Reticulate whipray

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Reticulate whipray
Honeycomb stingray at the California Academy of Sciences.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Superorder: Batoidea
Order: Myliobatiformes
Family: Dasyatidae
Genus: Himantura
Species:
H. uarnak
Binomial name
Himantura uarnak
(Forsskål, 1775) or (J. F. Gmelin, 1789)
Himantura uarnak rangemap.png
Range of the reticulate whipray
Synonyms

The reticulate whipray or honeycomb stingray (Himantura uarnak) is a species of stingray in the family Dasyatidae. It inhabits coastal waters in the western Indian Ocean including the Red Sea, Natal and the Arabian Sea; also a Lessepsian transmigrant in the eastern Mediterranean. A large species reaching 2 m (6.6 ft) in width, the reticulate whipray has a diamond-shaped pectoral fin disc and an extremely long tail without fin folds. Both its common and scientific names refer to its ornate dorsal color pattern of many small, close-set dark spots or reticulations on a lighter background. However, the reticulate whipray is only one of several large spotted stingrays in the Indo-Pacific which, coupled with the variability of its coloration with age and locality, has resulted in a great deal of taxonomic confusion.

Contents

Often encountered resting on the bottom during daytime, the reticulate whipray is a predator of bottom-dwelling invertebrates and bony fishes. Like other stingrays, it is aplacental viviparous, with the developing embryos nourished at first by yolk and later by histotroph ("uterine milk"). Females bear litters of up to five pups in summer, following a gestation period of a year. The reticulate whipray is fished in parts of the Indian Ocean for meat, skin, cartilage, and other purposes. It is highly susceptible to population depletion because of its large size, inshore habits, and low reproductive rate, and is additionally threatened by extensive habitat degradation. Consequently, the International Union for Conservation of Nature (IUCN) has assessed this species as Endangered.

Taxonomy and phylogeny

In 1775, Carsten Niebuhr published Descriptiones animalium – avium, amphibiorum, piscium, insectorum, vermium: quæ in itinere orientali observavit, the work of his late friend, the Swedish naturalist Peter Forsskål. Within Forsskål had described Raja sephen, now commonly known as the cowtail stingray, with a spotted variant he called uarnak ورنك, which is the Arabic name stingrays on the Red Sea shores. No type specimen was designated. [2] Forsskål's account formed the basis for two subsequent writings that named the spotted ray as a distinct species: Pierre Joseph Bonnaterre's Raia scherit in 1788, and Johann Friedrich Gmelin's Raja uarnak in 1789. Although Bonnaterre's name was published first and thus would have had precedence, later authorities have regarded Forsskål's uarnak as the earliest available name even though it was not initially part of a binomial. Consequently, modern sources give uarnak as the valid specific epithet and either Gmelin or Forsskål as the species authority. [3] [4] In 1837, Johannes Peter Müller and Friedrich Gustav Jakob Henle included the reticulate whipray in the newly created genus Himantura; David Starr Jordan and Barton Warren Evermann made it the type species in 1896. [5]

The reticulate whipray is closely related to H. undulata , H. leoparda as well as the recently described H. tutul (previously confused with H. leoparda [6] ) and H. australis . All five species are very similar in size and shape, and their colour patterns may overlap to some extent, [7] resulting in a long history of taxonomic confusion that only recently has begun to be untangled. [8] In 2004, Mabel Manjaji designated as the 'uarnak' species complex a group of genetically unrelated species including H. fai , H. gerrardi , H. jenkinsii , H. leoparda, H. toshi , H. uarnak, and H. undulata, plus three more undescribed species. [9] More recently, genera in the stingray family Dasyatidae were redefined based on molecular phylogenetics. Species previously under Himantura now belong to separate genera. [10] The reticulate whipray itself varies in appearance throughout its range, and further taxonomic comparisons are required to determine whether its spotted and reticulated color morphs in fact represent different species. [1] Alternate common names for this ray include coachwhip ray, leopard stingray, longtail stingray, and marbled stingray; some of these names are shared by other, similar species. [3]

Description

The reticulate whipray has a dorsal color pattern of many small dark spots. Himantura uarnak csiro-nfc.jpg
The reticulate whipray has a dorsal color pattern of many small dark spots.

The pectoral fin disc of the reticulate whipray is diamond-shaped and wider than long, with the leading margins almost straight and the snout and outer corners angular. In juveniles, the disc is about as wide as long, with a more obtuse snout and rounded corners. The eyes are small and immediately followed by the spiracles (paired respiratory openings). A short and wide curtain of skin with a minutely fringed rear margin is present between the long, thin nostrils. The mouth is relatively small, with a deep concavity at the center of the lower jaw and shallow furrows at the corners extending onto the lower jaw. [11] A row of 4–5 papillae (nipple-like structures) is found across the floor of the mouth. There are 26–40 upper tooth rows and 27–44 lower tooth rows. [12] The pelvic fins are small and triangular. The tail is whip-like and extremely thin, measuring 3–3.5 times as long as the disc when intact, and lacks fin folds. Usually one serrated stinging spine is located on the upper surface on the tail, some distance from the base. [11]

Adult rays have a wide band of flattened, heart-shaped dermal denticles that extend from between the eyes to the tail spine, increasing in density with age, along with two large pearl thorns at the center of the back. The tail behind the spine is covered by small thorns. The dorsal band of denticles is largely developed by the time the juveniles are 50 cm (20 in) across. The coloration of the reticulate whipray varies substantially with age and locality. Adults generally have a dorsal pattern of numerous closely spaced dark brown spots or reticulations on a beige to yellow-brown background, which becomes blackish past the spine with lighter bands on the sides. The underside is pale, without markings. Juveniles are yellowish above with tiny, densely packed dark spots, around seven spots in a line between the spiracles, and three rows of spots in front of the sting. [11] This large species has been reported to reach a disc width of 2 m (6.6 ft), a total length of 6 m (20 ft), and a weight of 120 kg (260 lb). [3] [12]

Distribution and habitat

The reticulate whipray frequents areas with fine sediment. Himantura uarnak egypt.jpg
The reticulate whipray frequents areas with fine sediment.

The reticulate whipray occurs in the western Indian Ocean, including the Red Sea, Natal in South Africa, and the Arabian Sea. [8] It also entered 30 years ago [13] the Mediterranean Sea through the Suez Canal and it is now common in the south eastern Mediterranean. [14] The recently-described H. australis formerly confused with H. uarnak seems to be common only in Australian waters, where it occurs from Shark Bay to Brisbane. [1] [11]

Bottom-dwelling in nature, the reticulate whipray is generally encountered over sandy flats near beaches, in lagoons, and around coral reefs, from the intertidal zone to offshore waters 50 m (160 ft) deep or more. [1] In Shark Bay, it frequents intertidal sand flats during the warm season, and shifts to slightly deeper seagrass patches in the cold season. [15] This species is tolerant of low salinities and has been known to enter estuaries and mangrove swamps, [16] though records from fresh water in Southeast Asia are unverified and may represent misidentifications. [17] Its preferred water temperatures are 23–26 °C (73–79 °F). [18]

Biology and ecology

Two reticulate whiprays and other fish being fed on a beach near Darwin, Australia. Himantura uarnak darwin2.jpg
Two reticulate whiprays and other fish being fed on a beach near Darwin, Australia.

During the day, the reticulate whipray is generally inactive and spends much time resting motionless on the sea floor, sometimes buried in sand. [16] In Shark Bay, Western Australia, this ray can be found resting singly or in small groups in very shallow water during high tide. Its lateral line extends to the tip of its extremely long tail, giving it advance warning of approaching predators such as bottlenose dolphins (Tursiops aduncus) and hammerhead sharks (Sphyrna). The cowtail stingray (Pastinachus sephen) prefers to rest with the reticulate whipray over others of its own species, because the whiprays' longer tails grant them superior predator detection. These mixed-species groups often settle into a "rosette" with their tails pointing radially outward for maximum predator awareness. [19]

The reticulate whipray preys on a variety of benthic and neritic organisms, including crabs, shrimps, mantis shrimps, bivalves, gastropods, worms, jellyfish, and bony fishes. [3] [20] In the western Indian Ocean, about two-thirds of its diet consists of fishes, in particular ponyfish and anchovies, with shrimps and other crustaceans making up most of the remainder. [21] By contrast, rays in Australian waters are apparently not piscivorous, and are known to consume penaeid prawns. [22] Known parasites of this species include Anthrobothrium loculatum, [23] Dendromonocotyle colorni, [24] Halysiorhynchus macrocephalus, [25] Monocotyle helicophallus, M. multiparous, and M. spiremae, [26] Thaumatocotyle australensis, [27] and Tylocephalum chiralensis. [28]

Like other stingrays, the reticulate whipray is aplacental viviparous: the developing embryos are initially sustained by yolk, which is later supplanted by histotroph ("uterine milk", enriched with proteins and lipids) produced by the mother. [1] Females give birth to up to five pups in the summer, after a year-long gestation period. [20] Off South Africa, the newborns measure 28–30 cm (11–12 in) across and sexual maturation is attained at a disc width of approximately 1 m (3.3 ft), which corresponds to an age of 4-5. [29] Off Australia, the newborns measure 21–28 cm (8.3–11.0 in) across, with males reported to mature at 82–84 cm (32–33 in) across. [11] The juveniles of this species, H. leoparda, and H. undulata differ in birth size, disc shape, denticle development, and amount of spotting, and are in fact more distinct from each other than are adults of the three species. [9] Shark Bay may serve as a nursery area for young rays. [15]

Human interactions

A reticulate whipray at the Palais de la Porte Doree Tropical Aquarium in Paris. Honeycomb stingray (Himantura uarnak).jpg
A reticulate whipray at the Palais de la Porte Dorée Tropical Aquarium in Paris.

The reticulate whipray fights strongly on hook-and-line and is thus popular with recreational anglers, who usually release it alive. [29] This species is caught by intensive artisanal and commercial fisheries operating in parts of the western Indian Ocean, using bottom trawls, gillnets and tangle nets, beach seines, and longlines. [1] The meat, skin, and cartilage are utilized, though this species is not a highly valued food fish. It also has applications in Chinese medicine, and its tail may be sold as a curio. [3]

The International Union for Conservation of Nature (IUCN) has listed the reticulate whipray as Endangered. Its large size, inshore habitat preferences, and slow reproductive rate render it susceptible to overfishing. Although specific data is lacking, significant declines in overall ray catches have been documented within its range. Habitat degradation also threatens this species, while pollution and destructive fishing practices may have also taken their toll.

The reticulate whipray is occasionally offered within the home aquarium trade. It is best avoided, however, because of its massive proportions. [30] It is also kept in some public aquariums such as the Aquarium of the Pacific (where it is one of the aquarium's largest inhabitants), [31] the Atlantis Dubai hotel aquarium, and the California Academy of Sciences' Steinhart Aquarium. [32]

Related Research Articles

<span class="mw-page-title-main">Short-tail stingray</span> Species of cartilaginous fish

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.

<span class="mw-page-title-main">Bluespotted ribbontail ray</span> Species of cartilaginous fish

The bluespotted ribbontail ray is a species of stingray in the family Dasyatidae. Found from the intertidal zone to a depth of 30 m (100 ft), this species is common throughout the tropical Indian and western Pacific Oceans in nearshore, coral reef-associated habitats. It is a fairly small ray, not exceeding 35 cm (14 in) in width, with a mostly smooth, oval pectoral fin disc, large protruding eyes, and a relatively short and thick tail with a deep fin fold underneath. It can be easily identified by its striking color pattern of many electric blue spots on a yellowish background, with a pair of blue stripes on the tail.

<span class="mw-page-title-main">Giant freshwater stingray</span> Species of cartilaginous fish

The giant freshwater stingray is a species of stingray in the family Dasyatidae. It is found in large rivers and estuaries in Southeast Asia and Borneo, though historically it may have been more widely distributed in South and Southeast Asia. The largest freshwater fish and the largest stingray in the world, this species grows up to 2.2 m (7.2 ft) across and can reach up to 300 kg (660 lb) in weight. It has a relatively thin, oval pectoral fin disc that is widest anteriorly, and a sharply pointed snout with a protruding tip. Its tail is thin and whip-like, and lacks fin folds. This species is uniformly grayish brown above and white below; the underside of the pectoral and pelvic fins bear distinctive wide, dark bands on their posterior margins.

<i>Himantura</i> Genus of cartilaginous fishes

Himantura is a genus of stingray in the family Dasyatidae that is native to the Indo-Pacific. In a 2016 taxonomic revision, many of the species formerly assigned to Himantura were reassigned to other genera.

<i>Himantura fluviatilis</i> Species of cartilaginous fish

Himantura fluviatilis, the Ganges whip ray, is an obscure species of stingray in the family Dasyatidae, apparently endemic to the Ganges River system and adjacent marine waters. With an oval pectoral fin disc and long projecting snout, it closely resembles, and may be the same species as Himantura chaophraya. Growing to 1.4 m (4.6 ft) across, it is a plain dark colour above, and lighter below with wide dark bands on the lateral disc margins. The International Union for Conservation of Nature (IUCN) has listed this species as Endangered, as it faces overfishing and habitat degradation within its heavily populated range.

<span class="mw-page-title-main">Marbled whipray</span> Species of cartilaginous fish

The marbled whipray is a little-known species of stingray in the family Dasyatidae, native to several freshwater rivers in Southeast Asia. This species has an oval pectoral fin disc with an elongated, pointed snout and a very long, whip-like tail without fin folds. It is characterized by numerous heart-shaped dermal denticles and tubercles on its upper surface, as well as a reticulated pattern of brown blotches on a light background. The maximum recorded disc width is 36 cm (14 in). The marbled whipray has been assessed as Endangered by the International Union for Conservation of Nature (IUCN); it is heavily threatened by fishing and habitat loss, degradation, and fragmentation.

<span class="mw-page-title-main">White-edge freshwater whipray</span> Species of cartilaginous fish

The white-edge freshwater whipray is an extremely rare species of stingray in the family Dasyatidae, native to four river systems in Southeast Asia. Measuring up to 60 cm (24 in) across, this ray has an oval pectoral fin disc and a very long, whip-like tail without fin folds. It can be identified by the presence of a sharply delineated white band running around the margin of its otherwise brown disc, as well as by its white tail and a band of dermal denticles along the middle of its back. This species feeds on benthic invertebrates and is aplacental viviparous. Its two long tail spines are potentially dangerous to humans. The International Union for Conservation of Nature (IUCN) has assessed the white-edge freshwater whipray as Endangered, as it is under heavy pressure from fishing and habitat loss, degradation, and fragmentation.

<span class="mw-page-title-main">Cowtail stingray</span> Species of cartilaginous fish

The cowtail stingray is a species of stingray in the family Dasyatidae, widespread in the Indo-Pacific region and occasionally entering freshwater habitats. Other common names include banana-tail ray, drab stingray, fantail ray, feathertail stingray, and frill tailed sting ray. This species is sometimes placed in the genus Dasyatis or Hypolophus. The most distinctive characteristic of the cowtail stingray is the large, flag-like ventral fold on its tail, which is especially prominent when the ray is swimming. This species is targeted by commercial fisheries as a source of high-quality shagreen, a type of leather, and its populations are now under threat from heavy exploitation.

<i>Pastinachus</i> Genus of cartilaginous fishes

Pastinachus is a genus of stingrays in the family Dasyatidae. This genus was long held to be monotypic, containing only the cowtail stingray. Recent studies of Indo-Pacific stingrays have revealed that there are in fact a number of different species of Pastinachus, and that some former junior synonyms may need to be resurrected.

<span class="mw-page-title-main">Mangrove whipray</span> Species of cartilaginous fish

The mangrove whipray or whitetail stingray, is a species of stingray in the family Dasyatidae. It is widely distributed in the Indo-Pacific region from the Red Sea to northern Australia and Micronesia. A benthic inhabitant of shallow inshore waters, juvenile mangrove whiprays favor mangrove and estuarine habitats, while adults favor sandy to rocky areas in lagoons and coral reefs. This species can be identified by its thick, oval pectoral fin disc that is dark gray above with numerous white flecks, and by its relatively short, whip-like tail that is white past the stinging spine. It grows up to 1.4 m (4.6 ft) across.

<span class="mw-page-title-main">Pink whipray</span> Species of stingray in the family Dasyatidae

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<span class="mw-page-title-main">Jenkins' whipray</span> Species of cartilaginous fish

The Jenkins' whipray is a species of stingray in the family Dasyatidae, with a wide distribution in the Indo-Pacific region from South Africa to the Malay Archipelago to northern Australia. This large species grows to 1.5 m (4.9 ft) across and has a broad, diamond-shaped pectoral fin disc and a whip-like tail without fin folds. It has a band of heart-shaped dermal denticles running from between the eyes to the tail on its upper surface, along with a characteristic row of large spear-like thorns along the midline. It is uniform yellowish brown above, becoming grayish on the tail past the stinging spine, and white below; there is apparently a spotted color variant that had previously been described as a different species, the dragon stingray.

<span class="mw-page-title-main">Freshwater whipray</span> Species of fish

The freshwater whipray is a little-known species of stingray in the family Dasyatidae, found in a number of large rivers and associated estuaries in northern Australia. Until recently, this species was regarded as a regional subpopulation of the similar-looking but much larger giant freshwater stingray of Southeast Asia. Typically reaching 1 m (3.3 ft) across, the freshwater whipray has a distinctively shaped, rounded pectoral fin disc, a projecting snout, and a thin tail without fin folds. It is plain brown above and white below with dark marginal bands. It may occasionally travel onto land and can "breathe" out of water for up to 7 minutes. The freshwater whipray is an active hunter of small fishes and shrimps, and does not currently face substantial conservation threats.

The tubemouth whipray is a little-known species of stingray in the family Dasyatidae, named for its distinctive, highly protrusible jaws. It is found in shallow, brackish water near mangrove forests and large river mouths along the coasts of southwestern Borneo and southern Sumatra. Measuring up to 1 m (3.3 ft) across, this species has a diamond-shaped pectoral fin disc with an elongated, pointed snout and broadly rounded outer corners. The upper surface of the disc is a plain grayish or brownish in color, and covered by small, flattened dermal denticles. The tubemouth whipray is relatively common at present, but is heavily pressured by habitat degradation and coastal fisheries. The International Union for Conservation of Nature (IUCN) has listed it as Vulnerable.

The Hortle's whipray is a little-known species of stingray in the family Dasyatidae, occurring in shallow estuaries and mud flats off southern New Guinea. This species, growing to 71 cm (28 in) across, has a heart-shaped pectoral fin disc with a long, pointed snout and minute eyes. It has a wide dorsal band of dermal denticles extending from in front of the eyes to the tail, as well as scattered sharp denticles on the snout. The underside of the disc is a distinctive bright yellow in color, sometimes with darker markings around the nostrils, mouth, and gill slits. The Hortle's whipray is threatened by extensive seine fisheries and habitat degradation, leading the International Union for Conservation of Nature (IUCN) to assess it as Near Threatened.

<span class="mw-page-title-main">Black-spotted whipray</span> Species of fish

The black-spotted whipray is a species of stingray in the family Dasyatidae, found in the coastal waters off southern New Guinea and northern Australia. Long thought to be a variant of the related brown whipray, this species has an angular, diamond-shaped pectoral fin disc and a whip-like tail without fin folds. It is characterized by its dorsal color pattern, which consists of a variably extensive covering of small, close-set dark, and sometimes also white spots, on a grayish-brown background. In addition, the tail has alternating light and dark saddles past the stinging spine. This species reaches a maximum recorded width of 80 cm (31 in).

<span class="mw-page-title-main">Brown whipray</span> Species of cartilaginous fish

The brown whipray is a species of stingray in the family Dasyatidae, common in inshore, muddy habitats along the northern coast of Australia. It has often been confused in literature for the honeycomb stingray and the black-spotted whipray, which until recently was thought to be the same species. This species has an angular, diamond-shaped pectoral fin disc and a long, very thin tail without fin folds. It is plain brown above, sometimes with white dots or flecks near the edge of the disc, and white below; the tail is dark all over, with alternating dark and light bands near the tip. The maximum recorded disc width is 74 cm (29 in).

<span class="mw-page-title-main">Leopard whipray</span> Species of cartilaginous fish

The leopard whipray is a little-known species of stingray in the family Dasyatidae, found in the Indian and Pacific Oceans from the Andaman Sea to the Coral Triangle. It is found close to shore at depths shallower than 70 m (230 ft), over soft substrates. Attaining a width of 1.8 m (5.9 ft), this species has a diamond-shaped pectoral fin disc with a pointed snout and an extremely long, whip-like tail without fin folds. Adult rays have a leopard-like dorsal pattern of dark brown rings on a yellowish brown background, as well as a row of enlarged, heart-shaped dermal denticles along the midline of the disc. Newborns and small juveniles have large, solid dark spots and few denticles. The leopard whipray is caught by fisheries in many parts of its range, primarily for meat.

<span class="mw-page-title-main">Honeycomb whipray</span> Species of cartilaginous fish

The honeycomb whipray is a species of stingray in the family Dasyatidae, found widely in the shallow coastal waters of the Indo-Pacific from India to the Malay Archipelago. This large species grows to 1.3 m (4.3 ft) across and has a diamond-shaped disc with rounded corners and a projecting, pointed snout. Its tail is long and whip-like, without fin folds. Adults have a striking dorsal color pattern consisting of large, dark brown rings and reticulations delineated by thin yellow lines, while juveniles have a pattern of large dark spots. This ray can also be distinguished from its similar relatives by an enlarged, pearl-like dermal denticle at the center of the back, which is followed by a few thorns. The International Union for Conservation of Nature (IUCN) has assessed the honeycomb whipray as endangered, as it faces heavy fishing pressure and habitat degradation across much of its range.

References

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  14. ↑ Atlas of Exotic Fishes in the Mediterranean Sea (Himantura uarnak). 2nd Edition. 2021. 366p. CIESM Publishers, Paris, Monaco.https://ciesm.org/atlas/fishes_2nd_edition/Himantura_uarnak.pdf
  15. 1 2 Vaudo, J.J. & M.R. Heithaus (2009). "Spatiotemporal variability in a sandflat elasmobranch fauna in Shark Bay, Australia". Marine Biology. 156 (12): 2579–2590. doi:10.1007/s00227-009-1282-2. S2CID   84230010.
  16. 1 2 Ferrari, A. & A. Ferrari (2002). Sharks . Firefly Books. p.  220. ISBN   978-1-55209-629-1.
  17. ↑ Compagno, L.J.V. & T.R. Roberts (1982). "Freshwater stingrays (Dasyatidae) of Southeast Asia and New Guinea, with description of a new species of Himantura and reports of unidentified species". Environmental Biology of Fishes. 7 (4): 321–339. doi:10.1007/BF00005567. S2CID   19692108.
  18. ↑ Last, P.R. & Compagno, L.J.V. (1999). "Myliobatiformes: Dasyatidae". In Carpenter, K.E. & Niem, V.H. (eds.). FAO identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Rome: Food and Agricultural Organization of the United Nations. pp. 1479–1505. ISBN   978-92-5-104302-8.
  19. ↑ Semeniuk, C.A.D. & L.M. Dill (2006). "Anti-Predator Benefits of Mixed-Species Groups of Cowtail Stingrays (Pastinachus sephen) and Whiprays (Himantura uarnak) at Rest". Ethology. 112: 33–43. doi:10.1111/j.1439-0310.2006.01108.x. S2CID   17529132.
  20. 1 2 Michael, S.W. (1993). Reef Sharks & Rays of the World. Sea Challengers. p. 87. ISBN   978-0-930118-18-1.
  21. ↑ Randall, J.E. & J.P. Hoover (1995). Coastal fishes of Oman. University of Hawaii Press. p. 46. ISBN   978-0-8248-1808-1.
  22. ↑ Salini, J.P.; S.J.M. Blaber & D.T. Brewer (Oct 1990). "Diets of piscivorous fishes in a tropical Australian estuary, with special reference to predation on penaeid prawns". Marine Biology. 105 (3): 363–374. doi:10.1007/BF01316307. S2CID   84232604.
  23. ↑ Vijayalakshmi, C. & S. Sarada (June 1993). "Studies on the new species Anthrobothrium loculatum parasite from Dasyatis (Himantura) uarnak (Forskal)". Boletín Chileno de Parasitología. 48 (1–2): 12–16. PMID   8110367.
  24. ↑ Chisholm, L.A.; I.D. Whittington & G.C. Kearn (2001). "Dendromonocotyle colorni sp. n. (Monogenea: Monocotylidae) from the skin of Himantura uarnak (Dasyatididae) from Israel and a new host record for D. octodiscus from the Bahamas". Folia Parasitologica. 48 (1): 15–20. doi: 10.14411/fp.2001.004 . PMID   11266131.
  25. ↑ Beveridge, I. & R.A. Campbell (June 1992). "Redescription of Halysiorhynchus macrocephalus (Cestoda: Trypanorhyncha), a genus newly recorded from the Australasian region". Systematic Parasitology. 22 (2): 151–157. doi:10.1007/BF00009607. S2CID   34400718.
  26. ↑ Measures, L.N.; M. Beverley-Burton & A. Williams (October 1990). "Three new species of Monocotyle (Monogenea: Monocotylidae) from the stingray, Himantura uarnak (Rajiformes: Dasyatidae) from the Great Barrier Reef: phylogenetic reconstruction, systematics and emended diagnoses". International Journal for Parasitology. 20 (6): 755–767. doi:10.1016/0020-7519(90)90009-C. PMID   2242960.
  27. ↑ Beverley-Burton, M. & A. Williams (1989). "Merizocotyle icopae, Sp-Nov, and Thaumatocotyle australensis, Sp-Nov, (Monogenea, Monocotylidae) From the Nasal Cavities of Rajiform Elasmobranchs of the Great-Barrier-Reef". Australian Journal of Zoology. 37 (1): 25–35. doi:10.1071/ZO9890025.
  28. ↑ Vijayalakshmi, C. & S. Sarada (July–December 1995). "Studies on the new species Tylocephalum chiralensis, parasite from Dasyatis (Himantura) uarnak (Förskal) from Chirala coast, Andhra Pradesh, India". Boletín Chileno de Parasitología. 50 (3–4): 73–75. PMID   8762670.
  29. 1 2 Heemstra, E. (2004). Coastal fishes of Southern Africa. NISC and SAIAB. p. 83. ISBN   978-1-920033-01-9.
  30. ↑ (2014): Elasmobranch Enthusiasts (Part 1): Modern Husbandry – Space | saltwatersmarts.com. Archived 2019-03-23 at the Wayback Machine . In: Saltwater Smarts.
  31. ↑
    Online Learning Center | Reticulate Whipray - Aquarium of the Pacific . In: Aquarium of the Pacific
  32. ↑
    Reef Lagoon Field Guide | California Academy of Sciences . In: California Academy of Sciences in San Francisco, CA

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