evolution|last=Kardong|first=Kenneth | name-list-style = vanc |publisher=McGraw-Hill Education|year=2015|isbn=978-0-07-802302-6|location=New York|pages=426}}</ref> These alternate ventilation organs are less efficient than the mouth, since spiracles are unable to pull the same volume of water. However, it is enough when the stingray is quietly waiting to ambush its prey.


The flattened bodies of stingrays allow them to effectively conceal themselves in their environments. Stingrays do this by agitating the sand and hiding beneath it. Because their eyes are on top of their bodies and their mouths on the undersides, stingrays cannot see their prey after capture; instead, they use smell and electroreceptors ([[ampullae of Lorenzini]]) similar to those of [[shark]]s.<ref>{{cite journal | vauthors = Bedore CN, Harris LL, Kajiura SM | title = Behavioral responses of batoid elasmobranchs to prey-simulating electric fields are correlated to peripheral sensory morphology and ecology | journal = Zoology | volume = 117 | issue = 2 | pages = 95–103 | date = April 2014 | pmid = 24290363 | doi = 10.1016/j.zool.2013.09.002 | bibcode = 2014Zool..117...95B }}</ref> Stingrays settle on the bottom while feeding, often leaving only their eyes and tails visible. [[Coral reef]]s are favorite feeding grounds and are usually shared with sharks during high tide.<ref>{{Cite web|title=Stingray City - Altering Stingray Behavior & Physiology?|url=https://www.divephotoguide.com/underwater-photography-scuba-ocean-news/stingray_city___altering_stingray_behavior___physiology/|access-date=2023-02-14|website=DivePhotoGuide
== As food ==
| first= Jason | last=Heller
Rays may be caught on a fishing line using small [[crab]]s as bait, and are often caught accidentally; they may also be speared from above. They are edible. Small rays may be cooked similarly to other fish, typically grilled or battered and fried. While not independently valuable as a food source, the stingray's capacity to damage shell fishing grounds can lead to bounties being placed on their removal. Also, they sometimes eat squid and crabs.
| date = 14 April 2009
|language=en}}</ref>


{{clear}}
Stingray recipes abound throughout the world, with dried forms of the wings being most common. For example, in [[Malaysia]] and [[Singapore]], stingray is commonly [[barbecue]]d over charcoal, then served with spicy ''[[sambal]]'' sauce. Generally, the most prized parts of the stingray are the wings, the "cheek" (the area surrounding the eyes) and the liver. The rest of the ray is considered too rubbery to have any culinary uses.


==Behavior==
In [[Iceland]], eating pickled stingray ("kæst skata") on December 23 is an old tradition.<ref>{{cite web |url=http://www.foodmuseum.com/ISyuletidelads.html |title=Iceland's Christmas Foods |accessdate=2006-12-23 |work=[http://www.foodmuseum.com/ The FOOD Museum }}</ref>
[[File:Dasyatis sabina - MC Gilbert 2019.jpg|thumb|{{center|Skeleton of an [[atlantic stingray]] (''Hypanus sabinus'')}}]]


== Viewing ==
===Reproduction===
[[File:Mobula breach 2.jpg|thumb|left|[[Mobula]] (devil rays) are thought to [[Cetacean surfacing behaviour|breach]] as a form of courtship.]]
[[Image:Stingray Ctiy, Grand Cayman.jpg|thumb|right|250px|Stingray City in Grand Cayman allows swimmers, snorkelers, and divers to swim with and feed the stingrays.]]
During the [[breeding season]], males of various stingray species such as the [[round stingray]] (''Urobatis halleri''), may rely on their [[ampullae of Lorenzini]] to sense certain electrical signals given off by mature females before potential [[copulation (zoology)|copulation]].<ref>{{cite journal |last1=Tricasa |first1=Timothy C. |last2=Michael |first2=Scott W. |last3=Sisneros |first3=Joseph A. |title=Electrosensory optimization to conspecific phasic signals for mating |journal=Neuroscience Letters |date=December 1995 |volume=202 |issue=1–2 |pages=129–132 |doi=10.1016/0304-3940(95)12230-3 |pmid=8787848 }}</ref> When a male is courting a female, he follows her closely, biting at her pectoral disc. He then places one of his two claspers into her valve.<ref>[http://www.wetwebmedia.com/fwsubwebindex/fwraybehfaq.htm FAQs on Freshwater Stingray Behavior] {{Webarchive|url=https://web.archive.org/web/20171002070831/http://www.wetwebmedia.com/fwsubwebindex/fwraybehfaq.htm |date=2017-10-02 }}. Wetwebmedia.com. Retrieved on 2012-07-17.</ref>
[[Image:stingray-buried-in-sand.jpg|thumb|right|250px|Stingrays can be seen burrowing into the [[sand]] just yards away from tourists at Stingray City.]]
[[Image:Dasyatis pastinaca(02).jpg|thumb|right|250px|A stingray's underside showing its mouth and [[gill]]s.]]
Stingrays are usually very docile creatures. The customary reaction of the stingray is to immediately flee the vicinity of a disturbance. Nevertheless, certain larger species are located in waters where they are easily excitable due to possible attacks from feeding sharks and should be approached with caution, as the stingray's defensive reflex and effort to flee may result in human contact with the stinger, resulting in serious injury or even (though rarely) death.


Reproductive ray behaviors are associated with their [[behavioral endocrinology]], for example, in species such as the [[atlantic stingray]] (''Hypanus sabinus''), social groups are formed first, then the sexes display complex [[courtship display|courtship]] behaviors that end in pair [[copulation (zoology)|copulation]] which is similar to the species ''Urobatis halleri.''<ref name = hormone>{{cite journal |last1=Tricas |first1=Timothy C. |last2=Rasmussen |first2=L. E. L. |last3=Maruska |first3=Karen P. |title=Annual Cycles of Steroid Hormone Production, Gonad Development, and Reproductive Behavior in the Atlantic Stingray |journal=General and Comparative Endocrinology |date=2000 |volume=118 |issue=2 |pages=209–25 |doi=10.1006/gcen.2000.7466 |pmid=10890563 }}</ref> Furthermore, their mating period is one of the longest recorded in elasmobranch fish. Individuals are known to mate for seven months before the females ovulate in March. During this time, the male stingrays experience increased levels of androgen hormones which has been linked to its prolonged mating periods.<ref name = hormone/> The behavior expressed among males and females during specific parts of this period involves aggressive social interactions.<ref name = hormone/> Frequently, the males trail females with their snout near the female vent then proceed to bite the female on her fins and her body.<ref name = hormone/> Although this mating behavior is similar to the species ''Urobatis halleri'', differences can be seen in the particular actions of ''Hypanus sabinus''. Seasonal elevated levels of serum androgens coincide with the expressed aggressive behavior, which led to the proposal that androgen steroids start, indorse and maintain aggressive sexual behaviors in the male rays for this species which drives the prolonged mating season. Similarly, concise elevations of serum androgens in females has been connected to increased aggression and improvement in [[mate choice]]. When their androgen steroid levels are elevated, they are able to improve their mate choice by quickly fleeing from tenacious males when undergoing ovulation succeeding impregnation. This ability affects the paternity of their offspring by refusing less qualified mates.<ref name = hormone/>
Dasyatids are not normally visible to swimmers, but divers and snorkelers may find them in shallow sandy waters, more so when the water is unseasonably warm.


Stingrays are [[ovoviviparous]], bearing live young in "litters" of five to thirteen. During this period, the female's behavior transitions to support of her future offspring. Females hold the embryos in the womb without a placenta. Instead, the embryos absorb nutrients from a [[yolk sac]] and after the sac is depleted, the mother provides uterine "milk".<ref>[http://www.flmnh.ufl.edu/fish/Gallery/Descript/AtlanticStingray/AtlanticStingray.html Florida Museum of Natural History Ichthyology Department: Atlantic Stingray] {{Webarchive|url=https://web.archive.org/web/20160104222940/https://www.flmnh.ufl.edu/fish/Gallery/Descript/AtlanticStingray/AtlanticStingray.html |date=2016-01-04 }}. Flmnh.ufl.edu. Retrieved on 2012-07-17.</ref> After birth, the offspring generally disassociate from the mother and swim away, having been born with the instinctual abilities to protect and feed themselves. In a very small number of species, like the [[giant freshwater stingray]] (''Urogymnus polylepis''), the mother "cares" for her young by having them swim with her until they are one-third of her size.<ref>{{cite web |last1=Seubert |first1=Curtis |title=How Do Stingrays Take Care of Their Young? |date=April 24, 2017 |url=https://sciencing.com/do-stingrays-care-young-8791629.html |access-date=December 14, 2018 |archive-date=December 16, 2018 |archive-url=https://web.archive.org/web/20181216031207/https://sciencing.com/do-stingrays-care-young-8791629.html |work=Sciencing|url-status=live }}</ref>
In the [[Cayman Islands]], there are several dive sites called [[Stingray City, Grand Cayman]], where divers and snorkelers can swim with large [[southern stingray]]s (''Dasyatis Americana'') and feed them by hand.


At the [[Sea Life London Aquarium]], two female stingrays delivered seven baby stingrays, although the mothers have not been near a male for two years. This suggests some species of rays can store sperm then give birth when they deem conditions to be suitable.<ref>{{cite news |url=https://www.smh.com.au/environment/conservation/stingrays-born-in-female-only-tank-20110810-1imik.html |title=Stingrays born in female only tank |date=2011-08-10 |work=[[The Sydney Morning Herald]] |access-date=2020-07-25 |archive-date=2020-07-25 |archive-url=https://web.archive.org/web/20200725012114/https://www.smh.com.au/environment/conservation/stingrays-born-in-female-only-tank-20110810-1imik.html |url-status=live }}</ref>
There is also a "Stingray City" in the sea surrounding the [[Caribbean]] island of [[Antigua]]. It consists of a large, shallow reserve where the rays live, and snorkelling is possible.


=== Locomotion ===
In [[Belize]] off the island of [[Ambergris Caye]] there is a popular marine sanctuary called [[Hol Chan]]. Here divers and snorkelers often gather to watch stingrays and [[nurse sharks]] that are drawn to the area by tour operators who feed the animals.
[[File:Stingray locomotion gif.gif|thumb|{{center|[[Atlantic stingray]] (''Hypanus sabinus'') undulation locomotion}}]]


The stingray uses its paired [[pectoral fin]]s for moving around. This is in contrast to sharks and most other fish, which get most of their swimming power from a single [[Caudal fin|caudal (tail) fin]].<ref>{{cite journal |last1=Wang |first1=Yangwei |last2=Tan |first2=Jinbo |last3=Zhao |first3=Dongbiao |title=Design and Experiment on a Biomimetic Robotic Fish Inspired by Freshwater Stingray |journal=Journal of Bionic Engineering |date=June 2015 |volume=12 |issue=2 |pages=204–216 |doi=10.1016/S1672-6529(14)60113-X }}</ref> Stingray pectoral fin [[Animal locomotion|locomotion]] can be divided into two categories, undulatory and oscillatory.<ref name="Fontanella">{{cite journal | vauthors = Fontanella J | date = 2013 | title = Two- and three-dimensional geometries of batoids in relation to locomotor mode|journal=Journal of Experimental Biology and Ecology|volume=446|pages=273–281 | doi = 10.1016/j.jembe.2013.05.016 | bibcode = 2013JEMBE.446..273F }}</ref> Stingrays that use undulatory locomotion have shorter thicker fins for slower motile movements in [[Benthic zone|benthic]] areas.<ref>{{cite journal |last1=Bottom II |first1=R. G. |last2=Borazjani |first2=I. |last3=Blevins |first3=E. L. |last4=Lauder |first4=G. V. |title=Hydrodynamics of swimming in stingrays: numerical simulations and the role of the leading-edge vortex |journal=Journal of Fluid Mechanics |date=10 February 2016 |volume=788 |pages=407–443 |doi=10.1017/jfm.2015.702 |bibcode=2016JFM...788..407B }}</ref> Longer thinner pectoral fins make for faster speeds in oscillation mobility in pelagic zones.<ref name="Fontanella" /> Visually distinguishable oscillation has less than one wave going, opposed to undulation having more than one wave at all times.<ref name="Fontanella" />
Many Tahitian island resorts regularly offer guests the chance to "feed the stingrays and sharks". This consists of taking a boat to the outer lagoon reefs then standing in waist-high water while habituated stingrays swarm around, pressing right up against you seeking food from your hand or tossed into the water. The boat owners also "call in" sharks, which when they arrive from the ocean swoop through the shallow water above the reef and snatch food offered to them.


=== Feeding behavior and diet ===
Most major aquariums feature stingrays, including the National [[Baltimore Aquarium]] and the [[Tennessee Aquarium]] in Chattanooga. Where there are stingray touch tanks where visitors can "pet" rays or when show divers routinely hand feed rays in giant saltwater exhibits, for diver and visitor safety the spines on the rays are snipped off with a pair of pliers. The tip of the spine is then presented as a harmless stub that can't penetrate the skin of visitors or divers who routinely handle the docile rays.
[[File:Bat-ray-NOAA-Tony-Chess.jpg|thumb|left|[[Bat ray]] (''Myliobatis californica'') in a feeding posture]]
Stingrays use a wide range of feeding strategies. Some have specialized jaws that allow them to crush hard mollusk shells,<ref>{{cite journal | vauthors = Kolmann MA, Huber DR, Motta PJ, Grubbs RD | title = Feeding biomechanics of the cownose ray, ''Rhinoptera bonasus'', over ontogeny | journal = Journal of Anatomy | volume = 227 | issue = 3 | pages = 341–51 | date = September 2015 | pmid = 26183820 | pmc = 4560568 | doi = 10.1111/joa.12342 }}</ref> whereas others use external mouth structures called cephalic lobes to guide plankton into their oral cavity.<ref>{{cite journal | vauthors = Dean MN, Bizzarro JJ, Summers AP | title = The evolution of cranial design, diet, and feeding mechanisms in batoid fishes | journal = Integrative and Comparative Biology | volume = 47 | issue = 1 | pages = 70–81 | date = July 2007 | pmid = 21672821 | doi = 10.1093/icb/icm034 | doi-access = free }}</ref> [[Benthic zone|Benthic]] stingrays (those that reside on the sea floor) are ambush hunters.<ref name="Curio">{{cite book |doi=10.1007/978-3-642-81028-2 |title=The Ethology of Predation |date=1976 |last1=Curio |first1=Eberhard |isbn=978-3-642-81030-5 }}{{pn|date=November 2024}}</ref> They wait until prey comes near, then use a strategy called "tenting".<ref>{{cite journal | vauthors = Wilga CD, Maia A, Nauwelaerts S, Lauder GV | title = Prey handling using whole-body fluid dynamics in batoids | journal = Zoology | volume = 115 | issue = 1 | pages = 47–57 | date = February 2012 | pmid = 22244456 | doi = 10.1016/j.zool.2011.09.002 | bibcode = 2012Zool..115...47W }}</ref> With pectoral fins pressed against the substrate, the ray will raise its head, generating a suction force that pulls the prey underneath the body. This form of whole-body suction is analogous to the buccal [[suction feeding]] performed by ray-finned fish. Stingrays exhibit a wide range of colors and patterns on their dorsal surface to help them camouflage with the sandy bottom. Some stingrays can even change color over the course of several days to adjust to new habitats. Since their mouths are on the underside of their bodies, they catch their prey, then crush and eat with their powerful jaws. Like its shark relatives, the stingray is outfitted with electrical sensors called ampullae of Lorenzini. Located around the stingray's mouth, these organs sense the natural electrical charges of potential prey. Many rays have jaw teeth to enable them to crush mollusks such as clams, oysters and mussels.


Most stingrays feed primarily on [[mollusca|mollusk]]s, [[crustacean]]s and, occasionally, on small fish. [[Potamotrygonidae|Freshwater stingrays]] in the Amazon feed on insects and break down their tough exoskeletons with mammal-like chewing motions.<ref>{{cite journal | vauthors = Kolmann MA, Welch KC, Summers AP, Lovejoy NR | title = Always chew your food: freshwater stingrays use mastication to process tough insect prey | journal = Proceedings. Biological Sciences | volume = 283 | issue = 1838 | pages = 20161392 | date = September 2016 | pmid = 27629029 | pmc = 5031661 | doi = 10.1098/rspb.2016.1392 }}</ref> Large [[Pelagic fish|pelagic]] rays like the [[Manta ray|manta]] use [[Aquatic feeding mechanisms|ram feeding]] to consume vast quantities of [[plankton]] and have been seen swimming in acrobatic patterns through plankton patches.<ref name="Notarbartolo-di-Sciara 607–614">{{Cite journal|last1=Notarbartolo-di-Sciara|first1=Giuseppe|last2=Hillyer|first2=Elizabeth V. | name-list-style = vanc |date=1989-01-01|title=Mobulid Rays off Eastern Venezuela (Chondrichthyes, Mobulidae)|journal=Copeia|volume=1989|issue=3|pages=607–614|doi=10.2307/1445487|jstor=1445487}}</ref>
The [[Atlantis Paradise Island]] Hotel houses many eagle rays, sting rays and one [[manta ray]]. The rays are often coexhibited with other marine life, such as the [[Caribbean reef shark]]. The [[Georgia Aquarium]] allows petting of southern stingrays in their "Georgia Explorer" exhibit. Similarly, visitors may use two fingers at a time to touch rays (with sting removed) and related [[guitarfish]] in outdoor exhibits at the [[Aquarium of the Pacific]] in [[Long Beach, California]]. Petting stingrays is also permitted in a special tank at the [[Blue Planet Aquarium]], [[Ellesmere Port]], [[England]]. Likewise, the [[Mote Marine Aquarium]] in [[Sarasota, Florida]] and the North Carolina Aquarium in Manteo, NC, allow visitors to pet a variety of rays in a controlled tank setting. [[Coral World]] Marine Park in St. Thomas, USVI, even allows supervised feeding of southern stingrays by visitors, as does [[Underwater Adventures Aquarium]] at the [[Mall of America]] in Bloomington, Minnesota. The [[London Aquarium]] in [[London]], [[England]], also has a touch tank where Aquarium visitors may touch a variety of small rays that live in a graduated-depth tank. The PPG Aquarium at the [[Pittsburgh Zoo]] also has a stingray tank where visitors can pet a variety of rays and small sharks, as well as crawl through a clear tunnel around the tank.


==Stingray injuries==
In 2006, the [[Tampa Bay Devil Rays]] added a 35 foot (10.7&nbsp;m), 10&nbsp;000 gallon (38&nbsp;000&nbsp;L), touch tank in their stadium where fans get a chance to interact with dozens of rays.<ref>{{cite news | url = http://www.sptimes.com/2006/07/13/Tampabay/Way_cool__but_can_the.shtml | title = Way cool, but can they pitch? | publisher = [[St. Petersburg Times]] | date = [[July 13]], [[2006]] }}</ref>
{{Main|Stingray injury}}
[[File:Stringray's sting.jpg|thumb|upright=1.5|The stinger of a stingray is known also as the spinal blade. It is located in the mid-area of the tail and can secrete venom. The ruler measures 10cm.]]


Stingrays are not usually aggressive and ordinarily attack humans only when provoked, such as when they are accidentally stepped on.<ref name="Slaughter">{{cite journal |last1=Slaughter |first1=Robin J. |last2=Beasley |first2=D. Michael G. |last3=Lambie |first3=Bruce S. |last4=Schep |first4=Leo J. |title=New Zealand's venomous creatures |journal=The New Zealand Medical Journal |date=27 February 2009 |volume=122 |issue=1290 |pages=83–97 |pmid=19319171 }}</ref> Stingrays can have one, two or three blades. Contact with the spinal blade or blades causes local trauma (from the cut itself), pain, swelling, muscle cramps from the venom and, later, may result in infection from bacteria or fungi.<ref>{{cite web|title=Stingray Injury Case Reports|url=http://www.toxinology.com/fusebox.cfm?staticaction=marine_vertebrates%2Fns-stingray_injuries.html|work=Clinical Toxicology Resources|publisher=University of Adelaide|access-date=22 October 2012|archive-date=4 April 2019|archive-url=https://web.archive.org/web/20190404015645/http://www.toxinology.com/fusebox.cfm?staticaction=marine_vertebrates%2Fns-stingray_injuries.html|url-status=live}}</ref> The injury is very painful, but rarely life-threatening unless the stinger pierces a vital area.<ref name="Slaughter"/> The blade is often deeply barbed and usually breaks off in the wound. Surgery may be required to remove the fragments.<ref>{{cite journal | vauthors = Flint DJ, Sugrue WJ | title = Stingray injuries: a lesson in debridement | journal = The New Zealand Medical Journal | volume = 112 | issue = 1086 | pages = 137–8 | date = April 1999 | pmid = 10340692 }}</ref>
==Danger to humans==
[[Image:4006 aquaimages.jpg|thumb|left|200px|A stingray buried in the sand in [[Saba]]. Stingrays can be hard to see when they cover themselves with [[substrate]].]]
[[Image:BLUESPOT 0001.jpg|thumb|right|200px|A stingray in dark waters. Stingrays are dangerous for humans because it's hard to see them when they're in dark waters.]]


Fatal stings are very rare.<ref name="Slaughter"/> The [[death of Steve Irwin]] in 2006 was only the second recorded in Australian waters since 1945.<ref>{{Cite web| url=https://scienceline.org/2006/09/ask-grant-irwin/|author=Hadhazy, Adam T. |work=Scienceline| title=I thought stingrays were harmless, so how did one manage to kill the "Crocodile Hunter?"| date=2006-09-11| access-date=2018-11-18| archive-date=2022-03-29| archive-url=https://web.archive.org/web/20220329163907/https://scienceline.org/2006/09/ask-grant-irwin/| url-status=live}}</ref> The stinger penetrated his [[thoracic wall]] and pierced his heart, causing massive trauma and bleeding.<ref>[http://animal.discovery.com/fansites/crochunter/steve-irwin/stingray/stingray.html Discovery Channel Mourns the Death of Steve Irwin] {{Webarchive|url=https://web.archive.org/web/20130107070254/http://animal.discovery.com/fansites/crochunter/steve-irwin/stingray/stingray.html |date=2013-01-07 }}. animal.discovery.com</ref>
The most famous stingray-related injury is the one that resulted in [[Steve Irwin#Death|Steve Irwin's death]]. On [[September 4]], [[2006]], he was pierced in the chest with a stingray barb while snorkeling in Australia. He was killed from severe loss of blood from the heart to the abdominal cavity, and was dead by the time his team had brought aboard Croc II.


=== Venom ===
{{Original research|date=October 2007}}
[[File:Wikipedia Project Anatomy Of Stingray Tail.svg|thumb|Posterior anatomy of a stingray. (1) Pelvic Fins (2) Caudal Tubercles (3) Stinger (4) Dorsal Fin (5) Claspers (6) Tail]]
The [[venom]] of the stingray has been relatively unstudied due to the mixture of venomous tissue secretions [[Cell (biology)|cells]] and [[mucous membrane]] cell products that occurs upon secretion from the spinal blade. The spine is covered with the epidermal skin layer. During secretion, the venom penetrates the [[epidermis]] and mixes with the mucus to release the venom on its victim. Typically, other venomous organisms create and store their venom in a [[gland]]. The stingray is notable in that it stores its venom within tissue cells. The toxins that have been confirmed to be within the venom are [[cystatin]]s, [[peroxiredoxin]] and [[galectin]].<ref>{{cite journal | vauthors = da Silva NJ, Ferreira KR, Pinto RN, Aird SD | title = A Severe Accident Caused by an Ocellate River Stingray (''Potamotrygon motoro'') in Central Brazil: How Well Do We Really Understand Stingray Venom Chemistry, Envenomation, and Therapeutics? | journal = Toxins | volume = 7 | issue = 6 | pages = 2272–88 | date = June 2015 | pmid = 26094699 | pmc = 4488702 | doi = 10.3390/toxins7062272 | doi-access = free }}</ref> Galectin induces cell death in its victims and cystatins inhibit defense enzymes. In humans, these toxins lead to increased blood flow in the superficial capillaries and cell death.<ref>{{cite journal | vauthors = Dos Santos JC, Grund LZ, Seibert CS, Marques EE, Soares AB, Quesniaux VF, Ryffel B, Lopes-Ferreira M, Lima C | title = Stingray venom activates IL-33 producing cardiomyocytes, but not mast cell, to promote acute neutrophil-mediated injury | journal = Scientific Reports | volume = 7 | issue = 1 | pages = 7912 | date = August 2017 | pmid = 28801624 | pmc = 5554156 | doi = 10.1038/s41598-017-08395-y | bibcode = 2017NatSR...7.7912D }}</ref> Despite the number of cells and toxins that are within the stingray, there is little relative energy required to produce and store the venom.


The venom is produced and stored in the secretory cells of the [[vertebral column]] at the mid-distal region. These secretory cells are housed within the ventrolateral grooves of the spine. The [[Cell (biology)|cells]] of both marine and freshwater stingrays are round and contain a great amount of [[Granule (cell biology)|granule]]-filled cytoplasm.<ref name = "Pedroso_2007">{{cite journal | vauthors = Pedroso CM, Jared C, Charvet-Almeida P, Almeida MP, Garrone Neto D, Lira MS, Haddad V, Barbaro KC, Antoniazzi MM | title = Morphological characterization of the venom secretory epidermal cells in the stinger of marine and freshwater stingrays | journal = Toxicon | volume = 50 | issue = 5 | pages = 688–97 | date = October 2007 | pmid = 17659760 | doi = 10.1016/j.toxicon.2007.06.004 | bibcode = 2007Txcn...50..688P }}</ref> The stinging cells of marine stingrays are located only within these lateral grooves of the stinger.<ref>{{cite journal | vauthors = Enzor LA, Wilborn RE, Bennett WA | date = December 2011 | title=Toxicity and metabolic costs of the Atlantic stingray (Dasyatis sabina) venom delivery system in relation to its role in life history |journal=Journal of Experimental Marine Biology and Ecology|volume=409|issue=1–2|pages=235–239|doi=10.1016/j.jembe.2011.08.026 | bibcode = 2011JEMBE.409..235E }}</ref> The stinging cells of freshwater stingray branch out beyond the lateral grooves to cover a larger surface area along the entire blade. Due to this large area and an increased number of proteins within the cells, the venom of freshwater stingrays has a greater toxicity than that of marine stingrays.<ref name = "Pedroso_2007" />
These chest-puncturing wounds are not typical, and most injuries that humans get from stingrays are because they step on one by accident. Stingrays are often semi-buried in the sand so they are easy to overlook. It is also possible to be bitten, but this is also a rare type of injury, since the mouth is on the bottom side of the ray. A good way to avoid stepping on a stingray while walking in shallow water is to slide your feet along the ground rather than taking steps (this is often referred to as the "stingray shuffle"). If a stingray feels something moving toward it, it will flee. If stepped on, however, its first reaction will be to sting.


== Species ==
==Human use==
===As food===
[[File:Eihire - stingray meat - Japanese pub food - September 2014.jpg|thumb|Dried strips of stingray meat served as food in Japan]]


Rays are edible, and may be caught as food using fishing lines or spears. Stingray recipes can be found in many coastal areas worldwide.<ref name="ADW">{{Cite web |date=2021-03-10 |title=Animal Diversity Web – Dasyatidae, Stingrays |url=https://animaldiversity.org/site/accounts/information/Dasyatidae.html |access-date=2021-03-10 |website=Animal Diversity Web |archive-date=2021-06-17 |archive-url=https://web.archive.org/web/20210617014811/https://animaldiversity.org/site/accounts/information/Dasyatidae.html |url-status=live }}</ref> For example, in [[Malaysia]] and [[Singapore]], stingray is commonly [[grilling|grill]]ed over charcoal, then served with spicy ''[[sambal]]'' sauce. In [[Goa]], and other [[India]]n states, it is sometimes used as part of spicy curries. Generally, the most prized parts of the stingray are the wings, the "cheek" (the area surrounding the eyes), and the liver. The rest of the ray is considered too rubbery to have any culinary uses.<ref>{{Cite web|title=The Delicious and Deadly Stingray. Nyonya. New York, NY. (Partially from the Archives.)|url=http://www.deependdining.com/2006/09/delicious-and-deadly-stingray-nyonya.html|access-date=2023-02-14|language=en
[[Image:Dasyatis sabina.jpg|thumb|right|An Atlantic stingray]]
| last = Lin | first = Eddie
| date = 2006
| website= Deep End Dining
| publisher= (blog)
}}</ref>


===Ecotourism===
[[Image:Dasyatis thetidis.jpg|thumb|right|A thorntail stingray]]
[[File:Stingray CIty in Grand Cayman.jpg|thumb|Divers can interact with [[Southern stingray|southern stingrays (''Hypanus americanus'')]] at [[Stingray City]] in the [[Cayman Islands]].]]
Stingrays are usually very docile and curious, their usual reaction being to flee any disturbance, but they sometimes brush their fins past any new object they encounter. Nevertheless, certain larger species may be more aggressive and should be approached with caution, as the stingray's defensive reflex (use of its venomous stinger) may result in serious injury or death.<ref>{{cite web | vauthors = Sullivan BN | date = May 2009 | url = http://therightblue.blogspot.com/2009/05/stingrays-dangerous-or-not.html | title = Stingrays: Dangerous or Not? | work = The Right Blue | access-date = 17 July 2012 | archive-date = 24 July 2012 | archive-url = https://web.archive.org/web/20120724162037/http://therightblue.blogspot.com/2009/05/stingrays-dangerous-or-not.html | url-status = live }}</ref>


===Other uses===
[[Image:Himantura kittipongi.jpg|thumb|right|An Himantura stingray]]
[[File:Stingray wallets.JPG|thumb|left|Stingray wallets]]
The skin of the ray is used as an under layer for the cord or leather wrap (known as ''samegawa'' in [[Japanese Language|Japanese]]) on [[Katana|Japanese swords]] due to its hard, rough texture that keeps the braided wrap from sliding on the handle during use.<ref name="Samegawa">{{Cite web |title=The Samegawa – Parts of a Japanese Katana |url=https://www.reliks.com/functional-swords/japanese-swords/samegawa/ |access-date=2021-03-10 |website=Reliks |archive-date=2021-02-26 |archive-url=https://web.archive.org/web/20210226180154/https://www.reliks.com/functional-swords/japanese-swords/samegawa/ |url-status=live }}</ref>


Several ethnological sections in museums,<ref>[http://www.flmnh.ufl.edu/fish/Gallery/Descript/DaisyStingray/DaisyStingray.html FLMNH Ichthyology Department: Daisy Stingray] {{Webarchive|url=https://web.archive.org/web/20160104222906/http://www.flmnh.ufl.edu/fish/Gallery/Descript/DaisyStingray/DaisyStingray.html |date=2016-01-04 }}. Flmnh.ufl.edu. Retrieved on 17 July 2012.</ref> such as the [[British Museum]], display arrowheads and spearheads made of stingray stingers, used in [[Micronesia]] and elsewhere.<ref>{{cite iucn |author=Séret, B. |author2=Couzens, G. |author3=Valenti, S.V. |year=2016 |title=''Hypanus rudis'' |volume=2016 |page=e.T161620A104133548 |doi=10.2305/IUCN.UK.2016-3.RLTS.T161620A104133548.en |access-date=2 November 2024}}</ref> [[Henry de Monfreid]] stated in his books that before [[World War II]], in the [[Horn of Africa]], [[whip]]s were made from the tails of big stingrays and these devices inflicted cruel cuts, so in [[Aden]], the British forbade their use on women and slaves. In former Spanish colonies, a stingray is called {{Lang|es|raya látigo}} ("whip ray").
There are about seventy species in six genera:


Some stingray species are commonly seen in public [[aquarium]] exhibits and more recently in home aquaria.<ref name="ADW" /><ref name="TFH">{{Cite magazine |last=Michael |first=Scott W. |date=September 2014 |title=Rays in the Home Aquarium |url=https://www.tfhmagazine.com/articles/saltwater/rays-in-the-home-aquarium-full-article |magazine=Tropical Fish Magazine |access-date=2021-03-10 |archive-date=2021-04-22 |archive-url=https://web.archive.org/web/20210422143732/https://www.tfhmagazine.com/articles/saltwater/rays-in-the-home-aquarium-full-article |url-status=live }}</ref>{{clear}}
* Genus ''[[Dasyatis]]''
** ''[[Dasyatis acutirostra]]'' <small>(Nishida & Nakaya, 1988)</small>.
** [[Red stingray]], ''[[Dasyatis akajei]]'' <small>([[Johannes Peter Müller|Müller]] & [[Friedrich Gustav Jakob Henle|Henle]], 1841)</small>.
** [[Southern stingray]], ''[[Southern stingray|Dasyatis americana]]'' <small>(Hildebrand & Schroeder, 1928)</small>.
** [[Plain maskray]], ''[[Dasyatis annotata]]'' <small>(Last, 1987)</small>.
** [[Bennett's stingray]], ''[[Dasyatis bennetti]]'' <small>([[Johannes Peter Müller|Müller]] & [[Friedrich Gustav Jakob Henle|Henle]], 1841)</small>.
** [[Short-tail stingray]] or [[bull ray]], ''[[Dasyatis brevicaudata]]'' <small>(Hutton, 1875)</small>.
** [[Whiptail stingray]], ''[[Dasyatis brevis]]'' <small>([[Samuel Garman|Garman]], 1880)</small>.
** [[Roughtail stingray]], ''[[Dasyatis centroura]]'' <small>(Mitchill, 1815)</small>.
** [[Blue stingray]], ''[[Dasyatis chrysonota]]'' <small>(Smith, 1828)</small>.
** [[Diamond stingray]], ''[[Dasyatis dipterura]]'' <small>([[David Starr Jordan|Jordan]] & Gilbert, 1880)</small>.
** [[Estuary stingray]], ''[[Dasyatis fluviorum]]'' <small>([[William Ogilby|Ogilby]], 1908)</small>.
** [[Smooth freshwater stingray]], ''[[Dasyatis garouaensis]]'' <small>(Stauch & Blanc, 1962)</small>.
** [[Sharpsnout stingray]], ''[[Dasyatis geijskesi]]'' <small>(Boeseman, 1948)</small>.
** [[Giant stumptail stingray]], ''[[Dasyatis gigantea]]'' <small>(Lindberg, 1930)</small>.
** [[Longnose stingray]], ''[[Dasyatis guttata]]'' <small>([[Marcus Elieser Bloch|Bloch]] & [[Johann Gottlob Schneider|Schneider]], 1801)</small>.
** ''[[Dasyatis hastata]]'' <small>(DeKay, 1842)</small>.
** [[Izu stingray]], ''[[Dasyatis izuensis]]'' <small>(Nishida & Nakaya, 1988)</small>.
** [[Bluespotted stingray]], ''[[Dasyatis kuhlii]]'' <small>([[Johannes Peter Müller|Müller]] & [[Friedrich Gustav Jakob Henle|Henle]], 1841)</small>.
** [[Yantai stingray]], ''[[Dasyatis laevigata]]'' <small>(Chu, 1960)</small>.
** [[Mekong stingray]], ''[[Dasyatis laosensis]]'' <small>(Roberts & Karnasuta, 1987)</small>.
** [[Brown stingray]], ''[[Dasyatis latus]]'' <small>([[Samuel Garman|Garman]], 1880)</small>.
** [[Painted maskray]], ''[[Dasyatis leylandi]]'' <small>(Last, 1987)</small>.
** [[Longtail stingray]], ''[[Dasyatis longa]]'' <small>([[Samuel Garman|Garman]], 1880)</small>.
** [[Daisy stingray]], ''[[Dasyatis margarita]]'' <small>([[Albert Günther|Günther]], 1870)</small>.
** [[Pearl stingray]], ''[[Dasyatis margaritella]]'' <small>(Compagno & Roberts, 1984)</small>.
** ''[[Dasyatis marianae]]'' <small>(Gomes, Rosa & Gadig, 2000)</small>.
** [[Marbled stingray]], ''[[Dasyatis marmorata]]'' <small>([[Franz Steindachner|Steindachner]], 1892)</small>.
** [[Pitted stingray]], ''[[Dasyatis matsubarai]]'' <small>(Miyosi, 1939)</small>.
** [[Smalleye stingray]], ''[[Dasyatis microps]]'' <small>(Annandale, 1908)</small>.
** [[Multispine giant stingray]], ''[[Dasyatis multispinosa]]'' <small>(Tokarev, 1959)</small>.
** [[Blackish stingray]], ''[[Dasyatis navarrae]]'' <small>([[Franz Steindachner|Steindachner]], 1892)</small>.
** [[Common stingray]], ''[[Dasyatis pastinaca]]'' <small>([[Carolus Linnaeus|Linnaeus]], 1758)</small>.
** [[Smalltooth stingray]], ''[[Dasyatis rudis]]'' <small>([[Albert Günther|Günther]], 1870)</small>.
** [[Atlantic stingray]], ''[[Dasyatis sabina]]'' <small>(Lesueur, 1824)</small>.
** [[Bluntnose stingray]], ''[[Dasyatis say]]'' <small>(Lesueur, 1817)</small>.
** [[Chinese stingray]], ''[[Dasyatis sinensis]]'' <small>([[Franz Steindachner|Steindachner]], 1892)</small>.
** [[Thorntail stingray]], ''[[Dasyatis thetidis]]'' <small>([[William Ogilby|Ogilby]], 1899)</small>.
** [[Tortonese's stingray]], ''[[Dasyatis tortonesei]]'' <small>(Capapé, 1975)</small>.
** [[Cow stingray]], ''[[Dasyatis ushiei]]'' <small>([[David Starr Jordan|Jordan]] & Hubbs, 1925)</small>.
** [[Pale-edged stingray]], ''[[Dasyatis zugei]]'' <small>([[Johannes Peter Müller|Müller]] & [[Friedrich Gustav Jakob Henle|Henle]], 1841)</small>.


==Gallery==
* Genus ''[[Himantura]]''
<gallery mode="packed" heights="140">
** [[Pale-spot whip ray]], ''[[Himantura alcockii]]'' <small>(Annandale, 1909)</small>.
File:Hexatrygon bickelli csiro-nfc.jpg|Unlike other [[Batoidea|rays]], [[sixgill stingray]]s (''Hexatrygon bickelli'') have six rather than five pairs of [[gill slits]].
** [[Bleeker's whipray]], ''[[Himantura bleekeri]]'' <small>(Blyth, 1860)</small>.
File:Plesiobatis daviesi cochin.jpg|[[Deepwater stingray]]s (''Plesiobatis daviesi'') are found on the upper continental slope throughout the [[Indo-Pacific]].
** [[Freshwater whipray]], ''[[Himantura chaophraya]]'' <small>(Monkolprasit & Roberts, 1990)</small>.
File:Urolophus gigas museum victoria.jpg|[[Spotted stingaree]]s (''Urolophus gigas'') are found along the [[Coastal regions of Western Australia|Western Australian coast]].
** [[Dragon stingray]], ''[[Himantura draco]]'' <small>(Compagno & Heemstra, 1984)</small>.
File:Round stingray.jpg|[[Round stingray]]s (''Urobatis halleri'') frequently sting beachgoers along the [[Western American]] coast.
** [[Pink whipray]], ''[[Himantura fai]]'' <small>([[David Starr Jordan|Jordan]] & Seale, 1906)</small>.
File:Himantura leoparda ala.jpg|[[Leopard whipray]]s (''Himantura leoparda'') are [[Vulnerable species|vulnerable]] from overfishing.
** [[Ganges stingray]], ''[[Himantura fluviatilis]]'' <small>(Hamilton, 1822)</small>.
File:Dasyatis sabina.jpg|[[Atlantic stingray]]s (''Hypanus sabinus'') are found in [[Ocean|marine]], [[brackish]], and [[freshwater]] environments along the [[Southeastern United States]] coast.
** [[Sharpnose stingray]], ''[[Himantura gerrardi]]'' <small>([[John Edward Gray|Gray]], 1851)</small>.
File:Smalleye Stingray (Dasyatis microps).jpeg|The [[smalleye stingray]] (''Megatrygon microps'') is a rare stingray distributed throughout the [[Indo-Pacific]].
** [[Mangrove whipray]], ''[[Himantura granulata]]'' <small>(Macleay, 1883)</small>.
File:Pelagic stingray fukushima.jpg|The [[pelagic stingray]] (''Pteroplatytrygon violacea'') is one of the few stingrays that primarily inhabit the [[open ocean]].
**''[[Himantura hortlei]]'' <small>Last, Manjaji-Matsumoto & Kailola, 2006</small>.<ref>{{cite journal | journal = Zootaxa | url = http://www.mapress.com/zootaxa/2006f/z01239p034f.pdf | volume = 1239 | year = 2006 | pages = 19-34 | title = Himantura hortlei n. sp., a new species of whipray (Myliobatiformes: Dasyatidae) from Irian Jaya, Indonesia | author = Last, Manjaji-Matsumoto & Kailola }}</ref>
File:Taeniura lymma edit.JPG|[[Bluespotted ribbontail ray]]s (''Taeniura lymma'')
** [[Scaly whipray]], ''[[Himantura imbricata]]'' <small>([[Marcus Elieser Bloch|Bloch]] & [[Johann Gottlob Schneider|Schneider]], 1801)</small>.
File:Himantura chaophraya (Monkolprasit & Roberts, 1990).jpg|[[Giant freshwater stingray]]s (''Urogymnus polylepis'') are amongst the largest freshwater fish.
** [[Pointed-nose stingray]], ''[[Himantura jenkinsii]]'' <small>(Annandale, 1909)</small>.
File:Ocellate river stingray, Boston Aquarium.jpg|[[Ocellate river stingray]]s (''Potamotrygon motoro'') are found in [[South American]] rivers.
** [[Kittipong's stingray]], ''[[Himantura kittipongi]]''
File:Gymnura altavela.jpg|[[Spiny butterfly ray]]s (''Gymnura altavela'') are [[Endangered species|endangered]] from [[overfishing]]. Found along the lower [[East Coast of the United States]] and the [[South American]] coast.
** [[Marbled freshwater whip ray]], ''[[Himantura krempfi]]'' <small>(Chabanaud, 1923)</small>.
File:Pteromylaeus bovinus valencia.jpg|[[Bull ray]]s (''Aetomylaeus bovinus'') are found along [[Europe]]an and [[Africa]]n coasts.
** ''[[Himantura lobistoma]]'' <small>Manjaji-Matsumoto & Last, 2006</small>.<ref>{{cite journal | journal = Ichthyological Research | volume = 53 | issue = 3 | year = 2006 | pages = 291ff | title = Himantura lobistoma, a new whipray (Rajiformes: Dasyatidae) from Borneo, with comments on the status of Dasyatis microphthalmus. | author = Manjaji-Matsumoto & Last}}</ref>
File:Corl0112 (28034475541).jpg|[[Giant oceanic manta ray]]s (''Mobula birostris'') are the largest of the stingrays.
** [[Blackedge whipray]], ''[[Himantura marginatus]]'' <small>(Blyth, 1860)</small>.
File:Rhinoptera steindachneri.jpg|[[Golden cownose ray]]s (''Rhinoptera steindachneri'') often migrate in large schools.
** [[Smalleye whip ray]], ''[[Himantura microphthalma]]'' <small>(Chen, 1948)</small>.
</gallery>
** [[Marbled whipray]], ''[[Himantura oxyrhyncha]]'' <small>(Sauvage, 1878)</small>.
** [[Pacific chupare]], ''[[Himantura pacifica]]'' <small>(Beebe & Tee-Van, 1941)</small>.
** ''[[Himantura pareh]]'' <small>([[Pieter Bleeker|Bleeker]], 1852)</small>.
** [[Round whip ray]], ''[[Himantura pastinacoides]]'' <small>([[Pieter Bleeker|Bleeker]], 1852)</small>.
** [[Chupare stingray]], ''[[Himantura schmardae]]'' <small>(Werner, 1904)</small>.
** [[White-edge freshwater whip ray]], ''[[Himantura signifer]]'' <small>(Compagno & Roberts, 1982)</small>.
** [[Black-spotted whipray]], ''[[Himantura toshi]]'' <small>(Whitley, 1939)</small>.
** [[Whitenose whip ray]], ''[[Himantura uarnacoides]]'' <small>([[Pieter Bleeker|Bleeker]], 1852)</small>.
** [[Honeycomb stingray]], ''[[Himantura uarnak]]'' <small>([[Peter Forsskål|Forsskål]], 1775)</small>.
** [[Leopard whipray]], ''[[Himantura undulata]]'' <small>([[Pieter Bleeker|Bleeker]], 1852)</small>.
** [[Dwarf whipray]], ''[[Himantura walga]]'' <small>([[Johannes Peter Müller|Müller]] & [[Friedrich Gustav Jakob Henle|Henle]], 1841)</small>.
* Genus ''[[Pastinachus]]''
** [[Cowtail stingray]], ''[[Pastinachus sephen]]'' <small>([[Peter Forsskål|Forsskål]], 1775)</small>. [http://www.sfu.ca/biology/faculty/dill/publications/417.pdf]
**''[[Pastinachus solocirostris]]'' <small>(Last, Manjaji & Yearsley, 2005)</small>.<ref>{{cite journal | journal = Zootaxa | url = http://www.mapress.com/zootaxa/2005f/z01040p016f.pdf | volume = 1040 | year = 2005 | pages = 1-16 | title = Pastinachus solocirostris sp. nov., a new species of Stingray (Elasmobranchii: Myliobatiformes) from the Indo-Malay Archipelago | author = Last, Manjaji & Yearsley }}</ref>
* Genus ''[[Pteroplatytrygon]]''
** [[Pelagic stingray]], ''[[Pteroplatytrygon violacea]]'' <small>([[Charles Lucien Jules Laurent Bonaparte|Bonaparte]], 1832)</small>.
* Genus ''[[Taeniura]]''
** [[Round stingray]], ''[[Taeniura grabata]]'' <small>([[Étienne Geoffroy Saint-Hilaire|É. Geoffroy Saint-Hilaire]], 1817)</small>.
** [[Bluespotted ribbontail ray]], ''[[Taeniura lymma]]'' <small>([[Peter Forsskål|Forsskål]], 1775)</small>.
** [[Blotched fantail ray]], ''[[Taeniura meyeni]]'' <small>([[Johannes Peter Müller|Müller]] & [[Friedrich Gustav Jakob Henle|Henle]], 1841)</small>.
* Genus ''[[Urogymnus]]''
** [[Porcupine ray]], ''[[Urogymnus asperrimus]]'' <small>([[Marcus Elieser Bloch|Bloch]] & [[Johann Gottlob Schneider|Schneider]], 1801)</small>.
** [[Thorny freshwater stingray]], ''[[Urogymnus ukpam]]'' <small>(Smith, 1863)</small>.


==References==
== See also ==
* [[List of threatened rays]]
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See http://en.wikipedia.org/wiki/Wikipedia:Footnotes for an explanation of how to generate footnotes using the <ref> and </ref> tags, and the template below.
== References ==
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==Bibliography==
==Bibliography==
* {{FishBase family | family = Dasyatidae | month = August | year = 2005}}
* {{FishBase family | family = Dasyatidae | month = August | year = 2005}}


==External links==
== External links ==
{{Commons category|Myliobatoidei|Stingray}}
{{commonscat|Dasyatidae}}
* {{cite journal |last1=Almagro |first1=Álvaro |last2=Barría |first2=Claudio |title=The end of silent predators: First cases of active sound production in batoids from the Mediterranean Sea and its potential implications |journal=Marine Biology |date=November 2024 |volume=171 |issue=11 |page=208 |doi=10.1007/s00227-024-04536-w |bibcode=2024MarBi.171..208A }}
*[http://www.pennhealth.com/ency/article/002853.htm University of Pennsylvania Health System] – Information on stingray poison.
* {{cite journal |last1=Barroil |first1=Adèle |last2=Deter |first2=Julie |last3=Holon |first3=Florian |last4=Bertucci |first4=Frédéric |title=Sound production in wild Mediterranean blonde ray Raja brachyura |journal=Ecology |date=7 October 2024 |volume=105 |issue=11 |pages=e4440 |doi=10.1002/ecy.4440 |pmid=39370952 }}
*[http://www.sheddaquarium.org/cownose_rays.html Cownose rays] and [http://www.sheddaquarium.org/freshwater_rays.html freshwater rays] at [[John G. Shedd Aquarium]], [[Chicago]]
* {{cite journal |last1=Fetterplace |first1=Lachlan C. |last2=Delgado Esteban |first2=J. Javier |last3=Pini-Fitzsimmons |first3=Joni |last4=Gaskell |first4=John |last5=Wueringer |first5=Barbara E. |title=Evidence of sound production in wild stingrays |journal=Ecology |date=November 2022 |volume=103 |issue=11 |pages=e3812 |doi=10.1002/ecy.3812 |pmid=35808819 |bibcode=2022Ecol..103E3812F |pmc=9786621 }}
*[https://books.google.com/books?id=zSADAAAAMBAJ&pg=-PA7 "Beware the Ugly Sting Ray."] ''Popular Science'', July 1954, pp.{{nbsp}}117–118/pp.{{nbsp}}224–228.


{{Taxonbar|from=Q5328202}}
[[Category:Rays]]
[[Category:Rajiformes]]
[[Category:Dasyatidae]]
[[Category:Venomous fish]]


[[Category:Myliobatiformes]]
[[ar:سمك الراي اللاسع]]
[[Category:Venomous fish]]
[[da:Pilrokke]]
[[Category:Articles containing video clips]]
[[de:Stechrochen]]
[[Category:Extant Early Cretaceous first appearances]]
[[et:Astelrai]]
[[es:Dasyatidae]]
[[fr:Dasyatidae]]
[[is:Stingskötur]]
[[it:Dasyatidae]]
[[he:טריגוניים]]
[[lt:Paprastosios dygliauodegės rajos]]
[[ms:Ikan Pari]]
[[nl:Pijlstaartroggen]]
[[no:Piggrokker]]
[[pl:Ogończowate]]
[[ru:Хвостоколовые]]
[[simple:Stingray]]
[[sv:Spjutrockor]]
[[th:วงศ์ปลากระเบนธง]]
[[yi:סטינגרעי]]
[[zh:魟科]]

Latest revision as of 16:20, 7 December 2024

Stingrays
Temporal range: Early Cretaceous to recent[1]
Southern stingray (Hypanus americanus)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Order: Myliobatiformes
Suborder: Myliobatoidei
Compagno, 1973
Families

Stingrays are a group of sea rays, a type of cartilaginous fish. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae (sixgill stingray), Plesiobatidae (deepwater stingray), Urolophidae (stingarees), Urotrygonidae (round rays), Dasyatidae (whiptail stingrays), Potamotrygonidae (river stingrays), Gymnuridae (butterfly rays) and Myliobatidae (eagle rays).[2][3] There are about 220 known stingray species organized into 29 genera.

Stingrays are common in coastal tropical and subtropical marine waters throughout the world. Some species, such as the thorntail stingray (Dasyatis thetidis), are found in warmer temperate oceans and others, such as the deepwater stingray (Plesiobatis daviesi), are found in the deep ocean. The river stingrays and a number of whiptail stingrays (such as the Niger stingray (Fontitrygon garouaensis)) are restricted to fresh water. Most myliobatoids are demersal (inhabiting the next-to-lowest zone in the water column), but some, such as the pelagic stingray and the eagle rays, are pelagic.[4]

Stingray species are progressively becoming threatened or vulnerable to extinction, particularly as the consequence of unregulated fishing.[5] As of 2013, 45 species have been listed as vulnerable or endangered by the IUCN. The status of some other species is poorly known, leading to their being listed as data deficient.[citation needed]

Evolution

Early Eocene fossil stingray Heliobatis radians

Stingrays diverged from their closest relatives, the panrays, during the Late Jurassic period, and diversified over the course of the Cretaceous into the different extant families today. The earliest stingrays appear to have been benthic, with the ancestors of the eagle rays becoming pelagic during the early Late Cretaceous.[6][7]

Fossils

The bizarre Lessiniabatis of Early Eocene Italy

Permineralized stingray teeth have been found in sedimentary deposits around the world as far back as the Early Cretaceous. The oldest known stingray taxon is "Dasyatis" speetonensis from the Hauterivian of England, whose teeth most closely resemble that of the extant sixgill stingray (Hexatrygon). Although stingray teeth are rare on sea bottoms compared to the similar shark teeth, scuba divers searching for the latter do encounter the teeth of stingrays.[6][8]

Full-body stingray fossils are very rare but are known from certain lagerstätte that preserve soft-bodied animals. The extinct Cyclobatis of the Cretaceous of Lebanon is thought to be a skate that had convergently evolved a highly stingray-like body plan, although its exact taxonomic placement is still uncertain.[9] True stingray fossils become more common in the Eocene, with the extinct freshwater stingrays Heliobatis and Asterotrygon known from the Green River Formation.[10] A diversity of stingray fossils is known from the Eocene Monte Bolca formation from Italy, including the early stingaree Arechia, as well as Dasyomyliobatis, which is thought to represent a transitional form between stingrays and eagle rays, and the highly unusual Lessiniabatis, which had an extremely short and slender tail with no sting.[6][7]

Anatomy

dorsal (topside) ←               → ventral (underside)
External anatomy of a male bluntnose stingray (Hypanus say)
Stingray jaw and teeth.
The teeth are modified placoid scales.
Like in other rays, the bluespotted ribbontail ray (Taeniura lymma) breathes though spiracles just behind the eyes when it hunts in seafloor sediment.

Jaw and teeth

The mouth of the stingray is located on the ventral side of the vertebrate. Stingrays exhibit hyostylic jaw suspension, which means that the mandibular arch is only suspended by an articulation with the hyomandibula. This type of suspensions allows for the upper jaw to have high mobility and protrude outward.[11] The teeth are modified placoid scales that are regularly shed and replaced.[12] In general, the teeth have a root implanted within the connective tissue and a visible portion of the tooth, is large and flat, allowing them to crush the bodies of hard shelled prey.[13] Male stingrays display sexual dimorphism by developing cusps, or pointed ends, to some of their teeth. During mating season, some stingray species fully change their tooth morphology which then returns to baseline during non-mating seasons.[14]

Spiracles

Spiracles are small openings that allow some fish and amphibians to breathe. Stingray spiracles are openings just behind its eyes. The respiratory system of stingrays is complicated by having two separate ways to take in water to use the oxygen. Most of the time stingrays take in water using their mouth and then send the water through the gills for gas exchange. This is efficient, but the mouth cannot be used when hunting because the stingrays bury themselves in the ocean sediment and wait for prey to swim by.[15] So the stingray switches to using its spiracles. With the spiracles, they can draw water free from sediment directly into their gills for gas exchange.[16] These alternate ventilation organs are less efficient than the mouth, since spiracles are unable to pull the same volume of water. However, it is enough when the stingray is quietly waiting to ambush its prey.

The flattened bodies of stingrays allow them to effectively conceal themselves in their environments. Stingrays do this by agitating the sand and hiding beneath it. Because their eyes are on top of their bodies and their mouths on the undersides, stingrays cannot see their prey after capture; instead, they use smell and electroreceptors (ampullae of Lorenzini) similar to those of sharks.[17] Stingrays settle on the bottom while feeding, often leaving only their eyes and tails visible. Coral reefs are favorite feeding grounds and are usually shared with sharks during high tide.[18]

Behavior

Skeleton of an atlantic stingray (Hypanus sabinus)

Reproduction

Mobula (devil rays) are thought to breach as a form of courtship.

During the breeding season, males of various stingray species such as the round stingray (Urobatis halleri), may rely on their ampullae of Lorenzini to sense certain electrical signals given off by mature females before potential copulation.[19] When a male is courting a female, he follows her closely, biting at her pectoral disc. He then places one of his two claspers into her valve.[20]

Reproductive ray behaviors are associated with their behavioral endocrinology, for example, in species such as the atlantic stingray (Hypanus sabinus), social groups are formed first, then the sexes display complex courtship behaviors that end in pair copulation which is similar to the species Urobatis halleri.[21] Furthermore, their mating period is one of the longest recorded in elasmobranch fish. Individuals are known to mate for seven months before the females ovulate in March. During this time, the male stingrays experience increased levels of androgen hormones which has been linked to its prolonged mating periods.[21] The behavior expressed among males and females during specific parts of this period involves aggressive social interactions.[21] Frequently, the males trail females with their snout near the female vent then proceed to bite the female on her fins and her body.[21] Although this mating behavior is similar to the species Urobatis halleri, differences can be seen in the particular actions of Hypanus sabinus. Seasonal elevated levels of serum androgens coincide with the expressed aggressive behavior, which led to the proposal that androgen steroids start, indorse and maintain aggressive sexual behaviors in the male rays for this species which drives the prolonged mating season. Similarly, concise elevations of serum androgens in females has been connected to increased aggression and improvement in mate choice. When their androgen steroid levels are elevated, they are able to improve their mate choice by quickly fleeing from tenacious males when undergoing ovulation succeeding impregnation. This ability affects the paternity of their offspring by refusing less qualified mates.[21]

Stingrays are ovoviviparous, bearing live young in "litters" of five to thirteen. During this period, the female's behavior transitions to support of her future offspring. Females hold the embryos in the womb without a placenta. Instead, the embryos absorb nutrients from a yolk sac and after the sac is depleted, the mother provides uterine "milk".[22] After birth, the offspring generally disassociate from the mother and swim away, having been born with the instinctual abilities to protect and feed themselves. In a very small number of species, like the giant freshwater stingray (Urogymnus polylepis), the mother "cares" for her young by having them swim with her until they are one-third of her size.[23]

At the Sea Life London Aquarium, two female stingrays delivered seven baby stingrays, although the mothers have not been near a male for two years. This suggests some species of rays can store sperm then give birth when they deem conditions to be suitable.[24]

Locomotion

Atlantic stingray (Hypanus sabinus) undulation locomotion

The stingray uses its paired pectoral fins for moving around. This is in contrast to sharks and most other fish, which get most of their swimming power from a single caudal (tail) fin.[25] Stingray pectoral fin locomotion can be divided into two categories, undulatory and oscillatory.[26] Stingrays that use undulatory locomotion have shorter thicker fins for slower motile movements in benthic areas.[27] Longer thinner pectoral fins make for faster speeds in oscillation mobility in pelagic zones.[26] Visually distinguishable oscillation has less than one wave going, opposed to undulation having more than one wave at all times.[26]

Feeding behavior and diet

Bat ray (Myliobatis californica) in a feeding posture

Stingrays use a wide range of feeding strategies. Some have specialized jaws that allow them to crush hard mollusk shells,[28] whereas others use external mouth structures called cephalic lobes to guide plankton into their oral cavity.[29] Benthic stingrays (those that reside on the sea floor) are ambush hunters.[30] They wait until prey comes near, then use a strategy called "tenting".[31] With pectoral fins pressed against the substrate, the ray will raise its head, generating a suction force that pulls the prey underneath the body. This form of whole-body suction is analogous to the buccal suction feeding performed by ray-finned fish. Stingrays exhibit a wide range of colors and patterns on their dorsal surface to help them camouflage with the sandy bottom. Some stingrays can even change color over the course of several days to adjust to new habitats. Since their mouths are on the underside of their bodies, they catch their prey, then crush and eat with their powerful jaws. Like its shark relatives, the stingray is outfitted with electrical sensors called ampullae of Lorenzini. Located around the stingray's mouth, these organs sense the natural electrical charges of potential prey. Many rays have jaw teeth to enable them to crush mollusks such as clams, oysters and mussels.

Most stingrays feed primarily on mollusks, crustaceans and, occasionally, on small fish. Freshwater stingrays in the Amazon feed on insects and break down their tough exoskeletons with mammal-like chewing motions.[32] Large pelagic rays like the manta use ram feeding to consume vast quantities of plankton and have been seen swimming in acrobatic patterns through plankton patches.[33]

Stingray injuries

The stinger of a stingray is known also as the spinal blade. It is located in the mid-area of the tail and can secrete venom. The ruler measures 10cm.

Stingrays are not usually aggressive and ordinarily attack humans only when provoked, such as when they are accidentally stepped on.[34] Stingrays can have one, two or three blades. Contact with the spinal blade or blades causes local trauma (from the cut itself), pain, swelling, muscle cramps from the venom and, later, may result in infection from bacteria or fungi.[35] The injury is very painful, but rarely life-threatening unless the stinger pierces a vital area.[34] The blade is often deeply barbed and usually breaks off in the wound. Surgery may be required to remove the fragments.[36]

Fatal stings are very rare.[34] The death of Steve Irwin in 2006 was only the second recorded in Australian waters since 1945.[37] The stinger penetrated his thoracic wall and pierced his heart, causing massive trauma and bleeding.[38]

Venom

Posterior anatomy of a stingray. (1) Pelvic Fins (2) Caudal Tubercles (3) Stinger (4) Dorsal Fin (5) Claspers (6) Tail

The venom of the stingray has been relatively unstudied due to the mixture of venomous tissue secretions cells and mucous membrane cell products that occurs upon secretion from the spinal blade. The spine is covered with the epidermal skin layer. During secretion, the venom penetrates the epidermis and mixes with the mucus to release the venom on its victim. Typically, other venomous organisms create and store their venom in a gland. The stingray is notable in that it stores its venom within tissue cells. The toxins that have been confirmed to be within the venom are cystatins, peroxiredoxin and galectin.[39] Galectin induces cell death in its victims and cystatins inhibit defense enzymes. In humans, these toxins lead to increased blood flow in the superficial capillaries and cell death.[40] Despite the number of cells and toxins that are within the stingray, there is little relative energy required to produce and store the venom.

The venom is produced and stored in the secretory cells of the vertebral column at the mid-distal region. These secretory cells are housed within the ventrolateral grooves of the spine. The cells of both marine and freshwater stingrays are round and contain a great amount of granule-filled cytoplasm.[41] The stinging cells of marine stingrays are located only within these lateral grooves of the stinger.[42] The stinging cells of freshwater stingray branch out beyond the lateral grooves to cover a larger surface area along the entire blade. Due to this large area and an increased number of proteins within the cells, the venom of freshwater stingrays has a greater toxicity than that of marine stingrays.[41]

Human use

As food

Dried strips of stingray meat served as food in Japan

Rays are edible, and may be caught as food using fishing lines or spears. Stingray recipes can be found in many coastal areas worldwide.[43] For example, in Malaysia and Singapore, stingray is commonly grilled over charcoal, then served with spicy sambal sauce. In Goa, and other Indian states, it is sometimes used as part of spicy curries. Generally, the most prized parts of the stingray are the wings, the "cheek" (the area surrounding the eyes), and the liver. The rest of the ray is considered too rubbery to have any culinary uses.[44]

Ecotourism

Divers can interact with southern stingrays (Hypanus americanus) at Stingray City in the Cayman Islands.

Stingrays are usually very docile and curious, their usual reaction being to flee any disturbance, but they sometimes brush their fins past any new object they encounter. Nevertheless, certain larger species may be more aggressive and should be approached with caution, as the stingray's defensive reflex (use of its venomous stinger) may result in serious injury or death.[45]

Other uses

Stingray wallets

The skin of the ray is used as an under layer for the cord or leather wrap (known as samegawa in Japanese) on Japanese swords due to its hard, rough texture that keeps the braided wrap from sliding on the handle during use.[46]

Several ethnological sections in museums,[47] such as the British Museum, display arrowheads and spearheads made of stingray stingers, used in Micronesia and elsewhere.[48] Henry de Monfreid stated in his books that before World War II, in the Horn of Africa, whips were made from the tails of big stingrays and these devices inflicted cruel cuts, so in Aden, the British forbade their use on women and slaves. In former Spanish colonies, a stingray is called raya látigo ("whip ray").

Some stingray species are commonly seen in public aquarium exhibits and more recently in home aquaria.[43][49]

See also

References

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Bibliography