Stingrays Temporal range: | |
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Southern stingray (Hypanus americanus) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Chondrichthyes |
Subclass: | Elasmobranchii |
Superorder: | Batoidea |
Order: | Myliobatiformes |
Suborder: | Myliobatoidei Compagno, 1973 |
Families | |
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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. [6]
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. [7] [8]
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. [7] [9]
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. [10] True stingray fossils become more common in the Eocene, with the extinct freshwater stingrays Heliobatis and Asterotrygon known from the Green River Formation. [11] 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. [7] [8]
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. [12] The teeth are modified placoid scales that are regularly shed and replaced. [13] 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. [14] 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. [15]
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. [16] 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. [17] 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. [18] 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. [19]
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. [20] 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. [21]
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. [22] 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. [22] The behavior expressed among males and females during specific parts of this period involves aggressive social interactions. [22] Frequently, the males trail females with their snout near the female vent then proceed to bite the female on her fins and her body. [22] 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. [22]
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". [23] 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. [24]
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. [25]
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. [26] [27] Stingray pectoral fin locomotion can be divided into two categories, undulatory and oscillatory. [28] Stingrays that use undulatory locomotion have shorter thicker fins for slower motile movements in benthic areas. [29] Longer thinner pectoral fins make for faster speeds in oscillation mobility in pelagic zones. [28] Visually distinguishable oscillation has less than one wave going, opposed to undulation having more than one wave at all times. [28]
Stingrays use a wide range of feeding strategies. Some have specialized jaws that allow them to crush hard mollusk shells, [30] whereas others use external mouth structures called cephalic lobes to guide plankton into their oral cavity. [31] Benthic stingrays (those that reside on the sea floor) are ambush hunters. [32] They wait until prey comes near, then use a strategy called "tenting". [33] 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. [34] 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. [35]
Stingrays are not usually aggressive and ordinarily attack humans only when provoked, such as when they are accidentally stepped on. [36] 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. [37] The injury is very painful, but rarely life-threatening unless the stinger pierces a vital area. [36] The blade is often deeply barbed and usually breaks off in the wound. Surgery may be required to remove the fragments. [38]
Fatal stings are very rare. [36] The death of Steve Irwin in 2006 was only the second recorded in Australian waters since 1945. [39] The stinger penetrated his thoracic wall and pierced his heart, causing massive trauma and bleeding. [40]
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. [41] 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. [42] 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. [43] The stinging cells of marine stingrays are located only within these lateral grooves of the stinger. [44] 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. [43]
Rays are edible, and may be caught as food using fishing lines or spears. Stingray recipes can be found in many coastal areas worldwide. [45] 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. [46]
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. [47]
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. [48]
Several ethnological sections in museums, [49] such as the British Museum, display arrowheads and spearheads made of stingray stingers, used in Micronesia and elsewhere. [50] 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. [45] [51]
The great hammerhead is the largest species of hammerhead shark, belonging to the family Sphyrnidae, attaining an average length of 4.6 m (15 ft) and reaching a maximum length of 6.2 m (20 ft). It is found in tropical and warm temperate waters worldwide, inhabiting coastal areas and the continental shelf. The great hammerhead can be distinguished from other hammerheads by the shape of its "hammer", which is wide with an almost straight front margin, and by its tall, sickle-shaped first dorsal fin. A solitary, strong-swimming apex predator, the great hammerhead feeds on a wide variety of prey ranging from crustaceans and cephalopods, to bony fish, to smaller sharks. Observations of this species in the wild suggest that the cephalofoil functions to immobilize stingrays, a favored prey. This species has a viviparous mode of reproduction, bearing litters of up to 50 pups every two years.
River stingrays or freshwater stingrays are Neotropical freshwater fishes of the family Potamotrygonidae in the order Myliobatiformes, one of the four orders of batoids, cartilaginous fishes related to sharks. They are found in rivers in tropical and subtropical South America. A single marine genus, Styracura, of the tropical West Atlantic and East Pacific are also part of Potamotrygonidae. They are generally brownish, greyish or black, often with a mottled, speckled or spotted pattern, have disc widths ranging from 31 to 200 centimetres (1.0–6.6 ft) and venomous tail stingers. River stingrays feed on a wide range of smaller animals and the females give birth to live young. There are more than 35 species in five genera.
The short-tail stingray or smooth stingray is a common species of stingray in the family Dasyatidae. It occurs off southern Africa, typically offshore at a depth of 180–480 m (590–1,570 ft), and off southern Australia and New Zealand, from the intertidal zone to a depth of 156 m (512 ft). It is mostly bottom-dwelling in nature and can be found across a range of habitats from estuaries to reefs, but also frequently will swim into open water. One of the largest stingrays in the world, this heavy-bodied species can grow upwards of 2.1 m (6.9 ft) across and 350 kg (770 lb) in weight. Its plain-colored, diamond-shaped pectoral fin disc is characterized by a lack of dermal denticles even in adults, and white pores beside the head on either side. The body can have colors as well as dark grey or black with rows of white spots along each wing. Its tail is usually shorter than the disc and thick at the base. It is armed with large tubercles and a midline row of large thorns in front of the stinging spine which has the dorsal and ventral fin folds behind.
The Atlantic stingray is a species of stingray in the family Dasyatidae, common along the Atlantic coast of North America from Chesapeake Bay to Mexico, including brackish and freshwater habitats. It may be distinguished from other stingrays in the area by its relatively elongated snout. This species is of little commercial importance, other than for sale in the aquarium industry.
The southern stingray is a whiptail stingray found in tropical and subtropical waters of the Western Atlantic Ocean from New Jersey to southern Brazil. It has a flat, diamond-shaped disc, with a mud brown, olive, and grey dorsal surface and white underbelly. The barb on its tail is serrated and covered in a venomous mucus, used for self-defense.
Kuhl's maskray, also known as the blue-spotted stingray, blue-spotted maskray, or Kuhl's stingray, is a species of stingray of the family Dasyatidae. It was recently changed from Dasyatis kuhlii in 2008 after morphological and molecular analyses showed that it is part of a distinct genus, Neotrygon. The body is rhomboidal and colored green with blue spots. Maximum disk width is estimated 46.5 cm (18.3 in). It is popular in aquaria, but usually not distinguished from the blue-spotted ribbontail ray. The ribbontail has a rounded body, is a brighter green with brighter blue and more vivid spots, but Kuhl's maskray is larger. The stingray's lifespan is estimated at 13 years for females and 10 years for males. The blue-spotted stingray preys on many fish and small mollusks. It is also generally found from Indonesia to Japan, and most of Australia. Kuhl's maskray also is targeted by many parasites, such as tapeworms, flatworms, and flukes.
The pelagic stingray is a species of stingray in the family Dasyatidae, and the sole member of its genus. It is characterized by the wedge-like shape of its pectoral fin disc, which is much wider than long, as well as by the pointed teeth in both sexes, whip-like tail with extremely long tail spine, and uniform violet to blue-green coloration. It generally reaches 59 cm (23 in) in width. The pelagic stingray has a worldwide distribution in waters warmer than 19 °C (66 °F), and migrates seasonally to spend the summer closer to the continental shelf and at higher latitudes. The only stingray that almost exclusively inhabits the open ocean, this species is typically found in surface waters down to a depth of 100 m (330 ft). As a consequence of its midwater habits, its swimming style has evolved to feature more of a flapping motion of the pectoral fins, as opposed to the disc margin undulations used by other, bottom-dwelling stingrays.
The red stingray is a species of stingray in the family Dasyatidae, found in the northwestern Pacific Ocean off Japan, Korea, and China, and possibly elsewhere. It primarily inhabits shallow, sandy habitats close to shore, and has been known to enter brackish water. The red stingray has a diamond-shaped pectoral fin disc and gains its common name from its bright orange-red underside; there may also be patches of orange at various spots on its upper surface. Most individuals are no more than 1 m (3.3 ft) long.
The plain maskray or brown stingray, is a species of stingray in the family Dasyatidae. It is found in shallow, soft-bottomed habitats off northern Australia. Reaching 24 cm (9.4 in) in width, this species has a diamond-shaped, grayish green pectoral fin disc. Its short, whip-like tail has alternating black and white bands with fin folds above and below. There are short rows of thorns on the back and the base of the tail, but otherwise the skin is smooth. While this species possesses the dark mask-like pattern across its eyes common to its genus, it is not ornately patterned like other maskrays.
The diamond stingray is a species of stingray in the family Dasyatidae. It is found in the coastal waters of the eastern Pacific Ocean from southern California to northern Chile, and around the Galápagos and Hawaiian Islands. This bottom-dweller generally inhabits sandy or muddy flats near rocky reefs and kelp forests, to a depth of 30 m (98 ft), though off Hawaii it may range considerably deeper. As its common name suggests, this species has an angular, diamond-shaped pectoral fin disc that is plain brown or gray above, with rows of tubercles along the midline and on the "shoulders". The long, whip-like tail has both dorsal and ventral fin folds, which distinguish this ray from the closely similar longtail stingray. It typically grows to 1 m (3.3 ft) across.
The estuary stingray, also called the estuary stingaree or brown stingray, is a species of stingray in the family Dasyatidae. Endemic to eastern Australia, it typically inhabits shallow, mangrove-lined tidal rivers, estuaries, and bays in southern Queensland and New South Wales. This yellow-brown to olive ray grows to at least 93 cm (37 in) across. It has a diamond-shaped pectoral fin disc and a mostly smooth, whip-like tail bearing both dorsal and ventral fin folds. It can additionally be identified by its long, narrow nostrils and the row of thorns along the midline of its back.
The Niger stingray or smooth freshwater stingray is a species of stingray in the family Dasyatidae, native to rivers in Nigeria and Cameroon. Attaining a width of 40 cm (16 in), this species can be distinguished by its thin, almost circular pectoral fin disk, slightly projecting snout tip, and mostly smooth skin with small or absent dermal denticles. The Niger stingray feeds on aquatic insect larvae and is ovoviviparous. The long stinging spine on the tail of this ray can inflict a painful wound. It has been assessed as Critically Endangered by the International Union for Conservation of Nature (IUCN), as its numbers are declining in some areas and it faces heavy fishing pressure and habitat degradation.
The longnose stingray is a species of stingray in the family Dasyatidae, native to the western Atlantic Ocean from the southern Gulf of Mexico to Brazil. Found in coastal waters no deeper than 36 m (118 ft), this demersal species favors muddy or sandy habitats. The longnose stingray is characterized by its angular, rhomboid pectoral fin disc, moderately projecting snout, and whip-like tail with a dorsal keel and ventral fin fold. It typically grows to 1.25 m (4.1 ft) across and is brownish above and light-colored below.
The Mekong freshwater stingray, Hemitrygon laosensis, is a species of stingray in the family Dasyatidae, restricted to the Mekong and Chao Phraya Rivers in Laos and Thailand; the occurrence in Chao Phraya is considered an introduction. Measuring up to 62 cm (24 in) across, this ray has an oval pectoral fin disc, a tail with both upper and lower fin folds, and a midline row of spine-like dermal denticles. A characteristic feature of this species is its bright orange underside. The Mekong freshwater stingray preys on invertebrates and is aplacental viviparous. It has been assessed as Endangered by the International Union for Conservation of Nature (IUCN), as it is threatened by overfishing and habitat degradation.
The round stingray or Haller's round ray and Little round stingray is a species of round ray, family Urotrygonidae, found in the coastal waters of the tropical and subtropical parts of the northeastern Pacific Ocean. It is a small, common ray that feeds mostly on benthic invertebrates. On the beaches of southern California, it is responsible for numerous injuries to bathers, who are stung when they accidentally step on the fish. The wound caused by its venomous spine can be painful, but is non-fatal.
The yellow stingray is a species of stingray in the family Urotrygonidae, found in the tropical western Atlantic Ocean from North Carolina to Trinidad. This bottom-dwelling species inhabits sandy, muddy, or seagrass bottoms in shallow inshore waters, commonly near coral reefs.
The common stingray is a species of stingray in the family Dasyatidae, found in the northeastern Atlantic Ocean and the Mediterranean and Black Seas. It typically inhabits sandy or muddy habitats in coastal waters shallower than 60 m (200 ft), often burying itself in sediment. Usually measuring 45 cm (18 in) across, the common stingray has a diamond-shaped pectoral fin disc slightly wider than long, and a whip-like tail with upper and lower fin folds. It can be identified by its plain coloration and mostly smooth skin, except for a row of tubercles along the midline of the back in the largest individuals.
The roughtail stingray is a species of stingray in the family Dasyatidae, with separate populations in coastal waters of the northwestern and southwestern Atlantic Ocean. This bottom-dwelling species typically inhabits sandy or muddy areas with patches of invertebrate cover, at a depth of 15–50 m (49–164 ft). It is seasonally migratory, overwintering in offshore waters and moving into coastal habitats for summer. The largest whip-tail stingray in the Atlantic, the roughtail stingray grows up to 2.6 m (8.5 ft) across and 360 kg (800 lb) in weight. It is plain in color, with an angular, diamond-shaped pectoral fin disc and a long, whip-like tail bearing a subtle fin fold underneath. The many thorns on its back and tail serve to distinguish it from other stingrays that share its range.
The daisy stingray, Fontitrygon margarita, is a little-known species of stingray in the family Dasyatidae, found in shallow waters along the coast of West Africa. This species typically grows to 60 cm (24 in) across and has a rounded pectoral fin disc and a wide band of dermal denticles over its back. It is characterized by a greatly enlarged, nacreous denticle in the middle of its back called a "pearl spine"; this feature is shared with the similar but much smaller pearl stingray, which has often been confused with this species. The daisy stingray feeds mainly on crustaceans and exhibits aplacental viviparity, with litters of 1–4 young. Heavily pressured by fisheries and possibly habitat degradation, this once-common species is declining and has been assessed as Vulnerable by the International Union for Conservation of Nature (IUCN).
Batoidea is a superorder of cartilaginous fishes, commonly known as rays. They and their close relatives, the sharks, compose the subclass Elasmobranchii. Rays are the largest group of cartilaginous fishes, with well over 600 species in 26 families. Rays are distinguished by their flattened bodies, enlarged pectoral fins that are fused to the head, and gill slits that are placed on their ventral surfaces.