Round Ribbontail Ray
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The round ribbontail ray (''Taeniurops meyeni'') is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of stingray in the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Dasyatidae, found throughout the nearshore waters of the
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
Indo-Pacific, as well as off islands in the eastern
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contine ...
. It is a bottom-dwelling inhabitant of
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') ...
s,
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environmen ...
, and
reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock o ...
s, generally at a depth of . Reaching across, this large ray is characterized by a thick, rounded
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
disc covered by small
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s on top, and a relatively short tail bearing a deep ventral fin fold. In addition, it has a variable but distinctive light and dark mottled pattern on its upper surface, and a black tail. Generally nocturnal, the round ribbontail ray can be solitary or gregarious, and is an active
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
of small, benthic molluscs,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s, and bony fishes. It is
aplacental viviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop ins ...
, with the embryos sustained by
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example ...
, and later histotroph ("uterine milk") secreted by the mother; up to seven pups are born at a time. Although not aggressive, if provoked the round ribbontail ray will defend itself with its
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a st ...
ous tail spine, and it has been responsible for at least one fatality. It is valued by
ecotourist Ecotourism is a form of tourism involving responsible travel (using sustainable transport) to natural areas, conserving the environment, and improving the well-being of the local people. Its purpose may be to educate the traveler, to provide fund ...
divers and recreational anglers. This slow-reproducing species is threatened by
commercial fishing Commercial fishing is the activity of catching fish and other seafood for commercial profit, mostly from wild fisheries. It provides a large quantity of food to many countries around the world, but those who practice it as an industry must often ...
, both targeted and as bycatch, and
habitat degradation Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
across much of its range. As a result, the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN) has assessed it as Vulnerable.


Taxonomy and phylogeny

As ''Taeniurops meyeni'', the round ribbontail ray was described by German biologists Johannes Peter Müller and Friedrich Gustav Jakob Henle in their 1841 ''Systematische Beschreibung der Plagiostomen'', based on two
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...
s collected from
Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label= Mauritian Creole, Moris ), officially the Republic of Mauritius, is an island nation in the Indian Ocean about off the southeast coast of the African continent, east of Madagascar. It ...
. However, this species is better known under the name ''Taeniura melanospila'' (or ''melanospilos''), which was applied by Dutch ichthyologist
Pieter Bleeker Pieter Bleeker (10 July 1819 – 24 January 1878) was a Dutch medical doctor, ichthyologist, and herpetologist. He was famous for the ''Atlas Ichthyologique des Indes Orientales Néêrlandaises'', his monumental work on the fishes of East Asia ...
to a juvenile specimen from
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mos ...
, in a 1953 volume of the scientific journal ''Natuurkundig Tijdschrift voor Nederlandsch Indië''. Other common names for the round ribbontail ray include black spotted ray, black-blotched stingray, black-spotted stingray, fantail ray, fantail stingray, giant reef ray, blotched fantail ray, and speckled stingray. In Australia, it is one of several species referred to as " bull ray". A minority of authors place this species with the river stingrays in the family
Potamotrygonidae River stingrays or freshwater stingrays are Neotropical freshwater fishes of the family Potamotrygonidae in the order Myliobatiformes, one of the four orders of batoids, cartilaginous fishes related to sharks. They are found in rivers in tropic ...
. Preliminary morphological examination has suggested that the round ribbontail ray is more related to ''
Dasyatis ''Dasyatis'' (Greek δασύς ''dasýs'' meaning rough or dense and βατίς ''batís'' meaning skate) is a genus of stingray in the family Dasyatidae that is native to the Atlantic, including the Mediterranean. In a 2016 taxonomic revisi ...
'' and Indo-Pacific ''
Himantura ''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 ('' Brevitrygon'', '' Fluvi ...
'' than to the congeneric bluespotted ribbontail ray (''T. lymma''), which is closer to the amphi-American ''Himantura'' ('' H. pacifica'' and '' H. schmardae'') and the river stingrays.


Etymology

The ray is named in honor of Franz Julius Ferdinand Meyen (1804-1840), a physician and a botanist, who collected or supplied the type specimens.


Description

The round ribbontail ray has a thick
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
disc wider than it is long, with a smoothly rounded outer margin. The eyes are of medium size and are followed by larger spiracles. There is a short and broad curtain of skin between the oval
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
s, with a finely fringed trailing margin. The mouth is wide and curved, with faint furrows at the corners. There is a row of seven
papilla Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...
e on the floor, with the outermost pair smaller and set apart from the others. There are 37–46 tooth rows in the upper jaw and 39–45 tooth rows in the lower jaw. The teeth are small with a deep groove across the crown and are arranged in a dense
quincunx A quincunx () is a geometric pattern consisting of five points arranged in a cross, with four of them forming a square or rectangle and a fifth at its center. The same pattern has other names, including "in saltire" or "in cross" in heraldry (d ...
pattern into flattened surfaces. The pelvic fins are small and narrow. The tail is relatively short, not exceeding the width of the disc, and bears one (rarely two) long, serrated stinging spine on the upper surface. The base of the tail is broad; past the spine, the tail rapidly thins, and bears a deep ventral fin fold that runs to the tail tip. The upper surface of the disc and tail are roughened by a uniform covering of small, widely spaced granules. There is also a midline row of sharp tubercles on the back, with two shorter rows alongside. The first of these tubercles develop at a length of around , over the "shoulders" and in the single midline row. The dorsal coloration is light to dark gray, brown-gray, or purplish, becoming most intense towards the fin margins, with a highly variable pattern of irregular darker mottling and white speckles or streaks. The tail past the spine, including the fin fold, is uniformly black, while the underside is creamy-white with darker fin margins and additional dots. Young rays are more plain in coloration than adults. One of the largest stingray species, the round ribbontail ray can grow to across, long, and in weight.


Distribution and habitat

The round ribbontail ray has a wide distribution in the Indo-Pacific region: it is found from KwaZulu-Natal in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. It is bounded to the south by of coastline that stretch along the Atlantic Ocean, South Atlantic and Indian Oceans; to the ...
northward along the East African coast to the
Red Sea The Red Sea ( ar, البحر الأحمر - بحر القلزم, translit=Modern: al-Baḥr al-ʾAḥmar, Medieval: Baḥr al-Qulzum; or ; Coptic: ⲫⲓⲟⲙ ⲛ̀ϩⲁϩ ''Phiom Enhah'' or ⲫⲓⲟⲙ ⲛ̀ϣⲁⲣⲓ ''Phiom ǹšari''; ...
, including
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
and the
Mascarenes The Mascarene Islands (, ) or Mascarenes or Mascarenhas Archipelago is a group of islands in the Indian Ocean east of Madagascar consisting of the islands belonging to the Republic of Mauritius as well as the French department of Réunion. Thei ...
; from there, its range extends eastward through the
Indian subcontinent The Indian subcontinent is a list of the physiographic regions of the world, physiographical region in United Nations geoscheme for Asia#Southern Asia, Southern Asia. It is situated on the Indian Plate, projecting southwards into the Indian O ...
to
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainlan ...
and
Micronesia Micronesia (, ) is a subregion of Oceania, consisting of about 2,000 small islands in the western Pacific Ocean. It has a close shared cultural history with three other island regions: the Philippines to the west, Polynesia to the east, and ...
, occurring as far north as
Korea Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...
and southern Japan, and as far south as Australia, where it is found from at least
Ningaloo Reef The Ningaloo Coast is a World Heritage Site located in the north west coastal region of Western Australia. The heritage-listed area is located approximately north of Perth, along the East Indian Ocean. The distinctive Ningaloo Reef that fring ...
off
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
to
Stradbroke Island Stradbroke Island, also known as Minjerribah, was a large sand island that formed much of the eastern side of Moreton Bay near Brisbane, Queensland until the late 19th century. Today the island is split into two islands: North Stradbroke Islan ...
off
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , establishe ...
, including
Lord Howe Island Lord Howe Island (; formerly Lord Howe's Island) is an irregularly crescent-shaped volcanic remnant in the Tasman Sea between Australia and New Zealand, part of the Australian state of New South Wales. It lies directly east of mainland P ...
. In the easternmost portion of its range, it has been reported from
Cocos Island Cocos Island ( es, Isla del Coco) is an island in the Pacific Ocean administered by Costa Rica, approximately southwest of the Costa Rican mainland. It constitutes the 11th of the 13 districts of Puntarenas Canton of the Province of Puntarena ...
and the
Galápagos Islands The Galápagos Islands (Spanish: , , ) are an archipelago of volcanic islands. They are distributed on each side of the equator in the Pacific Ocean, surrounding the centre of the Western Hemisphere, and are part of the Republic of Ecuador ...
, with individuals possibly dispersing as far as
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
. Bottom-dwelling in nature, the round ribbontail ray is typically found close to shore at a depth of , though it has been reported anywhere from the
surf zone As ocean surface waves approach shore, they get taller and break, forming the foamy, bubbly surface called ''surf''. The region of breaking waves defines the surf zone, or breaker zone. After breaking in the surf zone, the waves (now reduced i ...
to a depth of . It favors sand or rubble bottoms in shallow
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') ...
s or near
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and ...
and rocky
reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock o ...
s, and may also enter
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environmen ...
.


Biology and ecology

The round ribbontail ray has nocturnal habits and rests motionless for much of the day, often near vertical structures, in caves, or under ledges. It may be solitary, or form small to large groups. This ray is frequently shadowed by one or more jacks or
cobia The cobia (''Rachycentron canadum'') is a species of carangiform marine fish, the only extant representative of the genus ''Rachycentron'' and the family Rachycentridae. Other common names include black kingfish, black salmon, ling, lemonfis ...
(''Rachycentron canadum''); these smaller fishes may feed on food stirred up by the ray's activities, or use the ray's body as cover for approaching their own prey. The round ribbontail ray hunts for bivalves, crabs, shrimps, and small bony fishes on the bottom. When feeding, it adopts a characteristic posture in which it presses the margin of its disc against the bottom and takes in water through its spiracles, which it blows through its mouth to uncover prey buried in the sediment. This species may fall prey to larger fishes such as
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s, and
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their ...
s. When threatened, it raises its tail over its back so that the spine faces forward, and waves it back and forth. Known
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s of this species include the
monogenea Monogeneans are a group of ectoparasitic flatworms commonly found on the skin, gills, or fins of fish. They have a direct lifecycle and do not require an intermediate host. Adults are hermaphrodites, meaning they have both male and female repro ...
n '' Dasybatotrema spinosum'', '' Dendromonocotyle pipinna'', '' Neoentobdella garneri'' and '' N. taiwanensis'', and the nematode '' Echinocephalus overstreeti''. Little information is available on the life history of the round ribbontail ray. Like other stingrays, it is
aplacental viviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop ins ...
: the unborn embryos are initially sustained by
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example ...
, which is later supplemented by histotroph ("uterine milk", containing
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s,
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...
s, and
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
) produced by the mother. Reproductive aggregations numbering in the hundreds have been observed at Cocos Island shortly after the onset of ''
La Niña La Niña (; ) is an oceanic and atmospheric phenomenon that is the colder counterpart of as part of the broader El Niño–Southern Oscillation (ENSO) climate pattern. The name ''La Niña'' originates from Spanish for "the girl", by an ...
'', which brings cooler temperatures. During these periods a single female may be pursued by dozens of males. Females bear litters of up to seven pups, each measuring across and long. Off South Africa, birthing may take place in the summer. Males attain
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definit ...
at a disc width of ; the maturation size of females is unknown.


Human interactions

The round ribbontail ray is not aggressive and has been known to approach and investigate divers. However, if harassed it can inflict a severe wound with its
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a st ...
ous tail spine. This species has been responsible for at least one recorded fatality of a diver who was stabbed while attempting to ride the ray. The round ribbontail ray is popular with
ecotourist Ecotourism is a form of tourism involving responsible travel (using sustainable transport) to natural areas, conserving the environment, and improving the well-being of the local people. Its purpose may be to educate the traveler, to provide fund ...
divers because of its size and spectacular appearance. The
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN) has assessed the round ribbontail ray as Vulnerable. It cannot withstand heavy fishing pressure due to its low reproductive rate, and there is widespread degradation of its coral reef habitat, including from
agricultural runoff Agricultural pollution refers to biotic and abiotic byproducts of farming practices that result in contamination or degradation of the environment and surrounding ecosystems, and/or cause injury to humans and their economic interests. The pol ...
and destructive fishing practices such as
blast fishing Blast fishing, fish bombing, dynamite fishing or grenade fishing is a destructive fishing practice using explosives to stun or kill schools of fish for easy collection. This often illegal practice is extremely destructive to the surrounding e ...
. This species is caught by
commercial Commercial may refer to: * a dose of advertising conveyed through media (such as - for example - radio or television) ** Radio advertisement ** Television advertisement * (adjective for:) commerce, a system of voluntary exchange of products and s ...
and fisheries throughout its range, using line gear and
trawl Trawling is a method of fishing that involves pulling a fishing net through the water behind one or more boats. The net used for trawling is called a trawl. This principle requires netting bags which are towed through water to catch different speci ...
s. One region where it is heavily pressured is in Indonesian waters, where it and other large rays are taken intentionally and otherwise by tangle netters, longliners, and trawlers operating off Java, Bali,
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torr ...
, and
Lombok Lombok is an island in West Nusa Tenggara province, Indonesia. It forms part of the chain of the Lesser Sunda Islands, with the Lombok Strait separating it from Bali to the west and the Alas Strait between it and Sumbawa to the east. It is ...
. All landed individuals are brought to market for human consumption. Off South Africa, the round ribbontail ray is captured incidentally by
prawn Prawn is a common name for small aquatic crustaceans with an exoskeleton and ten legs (which is a member of the order decapoda), some of which can be eaten. The term "prawn"Mortenson, Philip B (2010''This is not a weasel: a close look at nature' ...
trawlers on offshore
banks A bank is a financial institution that accepts deposits from the public and creates a demand deposit while simultaneously making loans. Lending activities can be directly performed by the bank or indirectly through capital markets. Becaus ...
, but is not utilized. Because of its size and strength, it is also prized by sport anglers, who usually release it unharmed. South Africa sets a recreational bag limit of one ray per species per person per day and does not allow
spearfishing Spearfishing is a method of fishing that involves impaling the fish with a straight pointed object such as a spear, gig or harpoon. It has been deployed in artisanal fishing throughout the world for millennia. Early civilisations were familia ...
for this species. In Australian waters, this ray has been assessed as of
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
. Although it is caught (and discarded) by prawn trawlers, this mortality has been reduced by the mandatory installation of Turtle Excluder Devices (TEDs). Furthermore, a portion of its Australian range lies within the
Great Barrier Reef Marine Park The Great Barrier Reef Marine Park protects a large part of Australia's Great Barrier Reef from damaging activities. It is a vast multiple-use Marine Park which supports a wide range of uses, including commercial marine tourism, fishing, ports an ...
. This species has also been listed under Least Concern in the
Maldives Maldives (, ; dv, ދިވެހިރާއްޖެ, translit=Dhivehi Raajje, ), officially the Republic of Maldives ( dv, ދިވެހިރާއްޖޭގެ ޖުމްހޫރިއްޔާ, translit=Dhivehi Raajjeyge Jumhooriyyaa, label=none, ), is an archipelag ...
where, due to the tourist value of rays, the government has created protected marine reserves and banned the export of rays in 1995 and ray skins in 1996.


References


External links


"''Taeniura meyeni'', Round ribbontail ray" at FishBase
* ttp://fishesofaustralia.net.au/home/species/3530 Fishes of Australia : ''Taeniura meyeni''* {{Good article round ribbontail ray Fish of the Indian Ocean Marine fauna of East Africa Marine fish of Northern Australia Taxa named by Johannes Peter Müller Taxa named by Friedrich Gustav Jakob Henle round ribbontail ray