Pluridens
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''Pluridens'' ("many teeth") is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of marine lizard belonging to the
Mosasauridae Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains were discovered in ...
. ''Pluridens'' is placed in the subfamily
Halisaurinae The Halisaurinae are a subfamily of mosasaurs, a group of Late Cretaceous marine lizards. They were small to medium-sized, ranging from just under 3 meters in ''Eonatator sternbergi'' to as much as 8 or 9 meters in ''Pluridens serpentis''. The ...
with the genera '' Phosphorosaurus'', ''
Eonatator ''Eonatator'' is an extinct genus of marine lizard belonging to the mosasaur family. It is a close relative of '' Halisaurus'', and part of the same subfamily, the Halisaurinae. It is known from the Late Cretaceous of North America, Colombia and ...
'' and ''
Halisaurus ''Halisaurus'' is an extinct genus of mosasaur named by Othniel Charles Marsh in 1869. The holotype, consisting of an angular and a basicranium fragment discovered near Hornerstown, New Jersey, already revealed a relatively unique combination ...
''.Konishi, Takuya; Caldwell, Michael W.; Nishimura, Tomohiro; Sakurai, Kazuhiko; Tanoue, Kyo (2015). "A new halisaurine mosasaur (Squamata: Halisaurinae) from Japan: the first record in the western Pacific realm and the first documented insights into binocular vision in mosasaurs". Journal of Systematic Palaeontology: 1–31. doi:10.1080/14772019.2015.1113447. Compared to related halisaurines, ''Pluridens'' had longer jaws with more teeth, and smaller eyes. It also grew large size, measuring long and perhaps over in some individuals. The jaws in some specimens are robust, and sometimes show injuries suggestive of combat. The jaws may have been used for fighting over mates or territories. ''Pluridens'' lived in the shallow seas of
West Africa West Africa, also known as Western Africa, is the westernmost region of Africa. The United Nations geoscheme for Africa#Western Africa, United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Gha ...
during the late
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
-
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
. Three species of are known, ''P. walkeri'', ''P. calabaria'', and ''P. serpentis.'' ''P. walkeri'' is known from the Maastrichtian of southwest
Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...
,T. Lingham-Soliar. 1998. A new mosasaur ''Pluridens'' walkeri from the Upper Cretaceous, Maastrichtian of the Iullemmeden Basin, southwest Niger. Journal of Vertebrate Paleontology 18(4):709-707. whereas ''P. calabaria'' is found in slightly older (late Campanian) deposits in
Nigeria Nigeria, officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf of Guinea in the Atlantic Ocean to the south. It covers an area of . With Demographics of Nigeria, ...
.Nicholas R. Longrich (2016). "A new species of ''Pluridens'' (Mosasauridae: Halisaurinae) from the upper Campanian of Southern Nigeria". Cretaceous Research. in press. doi:10.1016/j.cretres.2016.03.013. ''P. serpentis'' was found in
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
deposits in
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
. ''Pluridens'' was briefly synonymized with ''
Halisaurus ''Halisaurus'' is an extinct genus of mosasaur named by Othniel Charles Marsh in 1869. The holotype, consisting of an angular and a basicranium fragment discovered near Hornerstown, New Jersey, already revealed a relatively unique combination ...
'' by Lindgren and Siverson (2005),Lindgren J, Siverson M. 2005.'' Halisaurus sternbergii'', a small mosasaur with an intercontinental distribution. Journal of Paleontology 79 (4): 763–773. but subsequent studies rejected the synonymy.Michael J. Polcyn, Johan Lindgren, Nathalie Bardet, Dirk Cornelissen, Louis Verding, and Anne S. Schulp (2012) Description of new specimens of ''Halisaurus arambourgi'' Bardet & Pereda Suberbiola, 2005 and the relationships of Halisaurinae. Bulletin de la Société Géologique de France 183:123-136 doi:10.2113/gssgfbull.183.2.123 http://bsgf.geoscienceworld.org/content/183/2/123.abstract


Description

''Pluridens'' was a moderately large mosasaur at roughly in length if the proportions of the dentary and the rest of the animal matches that of ''
Halisaurus ''Halisaurus'' is an extinct genus of mosasaur named by Othniel Charles Marsh in 1869. The holotype, consisting of an angular and a basicranium fragment discovered near Hornerstown, New Jersey, already revealed a relatively unique combination ...
''; the referred material suggests an even larger body size at more than in length. A feature that separates ''Pluridens'' not only from the rest of the
Halisaurinae The Halisaurinae are a subfamily of mosasaurs, a group of Late Cretaceous marine lizards. They were small to medium-sized, ranging from just under 3 meters in ''Eonatator sternbergi'' to as much as 8 or 9 meters in ''Pluridens serpentis''. The ...
, but from the rest of the
Mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
idae at large is its unusually high tooth count,
dentaries In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
assigned to the genus preserve almost twice the amount of teeth found in most other mosasaur genera. Additionally, the dentary of ''Pluridens'' also has one of the slenderest profiles among the mosasaurs; the only genus having a similarly narrow dentary being the mosasaurine ''
Plotosaurus ''Plotosaurus'' ("swimmer lizard") is an extinct genus of large mosasaurs which lived during the Upper Cretaceous (Maastrichtian) in what is now North America. The taxon was initially described by University of California, Berkeley, Berkeley pale ...
''. This combination of features indicate a feeding niche and lifestyle unique among the mosasaurs. The shallow dentary and small teeth of ''Pluridens'' was likely elongated relative to the rest of the head to allow for the massive battery of teeth. Due to force exerted by the lower jaw when biting against the upper jaw would decrease with the distance from the joint, the anterior teeth would have been ineffectual for biting and crushing. Lingham-Soliar (1998) likened the lifestyle and feeding of ''Pluridens'' with that of the
ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s, particularly early ichthyosaurs such as ''
Temnodontosaurus ''Temnodontosaurus'' (meaning "cutting-tooth lizard") is an extinct genus of large ichthyosaurs that lived during the Lower Jurassic in what is now Europe and possibly Chile. The first known fossil is a specimen consisting of a complete skull a ...
'' and ''
Leptonectes ''Leptonectes'' is a genus of ichthyosaur that lived in the Late Triassic to Early Jurassic (Rhaetian - Pliensbachian). Fossils have been found in Belgium, Germany, Spain and the United Kingdom The United Kingdom of Great Britain and ...
''. Lingham-Soliar (
1998 1998 was designated as the ''International Year of the Ocean''. Events January * January 6 – The ''Lunar Prospector'' spacecraft is launched into orbit around the Moon, and later finds evidence for Lunar water, frozen water, in soil i ...
) listed the following unambiguous character states for the genus: "Very long, slender dentary, anterior half cross-section almost circular extending to tip. Twenty eight-plus teeth in dentary, close-set. Tooth crowns short, no distinguishable carinae, broad, smooth anterior surface, hairline crenations posterior, slightly discernible broad facets, tips posteriorly recurved; teeth virtually uniform in size along most of the dental ramus; replacement teeth emerge medial to mature tooth crowns; resorption pits on mid-lingual side of tooth base".


Classification

The unusually high amount of teeth present in the dentaries of both species of ''Pluridens'' initially made it difficult to immediately assign a classification on the subfamily level. Lingham-Soliar (1998) remarked that the unique combination of features (e.g. the high tooth count and a unique tooth shape) would normally result in the creation of a completely new subfamily, but did not create a new subfamily due to the rather fragmentary nature of the remains. Lindgren and Siverson (2005) considered many of the unique features of ''Pluridens'' to be shared with ''
Halisaurus ''Halisaurus'' is an extinct genus of mosasaur named by Othniel Charles Marsh in 1869. The holotype, consisting of an angular and a basicranium fragment discovered near Hornerstown, New Jersey, already revealed a relatively unique combination ...
'' and reassigned ''P. walkeri'' as "''Halisaurus walkeri''". The genus synonymy was rejected by later authors, though similarities between the genera and the discovery of distinctly halisaurine vertebrae in
Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...
allowed the genus to be firmly placed within the
Halisaurinae The Halisaurinae are a subfamily of mosasaurs, a group of Late Cretaceous marine lizards. They were small to medium-sized, ranging from just under 3 meters in ''Eonatator sternbergi'' to as much as 8 or 9 meters in ''Pluridens serpentis''. The ...
. Thus, ''Pluridens'' is today considered a highly derived halisaurine.


Species and specimens

Three species are known, ''Pluridens walkeri'' from the early Maastrichtian of Niger, ''P. calabaria'' from the Campanian of Nigeria, and ''P. serpentis'' from the late Maastrichtian of Morocco. The type specimen of ''P. walkeri'' is BMNH R14153, an almost complete dentary recovered from the Maastrichtian Farin-Doutchi Formation near Mount Ilatarda,
Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...
. Lingham-Soliar (1998) also assigned BMNH R9804, a partial dentary with poorly preserved teeth (one out of which is almost complete) from the late Campanian of the Nkporo Formation near
Calabar Calabar (also referred to as Callabar, Calabari, Calbari, Cali and Kalabar) is the capital city of Cross River State, Nigeria. It was originally named Akwa Akpa, in the Efik language, as the Efik people dominate this area. The city is adjac ...
in southern
Nigeria Nigeria, officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf of Guinea in the Atlantic Ocean to the south. It covers an area of . With Demographics of Nigeria, ...
, to ''P. walkeri''. The Nigerian specimen BMNH R9804 was reassigned to its own species by Longrich (
2016 2016 was designated as: * International Year of Pulses by the sixty-eighth session of the United Nations General Assembly. * International Year of Global Understanding (IYGU) by the International Council for Science (ICSU), the Internationa ...
) as it lacks many of the derived features that characterize ''P. walkeri'', such as a high tooth count and extreme elongation of the dentary. The Calabar dentary also lack the extreme lateral protrusion and subcircular section of the dentary and the strong transverse expansion of the dental thecae, both of which are present in BMNH R14153. Longrich (2016) dubbed the new species ''Pluridens calabaria''. It may be a direct ancestor to ''P. walkeri''. A third species, ''Pluridens serpentis,'' was described from the late Maastrichtian phosphates of Morocco in 2021. It is the most completely known species, being represented by two skulls and a number of jaws. Longrich, N. R., Bardet, N., Khaldoune, F., Yazami, O. K., & Jalil, N.-E. (2021). ''Pluridens serpentis'', a new mosasaurid (Mosasauridae: Halisaurinae) from the Maastrichtian of Morocco and implications for mosasaur diversity. Cretaceous Research, 104882.


References

{{Taxonbar, from=Q23655541 Russellosaurins Mosasaurs of Africa Fossil taxa described in 1998 Campanian genus first appearances Maastrichtian genus extinctions Taxa_named_by_Nicholas_R._Longrich