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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Classis: Chondrichthyes
Subclassis: Elasmobranchii
Infraclassis: Euselachii
Division/Cohort: Neoselachii
Subdivision/Subcohort: Batoidea
Superordo: Batomorphii
Ordo: Myliobatiformes
Subordo: Myliobatoidei
Superfamilia: Dasyatoidea

Familia: Dasyatidae
Subfamilia: Urogymninae
Genus: Fluvitrygon
Species: Fluvitrygon signifer
Name

Fluvitrygon signifer (Compagno & Roberts, 1982)

References
Primary references

Leonard J. V. Compagno & Tyson R. Roberts 2008. Environmental Biology of Fishes 7(4): 321–339. [1]

The white-edge freshwater whipray (Fluvitrygon signifer) 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.
Taxonomy and phylogeny

The white-edge freshwater whipray was described by Leonard Compagno and Tyson Roberts in a 1982 issue of Environmental Biology of Fishes, with the specific epithet signifer (Latin for "sign-bearing") in reference to its distinctive coloration. The type specimen is an immature female 29 cm (11 in) across, collected from the mouth of the Sungai Ketungau off the Kapuas River in West Kalimantan, Indonesia.[2] Other common names for this species include freshwater stingray, pale whipray, and white-rimmed stingray.[3] Within its genus, the white-edge freshwater whipray most closely resembles Fluvitrygon kittipongi, described in 2005.[4] A 1999 phylogenetic analysis, based on cytochrome b sequences, found that it is closely related to Maculabatis gerrardi and Brevitrygon imbricata, which form a sister species pair.[5]
Distribution and habitat

One of the few members of its family that seems to exclusively inhabit fresh water, the white-edge freshwater whipray is known from four Southeast Asian river systems: the Kapuas River in Borneo, the Indragiri River in Sumatra, the Perak River in Peninsular Malaysia, and the Chao Phraya River in Thailand.[1] Although these rivers are now isolated from one another, when sea levels were low during the Pleistocene epoch (2.6 million to 12,000 years ago) they may have all been contiguous as part of the Central or North Sundaland drainage basin. At that time, this ray could have dispersed into its present range without having to enter salt water.[2] This bottom-dwelling species favors a sandy habitat.[3]
Description

The white-edge freshwater whipray has a thin and oval pectoral fin disc slightly wider than long. The anterior margins of the disc converge with a broad angle at the tip of the snout, which is slightly protruding. The eyes are small and immediately followed by much larger spiracles. There is a flap of skin between the nares with a fringed, subtly concave or three-lobed posterior margin. The mouth is gently bow-shaped, with a groove running from the nasal flap around its corners. There are 38–45 upper tooth rows and 37–46 lower tooth rows; the teeth are well-spaced and arranged in a quincunx pattern. Each tooth has a blunt conical crown with a transverse cutting edge; this edge is higher and serrated in males and lower and blunt in females. There is a row of 4–5 papillae across the floor of the mouth: a large inner pair, a smaller outer pair, and sometimes a central one of varying size.[2]

The tail is about 3.5 times as long as the disc and bears two stinging spines on top; behind the spines the tail becomes slender and whip-like, without any fin folds.[2] Two recorded individuals had tail spines 86 and 87 mm (3.4 and 3.4 in) long with 70 and 89 serrations respectively.[6] A band of skin running along the back from between the eyes to the tail base is roughened by small, flattened heart-shaped dermal denticles interspersed with small conical denticles. There are more denticles on the tail behind the spines, and small individuals also have a pearl spine in the center of the disc.[2] The disc is brown above, with mottling in the center and on the tail base, a white spot between the eye and spiracle, and a sharp white band running around its periphery. The underside is completely white, as is the tail behind the spines. This ray grows up to 60 cm (24 in) across and perhaps over 2 m (6.6 ft) long.[3][7]
Biology and ecology

Like the other freshwater and euryhaline members of its family, but unlike the specialized South American river stingrays of the family Potamotrygonidae, the white-edge freshwater whipray retains the ability to synthesize urea for osmoregulation and can survive in brackish water with a salinity of 20 ppt for at least two weeks.[8] Indeed, this ray might experience greater salinity-induced oxidative stress in freshwater than in brackish water, possibly related to its short history of freshwater invasion.[9] To this end, it has been suggested that there may be a possible relationship between the successful invasion of the freshwater environment by some euryhaline marine elasmobranchs and the ability of these elasmobranchs to increase the capacity of ascorbic acid synthesis to defend against hyposalinity stress. In addition, the ampullae of Lorenzini of F. signifer are intermediate in size and structure between the large, complex ampullae of its marine relatives and the small, simple ampullae of potamotrygonid stingrays, which may be a consequence of varying salinities in its environment and demands that would place on electroreception.[10] The white-edge freshwater whipray feeds mainly on small benthic organisms, including crustaceans, molluscs, and insect larvae. Reproduction is aplacental viviparous with the developing embryos sustained by maternally produced histotroph ("uterine milk"), as is the case in other stingrays. Newborns measure 10–12 cm (3.9–4.7 in) across; males and females mature sexually at 21–23 cm (8.3–9.1 in) and 25–26 cm (9.8–10.2 in) across respectively.[11]
Human interactions

The venomous stings of the white-edge freshwater whipray are capable of inflicting excruciating and even fatal wounds on humans.[3] This species is extremely rare, with only about 10 specimens having been deposited in museum collections. Most of its range is subject to heavy fishing pressure; this ray is infrequently caught using fish traps, spears, and bottom-set lines; it is sold for meat and the aquarium trade.[1][11] Habitat loss and degradation as a result of pollution, logging and dam construction are likely to pose an even a greater threat to the survival of this species. In Thailand, dams built on the Chao Phraya have fragmented the resident stingray population and effectively reduced its genetic diversity.[7] Because of diverse threats faced by this ray and the lack of exchange between the different rivers it inhabits, the International Union for Conservation of Nature (IUCN) has assessed the white-edge freshwater whipray as Endangered. The Thai government began a captive breeding program for this and other endangered freshwater stingrays at Chai Nat during the 1990s, but by 1996 the program had been placed on hold.[1]
References

Grant, I.; Rigby, C.L.; Bin Ali, A.; Fahmi, Hasan, V.; Sayer, C. (2021). "Fluvitrygon signifer". IUCN Red List of Threatened Species. 2021: e.T39411A2924238. doi:10.2305/IUCN.UK.2021-2.RLTS.T39411A2924238.en. Retrieved 14 November 2021.
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.
Froese, Rainer; Pauly, Daniel (eds.) (2010). "Himantura signifer" in FishBase. January 2010 version.
Vidthayanon, C. & T.R. Roberts (2005). "Himantura kittipongi, a new species of freshwater whiptail stingray from the Maekhlong River of Thailand (Elasmobranchii, Dasyatididae)". Natural History Bulletin of the Siam Society. 53 (1): 123–132.
Sezaki, K.; R.A. Begum; P. Wongrat; M.P. Srivastava; S. SriKantha; K. Kikuchi; H. Ishihara; S. Tanaka; T. Taniuchi & S. Watabe (1999). "Molecular Phylogeny of Asian Freshwater and Marine Stingrays Based on the DNA Nucleotide and Deduced Amino Acid Sequences of the Cytochrome b Gene". Fisheries Science. 65 (4): 563–570. doi:10.2331/fishsci.65.563.
Cuny, G. & C. Piyapong (March 2007). "Tail spine characteristics of stingrays (order Myliobatiformes): a comment to Schwartz (2005)". Electronic Journal of Ichthyology. 3 (1): 15–17.
Fowler, S.L.; R.D. Cavanagh; M. Camhi; G.H. Burgess; G.M. Cailliet; S.V. Fordham; C.A. Simpfendorfer & J.A. Musick (2005). Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes. International Union for Conservation of Nature and Natural Resources. pp. 351–352. ISBN 2-8317-0700-5.
Tam, W.L.; W.P. Wong; A.M. Loong; K.C. Hiong; S.F. Chew; J.S. Ballantyne & Y.K. Ip (September 2003). "The osmotic response of the Asian freshwater stingray (Himantura signifer) to increased salinity: a comparison with marine (Taeniura lymma) and Amazonian freshwater (Potamotrygon motoro) stingrays". Journal of Experimental Biology. 206 (17): 2931–2940. doi:10.1242/jeb.00510. PMID 12878662.
Wong, S.Z.H.; B. Ching; Y.R. Chng; W.P. Wong; S.F. Chew & Y.K. Ip (June 2013). "Ascorbic Acid Biosynthesis and Brackish Water Acclimation in the Euryhaline Freshwater White-Rimmed Stingray, Himantura signifer". PLOS ONE. 8 (6): e66691. Bibcode:2013PLoSO...866691W. doi:10.1371/journal.pone.0066691. PMC 3688944. PMID 23825042.
Raschi, W.; E.D. Keithan & W.C.H. Rhee (February 18, 1997). "Anatomy of the Ampullary Electroreceptor in the Freshwater Stingray, Himantura signifer". Copeia. 1997 (1): 101–107. doi:10.2307/1447844. JSTOR 1447844.
Last, P.R.; W.T. White; J.N. Caire; Dharmadi; Fahmi; K. Jensen; A.P.F. Lim; B. Mabel-Matsumoto; G.J.P. Naylor; J.J. Pogonoski; J.D. Stevens & G.K. Yearsley (2010). Sharks and Rays of Borneo. CSIRO Publishing. pp. 210–211. ISBN 978-1-921605-59-8.

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