Mosasaur

Mosasaurs (from Latin Mosa meaning the 'Meuse river', and Greek σαύρος sauros meaning 'lizard') comprise a group of extinct, large marine reptiles containing 38 genera in total. Their first fossil remains were discovered in a limestone quarry at Maastricht on the Meuse in 1764. Mosasaurs probably evolved from an extinct group of aquatic lizards[1] known as aigialosaurs in the Early Cretaceous. During the last 20 million years of the Cretaceous period (Turonian-Maastrichtian ages), with the extinction of the ichthyosaurs and pliosaurs, mosasaurs became the dominant marine predators. They became extinct as a result of the K-Pg event at the end of the Cretaceous period, about 66 million years ago.

Mosasaurs
Temporal range: Early-Late Cretaceous, 101–66 Ma
Platecarpus planifrons Clean
Mounted skeleton of a plioplatecarpine (Plesioplatecarpus planifrons), Rocky Mountain Dinosaur Resource Center
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Clade: Pythonomorpha
Superfamily: Mosasauroidea
Gervais, 1853
Subgroups

Description

Platecarpus tympaniticus
Life restoration of a mosasaur (Platecarpus tympaniticus) informed by fossil skin impressions

Mosasaurs breathed air, were powerful swimmers, and were well-adapted to living in the warm, shallow inland seas prevalent during the Late Cretaceous period. Mosasaurs were so well adapted to this environment that they gave birth to live young, rather than returning to the shore to lay eggs as sea turtles do.[2]

The smallest-known mosasaur was Dallasaurus turneri, which was less than 1 m (3.3 ft) long. Larger mosasaurs were more typical, with many species growing longer than 4 m (13 ft). Mosasaurus hoffmannii, the largest known species, may have reached up to 17 m (56 ft) in length.[3] Currently, the largest publicly exhibited mosasaur skeleton in the world is on display at the Canadian Fossil Discovery Centre in Morden, Manitoba. The specimen, nicknamed "Bruce", is just over 13 m (43 ft) long.[4]

Mosasaurs had a body shape similar to that of modern-day monitor lizards (varanids), but were more elongated and streamlined for swimming. Their limb bones were reduced in length and their paddles were formed by webbing between their long finger and toe bones. Their tails were broad, and supplied their locomotive power. Until recently, mosasaurs were assumed to have swum in a method similar to the one used today by conger eels and sea snakes, undulating their entire bodies from side to side. However, new evidence suggests that many advanced mosasaurs had large, crescent-shaped flukes on the ends of their tails, similar to those of sharks and some ichthyosaurs. Rather than use snake-like undulations, their bodies probably remained stiff to reduce drag through the water, while their tails provided strong propulsion.[5] These animals may have lurked and pounced rapidly and powerfully on passing prey, rather than chasing after it.[6]

Early reconstructions showed mosasaurs with dorsal crests running the length of their bodies, which were based on misidentified remains of tracheal cartilage. By the time this error was discovered, depicting mosasaurs with such crests in artwork had already become a trend.[7][8]

Paleobiology

Placenticeratidae - Placenticeras whitfieldi
Fossil shell of ammonite Placenticeras whitfieldi showing punctures caused by the bite of a mosasaur, Peabody Museum of Natural History, Yale
Mosasaur tooth
A tooth from a mosasaur

Mosasaurs had double-hinged jaws and flexible skulls (much like those of snakes), which enabled them to gulp down their prey almost whole. A skeleton of Tylosaurus proriger from South Dakota included remains of the diving seabird Hesperornis, a marine bony fish, a possible shark, and another, smaller mosasaur (Clidastes). Mosasaur bones have also been found with shark teeth embedded in them.

One of the food items of mosasaurs were ammonites, molluscs with shells similar to those of Nautilus, which were abundant in the Cretaceous seas. Holes have been found in fossil shells of some ammonites, mainly Pachydiscus and Placenticeras. These were once interpreted as a result of limpets attaching themselves to the ammonites, but the triangular shape of the holes, their size, and their presence on both sides of the shells, corresponding to upper and lower jaws, is evidence of the bite of medium-sized mosasaurs. Whether this behaviour was common across all size classes of mosasaurs is not clear.

Virtually all forms were active predators of fish and ammonites; a few, such as Globidens, had blunt, spherical teeth, specialized for crushing mollusk shells. The smaller genera, such as Platecarpus and Dallasaurus, which were about 1–6 m (3.3–19.7 ft) long, probably fed on fish and other small prey. The smaller mosasaurs may have spent some time in fresh water, hunting for food. The larger mosasaurs, such as Tylosaurus, Hainosaurus and Mosasaurus, reached sizes of 10–15 m (33–49 ft) long and were apex predators of the Late Cretaceous oceans, attacking other marine reptiles, as well as preying on large fish and ammonites.

Soft tissue

Tylosaurus skin
Scales of Tylosaurus proriger (KUVP-1075)

Despite the many mosasaur remains collected worldwide, knowledge of the nature of their skin coverings remains in its early stages. Few mosasaurid specimens collected from around the world retain fossilized scale imprints. This lack may be due to the delicate nature of the scales, which nearly eliminates the possibility of preservation, in addition to the preservation sediment types and the marine conditions under which the preservation occurred. Until the discovery of several mosasaur specimens with remarkably well-preserved scale imprints from late Maastrichtian deposits of the Muwaqqar Chalk Marl Formation of Harrana[9] in Jordan, knowledge of the nature of mosasaur integument was mainly based on very few accounts describing early mosasaur fossils dating back to the upper Santonian–lower Campanian, such as the famous Tylosaurus specimen (KUVP-1075) from Gove County, Kansas.[10] Material from Jordan has shown that the bodies of mosasaurs, as well as the membranes between their fingers and toes, were covered with small, overlapping, diamond-shaped scales resembling those of snakes. Much like those of modern reptiles, mosasaur scales varied across the body in type and size. In Harrana specimens, two types of scales were observed on a single specimen: keeled scales covering the upper regions of the body and smooth scales covering the lower.[9] As ambush predators, lurking and quickly capturing prey using stealth tactics,[11] they may have benefited from the nonreflective, keeled scales.[9]

Soft tissue structures in Platecarpus
Soft tissues in the head and neck of Platecarpus tympaniticus specimen LACM 128319: Tracheal rings are shown in the bottom three photographs.

More recently, a fossil of Platecarpus tympaniticus has been found that preserved not only skin impressions, but also internal organs. Several reddish areas in the fossil may represent the heart, lungs, and kidneys. The trachea is also preserved, along with part of what may be the retina in the eye. The placement of the kidneys is farther forward in the abdomen than it is in monitor lizards, and is more similar to those of cetaceans. As in cetaceans, the bronchi leading to the lungs run parallel to each other instead of splitting apart from one another as in monitors and other terrestrial reptiles. In mosasaurs, these features may be internal adaptations to fully marine lifestyles.[5]

Prognathodon tissue
Fibrous tissues and microstructures recovered from Prognathodon specimen IRSNB 1624

In 2011, collagen protein was recovered from a Prognathodon humerus dated to the Cretaceous.[12]

In 2005, a case study by A.S. Schulp, E.W.A Mulder, and K. Schwenk outlined the fact that mosasaurs had paired fenestrae in their palates. In monitor lizards and snakes, paired fenestrae are associated with a forked tongue, which is flicked in and out to detect chemical traces and provide a directional sense of smell. They therefore proposed that mosasaurs probably also had a sensitive forked tongue.[13]

Metabolism

A study published in 2016 by T. Lyn Harrell, Alberto Pérez-Huerta and Celina Suarez showed that mosasaurs were endothermic. The study contradicted findings published in 2010 indicating mosasaurs were ectothermic. The 2010 study did not use warm-blooded animals for comparison but analogous groups of common marine animals. Based on comparisons with modern warm-blooded animals and fossils of known cold-blooded animals from the same time period, the 2016 study found mosasaurs likely had body temperatures similar to those of contemporary seabirds and were able to internally regulate their temperatures to remain warmer than the surrounding water.[14]

Coloration

The coloration of mosasaurs was unknown until 2014, when the findings of Johan Lindgren of Lund University and colleagues revealed the pigment melanin in the fossilized scales of a mosasaur. Mosasaurs were likely countershaded, with dark backs and light underbellies, much like a great white shark or leatherback sea turtle, the latter of which had fossilized ancestors for which color was also determined. The findings were described in the journal Nature.[15]

Ontogeny and growth

Mosasaur growth is not well understood, as specimens of juveniles are rare, and many were mistaken for hesperornithine birds when discovered 100 years ago. However, the discovery of several specimens of juvenile and neonate-sized mosasaurs unearthed more than a century ago indicate that mosasaurs gave birth to live young, and that they spent their early years of life out in the open ocean, not in sheltered nurseries or areas such as shallow water as previously believed. Whether mosasaurs provided parental care, like other marine reptiles such as plesiosaurs, is currently unknown. The discovery of young mosasaurs was published in the journal Palaeontology.[16]

Environment

Paleotologists compared the taxonomic diversity and patterns of morphological disparity in mosasaurs with sea level, sea surface temperature, and stable carbon isotope curves for the Upper Cretaceous to explore factors that may have influenced their evolution. No single factor unambiguously accounts for all radiations, diversification, and extinctions; however, the broader patterns of taxonomic diversification and morphological disparity point to niche differentiation in a “fishing up” scenario under the influence of “bottom-up” selective pressures. The most likely driving force in mosasaur evolution was high productivity in the Late Cretaceous, driven by tectonically controlled sea levels and climatically controlled ocean stratification and nutrient delivery. When productivity collapsed at the end of the Cretaceous, coincident with bolide impact, mosasaurs became extinct.[17]

Fragment čelisti mosasaura z Dolního Újezda u Litomyšle
Fossil jaw fragment of a mosasaurid reptile from Dolní Újezd by Litomyšl, Czech Republic.

Sea levels were high during the Cretaceous period, causing marine transgressions in many parts of the world, and a great inland seaway in what is now North America. Mosasaur fossils have been found in the Netherlands, Belgium, Denmark, Portugal, Sweden, South Africa, Spain, France, Germany, Poland, the Czech Republic,[18] Bulgaria, the United Kingdom,[19][20] Russia, Ukraine, Kazakhstan, Azerbaijan,[21] Japan,[22] Egypt, Israel, Jordan, Syria,[23] Turkey,[24] Niger,[25][26] Angola, Morocco, Australia, New Zealand, and on Vega Island off the coast of Antarctica. Tooth taxon Globidens timorensis is known from the island of Timor; however, the phylogenetic placement of this species is uncertain and it might not even be a mosasaur.[27] Mosasaurs have been found in Canada in Manitoba and Saskatchewan[28] and in much of the contiguous United States. Complete or partial specimens have been found in Alabama, Mississippi, New Jersey, Tennessee, and Georgia, as well as in states covered by the Cretaceous seaway: Texas, southwest Arkansas, New Mexico, Kansas,[29] Colorado, Nebraska, South Dakota, Montana, Wyoming, and the Pierre Shale/Fox Hills formations of North Dakota.[30] Lastly, mosasaur bones and teeth are also known from California, Mexico, Colombia,[31] Brazil,[23] Peru, and Chile.[32]

Many of the so-called 'dinosaur' remains found on New Zealand are actually mosasaurs and plesiosaurs, both being Mesozoic predatory marine reptiles.

Discovery

Mosasaurus
The Mosasaurus hoffmannii skull found in Maastricht between 1770 and 1774

The first publicized discovery of a partial fossil mosasaur skull in 1764 by quarry workers in a subterranean gallery of a limestone quarry in Mount Saint Peter, near the Dutch city of Maastricht, preceded any major dinosaur fossil discoveries, but remained little known. However, a second find of a partial skull drew the Age of Enlightenment's attention to the existence of fossilized animals that were different from any known living creatures. When the specimen was discovered between 1770 and 1774, Johann Leonard Hoffmann, a surgeon and fossil collector, corresponded about it with the most influential scientists of his day, making the fossil famous. The original owner, though, was Godding, a canon of Maastricht cathedral.

When the French revolutionary forces occupied Maastricht in 1794, the carefully hidden fossil was uncovered, after a reward, it is said, of 600 bottles of wine, and transported to Paris. After it had been earlier interpreted as a fish, a crocodile, and a sperm whale, the first to understand its lizard affinities was the Dutch scientist Adriaan Gilles Camper in 1799. In 1808, Georges Cuvier confirmed this conclusion, although le Grand Animal fossile de Maëstricht was not actually named Mosasaurus ('Meuse reptile') until 1822 and not given its full species name, Mosasaurus hoffmannii, until 1829. Several sets of mosasaur remains, which had been discovered earlier at Maastricht but were not identified as mosasaurs until the 19th century, have been on display in the Teylers Museum, Haarlem, procured from 1790.

The Maastricht limestone beds were rendered so famous by the mosasaur discovery, they have given their name to the final six-million-year epoch of the Cretaceous, the Maastrichtian.

Relationships

Cladogram of the Mosasauridae modified from Simões et al. (2017):[33]

Mosasauridae
Russellosaurina
Tethysaurinae

Tethysaurus nopcsai

Pannoniasaurus osii

Yaguarasaurinae

Yaguarasaurus columbianus

Russellosaurus coheni

Romeosaurus fumanensis

Tylosaurinae

Tylosaurus nepaeolicus

Tylosaurus bernardi

Tylosaurus proriger

Taniwhasaurus oweni

Taniwhasaurus antarcticus

Plioplatecarpinae

Angolasaurus bocagei

Selmasaurus johnsoni

Ectenosaurus clidastoides

Plesioplatecarpus planifrons

Latoplatecarpus willistoni

Platecarpus tympaniticus

Plioplatecarpus

Halisaurinae

Halisaurus platyspondylus

Eonatator sternbergi

Mosasaurinae

Dallasaurus turneri

Clidastes liodontus

Clidastes moorevillensis

Clidastes propython

Globidens dakotensis

Globidens alabamaensis

Prognathodon rapax

Prognathodon overtoni

Prognathodon saturator

Prognathodon currii

Prognathodon solvayi

Mosasaurini

Prognathodon waiparaensis

Prognathodon kianda

Eremiasaurus heterodontus

Plesiotylosaurus crassidens

Mosasaurus conodon

Mosasaurus missouriensis

Mosasaurus hoffmannii

Plotosaurus bennisoni

Evolutionary history

Aigialosaurus bucchichi
Restoration of Opetiosaurus bucchichi, a basal mosasauroid.
GlobidensDB2
Life restoration of a mosasaurine, Globidens alabamaensis
Plotosaurus ben1DB
Life restoration of a mosasaurine, Plotosaurus bennisoni
Tylosaurus pembinensis 1DB
Restoration of a tylosaurine, Tylosaurus pembinensis

Based on features such as the double row of pterygoid ("flanged") teeth on the palate, the loosely hinged jaw, modified/reduced limbs and probable methods of locomotion, many researchers believe that snakes share a common marine ancestry with mosasaurs, a suggestion advanced in 1869 by Edward Drinker Cope, who coined the term Pythonomorpha to unite them. The idea lay dormant for more than a century, to be revived in the 1990s.[34][35] Recently, the discovery of Najash rionegrina, a fossorial snake from South America, cast doubt on the marine origin hypothesis.

The skeleton of Dallasaurus turneri, described by Bell and Polcyn (2005), has a mixture of features present in the skeletons of derived mosasaurs and in the skeletons of mosasaurid ancestors, such as aigialosaurids. Dallasaurus retains facultatively terrestrial limbs similar in their structure to the limbs of aigialosaurids and terrestrial squamates (plesiopedal limb condition), unlike derived mosasaurids, which evolved paddle-like limbs (hydropedal limb condition). However, the skeleton of Dallasaurus simultaneously had several characters that linked it with derived members of the subfamily Mosasaurinae; the authors of its description listed "invasion of the parietal by medial tongues from the frontal, teeth with smooth medial enamel surface, high coronoid buttress on surangular, interdigitate anterior scapulo-coracoid suture, humeral postglenoid process, elongate atlas synapophysis, sharp anterodorsal ridge on synapophyses, vertically oriented vertebral condyles, elongate posterior thoracic vertebrae, and fused haemal arches" as the characters uniting Dallasaurus with Mosasaurinae.[36] The phylogenetic analysis conducted by Bell and Polcyn indicated that hydropedal mosasaurids did not form a clade that wouldn't also include plesiopedal taxa, such as Dallasaurus, Yaguarasaurus, Russellosaurus, Tethysaurus, Haasiasaurus and Komensaurus (in 2005 only informally known as "Trieste aigialosaur"); the analysis indicated that hydropedal limb condition evolved independently in three different groups of mosasaurs (Halisaurinae, Mosasaurinae and the group containing the subfamilies Tylosaurinae and Plioplatecarpinae).[36][37] The result of this phylogenetic study was subsequently mostly confirmed by the analyses conducted by Caldwell and Palci (2007) and Leblanc, Caldwell and Bardet (2012);[38][39] the analysis conducted by Makádi, Caldwell and Ősi (2012) indicated that hydropedal limb condition evolved independently in two groups of mosasaurs (in Mosasaurinae and in the clade containing Halisaurinae, Tylosaurinae and Plioplatecarpinae).[40] Conrad et al. (2011), on the other hand, recovered hydropedal mosasaurs forming a clade that excluded their plesiopedal relatives.[41] If the hypothesis of Bell and Polcyn (2005) is correct, then mosasaurs in the traditional sense of the word, i.e. "lizards that evolved paddle-like limbs and radiated into aquatic environments in the late Mesozoic, going extinct at the end of that era",[37] are actually polyphyletic; Bell and Polcyn (2005) maintained monophyletic Mosasauridae by including Dallasaurus and other aforementioned plesiopedal taxa in the family as well,[36] while Caldwell (2012) suggested (though explicitly stated that it was not "a formal proposal of new nomenclature") to restrict Mosasauridae only to the genus Mosasaurus and its closest hydropedal relatives.[37]

The exact phylogenetic position of the clade containing mosasaurids and their closest relatives (aigialosaurids and dolichosaurs) within Squamata remains uncertain. Some cladistic analyses recovered them as the closest relatives of snakes,[42][43] taking into account similarities in jaw and skull anatomies;[42] however, this has been disputed[44][45][46] and the morphological analysis conducted by Conrad (2008) recovered them as varanoids closely related to terrestrial monitor lizards instead.[44] Subsequent analysis of anguimorph relationships conducted by Conrad et al. (2011) based on morphology alone recovered mosasaurids, aigialosaurids and dolichosaurs as anguimorphs lying outside the least inclusive clade containing monitor lizards and helodermatids; the analysis based on combined datasets of morphological and molecular data, on the other hand, found them more closely related to monitor lizards and the earless monitor lizard than to helodermatids and the Chinese crocodile lizard.[41] The large morphological analysis conducted by Gauthier et al. (2012) recovered mosasaurids, aigialosaurids and dolichosaurids in an unexpected position as basal members of the clade Scincogekkonomorpha (containing all taxa sharing a more recent common ancestor with Gekko gecko and Scincus scincus than with Iguana iguana[44]) that didn't belong to the clade Scleroglossa. The phylogenetic position of these taxa turned out to be highly dependent on which taxa were included in or excluded from the analysis. When mosasaurids were excluded from the analysis, dolichosaurs and aigialosaurids were recovered within Scleroglossa, forming a sister group to the clade containing snakes, amphisbaenians, dibamids and the American legless lizard. When mosasaurids were included in the analysis, and various taxa with reduced or absent limbs other than snakes (such as dibamids or amphisbaenians) were excluded, mosasaurids, aigialosaurids and dolichosaurs were recovered inside Scleroglossa forming the sister group to snakes.[47] Longrich, Bhullar and Gauthier (2012) conducted a morphological analysis of squamate relationships using a modified version of the matrix from the analysis of Gauthier et al. (2012); they found the phylogenetic position of the clade containing mosasaurs and their closest relatives within Squamata to be highly unstable, with the clade "variously being recovered outside Scleroglossa (as in Gauthier et al., 2012) or alongside the limbless forms".[48]

Distribution

Though no individual genus or subfamily is found worldwide, the Mosasauridae as a whole achieved global distribution during the Late Cretaceous with many locations typically having complex mosasaur faunas with multiple different genera and species in different ecological niches.

Two African countries are particularly rich in mosasaurs: Morocco[49] and Angola.[50][51]

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External links

Carinodens

Carinodens is an extinct genus of Cretaceous marine lizard belonging to the mosasaur 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 gadilidan scaphopod mollusk.Carinodens is widely considered a sister taxon to Globidens classified within the tribe Globidensini. 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, with the only known postcranial material being known from deposits of latest Maastrichtian age in Jordan.

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 exclusively known from the Santonian of Kansas and Texas.

The generic name means "Drawn-out lizard", from Greek ectenes ("drawn-out") and Greek sauros ("lizard") referencing the elongated muzzle.

Eremiasaurus

Eremiasaurus is a genus of extinct genus of Cretaceous marine lizard belonging to the mosasaur family. It is classified as part of the Mosasaurini tribe (within the Mosasaurinae) and is exclusively known from the Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco. Eremiasaurus means "desert lizard", referring to the arid climate of present-day Morocco where its fossils were recovered.

One species is known, E. heterodontus, whose specific name refers to high degree of heterodonty exhibited compared to other species of mosasaur.

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 are Maastrichtian in age. Its fossils were first described in the 1930s as Mosasaurus nigeriensis, but subsequent remains revealed a highly unique set of adaptations that prompted the species to be reclassified as the only species of the new genus Goronyosaurus in 1972. These unique adaptations have made Goronyosaurus notoriously difficult to classify within the Mosasauridae and it is often left out of phylogenetic analyses.

Goronyosaurus possesses unique teeth, which are unlike the teeth of any other mosasaur. Instead of the cutting teeth common among mosasaurs, Goronyosaurus has straight teeth with rounded apices adapted for smashing food.

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 of features and prompted a new genus to be described. It was named by Othniel Charles Marsh in 1869 and means "ocean lizard". It was renamed by Marsh to Baptosaurus in 1870, since he believed the name to already be preoccupied by the fish Halosaurus. According to modern rules, a difference of a letter is enough and the substitute name is unneeded, making "Baptosaurus" a junior synonym.

Since its description, more complete remains have been uncovered from fossil deposits throughout the world with particularly complete remains found in Morocco and the United States. The genus remains a key taxon in mosasaur systematics due to its unique set of features and as the most complete representative of its subfamily, the Halisaurinae.

With a length of 3–4 m (9.8–13.1 ft), Halisaurus was comparatively small by mosasaur standards. Though bigger than earlier and more basal mosasaurs, such as Dallasaurus, the sleek Halisaurus would have been dwarfed by many of its contemporaries, such as Tylosaurus and larger species of Clidastes.

Harrana

Harrana, named after Qasr Kharana, an archeological Umayyad desert palace in the area, is part of the Jordan eastern plateau some 60 kilometers southeast of Amman city.

The area is largely uninhabited except for seasonal bedouin sheep and camel herders who bring their livestock to the area during December through April.

Harrana's climate, much like the other parts of the country, is influenced by the moderate Mediterranean climate from the west, the very hot Sahara's climate from the east, and the cold European climate from the north.

Though arid, the area is rich in animal life. Birds, owls, rodents, rabbits, foxes, occasional wolves and hyenas, snakes, and lizards are some of the animals that take refuge in Harrana. A variety of flowering plants bloom during late winter–early spring months including mustard plants, oriental popies, and wild irisis. Cistanche tubulosa, or the desert broomrape, is also another beautiful resident in Harrana blooming towards the end of spring and beginning of summer.

Harrana is significant for its fossil deposits preserved in gigantic limestone concretions that date back to the latest Maastrichtian some 66–67 million years ago, a period notably close to the end-Cretaceous extinction events when many groups of animals such as dinosaurs and as much as 65–70% of all marine animal species became extinct. Mosasaur specimens along with their remarkably well preserved scale imprints have been discovered from late Maastrichtian deposits of the Muwaqqar Chalk Marl Formation of Harrana

The best preserved and complete specimens of the extinct teleostean fish genus Saurocephalus and the most complete mosasaur Carinodens remains come from the latest Maastrichtian of Harrana.

Kourisodon

Kourisodon ("razor tooth") is an extinct genus of mosasaur. Fossils have been found from Vancouver Island in British Columbia, Canada, as well as from the Izumi Group of Japan. These finds date back to the late Santonian stage and the late Campanian to the late Maastrichtian, respectively, of the Late Cretaceous. Kourisodon was originally described as a member of the "Leiodontini", more recently as a "Clidastine".

Liodon

Liodon ("smooth tooth") is a dubious genus of mosasaur from the Late Cretaceous. Remains assigned to this genus have been found in Africa, Asia, Europe, North and South America, and New Zealand. Though dubious and of uncertain phylogenetic affinities, Liodon was historically a highly important taxon in mosasaur systematics, being one of the genera on which the family Mosasauridae was based.

List of mosasaur-bearing stratigraphic units

This is a list of stratigraphic units from which mosasaur body fossils have been recovered. Units listed are all either formation rank or higher (e.g. group). Formations are listed by continent, and alphabetically within the individual lists.

List of mosasaur genera

This list of mosasaurs is a comprehensive listing of all genera that have ever been included in the family Mosasauridae or the parent clade Mosasauroidea, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered invalid, doubtful (nomen dubium), or were not formally published (nomen nudum), as well as junior synonyms of more established names, and genera that are no longer considered mosasauroid. Non-mosasaurid mosasauroids shall be noted as such. The list currently includes 78 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 in the United States and a possible specimen in Belgium and Africa. A well-preserved specimen of Platecarpus shows that it fed on moderate-sized fish, and it has been hypothesized to have fed on squid, and ammonites as well. Like other mosasaurs, it was initially thought to have swum in an eel-like fashion, although another study suggests that it swam more like modern sharks. An exceptionally well-preserved specimen of P. tympaniticus known as LACM 128319 shows skin impressions, pigments around the nostrils, bronchial tubes, and the presence of a high-profile tail fluke, showing that it and other mosasaurs did not necessarily have an eel-like swimming method, but were more powerful, fast swimmers. It is held in the Natural History Museum of Los Angeles County.

Plesiotylosaurus

Plesiotylosaurus, meaning "near Tylosaurus", 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 Prognathodon. The genus contains one species, Plesiotylosaurus crassidens, recovered from deposits of Middle Maastrichtian age in the Moreno Formation in California.Though it is classified as a mosasaurine mosasaur, and not closely related to Tylosaurus, the name is not entirely misplaced as a number of cranial features found in the relatively intact holotype skull suggest some degree of convergent evolution with tylosaurine mosasaurs.

Plioplatecarpus

Plioplatecarpus is a genus of mosasaur lizard. Like all mosasaurs, it lived in the late Cretaceous period, about 73-68 million years ago.

Plotosaurus

Plotosaurus ("swimmer lizard") is an extinct genus of mosasaur from the Upper Cretaceous (Maastrichtian) of Fresno County, California. Originally named Kolposaurus (meaning "bay lizard") by Berkeley paleontologist Charles Lewis Camp in 1942, it was changed to Plotosaurus in 1951 when Camp discovered the name had already been assigned to a type of nothosaur.

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 deposits ranging in age from the Campanian to the Maastrichtian in the Middle East, Europe and North America.Prognathodon means "forejaw tooth", which originates from the Latin pro- ("earlier" or "prior"), Greek gnathos ("jaw") and odṓn ("tooth"). Twelve nominal species of Prognathodon are recognised, from North America, northern and western Africa, the Middle East, western Europe and New Zealand. Due to the sometimes clear differences between them and the incomplete nature of many of the specimens, the systematics of the genus and which species should properly be considered Prognathodon is controversial.Prognathodon is known for its massively built jaws and teeth. Its distinct feeding adaptations have generated much interest in its ecology ever since its discovery, though direct evidence of its diet, such as gastric residues, is rare.

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 and S. johnsoni, both are exclusively known from Santonian deposits in the United States.

Selmasaurus is unique among the mosasaurs in that its skull is unusually akinetic, meaning that it is incapable of widening to swallow larger prey. Most mosasaurs have skulls which possess "coupled kinesis" (mesokinesis and streptostyly), that is, parts of the jaw can open widely to accommodate large prey.

Taniwhasaurus

Taniwhasaurus (from the Maori Taniwha, a supernatural, aquatic creature, and the Greek σαυρος/sauros, meaning lizard) is an extinct genus of mosasaur (carnivorous marine lizards) which inhabited New Zealand, Japan and Antarctica. The genus was a close relative of the genera Tylosaurus and Hainosaurus.

Timeline of mosasaur research

This timeline of mosasaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of mosasaurs, a group of giant marine lizards that lived during the Late Cretaceous Epoch. Although mosasaurs went extinct millions of years before humans evolved, humans have coexisted with mosasaur fossils for millennia. Before the development of paleontology as a formal science, these remains would have been interpreted through a mythological lens. Myths about warfare between serpentine water monsters and aerial thunderbirds told by the Native Americans of the modern western United States may have been influenced by observations of mosasaur fossils and their co-occurrence with creatures like Pteranodon and Hesperornis.The scientific study of mosasaurs began in the late 18th century with the serendipitous discovery of a large fossilized skeleton in a limestone mine near Maastricht in the Netherlands. The fossils were studied by local scholar Adriaan Gilles Camper, who noted a resemblance to modern monitor lizards in correspondence with renowned French anatomist Georges Cuvier. Nevertheless, the animal was not scientifically described until the English Reverend William Daniel Conybeare named it Mosasaurus, after the river Meuse located near the site of its discovery.By this time the first mosasaur fossils from the United States were discovered by the Lewis and Clark expedition, and the first remains in the country to be scientifically described were reported slightly later from New Jersey. This was followed by an avalanche of discoveries by the feuding Bone War paleontologists Edward Drinker Cope and Othniel Charles Marsh in the Smoky Hill Chalk of Kansas. By the end of the century a specimen of Tylosaurus would be found that preserved its scaley skin. Later Samuel Wendell Williston mistook fossilized tracheal rings for the remains of a fringe of skin running down the animal’s back, which subsequently became a common inaccuracy in artistic restorations.The 20th century soon saw the discovery in Alabama of a strange mosasaur called Globidens, with rounded teeth suited to crushing shells. Mosasaur remains were also discovered in Africa and California. In 1967 Dale Russell published a scientific monograph dedicated to mosasaurs. Embryonic remains in the 1990s confirmed that mosasaurs gave live birth like in ichthyosaurs. The 1990s also saw a revival and escalation of a debate regarding whether or not some supposed mosasaur toothmarks in ammonoid shells were actually made by limpets. By the end of the century, the evolutionary relationship between mosasaurs and snakes as well as the possible involvement of mosasaurs in the extinction of the aforementioned ichthyosaurs became hot button controversies.The debates regarding snakes, toothmarks, and ichthyosaurs spilled over into the early 21st century. These discussions were also accompanied by the discovery of many new taxa, including new species of Globidens, Mosasaurus, and Tylosaurus as well as entirely new genera like Yaguarasaurus and Tethysaurus. In 2013, Lindgren, Kaddumi, and Polcyn reported the discovery of a Prognathodon specimen from Jordan that preserved the soft tissues of its scaley skin, flippers and tail. Significantly, the tail resembled those of modern carcharinid sharks, although the bottom lobe of the tail fin was longest in the mosasaur whereas shark tails have longer upper lobes.

Tylosaurus

Tylosaurus (Greek τυλος/tylos "protuberance, knob" + Greek σαυρος/sauros "lizard") was a mosasaur, a large, predatory marine reptile closely related to modern monitor lizards and to snakes, from the Late Cretaceous.

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Mosasaurinae
Halisaurinae
Russellosaurina
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