Angolasaurus
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''Angolasaurus'' ("
Angola , national_anthem = " Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordina ...
lizard") is an extinct genus of mosasaur. Definite remains from this genus have been recovered from the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
and
Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by t ...
of
Angola , national_anthem = " Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordina ...
, and possibly the
Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by t ...
of the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
, the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
of
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
of
Niger ) , official_languages = , languages_type = National languagesPlatecarpus ''Platecarpus'' ("flat wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
'', recent phylogenetic analyses have placed it between the (then) plioplatecarpines ''
Ectenosaurus ''Ectenosaurus'' is an extinct genus of marine lizard belonging to the mosasaur family. It is classified as part of the Plioplatecarpinae subfamily alongside genera like ''Angolasaurus'' and ''Platecarpus''. ''Ectenosaurus'' is known from the Sa ...
'' and ''
Selmasaurus ''Selmasaurus'' is an extinct genus of marine lizard belonging to the mosasaur family. It is classified as part of the Plioplatecarpinae subfamily alongside genera like '' Angolasaurus'' and ''Platecarpus''. Two species are known, ''S. russelli'' ...
'', maintaining a basal position within the plioplatecarpinae. Its wide geographic range make it the one of the only
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...
s with a transatlantic range.


Description

''Angolasaurus'' was a small mosasaur, with a total length of about 4 meters (13 feet). It shared much of a body plan with its relative ''
Platecarpus ''Platecarpus'' ("flat wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
'', but with a slightly longer skull relative to body length. Its skull housed 11 maxillary teeth, 4 premaxillary teeth, and 12 dentary teeth. The phylogenetic relationship of ''Angolasaurus'' indicates that individuals of this genus possessed a tail fluke, more forward-lying nostrils, and keeled scales for hydrodynamic efficiency. Due to declining sea temperatures in the area that ''Angolasaurus'' inhabited, as well as the later Bientiaba locality, it has been hypothesized that it and the other mosasaurs inhabiting its region may have had an increased coverage of dark patterning on its dorsal surface to aid in thermoregulation.


History of discovery

First named in 1964 by Miguel Telles Antunes on the basis of a partial skull and skeleton, ''Angolasaurus'' was reassigned in 1994 to the genus ''
Platecarpus ''Platecarpus'' ("flat wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
''. This placement was subsequently struck down in studies in 2005, which nested it within the
plioplatecarpinae Plioplatecarpinae is a subfamily of mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the subfamily are informally and collectively known as "plioplatecarpines" and have been recovered from all continents, though the occu ...
. Fieldwork between 2005 and 2009 recovered at least two new ''Angolasaurus'' skeletons. One new skull as well as the type skull were CT scanned to reveal intricate details of the interior braincase that allowed for a more concrete placement within the
plioplatecarpinae Plioplatecarpinae is a subfamily of mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the subfamily are informally and collectively known as "plioplatecarpines" and have been recovered from all continents, though the occu ...
. The most recent major phylogenetic analysis conducted on the
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...
idae placed ''Angolasaurus'' as basal to the clade composed of the russellosaurine subfamilies
Tylosaurinae The Tylosaurinae are a subfamily of mosasaurs,Williston, S. W. 1897. Range and distribution of the mosasaurs with remarks on synonymy. ''Kansas University Quarterly'' 4(4):177-185. a diverse group of Late Cretaceous marine squamates. Members of t ...
and Plioplatecarpinae, and part of neither. In 2007, two individuals from the
Eagle Ford Formation The Eagle Ford Group (also called the Eagle Ford Shale) is a sedimentary rock formation deposited during the Cenomanian and Turonian ages of the Late Cretaceous over much of the modern-day state of Texas. The Eagle Ford is predominantly compos ...
of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
were described as belonging to the genus ''Angolasaurus'', one of which preserved part of the
hyoid apparatus The hyoid apparatus is the collective term used in veterinary anatomy for the bones which suspend the tongue and larynx. It consists of pairs of stylohyoid, thyrohyoid, epihyoid and ceratohyoid bones, and a single basihyoid bone. The hyoid appara ...
. The same abstract assigned two teeth previously assigned to ''
Platecarpus ''Platecarpus'' ("flat wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
'' to ''Angolasaurus''. These teeth came from the
Sergipe Sergipe (), officially State of Sergipe, is a state of Brazil. Located in the Northeast Region along the Atlantic coast of the country, Sergipe is the smallest state in Brazil by geographical area at , larger only than the Federal District. Serg ...
Basin of
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and are virtually indistinguishable from those found in the holotype of ''Angolasaurus bocagei''. These discoveries made ''Angolasaurus'' the first known
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...
genus with a transatlantic distribution.


Paleoecology


Angola

''Angolasaurus'' ''bocagei'', recovered only from the
Itombe Formation The Itombe Formation is a geological formation of the Kwanza Basin in Angola dated to the Coniacian stage of the Late Cretaceous. The environment of deposition is shallow marine. Reptile fossils have been recovered from the Tadi beds locality wi ...
, shared its habitat with the tylosaurine species ''
Tylosaurus ''Tylosaurus'' (from the ancient Greek (') 'protuberance, knob' + Greek (') 'lizard') is a genus of mosasaur, a large, predatory marine reptile closely related to modern monitor lizards and to snakes, from the Late Cretaceous. Description A ...
'' (formerly ''Mosasaurus'') ''iembeensis'' and the durophagous shallow-water
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
''
Angolachelys ''Angolachelys'' is an extinct genus of African eucryptodiran turtle which existed in Angola during the Turonian stage of the Late Cretaceous. The type species is ''Angolachelys mbaxi''. The type MGUAN-PA includes skull, jaw, and postcranial frag ...
''. Indeterminate halisaurine and
plesiosaur The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...
remains have also been recovered from this region. Terrestrial fauna consisted solely of the sauropod ''
Angolatitan ''Angolatitan'' (meaning "Angolan giant") is a genus of titanosauriform sauropod dinosaur from the Upper Cretaceous. It is also the first non-avian dinosaur discovered in Angola. The genus contains a single species, ''Angolatitan adamastor'', kn ...
''.


Niger

Known from the
Dukamaje Formation The Dukamaje Formation is a geological formation in Niger whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, David B; et al. (2004). "Dinosaur distribution ( ...
on the basis of a few vertebrae of varying ontogenetic stages, ''Angolasaurus'' coexisted here with fellow plioplatecarpine genera ''
Platecarpus ''Platecarpus'' ("flat wrist") is an extinct genus of aquatic lizards belonging to the mosasaur family, living around 84–81 million years ago during the middle Santonian to early Campanian, of the Late Cretaceous period. Fossils have been found ...
'' and ''
Plioplatecarpus ''Plioplatecarpus'' is a genus of mosasaur lizard. Like all mosasaurs, it lived in the late Cretaceous period, about 73-68 million years ago. Description ''Plioplatecarpus'' was a medium-sized mosasaur, measuring long and weighing . The eyes ...
,'' the globidensine genus ''
Igdamanosaurus ''Igdamanosaurus'', meaning "lizard from Igdaman", is an extinct genus of Cretaceous marine lizard belonging to the mosasaur family. It is classified as part of the Globidensini tribe (within the Mosasaurinae), and is like the other members of t ...
,'' the halisaurine genus ''
Halisaurus ''Halisaurus'' is an extinct genus of marine reptile belonging to the mosasaur family. The holotype, consisting of an angular and a basicranium fragment discovered near Hornerstown, New Jersey, already revealed a relatively unique combination ...
'', the mosasaurine genus ''
Mosasaurus ''Mosasaurus'' (; "lizard of the Meuse River") is the type genus (defining example) of the mosasaurs, an extinct group of aquatic squamate reptiles. It lived from about 82 to 66 million years ago during the Campanian and Maastrichtian stages o ...
'', and the
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...
id genus ''
Goronyosaurus ''Goronyosaurus'' is an extinct genus of marine lizard belonging to the mosasaur family. Fossils of ''Goronyosaurus'' are exclusively known from the Dukamaje Formation of Niger and Nigeria and also the Rima Formation of Nigeria, and are both Ma ...
''.


United States

''Angolasaurus'' is known from the
Eagle Ford Formation The Eagle Ford Group (also called the Eagle Ford Shale) is a sedimentary rock formation deposited during the Cenomanian and Turonian ages of the Late Cretaceous over much of the modern-day state of Texas. The Eagle Ford is predominantly compos ...
of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
. Other Turonian aquatic reptiles from the Eagle Ford Formation include the
plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...
''
Polyptychodon ''Polyptychodon'' (meaning 'shaped fin tooth') is a genus of pliosaurid found in Middle-Late Cretaceous marine deposits in southern England, France and Argentina. It has been considered a '' nomen dubium'' in a 2016 review. History of discovery ...
,
Libonectes ''Libonectes'' is an extinct genus of sauropterygian reptile belonging to the plesiosaur order. It is known from specimens found in the Britton Formation of Texas (USA) and the Akrabou Formation of Morocco, which have been dated to the lower Tur ...
,
Cimoliasaurus ''Cimoliasaurus'' was a plesiosaur that lived during the Late Cretaceous (Maastrichtian) of New Jersey. It grew up to long and weighed up to . Etymology The name is derived from the Greek , meaning "white chalk", and , meaning "lizard", in ...
,'' and ''
Plesiosaurus ''Plesiosaurus'' (Greek: ' ('), near to + ' ('), lizard) is a genus of extinct, large marine sauropterygian reptile that lived during the Early Jurassic. It is known by nearly complete skeletons from the Lias Group, Lias of England. It is disting ...
'', and the mosasaur ''Clidastes''. Indeterminate mosasaur and plesiosaur remains are also known from here.


References

{{Taxonbar, from=Q4763802 Russellosaurins Maastrichtian genus extinctions Mosasaurs of Africa Mosasaurs of North America Cretaceous Niger Fossils of Niger Fossil taxa described in 1964 Taxa named by Miguel Telles Antunes Mosasaurs of South America Turonian genus first appearances Coniacian genera Santonian genera Campanian genera