Carinodens Fraasi
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''Carinodens'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
marine lizard belonging to 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 ...
family. "''Carinodens''" means "keel teeth" and was named in 1969 as a replacement name for ''Compressidens'', "compressed teeth", which was already in use for a
gadilida Gadilida is an order of very small tusk shells, marine scaphopod molluscs. The species within the Gadilida are usually very much smaller than those within the other order of scaphopods, the Dentaliida Dentaliida is one of the two orders of ...
n scaphopod mollusk. ''Carinodens'' is widely considered a
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to ''
Globidens ''Globidens'' ("Globe teeth") is an extinct genus of mosasaur lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. ''Globidens alabamaensis'' was the first species of ''Globidens'' described, in a publication b ...
'' classified within the tribe
Globidensini The Globidensini or Globidentatini are a tribe of mosasaurine mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the tribe, known as "globidensins" or "globidensine mosasaurs", have been recovered from North America, Eu ...
. Like its close relative, ''Carinodens'' also possesses distinctive round, blunt teeth for crushing primitive clams and oysters. Most of the cranial elements known from the genus have been recovered from deposits in the
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
, with the only known
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l material being known from deposits of latest
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 ...
age in
Jordan Jordan ( ar, الأردن; tr. ' ), officially the Hashemite Kingdom of Jordan,; tr. ' is a country in Western Asia. It is situated at the crossroads of Asia, Africa, and Europe, within the Levant region, on the East Bank of the Jordan Rive ...
.Kaddumi H. F. 2009. The first and most complete ''Carinodens'' (Squamata: Mosasauridae) skeleton yet with a description of a new species from the Harrana Fauna. In: Fossils of the Harrana Fauna and the Adjacent Areas. Publications of the Eternal River Museum of Natural History, Amman, pp 49-64 Fragmentary fossils belonging to an unknown species of ''Carinodens'' have been found in
Crimea Crimea, crh, Къырым, Qırım, grc, Κιμμερία / Ταυρική, translit=Kimmería / Taurikḗ ( ) is a peninsula in Ukraine, on the northern coast of the Black Sea, that has been occupied by Russia since 2014. It has a pop ...
.


Description

''Carinodens'' measured approximately 3.5 metres (11.5 ft) in length and is one of the smallest known mosasaurs. It was closely related to ''
Globidens ''Globidens'' ("Globe teeth") is an extinct genus of mosasaur lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. ''Globidens alabamaensis'' was the first species of ''Globidens'' described, in a publication b ...
'', though is scantly known in comparison. The holotype specimen consists of an incomplete right
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
and most subsequently referred fossils are isolated teeth. The holotype dentary only preserves the posteriormost teeth, meaning that until recently when more comprehensive material was recovered, most of the dentition of the genus (its most distinctive feature) was unknown. ''Carinodens'' can easily be distinguished from the closely related ''Globidens'' by the compressed nature of its teeth and its relatively delicate dentary. Russell (1967) offered a brief diagnosis (due to the fragmentary nature of the fossils) of the genus, then known as ''Compressidens'': "Small projection of dentary anterior to first dentary tooth. Median dentary teeth bilaterally compressed, bicarinate, subrectangular in lateral view and with pointed apices. Anterior teeth circular in cross-section with strongly recurved pointed apices".


Dentition

By mosasaur standards, the teeth of ''Carinodens'' are unusually
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example, ...
, both in morphology and size. The alveoli show a marked size decrease between teeth #8 and #7, and the teeth themselves change dramatically in both size and morphology between #8 and #7. This is similar to the maxillary teeth of ''
Globidens dakotensis ''Globidens'' ("Globe teeth") is an extinct genus of mosasaur lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. ''Globidens alabamaensis'' was the first species of ''Globidens'' described, in a publication by ...
'' (between positions #5 and #6, though this is less pronounced than in ''Carinodens'') and in ''
Globidens alabamaensis ''Globidens'' ("Globe teeth") is an extinct genus of mosasaur lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. ''Globidens alabamaensis'' was the first species of ''Globidens'' described, in a publication by Cha ...
''.


Diet

''Carinodens'', is like its close relative ''
Globidens ''Globidens'' ("Globe teeth") is an extinct genus of mosasaur lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. ''Globidens alabamaensis'' was the first species of ''Globidens'' described, in a publication b ...
'', considered to have been a durophagous mosasaur. Because the anteriormost part of the dentary of ''Carinodens'' is relatively slender with small pointed tooth crowns, only the posteriormost five teeth actually functioned for crushing food. The anteriormost portion of the dentary was thus likely used for acquiring and handling food rather than crushing it, an idea already suggested by Dollo (1913) during the description of the type species. The maxilla of ''Carinodens'' is unknown, which hinders knowledge on the interaction between the lower and upper jaw. Dollo (1913, 1924) suggested a diet dominated by
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s, whereas Lingham-Soliar (1990, 1999) listed a wide array of potential prey items, including
belemnites Belemnites may refer to: *Belemnitida Belemnitida (or the belemnite) is an extinct order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous. Unlike squid, belemnites had an internal skeleton that made up the cone. ...
, nautilids,
bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
,
gastropods The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
, scaphopods,
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s, echinoderms and
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. These groups were abundant in the late Cretaceous seas around
Maastricht Maastricht ( , , ; li, Mestreech ; french: Maestricht ; es, Mastrique ) is a city and a municipality in the southeastern Netherlands. It is the capital and largest city of the province of Limburg. Maastricht is located on both sides of the ...
, meaning that their population numbers cannot explain the rarity of ''Carinodens''. It is possible that ''Carinodens'' spent most of its life in deep waters, only rarely swimming in shallow seas.


Classification

''Carinodens fraasi'' was first described and illustrated by Louis Dollo in 1913 as "''
Globidens ''Globidens'' ("Globe teeth") is an extinct genus of mosasaur lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. ''Globidens alabamaensis'' was the first species of ''Globidens'' described, in a publication b ...
fraasi''". Dollo later erected a separate genus, "''Compressidens''" for the species in 1924, recognising the more compressed nature of the teeth in comparison to those of ''Globidens''. Dollo also assigned ''
Bottosaurus ''Bottosaurus'' is an extinct genus of alligatorid from the Late Cretaceous-Early Paleocene of New Jersey, Texas, and possibly North Carolina and South Carolina. Two species are currently accepted, with a third requiring re-evaluation. Taxonomy ...
belgicus'', previously misinterpreted as a species of crocodilian, to the genus as ''Compressidens belgicus''. With the name ''Compressidens'' being preoccupied by a scaphopod mollusk, Thurmond (1969) proposed the substitute name ''Carinodens''. ''Carinodens'' is most frequently recovered as a sister taxon to ''Globidens'' within the
Globidensini The Globidensini or Globidentatini are a tribe of mosasaurine mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the tribe, known as "globidensins" or "globidensine mosasaurs", have been recovered from North America, Eu ...
tribe in the
Mosasaurinae The Mosasaurinae are a subfamily of mosasaurs, a diverse group of Late Cretaceous marine squamates. Members of the subfamily are informally and collectively known as "mosasaurines" and their fossils have been recovered from every continent except ...
. The cladogram below, covering the Globidensini, is based upon a summary of evolutionary adaptations in the Globidensini featured in Schulp et al. (2004): It is worth noting that placing ''
Prognathodon ''Prognathodon'' is an extinct genus of marine lizard belonging to the mosasaur family. It is classified as part of the Mosasaurinae subfamily, alongside genera like ''Mosasaurus'' and ''Clidastes''. ''Prognathodon'' has been recovered from depos ...
'' within the Globidensini is controversial, and it is most often seen as either a more basal mosasaurine or as part of its own tribe, the Prognathodontini. The primary feature distinguishing the two recognised species, ''C. fraasi'' and ''C. belgicus'' is found in their dentition. The teeth of ''C. fraasi'' are unicuspid and the teeth of ''C. belgicus'' are tricuspid. ''Carinodens'' fossils from Jordan, consisting of an almost complete skull with at least 24 teeth still occupying their natural locations, a complete neck
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
l series as well as several back vertebrae, and front paddles were reported by Kaddumi (2009). In addition to the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
,
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
ry, and
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
ry teeth, several small
pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medial ...
teeth were also recovered from the same specimen. Kaddumi (2009) fully described the remains and referred them to a new species of ''Carinodens''. Based on the remarkable dental
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example, ...
y exhibited in the new species from Jordan, several previously not considered prey items may be postulated for ''Carinodens'' (Kaddumi 2009).


References

{{Taxonbar, from=Q1622418 Mosasaurines Mosasaurs of Europe Mosasaurs of Asia Fossil taxa described in 1969