Russellosaurina

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 the Meuse in 1764. They belong to the order Squamata, which includes lizards and snakes.

Mosasaurs probably evolved from an extinct group of aquatic lizards known as aigialosaurs in the Earliest Late Cretaceous with 42 described genera. 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 themselves became extinct as a result of the K-Pg event at the end of the Cretaceous period, about 66 million years ago.

Description

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 most likely gave birth to live young, rather than returning to the shore to lay eggs as sea turtles do.

The smallest-known mosasaur was ''Dallasaurus turneri'', which was less than long. Larger mosasaurs were more typical, with many species growing longer than . ''Mosasaurus hoffmannii'', the largest known species reached up to , but it has been considered to be probably overestimated by Cleary ''et al.'' (2018). 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 long, but this might be an overestimate as the skeleton was assembled for display prior to a 2010 reassessment of the species that found its original number of vertebrae to be exaggerated, implying that the actual size of the animal was likely smaller.

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. These animals may have lurked and pounced rapidly and powerfully on passing prey, rather than chasing after it.

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.

Paleobiology

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 long, probably fed on fish and other small prey. The smaller mosasaurs may have spent some time in fresh water, hunting for food. The largest mosasaur ''Mosasaurus hoffmannii'' was the apex predator of the Late Cretaceous oceans, reaching more than in length and weighing up to in body mass.

Soft tissue

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 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.

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. As ambush predators, lurking and quickly capturing prey using stealth tactics, they may have benefited from the nonreflective, keeled scales. Additionally, mosasaurs had large pectoral girdles, and such genera as ''Plotosaurus'' may have used their front flippers in a breaststroke motion to gain added bursts of speed during an attack on prey.

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.

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

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.

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.

Coloration

The coloration of mosasaurs was unknown until 2014, when the findings of Johan Lindgren of Lund University

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