Shovelnose Ray
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The guitarfish, also referred to as shovelnose rays, are a family, Rhinobatidae, of
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
. The guitarfish are known for an elongated body with a flattened head and trunk and small, ray-like wings. The combined range of the various species is tropical, subtropical, and warm temperate waters worldwide.


Names

In
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
, guitarfish are commonly referred to as shovelnose rays or shovelnose sharks.


Description

Guitarfish have a body from intermediate between those of sharks and rays. The tail has a typical shark-like form, but in many species, the head has a triangular, or
guitar The guitar is a fretted musical instrument that typically has six strings. It is usually held flat against the player's body and played by strumming or plucking the strings with the dominant hand, while simultaneously pressing selected stri ...
-like shape, rather than the disc-shape formed by fusion with the
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 ...
s found in other rays.


Reproduction

Guitarfish are
ovoviviparous 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 insi ...
; the embryo matures inside an egg within the mother until it is ready to hatch. This is typical of rays.


Habitat

Guitarfish are
bottom feeder A bottom feeder is an aquatic animal that feeds on or near the bottom of a body of water. Biologists often use the terms ''benthos''—particularly for invertebrates such as shellfish, crabs, crayfish, sea anemones, starfish, snails, bri ...
s that bury themselves in mud or sand and eat worms, crabs, and clams. Some can tolerate salt, fresh, and
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
water. They generally live close to the beach/coastline or in estuaries.


Evolution

Rays, including guitarfish, belong to the ancient lineage of
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...
es. Fossil
denticle A denticle is any small tooth-like or bristle-like structure. "Denticle" may refer to: * Denticle (tooth feature), serrations on the teeth of dinosaurs, lizards, sharks, and mammals * Dermal denticles or placoid scales, in cartilaginous fishes * ...
s (tooth-like scales in the skin) resembling that of today's
chondrichthyan Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. C ...
s date at least as far back as the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
, with the oldest unambiguous fossils of
cartilaginous fish Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...
dating from the middle
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
. A
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
within this diverse family, the
Neoselachii Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gi ...
, emerged by the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, with the best-understood neoselachian fossils dating from the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
. This
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
is represented today by
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 Selachimo ...
s,
sawfish Sawfish, also known as carpenter sharks, are a family of rays characterized by a long, narrow, flattened rostrum, or nose extension, lined with sharp transverse teeth, arranged in a way that resembles a saw. They are among the largest fish wi ...
, rays and
skate Skate or Skates may refer to: Fish *Skate (fish), several genera of fish belonging to the family Rajidae * Pygmy skates, several genera of fish belonging to the family Gurgesiellidae * Smooth skates or leg skates, several genera of fish belongin ...
s.


Classification

Nelson's 2006 ''
Fishes of the World ''Fishes of the World'' by the American ichthyologist Joseph S. Nelson (1937–2011) is a standard reference for fish systematics. Now in its fifth edition (2016), the work is a comprehensive overview of the diversity and classification of the ...
'' recognized four genera in this family: ''
Aptychotrema ''Aptychotrema'' is a genus of guitarfish, belonging to the family Rhinobatidae. They are found around Australia. Species There are three living species, and one extinct species only known from fossil remains of Cenomanian age.Bernárdez, E. ( ...
'', ''
Rhinobatos ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'', '' Trygonorrhina'', and ''
Zapteryx ''Zapteryx'' is a genus of fish in the Rhinobatidae family found in coastal parts of the Americas. Species * †''Zapteryx bichuti'' Signeux, 1961 * ''Zapteryx brevirostris'' J. P. Müller & Henle, 1841 (Shortnose guitarfish) * ''Zapteryx e ...
''; other taxa once placed in the Rhinobatidae, such as '' Platyrhinoidis'' and '' Rhina'', have since been moved to their own families. Recently, the genus ''
Glaucostegus ''Glaucostegus'', also known as giant guitarfishes, is a genus of large Indo-Pacific rays, with a single species, ''Glaucostegus cemiculus'', in the East Atlantic, and Mediterranean. They were formerly classified in the family Rhinobatidae but a ...
'' has again become recognized as distinct from ''Rhinobatos''. ''
Rhinobatos ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' has been split in three genera based on genetic and morphological considerations: ''Rhinobatos'', ''
Acroteriobatus ''Acroteriobatus'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to '' Rhinobatos'', recent authors treat it as distinct. They are found near the shore off southern Africa and in the no ...
'' and ''
Pseudobatos ''Pseudobatos'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found in warmer coastal parts of the Americas, ranging fro ...
''. '' Tarsistes'' is dubious and may be a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of ''Pseudobatos'', and other genera formerly included in Rhinobatidae have been moved to Glaucostegidae, Rhinidae and
Trygonorrhinidae Trygonorrhinidae, the banjo rays, is a family of rays, comprising eight species in three genera. They were formerly classified in the family Rhinobatidae. Taxonomy * '' Aptychotrema'' Norman, 1926 ** '' Aptychotrema rostrata'' Shaw, 1794 (East ...
. A 2021 re-evaluation of almost complete and articulated material from the Konservat-Lagerstätten of Bolca in Italy suggested that ''"Rhinobatos" dezignii'' and ''"Rhinobatos" primaevus'' should be excluded from ''Rhinobatos'' and assigned to the new genera †''Pseudorhinobatos'' gen. nov. and †''Eorhinobatos'' gen. nov. respectively. * Genus ''
Acroteriobatus ''Acroteriobatus'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to '' Rhinobatos'', recent authors treat it as distinct. They are found near the shore off southern Africa and in the no ...
'' Giltay, 1928 ** '' Acroteriobatus andysabini'' (2021) (Malagasy blue-spotted guitarfish) ** ''
Acroteriobatus annulatus The lesser guitarfish or lesser sandshark (''Acroteriobatus annulatus'') is a species of fish in the Rhinobatidae family. It is found in Angola, Namibia, South Africa, and possibly Mozambique. Its natural habitats are shallow seas and estuarine ...
'' ( J. P. Müller &
Henle Henle can refer to: * Friedrich Gustav Jakob Henle, a German physician, pathologist and anatomist (1809–1885) ** Loop of Henle in the kidney, named after Henle *Fritz Henle, a photographer, known as "Mr. Rollei" for his use of the 2.25" square for ...
, 1841)
(Lesser guitarfish) ** ''
Acroteriobatus blochii The bluntnose guitarfish or fiddlefish (''Rhinobatos blochii'') is a species of fish in the Rhinobatidae family. It is found in Namibia and South Africa. Its natural habitats are shallow seas and estuarine waters. It is threatened by habitat lo ...
'' ( J. P. Müller &
Henle Henle can refer to: * Friedrich Gustav Jakob Henle, a German physician, pathologist and anatomist (1809–1885) ** Loop of Henle in the kidney, named after Henle *Fritz Henle, a photographer, known as "Mr. Rollei" for his use of the 2.25" square for ...
, 1841)
(Bluntnose guitarfish) ** '' Acroteriobatus leucospilus'' (
Norman Norman or Normans may refer to: Ethnic and cultural identity * The Normans, a people partly descended from Norse Vikings who settled in the territory of Normandy in France in the 10th and 11th centuries ** People or things connected with the Norm ...
, 1926)
(Grayspotted guitarfish) ** '' Acroteriobatus ocellatus'' (
Norman Norman or Normans may refer to: Ethnic and cultural identity * The Normans, a people partly descended from Norse Vikings who settled in the territory of Normandy in France in the 10th and 11th centuries ** People or things connected with the Norm ...
, 1926)
(Speckled guitarfish) ** ''
Acroteriobatus omanensis ''Acroteriobatus'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found near the shore off southern Africa and in the nor ...
'' Last, Hendeson & Naylor, 2016 (Oman guitarfish) ** ''
Acroteriobatus salalah The Salalah guitarfish (''Rhinobatos salalah'') is a species of fish in the Rhinobatidae family. It is nearly endemic to the waters off Oman, with a few records off Pakistan. Its natural habitat is open seas. Salalah (صَلَالَة Ṣalālah ...
'' ( J. E. Randall & Compagno, 1995) (Salalah guitarfish) ** ''
Acroteriobatus stehmanni ''Acroteriobatus'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found near the shore off southern Africa and in the nor ...
'' (2021) (Socotra blue-spotted guitarfish) ** ''
Acroteriobatus variegatus The stripenose guitarfish (''Acroteriobatus variegatus'') is a species of fish in the Rhinobatidae family endemic to the eastern coast of India, and Sri Lanka. Its natural habitat is in the neritic zone.Kyne, P.M., Simpfendorfer, C., Bineesh, ...
'' (
Nair The Nair , also known as Nayar, are a group of Indian Hindu castes, described by anthropologist Kathleen Gough as "not a unitary group but a named category of castes". The Nair include several castes and many subdivisions, not all of whom histor ...
& Lal Mohan, 1973)
(Stripenose guitarfish) ** ''
Acroteriobatus zanzibarensis The Zanzibar guitarfish (''Rhinobatos zanzibarensis'') is a species of fish in the Rhinobatidae family endemic to Tanzania. Its natural habitats are open sea The sea, connected as the world ocean or simply the ocean, is the body of salty ...
'' (
Norman Norman or Normans may refer to: Ethnic and cultural identity * The Normans, a people partly descended from Norse Vikings who settled in the territory of Normandy in France in the 10th and 11th centuries ** People or things connected with the Norm ...
, 1926)
(Zanzibar guitarfish) * Genus '' Eorhinobatos'' gen. nov. ** '' Eorhinobatos primaevus'' (De Zigno, 1874) comb. nov. * Genus ''
Pseudobatos ''Pseudobatos'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found in warmer coastal parts of the Americas, ranging fro ...
'' Last, Seret, and Naylor, 2016 ** ''
Pseudobatos buthi ''Pseudobatos buthi'', the spadenose guitarfish, is a ray in the family Rhinobatidae. Described in 2019 based on 82 museum specimens, little is known about this species in the wild. All 82 museum specimens were collected in the 1940s and 1950s ...
'' K.M. Rutledge, 2019 (Spadenose guitarfish) ** ''
Pseudobatos glaucostigmus ''Pseudobatos'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found in warmer coastal parts of the Americas, ranging fro ...
'' ( D. S. Jordan & C. H. Gilbert, 1883) (Speckled guitarfish) ** ''
Pseudobatos horkelii The Brazilian guitarfish (''Rhinobatos horkelii'') is a species of fish in the family Rhinobatidae. It is endemic to Brazil, where its natural habitat is coastal waters on the continental shelf. This fish is viviparous and has a long gestation p ...
'' ( J. P. Müller &
Henle Henle can refer to: * Friedrich Gustav Jakob Henle, a German physician, pathologist and anatomist (1809–1885) ** Loop of Henle in the kidney, named after Henle *Fritz Henle, a photographer, known as "Mr. Rollei" for his use of the 2.25" square for ...
, 1841)
(Brazilian guitarfish) ** ''
Pseudobatos lentiginosus ''Pseudobatos'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found in warmer coastal parts of the Americas, ranging fro ...
'' (
Garman Garman is a surname or first name. Notable people with the name include: Sports * Ann Garman, All-American Girls Professional Baseball League player * Judi Garman (born 1954), American softball coach * Mike Garman (born 1949), American baseball pla ...
, 1880)
(Atlantic guitarfish) ** ''
Pseudobatos leucorhynchus ''Pseudobatos'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to ''Rhinobatos'', recent authors treat it as distinct. They are found in warmer coastal parts of the Americas, ranging fro ...
'' ( Günther, 1867) (Whitesnout guitarfish) ** '' Pseudobatos percellens'' (
Walbaum Walbaum is a German surname. Notable people with the surname include: *Johann Julius Walbaum (1724–1799), German physician, naturalist and taxonomist * Johannes Walbaum (born 1987), German footballer * Justus Erich Walbaum (1768-1837), German typ ...
, 1792)
(Chola guitarfish) ** ''
Pseudobatos planiceps The flathead guitarfish or Pacific guitarfish (''Rhinobatos planiceps'') is a species of fish in the Rhinobatidae family. It is found in Chile, Ecuador, Peru, and possibly Nicaragua. Its natural habitats are open sea The sea, connected as ...
'' (
Garman Garman is a surname or first name. Notable people with the name include: Sports * Ann Garman, All-American Girls Professional Baseball League player * Judi Garman (born 1954), American softball coach * Mike Garman (born 1949), American baseball pla ...
, 1880)
(Pacific guitarfish) ** ''
Pseudobatos prahli ''Pseudobatos prahli'', also known by its common name Gorgona guitarfish is a species from the genus ''Pseudobatos ''Pseudobatos'' is a genus of fish in the Rhinobatidae family. Although its constituent species were previously assigned to '' R ...
'' ( Acero P & Franke, 1995) (Gorgona guitarfish) ** ''
Pseudobatos productus The shovelnose guitarfish, ''Rhinobatos productus'', is a ray in the family Rhinobatidae. It becomes mature at an estimated seven to eight years old. Males are between 90 and 100 cm long, while females are around 99 cm at that age.
'' ( Ayres, 1854) (Shovelnose guitarfish) * Genus '' Pseudorhinobatos'' gen. nov. ** '' Pseudorhinobatos dezignii'' (Heckel, 1853) comb. nov. * Genus ''
Rhinobatos ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' H. F. Linck, 1790 ** ''
Rhinobatos albomaculatus The white-spotted guitarfish (''Rhinobatos albomaculatus'') is a type of ray. It is found in the eastern Atlantic from the Gulf of Guinea to Angola , national_anthem = "Angola Avante"() , image_map = , map_caption ...
''
Norman Norman or Normans may refer to: Ethnic and cultural identity * The Normans, a people partly descended from Norse Vikings who settled in the territory of Normandy in France in the 10th and 11th centuries ** People or things connected with the Norm ...
, 1930
(white-spotted guitarfish) ** ''
Rhinobatos annandalei ''Rhinobatos annandalei'', or Annandale's guitarfish, is a type of ray. It is found in the Indian ocean around India, Pakistan, Sri Lanka and possibly the Persian Gulf The Persian Gulf ( fa, خلیج فارس, translit=xalij-e fârs, lit ...
'' Norman, 1926 (Annandale's guitarfish) ** ''
Rhinobatos borneensis ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Last, Séret & Naylor, 2016 (Borneo guitarfish) ** ''
Rhinobatos holcorhynchus ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Norman, 1922 (slender guitarfish) ** ''
Rhinobatos hynnicephalus The ringstreaked guitarfish (''Rhinobatos hynnicephalus'') is a species of fish in the Rhinobatidae family found in China, Japan, South Korea, Taiwan, and Vietnam. Its natural habitats are open seas, shallow seas, coral reefs, and estuarine A ...
'' J. Richardson, 1846 (Ringstreaked guitarfish) ** ''
Rhinobatos irvinei ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Norman, 1931 (spineback guitarfish) ** ''
Rhinobatos jimbaranensis ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Last, W. T. White & Fahmi, 2006 (Jimbaran shovelnose ray) ** ''
Rhinobatos lionotus ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Norman, 1926 (smoothback guitarfish) ** ''
Rhinobatos nudidorsalis The bareback shovelnose ray (''Rhinobatos nudidorsalis'') or nakedback guitarfish, is a species of fish in the Rhinobatidae family. It is found in Seychelles and Mauritius. Its natural habitat is open sea The sea, connected as the world oc ...
'' Last, Compagno & Nakaya, 2004 (Bareback shovelnose ray) ** ''
Rhinobatos penggali ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Last, W. T. White & Fahmi, 2006 (Indonesian shovelnose ray) ** ''
Rhinobatos punctifer ''Rhinobatos'' is a genus of fish in the Rhinobatidae family. Although previously used to encompass all guitarfishes, it was found to be polyphyletic, and recent authorities have transferred many species included in the genus to ''Acroteriobatu ...
'' Compagno & Randall, 1987 (spotted guitarfish) ** '' Rhinobatos rhinobatos''
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
,
1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of Systema Naturae, 10th edition of ''Systema Naturae'', the sta ...
(common guitarfish) ** ''
Rhinobatos sainsburyi The goldeneye shovelnose ray (''Rhinobatos sainsburyi'') is a species of fish in the Rhinobatidae family. It is endemic to north-western Australia (Monte Bello Islands to Melville Island). Its natural habitat In ecology, the term habi ...
'' Last, 2004 (goldeneye shovelnose ray) ** '' Rhinobatos schlegelii'' J. P. Müller & Henle, 1841 (brown guitarfish) ** '' Rhinobatos whitei''
Last A last is a mechanical form shaped like a human foot. It is used by shoemakers and cordwainers in the manufacture and repair of shoes. Lasts typically come in pairs and have been made from various materials, including hardwoods, cast iron, and ...
, Corrigan & Naylor, 2014
(Philippine guitarfish) * Genus †''
Myledaphus ''Myledaphus'' is a genus of Late Cretaceous cartilaginous fish whose fossils are known from Canada, the Midwest of the United States, Olmos Formation of the Difunta Group of Mexico, and the Beshtyubin and Bissekty Formations of Uzbekistan. ...
''
Cope The cope (known in Latin as ''pluviale'' 'rain coat' or ''cappa'' 'cape') is a liturgical vestment, more precisely a long mantle or cloak, open in front and fastened at the breast with a band or clasp. It may be of any liturgical colours, litu ...
, 1876
** '' Myledaphus araucanus'' Otero, 2019 ** †''
Myledaphus bipartitus ''Myledaphus'' is a genus of Late Cretaceous cartilaginous fish whose fossils are known from Canada, the Midwest of the United States, Olmos Formation of the Difunta Group of Mexico, and the Beshtyubin and Bissekty Formations of Uzbekistan. ...
'' Cope, 1876


References

{{Taxonbar, from=Q829343 .01 Ray families Rhinopristiformes Late Jurassic fish Extant Late Jurassic first appearances Taxa named by Friedrich Gustav Jakob Henle