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| status2_system = CITES
| status2_system = CITES
| status2_ref = <ref>{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/appendices.php|access-date=2022-01-14|website=cites.org}}</ref>
| status2_ref = <ref>{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/appendices.php|access-date=2022-01-14|website=cites.org}}</ref>
| grandparent_authority = [[Johannes Peter Müller|J. P. Müller]] and [[Friedrich Gustav Jakob Henle|Henle]], 1841
| parent_authority = [[Marcus Elieser Bloch|Bloch]] & [[Johann Gottlob Theaenus Schneider|J. G. Schneider]], 1801
| parent_authority = [[Marcus Elieser Bloch|Bloch]] & [[Johann Gottlob Theaenus Schneider|J. G. Schneider]], 1801
| taxon = Rhina ancylostoma
| taxon = Rhina ancylostoma
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}}
}}


'''''Rhina ancylostoma''''', the '''bowmouth guitarfish''', '''shark ray''' or '''mud skate''', is a [[species]] of [[Batoidea|ray]] and a member of the [[family (biology)|family]] [[Rhinidae]]. Its evolutionary affinities are not fully resolved, though it may be related to [[Rhinobatidae|true guitarfish]]es and [[Skate (fish)|skate]]s.{{citation needed|date=May 2023}} This rare species occurs widely in the [[tropical]] coastal waters of the western [[Indo-Pacific]], at depths of up to {{convert|90|m|ft|abbr=on}}. Highly distinctive in appearance, ''Rhina ancylostoma'' has a wide and thick body with a rounded snout and large shark-like [[dorsal fin|dorsal]] and [[caudal fin|tail fins]]. Its mouth forms a W-shaped undulating line, and there are multiple thorny ridges over its head and back. It has a dorsal color pattern of many white spots over a bluish gray to brown background, with a pair of prominent black markings over the [[pectoral fin]]s. This large species can reach a length of {{convert|2.7|m|ft|abbr=on}} and weight of {{convert|135|kg|lb|abbr=on}}.{{citation needed|date=May 2023}}
'''''Rhina ancylostoma''''', also known as the '''bowmouth guitarfish''', '''shark ray''' or '''mud skate''', is a [[species]] of [[Batoidea|ray]] and a member of the [[family (biology)|family]] [[Rhinidae]]. Its evolutionary affinities are not fully resolved, though it may be related to [[Rhinobatidae|true guitarfish]]es and [[Skate (fish)|skate]]s.{{citation needed|date=May 2023}} This rare species occurs widely in the [[tropical]] coastal waters of the western [[Indo-Pacific]], at depths of up to {{convert|90|m|ft|abbr=on}}. Highly distinctive in appearance, ''Rhina ancylostoma'' has a wide and thick body with a rounded snout and large shark-like [[dorsal fin|dorsal]] and [[caudal fin|tail fins]]. Its mouth forms a W-shaped undulating line, and there are multiple thorny ridges over its head and back. It has a dorsal color pattern of many white spots over a bluish gray to brown background, with a pair of prominent black markings over the [[pectoral fin]]s. This large species can reach a length of {{convert|2.7|m|ft|abbr=on}} and weight of {{convert|135|kg|lb|abbr=on}}.{{citation needed|date=May 2023}}


Usually found near the [[sea floor]], ''Rhina ancylostoma'' prefers sandy or muddy areas near underwater structures. It is a strong-swimming [[predator]] of [[bony fish]]es, [[crustacean]]s, and [[mollusc]]s. This species [[viviparous|gives live birth]] to litters of two to eleven pups, which are nourished during [[gestation]] by [[yolk]]. The [[International Union for Conservation of Nature]] (IUCN) has assessed ''Rhina ancylostoma'' as [[Critically Endangered species|Critically Endangered]] because it is widely caught by [[artisan fishing|artisanal]] and [[commercial fishing|commercial fisheries]] for its valuable fins and meat. It is viewed as a nuisance by [[trawl]]ers, however, because its bulk and thorny skin cause it to damage netted catches. [[Habitat degradation]] and [[habitat destruction|destruction]] pose an additional, significant challenge to this ray's survival. ''Rhina ancylostoma'' adapts well to captivity and is displayed in [[public aquarium]]s.
Usually found near the [[sea floor]], the bowmouth guitarfish prefers sandy or muddy areas near underwater structures. It is a strong-swimming [[predator]] of [[bony fish]]es, [[crustacean]]s, and [[mollusc]]s. This species [[viviparous|gives live birth]] to litters of two to eleven pups, which are nourished during [[gestation]] by [[yolk]]. The [[International Union for Conservation of Nature]] (IUCN) has assessed ''Rhina ancylostoma'' as [[Critically Endangered species|Critically Endangered]] because it is widely caught by [[artisan fishing|artisanal]] and [[commercial fishing|commercial fisheries]] for its valuable fins and meat. It is viewed as a nuisance by [[trawl]]ers, however, because its bulk and thorny skin cause it to damage netted catches. [[Habitat degradation]] and [[habitat destruction|destruction]] pose an additional, significant challenge to this ray's survival. The ''bowmouth guitarfish'' adapts well to captivity and is displayed in [[public aquarium]]s.


==Taxonomy and phylogeny==
==Taxonomy and phylogeny==
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==Description==
==Description==
[[File:Rhina ancylostoma georgia.jpg|thumb|upright=1.2|left|The rounded head, humpbacked profile, and large fins of ''Rhina ancylostoma'' give it a unique appearance.]]
[[File:Rhina ancylostoma georgia.jpg|thumb|left|The rounded head, humpbacked profile, and large fins of ''Rhina ancylostoma'' give it a unique appearance.|220x220px]]
''Rhina ancylostoma'' is a heavily built fish growing to {{convert|2.7|m|ft|abbr=on}} long and {{convert|135|kg|lb|abbr=on}} in weight.<ref name="last and stevens"/><ref name="fishbase"/>
⚫
''Rhina ancylostoma'' is a heavily built fish growing to {{convert|2.7|m|ft|abbr=on}} long and {{convert|135|kg|lb|abbr=on}} in weight.<ref name="last and stevens"/><ref name="fishbase"/> The head is short, wide, and flattened with an evenly rounded snout; the front portion of the head, including the medium-sized eyes and large [[Spiracle (vertebrates)|spiracle]]s, is clearly distinct from the body. The long nostrils are transversely oriented and have well-developed skin flaps on their anterior margins. The lower jaw has three protruding lobes that fit into corresponding depressions in the upper jaw.<ref name="last and stevens"/><ref name="randall and hoover"/> There are around 47 upper and 50 lower tooth rows arranged in winding bands; the teeth are low and blunt with ridges on the crown. The five pairs of ventral [[gill slit]]s are positioned close to the lateral margins of the head.<ref name="last and stevens"/><ref name="smith et al"/>


⚫
The head is short, wide, and flattened with an evenly rounded snout; the front portion of the head, including the medium-sized eyes and large [[Spiracle (vertebrates)|spiracle]]s, is clearly distinct from the body. The long nostrils are transversely oriented and have well-developed skin flaps on their anterior margins.<ref name="randall and hoover" /><ref name="last and stevens" />
⚫
The body is deepest in front of the two tall and falcate (sickle-shaped) [[dorsal fin]]s. The first dorsal fin is about a third larger than the second and originates over the [[pelvic fin]] origins. The second dorsal fin is located midway between the first dorsal and the [[caudal fin]]. The broad and triangular [[pectoral fin]]s have a deep indentation where their leading margins meet the head. The [[pelvic fin]]s are much smaller than the pectoral fins, and the [[anal fin]] is absent. The tail is much longer than the body and ends in a large, crescent-shaped [[caudal fin]]; the lower caudal fin lobe is more than half the length of the upper.<ref name="last and stevens"/><ref name="compagno and last"/><ref name="randall and hoover"/>

⚫
The lower jaw has three protruding lobes that fit into corresponding depressions in the upper jaw.<ref name="last and stevens" /><ref name="randall and hoover" /> There are around 47 upper and 50 lower tooth rows arranged in winding bands; the teeth are low and blunt with ridges on the crown. The five pairs of ventral [[gill slit]]s are positioned close to the lateral margins of the head.<ref name="last and stevens" /><ref name="smith et al" />[[File:Jaw of Rhina ancylostoma.jpg|thumb|Jaws|left]]The body is deepest in front of the two tall and falcate (sickle-shaped) [[dorsal fin]]s. The first dorsal fin is about a third larger than the second and originates over the [[pelvic fin]] origins. The second dorsal fin is located midway between the first dorsal and the [[caudal fin]]. The broad and triangular [[pectoral fin]]s have a deep indentation where their leading margins meet the head. The [[pelvic fin]]s are much smaller than the pectoral fins, and the [[anal fin]] is absent. The tail is much longer than the body and ends in a large, crescent-shaped [[caudal fin]]; the lower caudal fin lobe is more than half the length of the upper.<ref name="last and stevens"/><ref name="compagno and last"/><ref name="randall and hoover"/>


The entire dorsal surface has a grainy texture from a dense covering of tiny [[dermal denticle]]s. A thick ridge is present along the midline of the back, which bears a band of sharp, robust thorns. There are also a pair of thorn-bearing ridges in front of the eyes, a second pair running from above the eyes to behind the spiracles, and a third pair on the "shoulders". This species is bluish to brownish gray above, lightening towards the margins of the head and over the pectoral fins. There are prominent white spots scattered over the body and fins, a white-edged black marking above each pectoral fin, and two dark transverse bands atop the head between the eyes. The underside is light gray to white. Young rays are more vividly colored than adults, which are browner with fainter patterning and proportionately smaller spots.<ref name="last and stevens"/>
The entire dorsal surface has a grainy texture from a dense covering of tiny [[dermal denticle]]s. A thick ridge is present along the midline of the back, which bears a band of sharp, robust thorns. There are also a pair of thorn-bearing ridges in front of the eyes, a second pair running from above the eyes to behind the spiracles, and a third pair on the "shoulders". This species is bluish to brownish gray above, lightening towards the margins of the head and over the pectoral fins. There are prominent white spots scattered over the body and fins, a white-edged black marking above each pectoral fin, and two dark transverse bands atop the head between the eyes. The underside is light gray to white. Young rays are more vividly colored than adults, which are browner with fainter patterning and proportionately smaller spots.<ref name="last and stevens"/>
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While uncommon, ''Rhina ancylostoma'' is widely distributed in the coastal [[tropical]] waters of the western [[Indo-Pacific]]. In the Indian Ocean, it is found from [[KwaZulu-Natal]] in South Africa to the [[Red Sea]] (including the [[Seychelles]]), across the [[Indian subcontinent]] and [[Southeast Asia]] (including the [[Maldives]]), to [[Shark Bay]] in [[Western Australia]]. Its Pacific range extends northward to Korea and southern Japan, eastward to New Guinea, and southward to [[New South Wales]].<ref name="last and stevens"/><ref name="randall and hoover"/> Found between {{convert|3|and|90|m|ft|abbr=on|-1}} deep, this ray spends most of its time near the [[sea floor]] but can occasionally be seen swimming in midwater. It favors sandy or muddy [[habitat]]s, and can also be found in the vicinity of rocky and [[coral reef]]s and [[shipwreck]]s.<ref name="last and stevens"/><ref name="michael"/>
While uncommon, ''Rhina ancylostoma'' is widely distributed in the coastal [[tropical]] waters of the western [[Indo-Pacific]]. In the Indian Ocean, it is found from [[KwaZulu-Natal]] in South Africa to the [[Red Sea]] (including the [[Seychelles]]), across the [[Indian subcontinent]] and [[Southeast Asia]] (including the [[Maldives]]), to [[Shark Bay]] in [[Western Australia]]. Its Pacific range extends northward to Korea and southern Japan, eastward to New Guinea, and southward to [[New South Wales]].<ref name="last and stevens"/><ref name="randall and hoover"/> Found between {{convert|3|and|90|m|ft|abbr=on|-1}} deep, this ray spends most of its time near the [[sea floor]] but can occasionally be seen swimming in midwater. It favors sandy or muddy [[habitat]]s, and can also be found in the vicinity of rocky and [[coral reef]]s and [[shipwreck]]s.<ref name="last and stevens"/><ref name="michael"/>


==Ecology==
==Biology and ecology==
[[File:Tigershark2.jpg|thumb|The tiger shark preys on ''Rhina ancylostoma''.]]
[[File:Tigershark2.jpg|thumb|The tiger shark preys on ''Rhina ancylostoma''.]]
''Rhina ancylostoma'' is a strong swimmer that propels itself with its tail like a shark. It is [[nocturnal|more active at night]] and is not known to be territorial.<ref name="ferrari and ferrari"/> This species feeds mainly on [[demersal fish|demersal]] [[bony fish]]es such as [[Sciaenidae|croakers]] and [[crustacean]]s such as [[crab]]s and [[shrimp]]; [[bivalve]]s and [[cephalopod]]s are also consumed. Its bands of flattened teeth allow it to crush hard-shelled prey.<ref name="compagno and last"/><ref name="raje"/> Curiously, two ''Rhina ancylostoma'' examined in a 2011 [[stable isotope]] study were found to have fed on [[pelagic zone|pelagic]] rather than demersal animals, in contrast to previous observations.<ref name="borrell et al"/>
''Rhina ancylostoma'' is a strong swimmer that propels itself with its tail like a shark. It is [[nocturnal|more active at night]] and is not known to be territorial.<ref name="ferrari and ferrari"/> This species feeds mainly on [[demersal fish|demersal]] [[bony fish]]es such as [[Sciaenidae|croakers]] and [[crustacean]]s such as [[crab]]s and [[shrimp]]; [[bivalve]]s and [[cephalopod]]s are also consumed. Its bands of flattened teeth allow it to crush hard-shelled prey.<ref name="compagno and last"/><ref name="raje"/> Curiously, two ''Rhina ancylostoma'' examined in a 2011 [[stable isotope]] study were found to have fed on [[pelagic zone|pelagic]] rather than demersal animals, in contrast to previous observations.<ref name="borrell et al"/>


The [[tiger shark]] (''Galeocerdo cuvier'') is known to prey on ''Rhina ancylostoma''.<ref name="simpfendorfer et al"/> The ray is protected by the thorns on its head and back, and it may ram perceived threats.<ref name="fishbase"/> [[Parasite]]s documented from this species include the [[tapeworm]]s ''[[Carpobothrium rhinei]]'',<ref name="sarada et al"/> ''[[Dollfusiella michiae]]'',<ref name="campbell and beveridge"/> ''[[Nybelinia southwelli]]'',<ref name="palm and walter"/> ''[[Stoibocephalum arafurense]]'',<ref name="cielocha and jensen"/> and ''[[Tylocephalum carnpanulatum]]'',<ref name="butler"/> the [[leech]] ''[[Pontobdella macrothela]]'',<ref name="de silva"/> the [[trematode]] ''[[Melogonimus rhodanometra]]'',<ref name="bray et al"/> the [[monogenea]]ns ''[[Branchotenthes robinoverstreeti]]''<ref name="bullard and dippenaar"/> and ''[[Monocotyle ancylostomae]]'',<ref name="zhang et al"/> and the [[copepod]]s ''[[Nesippus vespa]]'',<ref name="dippenaar et al"/> ''[[Pandarus cranchii]]'', and ''[[Pandarus smithii|P. smithii]]''.<ref name="izawa"/> There is a record of a ''Rhina ancylostoma'' being cleaned by [[bluestreak cleaner wrasse]]s (''Labroides dimidiatus'').<ref name="michael"/>
The [[tiger shark]] (''Galeocerdo cuvier'') is known to prey on ''Rhina ancylostoma''.<ref name="simpfendorfer et al"/> The ray is protected by the thorns on its head and back, and it may ram perceived threats.<ref name="fishbase"/>
=== Parasites ===
[[Parasite]]s documented from this species include the [[tapeworm]]s ''[[Carpobothrium rhinei]]'',<ref name="sarada et al" /> ''[[Dollfusiella michiae]]'',<ref name="campbell and beveridge" /> ''[[Nybelinia southwelli]]'',<ref name="palm and walter" /> ''[[Stoibocephalum arafurense]]'',<ref name="cielocha and jensen" /> and ''[[Tylocephalum carnpanulatum]]'',<ref name="butler" /> the [[leech]] ''[[Pontobdella macrothela]]'',<ref name="de silva" /> the [[trematode]] ''[[Melogonimus rhodanometra]]'',<ref name="bray et al" /> the [[monogenea]]ns ''[[Branchotenthes robinoverstreeti]]''<ref name="bullard and dippenaar" /> and ''[[Monocotyle ancylostomae]]'',<ref name="zhang et al" /> and the [[copepod]]s ''[[Nesippus vespa]]'',<ref name="dippenaar et al" /> ''[[Pandarus cranchii]]'', and ''[[Pandarus smithii|P. smithii]]''.<ref name="izawa" /> There is a record of a ''Rhina ancylostoma'' being cleaned by [[bluestreak cleaner wrasse]]s (''Labroides dimidiatus'').<ref name="michael" />


== Reproduction ==
Reproduction in ''Rhina ancylostoma'' is [[viviparous]], with the developing [[embryo]]s sustained to term by [[yolk]]. Adult females have a single functional [[ovary]] and [[uterus]]. The litter size varies between two and eleven pups, and newborns measure {{convert|45|-|51|cm|in|abbr=on}} long.<ref name="michael"/><ref name="devadoss and matcha"/><ref name="last et al"/> [[Sexual maturity]] is attained at lengths of {{convert|1.5|-|1.8|m|ft|abbr=on}} for males and over {{convert|1.8|m|ft|abbr=on}} in females. Females grow larger than males.<ref name="last and stevens"/><ref name="raje"/>
Reproduction in ''Rhina ancylostoma'' is [[viviparous]], with the developing [[embryo]]s sustained to term by [[yolk]]. Adult females have two [[ovaries]] and two [[uterine horns]].<ref>{{cite journal |last1=Purushottama |first1=G. B. |last2=Thomas |first2=Sujitha |last3=Kizhakudan |first3=Shoba Joe |last4=Zacharia |first4=P. U. |title=Catch composition, reproductive biology and diet of the bowmouth guitarfish <em>Rhina ancylostomus</em> Bloch and Schneider, 1801 (Batoidea: Rhinidae) in the eastern Arabian Sea, India |journal=Indian J. Fish. |date=2022 |volume=69 |issue=3 |page=1-6}}</ref> The litter size varies between two and eleven pups, and newborns measure {{convert|45|-|51|cm|in|abbr=on}} long.<ref name="michael"/><ref name="devadoss and matcha"/><ref name="last et al"/> [[Sexual maturity]] is attained at lengths of {{convert|1.5|-|1.8|m|ft|abbr=on}} for males and over {{convert|1.8|m|ft|abbr=on}} in females. Females grow larger than males.<ref name="last and stevens"/><ref name="raje"/>


==Human interactions==
==Human interactions==
[[File:shark ray newport.jpg|thumb|"Sweet Pea", a female ''Rhina ancylostoma'' at the Newport Aquarium.]]
[[File:shark ray newport.jpg|thumb|"Sweet Pea", a female bowmouth guitarfish at the [[Newport Aquarium]].]]
Throughout its range, ''Rhina ancylostoma'' is caught incidentally or intentionally by [[artisan fishing|artisanal]] and [[commercial fishing|commercial fisheries]] using [[trawl]]s, [[gillnet]]s, and line gear.<ref name="iucn status 18 November 2021" /> The fins are extremely valuable due to their use in [[shark fin soup]], and are often the only parts of the fish kept and brought to market. However, the meat may also be sold fresh or dried and salted, and it is highly sought after in India.<ref name="fishbase"/><ref name="raje"/> When caught as [[bycatch]] in trawls, ''Rhina ancylostoma'' is considered a nuisance because its strength and rough skin make it difficult to handle, and as the heavy ray thrashes in the net it can damage the rest of the catch.<ref name="last and stevens"/> In Thailand, the enlarged thorns of this species are used to make bracelets.<ref name="fowler et al"/>
Throughout its range, the ''bowmouth guitarfish'' is caught incidentally or intentionally by [[artisan fishing|artisanal]] and [[commercial fishing|commercial fisheries]] using [[trawl]]s, [[gillnet]]s, and line gear.<ref name="iucn status 18 November 2021" /> The fins are extremely valuable due to their use in [[shark fin soup]], and are often the only parts of the fish kept and brought to market. However, the meat may also be sold fresh or dried and salted, and it is highly sought after in India.<ref name="fishbase"/><ref name="raje"/> When caught as [[bycatch]] in trawls, ''Rhina ancylostoma'' is considered a nuisance because its strength and rough skin make it difficult to handle, and as the heavy ray thrashes in the net it can damage the rest of the catch.<ref name="last and stevens"/> In Thailand, the enlarged thorns of this species are used to make bracelets.<ref name="fowler et al"/>


The [[International Union for Conservation of Nature]] (IUCN) has assessed ''Rhina ancylostoma'' as [[Critically Endangered species|Critically endangered]] in 2019, delisted from the [[vulnerable species|vulnerable]] category, along with many other guitarfish species. It is threatened by fishing and by [[habitat destruction]] and [[habitat degradation|degradation]], particularly from [[blast fishing]], [[coral bleaching]], and [[siltation]]. Its numbers are known to have declined substantially in Indonesian waters, where it is one of the large rays targeted by a mostly unregulated gillnet fishery. The IUCN has given this species a regional assessment of [[Near Threatened]] in Australian waters, where it is not a targeted species but is taken as bycatch in bottom trawls. The installation of [[turtle excluder device]]s on some Australian trawlers has benefited this species.<ref name="iucn status 18 November 2021" /> Since it is rare and faces many conservation threats, ''Rhina ancylostoma'' has been called "the [[panda]] of the aquatic world".<ref name="underwatertimes"/>
The [[International Union for Conservation of Nature]] (IUCN) has assessed ''Rhina ancylostoma'' as [[Critically Endangered species|critically endangered]] in 2019, delisted from the [[vulnerable species|vulnerable]] category, along with many other guitarfish species. It is threatened by fishing and by [[habitat destruction]] and [[habitat degradation|degradation]], particularly from [[blast fishing]], [[coral bleaching]], and [[siltation]]. Its numbers are known to have declined substantially in Indonesian waters, where it is one of the large rays targeted by a mostly unregulated gillnet fishery. The IUCN has given this species a regional assessment of [[Near Threatened]] in Australian waters, where it is not a targeted species but is taken as bycatch in bottom trawls. The installation of [[turtle excluder device]]s on some Australian trawlers has benefited this species.<ref name="iucn status 18 November 2021" /> Since it is rare and faces many conservation threats, the ''bowmouth guitarfish'' has been called "the [[panda]] of the aquatic world".<ref name="underwatertimes"/>


It is a popular subject of [[public aquarium]]s and fares relatively well, with one individual having lived for seven years in captivity.<ref name="last and stevens"/><ref name="michael"/> In 2007, the [[Newport Aquarium]] in [[Kentucky]] initiated the world's first [[captive breeding]] program for this species.<ref name="underwatertimes"/> Newport Aquarium announced in January 2014 that the female, "Sweet Pea", had become pregnant and given birth to seven pups.<ref name="aquariumworks"/> By February 2014, all seven pups had died.<ref>{{cite web | url=http://local12.com/news/features/top-stories/stories/newport-aquarium-says-shark-ray-pups-died-8807.shtml | title=Newport Aquarium says shark ray pups died | publisher=WKRC-TV | date=26 February 2014 | access-date=26 February 2014 | url-status=dead | archive-url=https://archive.today/20140226160245/http://local12.com/news/features/top-stories/stories/newport-aquarium-says-shark-ray-pups-died-8807.shtml | archive-date=26 February 2014 }}</ref> On January 7, 2016, Sweet Pea gave birth to nine shark pups<ref>{{cite web | url=http://www.fox19.com/story/30910069/newport-aquarium-shark-ray-gives-birth-to-nine-pups | title=Newport Aquarium shark ray gives birth to nine pups | publisher=FOX19NOW | date=7 January 2016 | access-date=10 February 2016}}</ref> which were eating on their own and still gaining weight by February 10, 2016.<ref>{{cite web | url=http://aquariumworks.org/2016/02/10/shark-ray-pups-reach-milestones/ | title=Shark Ray Pups Reach Milestones | publisher=aquariumworks.org | date=10 February 2016 | access-date=10 February 2016}}</ref> Newport Aquarium later announced that the pups would be moved into a coral reef exhibit where they can be viewed by the public starting on June 24.<ref>{{Cite web|url=http://www.wlwt.com/news/rare-shark-ray-pups-to-move-to-exhibit-at-newport-aquarium/40194054|title=Rare shark ray pups to move to exhibit at Newport Aquarium|last=Ferrell|first=Nikki|date=23 June 2016|access-date=2016-06-25}}</ref> The species also bred at the [[Marine Life Park|S.E.A. Aquarium]] in [[Singapore]] in 2015.<ref>{{cite news | url=http://www.straitstimes.com/singapore/sea-aquarium-successfully-breeds-shark-ray-pup-a-vulnerable-species/ | title=S.E.A. Aquarium successfully breeds shark ray pup, a vulnerable species | newspaper=[[The Straits Times]] | date=5 May 2016 | access-date=12 August 2016}}</ref>
It is a popular subject of [[public aquarium]]s and fares relatively well, with one individual having lived for seven years in captivity.<ref name="last and stevens"/><ref name="michael"/> In 2007, the [[Newport Aquarium]] in [[Kentucky]] initiated the world's first [[captive breeding]] program for this species.<ref name="underwatertimes"/> Newport Aquarium announced in January 2014 that the female, "Sweet Pea", had become pregnant and given birth to seven pups.<ref name="aquariumworks"/> By February 2014, all seven pups had died.<ref>{{cite web | url=http://local12.com/news/features/top-stories/stories/newport-aquarium-says-shark-ray-pups-died-8807.shtml | title=Newport Aquarium says shark ray pups died | publisher=WKRC-TV | date=26 February 2014 | access-date=26 February 2014 | url-status=dead | archive-url=https://archive.today/20140226160245/http://local12.com/news/features/top-stories/stories/newport-aquarium-says-shark-ray-pups-died-8807.shtml | archive-date=26 February 2014 }}</ref> On January 7, 2016, Sweet Pea gave birth to nine shark pups<ref>{{cite web | url=http://www.fox19.com/story/30910069/newport-aquarium-shark-ray-gives-birth-to-nine-pups | title=Newport Aquarium shark ray gives birth to nine pups | publisher=FOX19NOW | date=7 January 2016 | access-date=10 February 2016}}</ref> which were eating on their own and still gaining weight by February 10, 2016.<ref>{{cite web | url=http://aquariumworks.org/2016/02/10/shark-ray-pups-reach-milestones/ | title=Shark Ray Pups Reach Milestones | publisher=aquariumworks.org | date=10 February 2016 | access-date=10 February 2016}}</ref> Newport Aquarium later announced that the pups would be moved into a coral reef exhibit where they can be viewed by the public starting on June 24.<ref>{{Cite web|url=http://www.wlwt.com/news/rare-shark-ray-pups-to-move-to-exhibit-at-newport-aquarium/40194054|title=Rare shark ray pups to move to exhibit at Newport Aquarium|last=Ferrell|first=Nikki|date=23 June 2016|access-date=2016-06-25}}</ref> The species also bred at the [[Marine Life Park|S.E.A. Aquarium]] in [[Singapore]] in 2015.<ref>{{cite news | url=http://www.straitstimes.com/singapore/sea-aquarium-successfully-breeds-shark-ray-pup-a-vulnerable-species/ | title=S.E.A. Aquarium successfully breeds shark ray pup, a vulnerable species | newspaper=[[The Straits Times]] | date=5 May 2016 | access-date=12 August 2016}}</ref>
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[[Category:Rajiformes]]
[[Category:Rajiformes]]
[[Category:Fish of the Red Sea]]
[[Category:Fish of the Red Sea]]
[[Category:Marine fauna of East Africa]]
[[Category:Marine fish of East Africa]]
[[Category:Fish described in 1801]]
[[Category:Fish described in 1801]]
[[Category:Taxa named by Marcus Elieser Bloch]]
[[Category:Taxa named by Marcus Elieser Bloch]]

Latest revision as of 07:35, 30 October 2024

"Shark ray" redirects here. For the species from the East Atlantic, see false shark ray.

Rhina ancylostoma
CITES Appendix II (CITES)[2]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Order: Rhinopristiformes
Family: Rhinidae
Genus: Rhina
Bloch & J. G. Schneider, 1801
Species:
R. ancylostoma
Binomial name
Rhina ancylostoma
Range of Rhina ancylostoma[3]
Synonyms

Rhina cyclostomus Swainson, 1839

Rhina ancylostoma, also known as the bowmouth guitarfish, shark ray or mud skate, is a species of ray and a member of the family Rhinidae. Its evolutionary affinities are not fully resolved, though it may be related to true guitarfishes and skates.[citation needed] This rare species occurs widely in the tropical coastal waters of the western Indo-Pacific, at depths of up to 90 m (300 ft). Highly distinctive in appearance, Rhina ancylostoma has a wide and thick body with a rounded snout and large shark-like dorsal and tail fins. Its mouth forms a W-shaped undulating line, and there are multiple thorny ridges over its head and back. It has a dorsal color pattern of many white spots over a bluish gray to brown background, with a pair of prominent black markings over the pectoral fins. This large species can reach a length of 2.7 m (8.9 ft) and weight of 135 kg (298 lb).[citation needed]

Usually found near the sea floor, the bowmouth guitarfish prefers sandy or muddy areas near underwater structures. It is a strong-swimming predator of bony fishes, crustaceans, and molluscs. This species gives live birth to litters of two to eleven pups, which are nourished during gestation by yolk. The International Union for Conservation of Nature (IUCN) has assessed Rhina ancylostoma as Critically Endangered because it is widely caught by artisanal and commercial fisheries for its valuable fins and meat. It is viewed as a nuisance by trawlers, however, because its bulk and thorny skin cause it to damage netted catches. Habitat degradation and destruction pose an additional, significant challenge to this ray's survival. The bowmouth guitarfish adapts well to captivity and is displayed in public aquariums.

Taxonomy and phylogeny

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German naturalists Marcus Elieser Bloch and Johann Gottlob Schneider described Rhina ancylostoma in their 1801 Systema Ichthyologiae. Their account was based on a 51 cm (20 in) long specimen, now lost, collected off the Coromandel Coast of India. The genus name Rhina comes from the Greek rhinos ("snout"); the specific epithet ancylostoma is derived from the Greek ankylos ("curved" or "crooked") and stoma ("mouth").[4][5] Although Block and Schneider wrote the epithet as ancylostomus and that form appears in some literature, most modern sources regard the correct form to be ancylostoma.[6] Other common names for this species include shark ray, mud skate, shortnose mud skate, bow-mouthed angel fish, and bow-mouthed angel shark.[7]

The evolutionary relationships between Rhina ancylostoma and other rays are debated. Morphological evidence generally points to a close relationship between Rhina, Rhynchobatus and Rhynchorhina, which are a group of rays known as the wedgefishes that also have large, shark-like fins. Morphological analyses have tended to place these two genera basally among rays, though some have them as basal to just the guitarfishes (Rhinobatidae) and skates (Rajidae) while others have them basal to all other rays except sawfishes (Pristidae).[8][9][10] A 2012 study based on mitochondrial DNA upheld Rhina and Rhynchobatus as sister taxa related to the guitarfishes, but also unexpectedly found that they formed a clade with the sawfishes rather than the skates.[11] Following the description of Rhynchorhina in 2016, a study of mtDNA found that it is part of the same group and their phylogenetic relationship is ((Rhynchobatus+Rhynchorhina)+Rhina).[12]

In terms of classification, Bloch and Schneider originally placed the bowmouth guitarfish in the order Abdominales, a now-obsolete grouping of fishes defined by the positioning of their pelvic fins directly behind the pectoral fins.[4] Modern sources have included it variously in the order Rajiformes, Rhinobatiformes, Rhiniformes, or the newly proposed Rhinopristiformes.[9][11] The placement of the bowmouth guitarfish in the family Rhinidae originates from the group "Rhinae", consisting of Rhina and Rhynchobatus, in Johannes Müller and Jakob Henle's 1841 Systematische Beschreibung der Plagiostomen.[13] Later authors have also assigned this species to the family Rhinobatidae or Rhynchobatidae.[9][14] Joseph Nelson, in the 2006 fourth edition of Fishes of the World, placed this species as the sole member of Rhinidae in the order Rajiformes, which is supported by morphological but not molecular data.[10][15] More recent authorities have placed it in Rhinidae together with Rhynchobatus and Rhynchorhina, reflecting both genetic data and the morphologically intermediate position of Rhynchobatus between Rhina and Rhynchorhina.[12][16]

Description

[edit]
The rounded head, humpbacked profile, and large fins of Rhina ancylostoma give it a unique appearance.

Rhina ancylostoma is a heavily built fish growing to 2.7 m (8.9 ft) long and 135 kg (298 lb) in weight.[3][7]

The head is short, wide, and flattened with an evenly rounded snout; the front portion of the head, including the medium-sized eyes and large spiracles, is clearly distinct from the body. The long nostrils are transversely oriented and have well-developed skin flaps on their anterior margins.[17][3]

The lower jaw has three protruding lobes that fit into corresponding depressions in the upper jaw.[3][17] There are around 47 upper and 50 lower tooth rows arranged in winding bands; the teeth are low and blunt with ridges on the crown. The five pairs of ventral gill slits are positioned close to the lateral margins of the head.[3][18]

Jaws

The body is deepest in front of the two tall and falcate (sickle-shaped) dorsal fins. The first dorsal fin is about a third larger than the second and originates over the pelvic fin origins. The second dorsal fin is located midway between the first dorsal and the caudal fin. The broad and triangular pectoral fins have a deep indentation where their leading margins meet the head. The pelvic fins are much smaller than the pectoral fins, and the anal fin is absent. The tail is much longer than the body and ends in a large, crescent-shaped caudal fin; the lower caudal fin lobe is more than half the length of the upper.[3][14][17]

The entire dorsal surface has a grainy texture from a dense covering of tiny dermal denticles. A thick ridge is present along the midline of the back, which bears a band of sharp, robust thorns. There are also a pair of thorn-bearing ridges in front of the eyes, a second pair running from above the eyes to behind the spiracles, and a third pair on the "shoulders". This species is bluish to brownish gray above, lightening towards the margins of the head and over the pectoral fins. There are prominent white spots scattered over the body and fins, a white-edged black marking above each pectoral fin, and two dark transverse bands atop the head between the eyes. The underside is light gray to white. Young rays are more vividly colored than adults, which are browner with fainter patterning and proportionately smaller spots.[3]

Distribution and habitat

[edit]

While uncommon, Rhina ancylostoma is widely distributed in the coastal tropical waters of the western Indo-Pacific. In the Indian Ocean, it is found from KwaZulu-Natal in South Africa to the Red Sea (including the Seychelles), across the Indian subcontinent and Southeast Asia (including the Maldives), to Shark Bay in Western Australia. Its Pacific range extends northward to Korea and southern Japan, eastward to New Guinea, and southward to New South Wales.[3][17] Found between 3 and 90 m (10 and 300 ft) deep, this ray spends most of its time near the sea floor but can occasionally be seen swimming in midwater. It favors sandy or muddy habitats, and can also be found in the vicinity of rocky and coral reefs and shipwrecks.[3][19]

Ecology

[edit]
The tiger shark preys on Rhina ancylostoma.

Rhina ancylostoma is a strong swimmer that propels itself with its tail like a shark. It is more active at night and is not known to be territorial.[20] This species feeds mainly on demersal bony fishes such as croakers and crustaceans such as crabs and shrimp; bivalves and cephalopods are also consumed. Its bands of flattened teeth allow it to crush hard-shelled prey.[14][21] Curiously, two Rhina ancylostoma examined in a 2011 stable isotope study were found to have fed on pelagic rather than demersal animals, in contrast to previous observations.[22]

The tiger shark (Galeocerdo cuvier) is known to prey on Rhina ancylostoma.[23] The ray is protected by the thorns on its head and back, and it may ram perceived threats.[7]

Parasites

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Parasites documented from this species include the tapeworms Carpobothrium rhinei,[24] Dollfusiella michiae,[25] Nybelinia southwelli,[26] Stoibocephalum arafurense,[27] and Tylocephalum carnpanulatum,[28] the leech Pontobdella macrothela,[29] the trematode Melogonimus rhodanometra,[30] the monogeneans Branchotenthes robinoverstreeti[31] and Monocotyle ancylostomae,[32] and the copepods Nesippus vespa,[33] Pandarus cranchii, and P. smithii.[34] There is a record of a Rhina ancylostoma being cleaned by bluestreak cleaner wrasses (Labroides dimidiatus).[19]

Reproduction

[edit]

Reproduction in Rhina ancylostoma is viviparous, with the developing embryos sustained to term by yolk. Adult females have two ovaries and two uterine horns.[35] The litter size varies between two and eleven pups, and newborns measure 45–51 cm (18–20 in) long.[19][36][37] Sexual maturity is attained at lengths of 1.5–1.8 m (4.9–5.9 ft) for males and over 1.8 m (5.9 ft) in females. Females grow larger than males.[3][21]

Human interactions

[edit]
"Sweet Pea", a female bowmouth guitarfish at the Newport Aquarium.

Throughout its range, the bowmouth guitarfish is caught incidentally or intentionally by artisanal and commercial fisheries using trawls, gillnets, and line gear.[1] The fins are extremely valuable due to their use in shark fin soup, and are often the only parts of the fish kept and brought to market. However, the meat may also be sold fresh or dried and salted, and it is highly sought after in India.[7][21] When caught as bycatch in trawls, Rhina ancylostoma is considered a nuisance because its strength and rough skin make it difficult to handle, and as the heavy ray thrashes in the net it can damage the rest of the catch.[3] In Thailand, the enlarged thorns of this species are used to make bracelets.[38]

The International Union for Conservation of Nature (IUCN) has assessed Rhina ancylostoma as critically endangered in 2019, delisted from the vulnerable category, along with many other guitarfish species. It is threatened by fishing and by habitat destruction and degradation, particularly from blast fishing, coral bleaching, and siltation. Its numbers are known to have declined substantially in Indonesian waters, where it is one of the large rays targeted by a mostly unregulated gillnet fishery. The IUCN has given this species a regional assessment of Near Threatened in Australian waters, where it is not a targeted species but is taken as bycatch in bottom trawls. The installation of turtle excluder devices on some Australian trawlers has benefited this species.[1] Since it is rare and faces many conservation threats, the bowmouth guitarfish has been called "the panda of the aquatic world".[39]

It is a popular subject of public aquariums and fares relatively well, with one individual having lived for seven years in captivity.[3][19] In 2007, the Newport Aquarium in Kentucky initiated the world's first captive breeding program for this species.[39] Newport Aquarium announced in January 2014 that the female, "Sweet Pea", had become pregnant and given birth to seven pups.[40] By February 2014, all seven pups had died.[41] On January 7, 2016, Sweet Pea gave birth to nine shark pups[42] which were eating on their own and still gaining weight by February 10, 2016.[43] Newport Aquarium later announced that the pups would be moved into a coral reef exhibit where they can be viewed by the public starting on June 24.[44] The species also bred at the S.E.A. Aquarium in Singapore in 2015.[45]

References

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  31. ^ Bullard, S.A.; Dippenaar, S.M. (June 2003). "Branchotenthes robinoverstreeti n. gen. and n. sp. (Monogenea: Hexabothriidae) from Gill Filaments of the Bowmouth Guitarfish, Rhina ancylostoma (Rhynchobatidae), in the Indian Ocean". The Journal of Parasitology. 89 (3): 595–601. doi:10.1645/0022-3395(2003)089[0595:BRNGAN]2.0.CO;2. PMID 12880262. S2CID 21951837.
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  33. ^ Dippenaar, S.M.; Mathibela, R.B.; Bloomer, P. (2010). "Cytochrome oxidase I sequences reveal possible cryptic diversity in the cosmopolitan symbiotic copepod Nesippus orientalis Heller, 1868 (Pandaridae: Siphonostomatoida) on elasmobranch hosts from the KwaZulu-Natal coast of South Africa". Experimental Parasitology. 125 (1): 42–50. doi:10.1016/j.exppara.2009.08.017. hdl:2263/11615. PMID 19723521.
  34. ^ Izawa, K. (2010). "Redescription of eight species of parasitic copepods (Siphonostomatoida, Pandaridae) infecting Japanese elasmobranchs". Crustaceana. 83 (3): 313–341. doi:10.1163/001121609X12591347509329.
  35. ^ Purushottama, G. B.; Thomas, Sujitha; Kizhakudan, Shoba Joe; Zacharia, P. U. (2022). "Catch composition, reproductive biology and diet of the bowmouth guitarfish Rhina ancylostomus Bloch and Schneider, 1801 (Batoidea: Rhinidae) in the eastern Arabian Sea, India". Indian J. Fish. 69 (3): 1-6.
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  38. ^ Fowler, S.L.; Reed, T.M.; Dipper, F.A., eds. (2002). Elasmobranch Biodiversity, Conservation and Management: Proceedings of the International Seminar and Workshop, Sabah, Malaysia, July 1997. IUCN SSC Shark Specialist Group. p. 112. ISBN 2-8317-0650-5.
  39. ^ a b "Newport Aquarium Launches World's First Shark Ray Breeding Program, Adds Rare Male Shark Ray". UnderwaterTimes. February 1, 2007. Retrieved January 22, 2009.
  40. ^ "Newport Aquarium's Sweet Pea, the First Documented Shark Ray to Breed in Captivity, Gives Birth to Seven Pups". Aquarium Works. Newport Aquarium. January 29, 2014. Retrieved February 1, 2014.
  41. ^ "Newport Aquarium says shark ray pups died". WKRC-TV. 26 February 2014. Archived from the original on 26 February 2014. Retrieved 26 February 2014.
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  43. ^ "Shark Ray Pups Reach Milestones". aquariumworks.org. 10 February 2016. Retrieved 10 February 2016.
  44. ^ Ferrell, Nikki (23 June 2016). "Rare shark ray pups to move to exhibit at Newport Aquarium". Retrieved 2016-06-25.
  45. ^ "S.E.A. Aquarium successfully breeds shark ray pup, a vulnerable species". The Straits Times. 5 May 2016. Retrieved 12 August 2016.

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