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Porcupine ray

The porcupine ray (Urogymnus asperrimus) is a rare species of stingray in the family Dasyatidae. This bottom-dweller is found throughout the tropical Indo-Pacific, as well as off West Africa. It favors sand, coral rubble, and seagrass habitats in inshore waters to a depth of 30 m (100 ft). A large and heavy-bodied species reaching 1.2–1.5 m (3.9–4.9 ft) in width, the porcupine ray has a nearly circular, plain-colored pectoral fin disc and a thin tail without any fin folds. Uniquely within its family, it lacks a venomous stinging spine. However, an adult ray can still defend itself ably with the many large, sharp thorns found over its disc and tail.

Porcupine ray
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Superorder: Batoidea
Order: Myliobatiformes
Family: Dasyatidae
Genus: Urogymnus
Species:
U. asperrimus
Binomial name
Urogymnus asperrimus
Range of the porcupine ray[2]
Synonyms
  • Raja africana Bloch & Schneider, 1801
  • Raja asperrima Bloch & Schneider, 1801
  • Urogymnus asperrimus solanderi Whitley, 1939
  • Urogymnus rhombeus Klunzinger, 1871

The diet of the porcupine ray consists mainly of benthic invertebrates and bony fishes, which it digs up from the sea floor. It is aplacental viviparous, in which the developing embryos are nourished by histotroph ("uterine milk") produced by the mother. The porcupine ray has long been valued for its rough and durable skin, which was made into a shagreen leather once used for various utilitarian and ornamental purposes, such as to cover sword hilts and shields. It is caught incidentally by coastal fisheries. Because it must be handled carefully due to its thorns, its commercial significance is limited. Unregulated fishing has led to this species declining in many parts of its range, thus it has been listed as Vulnerable by the International Union for Conservation of Nature.

Taxonomy edit

 
Early pictures of a porcupine ray by Nelson Annandale

German naturalists Marcus Elieser Bloch and Johann Gottlob Schneider described the porcupine ray in their 1801 work Systema Ichthyologiae, based on a partial dried skin obtained from Mumbai, India. They placed it in the genus Raja and named it asperrima, meaning "roughest" in Latin. In the same work, they also described a West African form, Raja africana.[3] Later authors have regarded the two as synonymous. However, since the two names were published simultaneously, uncertainty exists about which name has priority. Some works give the ray's specific epithet as asperrimus, and others as africanus.[4][5]

In 1837, Johannes Peter Müller and Friedrich Gustav Jakob Henle placed the porcupine ray in a new genus, Gymnura. As the name Gymnura was already in use, referring to the butterfly rays, so later that year, Müller and Henle replaced it with Urogymnus.[6] Both names are derived from the Ancient Greek oura ("tail") and gymnos ("naked" or "unarmed"), referring to the lack of a tail sting.[7] Urogymnus has traditionally been considered monotypic (only containing U. asperrimus), but several other species were moved to this genus from Himantura in 2016 based on morphology and molecular evidence.[8]

Other common names for this species include black spotted ray, rough-skinned ray, roughback stingaree, Solander's ray, and thorny ray.[9] More than one species of porcupine ray may exist, as is currently recognized.[10]

Distribution and habitat edit

 
Camouflaged porcupine ray in Lakshadweep, India

The porcupine ray is widely distributed, but uncommon compared to other stingrays that share its range. It is found all along the continental periphery of the Indian Ocean, from South Africa to the Arabian Peninsula to Southeast Asia to Ningaloo Reef off western Australia, including Madagascar, the Seychelles, and Sri Lanka; it has colonized the eastern Mediterranean Sea through the Suez Canal. In the Pacific Ocean, its range continues through Indonesia and New Guinea, north to the Philippines, east to the Gilbert Islands and Fiji, and south to Heron Island off eastern Australia.[1][2] This species is also found in the eastern Atlantic Ocean off Senegal, Guinea, and Côte d'Ivoire.[9] Bottom-dwelling in nature, the porcupine ray is found close to shore at depths of 1–30 m (3–100 ft). It inhabits sandy flats, coral rubble, and seagrass beds, often near reefs, and also enters brackish water.[11][12]

Description edit

 
Distinctive traits of the porcupine ray include its thick, rounded shape, and dorsal covering of sharp thorns.

The pectoral fin disc of the porcupine ray is evenly oval, almost as wide as long, and very thick at the center, giving it a domed appearance. The tip of the snout is rounded and barely protruding. The small eyes are closely followed by much larger spiracles. Between the narrow nostrils is a skirt-shaped curtain of skin; the posterior margin of the curtain is strongly fringed and overhangs the mouth. The mouth bears three to five papillae on the floor and prominent furrows at the corners. The area around the mouth, including the curtain, are heavily covered by more papillae.[2][13] Around 48 tooth rows are found in either jaw.[10] The teeth are small and flattened. Five pairs of gill slits are located on the underside of the disc.[14]

The pelvic fins are small and narrow. The thin, rapidly tapering tail is about equal to the disc in length and has an almost cylindrical cross-section, without fin folds. The tail also has no venomous stinging spine, unlike other members of the family. A dense patch of flattened, heart-shaped dermal denticles covers the center of the disc and extends onto the tail. Larger individuals additionally have numerous tall, sharp thorns over the entire upper surface of the disc. The porcupine ray is plain light to dark gray or brown above, darkening to blackish towards the tail tip, and white below.[9][13][14] This large species grows to at least 1.2 m (3.9 ft) across and 2.2 m (7.2 ft) long, and may reach 1.5 m (4.9 ft) across.[2][14]

Biology and ecology edit

 
Porcupine ray digging itself into the sand of the lagoon, eating what it finds

The porcupine ray can sometimes be observed lying still on the bottom in the open or inside caves.[5] It is known to form groups at Ningaloo Reef.[2] Its diet consists primarily of sipunculids, polychaete worms, crustaceans, and bony fishes.[12][13] When feeding, it plows deeply into the bottom, expelling excess sediment from its spiracles in a plume visible from a long distance away.[2] Parasites documented from this ray include the tapeworm Rhinebothrium devaneyi, the nematode Echinocephalus overstreeti,[15] and the capsalid monogeneans Dendromonocotyle urogymni[16] and Neoentobdella baggioi.[17] The porcupine ray is aplacental viviparous, with the developing embryos sustained to term by histotroph ("uterine milk") secreted by the mother.[9] Mangrove forests serve as important habitat for juvenile rays.[18] Males and females mature sexually at about 90 and 100 cm (35 and 39 in) across, respectively.[2]

Human interactions edit

 
Close-up on the tail of a porcupine ray (Landaa Giraavaru, Maldives)

Despite not having a sting, the porcupine ray is capable of injuring humans with its many sharp thorns.[9] It is reportedly bold and tolerant of being approached closely underwater.[19] The tough, thorny skin of this ray, made into a form of leather called shagreen, had many historical uses. In particular, it was used to cover the hilts of various melee weapons, as its extremely rough texture prevented slippage during battle. For example, the Japanese deemed it the only species whose skin was acceptable for covering sword grips.[20][21] The Malayans used it to cover shields.[22] The skin was also used ornamentally, such as by the Chinese, who dyed it and ground down the thorns to yield a mottled pattern.[21][23] The native inhabitants of Funafuti Atoll used dried portions of the ray's tail as a rasp-like tool.[24]

Presently, the porcupine ray is caught incidentally in trawls, tangle nets, and beach seines. Its skin continues to be highly valued, while the meat and cartilage may also be used. In the Farasan Islands and some other places in the Red Sea, its liver is eaten as a seasonal dish.[9][12] However, the economic importance of this ray is limited by how difficult it is to handle.[14] The multispecies coastal fisheries that catch the porcupine ray are largely unregulated, which seems to have resulted in its dramatic decline or local extinction in the Bay of Bengal, the Gulf of Thailand, and likely elsewhere in its range. Potential additional threats to this species include habitat degradation from coastal development, and depletion of its food supply from overfishing. As a result, the International Union for Conservation of Nature has assessed it as vulnerable.[25]

References edit

  1. ^ a b Chin, A.; Compagno, L.J.V. (2016). "Urogymnus asperrimus". IUCN Red List of Threatened Species. 2016: e.T39413A68648645. doi:10.2305/IUCN.UK.2016-1.RLTS.T39413A68648645.en. Retrieved 15 November 2021.
  2. ^ a b c d e f g Last, P.R.; Stevens, J.D. (2009). Sharks and Rays of Australia (second ed.). Harvard University Press. pp. 461–462. ISBN 978-0-674-03411-2.
  3. ^ Bloch, M.E.; Schneider, J.G. (1801). Systema Ichthyologiae iconibus cx illustratum. Berolini: Sumtibus auctoris impressum et Bibliopolio Sanderiano commissum. p. 367.
  4. ^ asperrima, Raja 2012-03-19 at the Wayback Machine in: Eschmeyer, W.N.; Fricke, R., eds. Catalog of Fishes electronic version (29 March 2011).
  5. ^ a b Randall, J.E.; Allen, G.R.; Steene, R.C. (1997). Fishes of the Great Barrier Reef and Coral Sea. University of Hawaii Press. p. 30. ISBN 978-0-8248-1895-1.
  6. ^ Urogymnus 2012-03-19 at the Wayback Machine in: Eschmeyer, W.N.; Fricke, R., eds. Catalog of Fishes electronic version (29 March 2011).
  7. ^ Hunter, R. (1895). Lloyd's Encyclopædic Dictionary. p. 387.
  8. ^ Last, P.R.; Naylor, G.J.; Manjaji-Matsumoto, B.M. (2016). "A revised classification of the family Dasyatidae (Chondrichthyes: Myliobatiformes) based on new morphological and molecular insights". Zootaxa. 4139 (3): 345–368. doi:10.11646/zootaxa.4139.3.2. PMID 27470808.
  9. ^ a b c d e f Froese, Rainer; Pauly, Daniel (eds.) (2011). "Urogymnus asperrimus" in FishBase. April 2011 version.
  10. ^ a b Smith, J.L.B.; Smith, M.M.; Heemstra, P. (2003). Smiths' Sea Fishes. Struik. p. 141. ISBN 978-1-86872-890-9.
  11. ^ Hennemann, R.M. (2001). Sharks & Rays: Elasmobranch Guide of the World (second ed.). IKAN – Unterwasserarchiv. p. 260. ISBN 978-3-925919-33-6.
  12. ^ a b c Bonfil, R.; Abdallah, M. (2004). FAO species identification guide for fishery purposes: Field identification guide to the sharks and rays of the Red Sea and Gulf of Aden. Food and Agriculture Organization of the United Nations. p. 52. ISBN 978-92-5-105045-3.
  13. ^ a b c Randall, J.E.; Hoover, J.P. (1995). Coastal fishes of Oman. University of Hawaii Press. p. 47. ISBN 978-0-8248-1808-1.
  14. ^ a b c d Last, P.R.; Compagno, L.J.V. (1999). "Myliobatiformes: Dasyatidae". In Carpenter, K.E.; Niem, V.H. (eds.). FAO species identification guide for fishery purposes: The living marine resources of the Western Central Pacific. Food and Agricultural Organization of the United Nations. p. 1497. ISBN 978-92-5-104302-8.
  15. ^ Brooks, D.R.; Deardorff, T.L. (June 1988). "Rhinebothrium devaneyi n. sp. (Eucestoda: Tetraphyllidea) and Echinocephalus overstreeti Deardorff and Ko, 1983 (Nematoda: Gnathostomatidae) in a Thorny Back Ray, Urogymnus asperrimus, from Enewetak Atoll, with Phylogenetic Analysis of Both Species Groups". The Journal of Parasitology. 74 (3): 459–465. doi:10.2307/3282056. JSTOR 3282056. PMID 3379526. S2CID 23529337.
  16. ^ Chisholm, L.; Whittington, I. (June 2009). "Dendromonocotyle urogymni sp. nov (Monogenea, Monocotylidae) from Urogymnus asperrimus (Elasmobranchii, Dasyatidae) off eastern Australia". Acta Parasitologica. 54 (2): 113–118. doi:10.2478/s11686-009-0023-5.
  17. ^ Whittington, I.D.; Kearn, G.C. (March 2009). "Two new species of Neoentobdella (Monogenea: Capsalidae: Entobdellinae) from the skin of Australian stingrays (Dasyatidae)". Folia Parasitologica. 56 (1): 29–35. doi:10.14411/fp.2009.005. PMID 19391329.
  18. ^ White, W.T.; Sommerville, E. (2010). "Elasmobranchs of Tropical Marine Ecosystems". In Carrier, J.C.; Musick, J.A.; Heithaus, M.R. (eds.). Sharks and Their Relatives 2. CRC Press. pp. 159–240. ISBN 978-1-4200-8047-6.
  19. ^ Michael, S.W. (1993). Reef Sharks & Rays of the World. Sea Challengers. p. 83. ISBN 978-0-930118-18-1.
  20. ^ Stone, G.C.; LaRocca, D.J. (1999). A Glossary of the Construction, Decoration and Use of Arms and Armor in All Countries and in All Times. Courier Dover Publications. p. 537. ISBN 978-0-486-40726-5.
  21. ^ a b Carpenter, G.H.; Praeger, R.L., eds. (1896). The Irish Naturalist. Eason & Son. p. 167. ISBN 978-0-00-715097-7.
  22. ^ Sterndale, R.A.; Aitken, E.H., eds. (1887). Journal of the Bombay Natural History Society, Volume 2. Bombay Natural History Society. p. 154.
  23. ^ Kingsley, J.S. (1888). The Riverside Natural History. Kegan Paul, Trench. p. 89.
  24. ^ Waite, E.R. (1900). "The Mammals, Reptiles, and Fishes of Funafuti". Memoir III – Australian Museum, Sydney – The Atolls of Funafuti, Ellice Group: Its Zoology, Botany, Ethnology, and General Structure. The Trustees. pp. 165–202.
  25. ^ Fowler, S.L.; Cavanagh, R.D.; Camhi, M.; Burgess, G.H.; Cailliet, G.M.; Fordham, S.V.; Simpfendorfer, C.A.; Musick, J.A. (2005). Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes. International Union for Conservation of Nature and Natural Resources. pp. 352–353. ISBN 978-2-8317-0700-6.

External links edit

  • Photos of Porcupine ray on Sealife Collection
  • Porcupine Ray @ Fishes of Australia

porcupine, confused, with, porcupine, river, stingray, porcupine, urogymnus, asperrimus, rare, species, stingray, family, dasyatidae, this, bottom, dweller, found, throughout, tropical, indo, pacific, well, west, africa, favors, sand, coral, rubble, seagrass, . Not to be confused with porcupine river stingray The porcupine ray Urogymnus asperrimus is a rare species of stingray in the family Dasyatidae This bottom dweller is found throughout the tropical Indo Pacific as well as off West Africa It favors sand coral rubble and seagrass habitats in inshore waters to a depth of 30 m 100 ft A large and heavy bodied species reaching 1 2 1 5 m 3 9 4 9 ft in width the porcupine ray has a nearly circular plain colored pectoral fin disc and a thin tail without any fin folds Uniquely within its family it lacks a venomous stinging spine However an adult ray can still defend itself ably with the many large sharp thorns found over its disc and tail Porcupine rayConservation statusVulnerable IUCN 3 1 1 Scientific classificationDomain EukaryotaKingdom AnimaliaPhylum ChordataClass ChondrichthyesSubclass ElasmobranchiiSuperorder BatoideaOrder MyliobatiformesFamily DasyatidaeGenus UrogymnusSpecies U asperrimusBinomial nameUrogymnus asperrimus Bloch amp J G Schneider 1801 Range of the porcupine ray 2 SynonymsRaja africana Bloch amp Schneider 1801 Raja asperrima Bloch amp Schneider 1801 Urogymnus asperrimus solanderi Whitley 1939 Urogymnus rhombeus Klunzinger 1871The diet of the porcupine ray consists mainly of benthic invertebrates and bony fishes which it digs up from the sea floor It is aplacental viviparous in which the developing embryos are nourished by histotroph uterine milk produced by the mother The porcupine ray has long been valued for its rough and durable skin which was made into a shagreen leather once used for various utilitarian and ornamental purposes such as to cover sword hilts and shields It is caught incidentally by coastal fisheries Because it must be handled carefully due to its thorns its commercial significance is limited Unregulated fishing has led to this species declining in many parts of its range thus it has been listed as Vulnerable by the International Union for Conservation of Nature Contents 1 Taxonomy 2 Distribution and habitat 3 Description 4 Biology and ecology 5 Human interactions 6 References 7 External linksTaxonomy edit nbsp Early pictures of a porcupine ray by Nelson AnnandaleGerman naturalists Marcus Elieser Bloch and Johann Gottlob Schneider described the porcupine ray in their 1801 work Systema Ichthyologiae based on a partial dried skin obtained from Mumbai India They placed it in the genus Raja and named it asperrima meaning roughest in Latin In the same work they also described a West African form Raja africana 3 Later authors have regarded the two as synonymous However since the two names were published simultaneously uncertainty exists about which name has priority Some works give the ray s specific epithet as asperrimus and others as africanus 4 5 In 1837 Johannes Peter Muller and Friedrich Gustav Jakob Henle placed the porcupine ray in a new genus Gymnura As the name Gymnura was already in use referring to the butterfly rays so later that year Muller and Henle replaced it with Urogymnus 6 Both names are derived from the Ancient Greek oura tail and gymnos naked or unarmed referring to the lack of a tail sting 7 Urogymnus has traditionally been considered monotypic only containing U asperrimus but several other species were moved to this genus from Himantura in 2016 based on morphology and molecular evidence 8 Other common names for this species include black spotted ray rough skinned ray roughback stingaree Solander s ray and thorny ray 9 More than one species of porcupine ray may exist as is currently recognized 10 Distribution and habitat edit nbsp Camouflaged porcupine ray in Lakshadweep IndiaThe porcupine ray is widely distributed but uncommon compared to other stingrays that share its range It is found all along the continental periphery of the Indian Ocean from South Africa to the Arabian Peninsula to Southeast Asia to Ningaloo Reef off western Australia including Madagascar the Seychelles and Sri Lanka it has colonized the eastern Mediterranean Sea through the Suez Canal In the Pacific Ocean its range continues through Indonesia and New Guinea north to the Philippines east to the Gilbert Islands and Fiji and south to Heron Island off eastern Australia 1 2 This species is also found in the eastern Atlantic Ocean off Senegal Guinea and Cote d Ivoire 9 Bottom dwelling in nature the porcupine ray is found close to shore at depths of 1 30 m 3 100 ft It inhabits sandy flats coral rubble and seagrass beds often near reefs and also enters brackish water 11 12 Description edit nbsp Distinctive traits of the porcupine ray include its thick rounded shape and dorsal covering of sharp thorns The pectoral fin disc of the porcupine ray is evenly oval almost as wide as long and very thick at the center giving it a domed appearance The tip of the snout is rounded and barely protruding The small eyes are closely followed by much larger spiracles Between the narrow nostrils is a skirt shaped curtain of skin the posterior margin of the curtain is strongly fringed and overhangs the mouth The mouth bears three to five papillae on the floor and prominent furrows at the corners The area around the mouth including the curtain are heavily covered by more papillae 2 13 Around 48 tooth rows are found in either jaw 10 The teeth are small and flattened Five pairs of gill slits are located on the underside of the disc 14 The pelvic fins are small and narrow The thin rapidly tapering tail is about equal to the disc in length and has an almost cylindrical cross section without fin folds The tail also has no venomous stinging spine unlike other members of the family A dense patch of flattened heart shaped dermal denticles covers the center of the disc and extends onto the tail Larger individuals additionally have numerous tall sharp thorns over the entire upper surface of the disc The porcupine ray is plain light to dark gray or brown above darkening to blackish towards the tail tip and white below 9 13 14 This large species grows to at least 1 2 m 3 9 ft across and 2 2 m 7 2 ft long and may reach 1 5 m 4 9 ft across 2 14 Biology and ecology edit nbsp Porcupine ray digging itself into the sand of the lagoon eating what it findsThe porcupine ray can sometimes be observed lying still on the bottom in the open or inside caves 5 It is known to form groups at Ningaloo Reef 2 Its diet consists primarily of sipunculids polychaete worms crustaceans and bony fishes 12 13 When feeding it plows deeply into the bottom expelling excess sediment from its spiracles in a plume visible from a long distance away 2 Parasites documented from this ray include the tapeworm Rhinebothrium devaneyi the nematode Echinocephalus overstreeti 15 and the capsalid monogeneans Dendromonocotyle urogymni 16 and Neoentobdella baggioi 17 The porcupine ray is aplacental viviparous with the developing embryos sustained to term by histotroph uterine milk secreted by the mother 9 Mangrove forests serve as important habitat for juvenile rays 18 Males and females mature sexually at about 90 and 100 cm 35 and 39 in across respectively 2 Human interactions edit nbsp Close up on the tail of a porcupine ray Landaa Giraavaru Maldives Despite not having a sting the porcupine ray is capable of injuring humans with its many sharp thorns 9 It is reportedly bold and tolerant of being approached closely underwater 19 The tough thorny skin of this ray made into a form of leather called shagreen had many historical uses In particular it was used to cover the hilts of various melee weapons as its extremely rough texture prevented slippage during battle For example the Japanese deemed it the only species whose skin was acceptable for covering sword grips 20 21 The Malayans used it to cover shields 22 The skin was also used ornamentally such as by the Chinese who dyed it and ground down the thorns to yield a mottled pattern 21 23 The native inhabitants of Funafuti Atoll used dried portions of the ray s tail as a rasp like tool 24 Presently the porcupine ray is caught incidentally in trawls tangle nets and beach seines Its skin continues to be highly valued while the meat and cartilage may also be used In the Farasan Islands and some other places in the Red Sea its liver is eaten as a seasonal dish 9 12 However the economic importance of this ray is limited by how difficult it is to handle 14 The multispecies coastal fisheries that catch the porcupine ray are largely unregulated which seems to have resulted in its dramatic decline or local extinction in the Bay of Bengal the Gulf of Thailand and likely elsewhere in its range Potential additional threats to this species include habitat degradation from coastal development and depletion of its food supply from overfishing As a result the International Union for Conservation of Nature has assessed it as vulnerable 25 References edit a b Chin A Compagno L J V 2016 Urogymnus asperrimus IUCN Red List of Threatened Species 2016 e T39413A68648645 doi 10 2305 IUCN UK 2016 1 RLTS T39413A68648645 en Retrieved 15 November 2021 a b c d e f g Last P R Stevens J D 2009 Sharks and Rays of Australia second ed Harvard University Press pp 461 462 ISBN 978 0 674 03411 2 Bloch M E Schneider J G 1801 Systema Ichthyologiae iconibus cx illustratum Berolini Sumtibus auctoris impressum et Bibliopolio Sanderiano commissum p 367 asperrima Raja Archived 2012 03 19 at the Wayback Machine in Eschmeyer W N Fricke R eds Catalog of Fishes electronic version 29 March 2011 a b Randall J E Allen G R Steene R C 1997 Fishes of the Great Barrier Reef and Coral Sea University of Hawaii Press p 30 ISBN 978 0 8248 1895 1 Urogymnus Archived 2012 03 19 at the Wayback Machine in Eschmeyer W N Fricke R eds Catalog of Fishes electronic version 29 March 2011 Hunter R 1895 Lloyd s Encyclopaedic Dictionary p 387 Last P R Naylor G J Manjaji Matsumoto B M 2016 A revised classification of the family Dasyatidae Chondrichthyes Myliobatiformes based on new morphological and molecular insights Zootaxa 4139 3 345 368 doi 10 11646 zootaxa 4139 3 2 PMID 27470808 a b c d e f Froese Rainer Pauly Daniel eds 2011 Urogymnus asperrimus in FishBase April 2011 version a b Smith J L B Smith M M Heemstra P 2003 Smiths Sea Fishes Struik p 141 ISBN 978 1 86872 890 9 Hennemann R M 2001 Sharks amp Rays Elasmobranch Guide of the World second ed IKAN Unterwasserarchiv p 260 ISBN 978 3 925919 33 6 a b c Bonfil R Abdallah M 2004 FAO species identification guide for fishery purposes Field identification guide to the sharks and rays of the Red Sea and Gulf of Aden Food and Agriculture Organization of the United Nations p 52 ISBN 978 92 5 105045 3 a b c Randall J E Hoover J P 1995 Coastal fishes of Oman University of Hawaii Press p 47 ISBN 978 0 8248 1808 1 a b c d Last P R Compagno L J V 1999 Myliobatiformes Dasyatidae In Carpenter K E Niem V H eds FAO species identification guide for fishery purposes The living marine resources of the Western Central Pacific Food and Agricultural Organization of the United Nations p 1497 ISBN 978 92 5 104302 8 Brooks D R Deardorff T L June 1988 Rhinebothrium devaneyi n sp Eucestoda Tetraphyllidea and Echinocephalus overstreeti Deardorff and Ko 1983 Nematoda Gnathostomatidae in a Thorny Back Ray Urogymnus asperrimus from Enewetak Atoll with Phylogenetic Analysis of Both Species Groups The Journal of Parasitology 74 3 459 465 doi 10 2307 3282056 JSTOR 3282056 PMID 3379526 S2CID 23529337 Chisholm L Whittington I June 2009 Dendromonocotyle urogymni sp nov Monogenea Monocotylidae from Urogymnus asperrimus Elasmobranchii Dasyatidae off eastern Australia Acta Parasitologica 54 2 113 118 doi 10 2478 s11686 009 0023 5 Whittington I D Kearn G C March 2009 Two new species of Neoentobdella Monogenea Capsalidae Entobdellinae from the skin of Australian stingrays Dasyatidae Folia Parasitologica 56 1 29 35 doi 10 14411 fp 2009 005 PMID 19391329 White W T Sommerville E 2010 Elasmobranchs of Tropical Marine Ecosystems In Carrier J C Musick J A Heithaus M R eds Sharks and Their Relatives 2 CRC Press pp 159 240 ISBN 978 1 4200 8047 6 Michael S W 1993 Reef Sharks amp Rays of the World Sea Challengers p 83 ISBN 978 0 930118 18 1 Stone G C LaRocca D J 1999 A Glossary of the Construction Decoration and Use of Arms and Armor in All Countries and in All Times Courier Dover Publications p 537 ISBN 978 0 486 40726 5 a b Carpenter G H Praeger R L eds 1896 The Irish Naturalist Eason amp Son p 167 ISBN 978 0 00 715097 7 Sterndale R A Aitken E H eds 1887 Journal of the Bombay Natural History Society Volume 2 Bombay Natural History Society p 154 Kingsley J S 1888 The Riverside Natural History Kegan Paul Trench p 89 Waite E R 1900 The Mammals Reptiles and Fishes of Funafuti Memoir III Australian Museum Sydney The Atolls of Funafuti Ellice Group Its Zoology Botany Ethnology and General Structure The Trustees pp 165 202 Fowler S L Cavanagh R D Camhi M Burgess G H Cailliet G M Fordham S V Simpfendorfer C A Musick J A 2005 Sharks Rays and Chimaeras The Status of the Chondrichthyan Fishes International Union for Conservation of Nature and Natural Resources pp 352 353 ISBN 978 2 8317 0700 6 External links editPhotos of Porcupine ray on Sealife Collection Porcupine Ray Fishes of Australia Retrieved from https en wikipedia org w index php title Porcupine ray amp oldid 1191122837, wikipedia, wiki, book, books, library,

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