Mosasaur

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[13] 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.[14]

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.[13] As ambush predators, lurking and quickly capturing prey using stealth tactics,[15] they may have benefited from the nonreflective, keeled scales.[13] 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.[16]

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.[8]

Fibrous tissues and microstructures recovered from Prognathodon specimen IRSNB 1624

In 2011, collagen protein was recovered from a Prognathodon humerus dated to the Cretaceous.[17]

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.[18]

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.[19]

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 Nature.[20]

Mosasaurs possessed a thecodont dentiton, meaning that the roots were cemented deeply into the jaw bone. Mosasaurs did not use permanent teeth but instead constantly shed them. Replacement teeth developed within a pit inside the roots of the original tooth called the resorption pit. This is done through a distinctively unique eight-stage process. The first stage was characterized by the mineralization of a small tooth crown developed elsewhere that descended into the resorption pit by the second stage. In the third stage, the developing crown firmly cemented itself within the resorption pit and grew in size; by the fourth stage, it would be of the same size as the crown in the original tooth. Stages five and six were characterized by the development of the replacement tooth's root: in stage five the root developed vertically, and in stage six the root expanded in all directions to the point that the replacement tooth became exposed and actively pushed on the original tooth. In the seventh stage, the original tooth was shed and the now-independent replacement tooth began to anchor itself into the vacancy. In the eighth and final stage, the replacement tooth has grown to firmly anchor itself.[21]

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.[22]

In late 2014 Guinness World Records awarded the museum with a record for Largest Publicly Displayed Mosasaur – Bruce. The record was added to the 2016 print edition of Guinness World Records.[23]

A 2020 study published in Nature described a large fossilized hatched egg from Antarctica from the very end of the Cretaceous, about 68 million years ago. The egg is considered one of the largest amniote eggs ever known, rivalling that of the elephant bird, and due to its soft, thin, folded texture, it likely belonged to a marine animal. While the organism that produced it remains unknown, the egg's pore structure is very similar to that of extant lepidosaurs such as lizards and snakes, and presence of mosasaur fossils nearby indicates that it may have been a mosasaur egg. It is unknown whether the egg was laid on land or in the water. The egg was assigned to the newly described oospecies Antarcticoolithus bradyi.[24][25][26]

Scientists continue to debate on whether monitor lizards (left) or snakes (right) are the closest living relatives of mosasaurs.

The specific placement of mosasaurs within the Squamata has been controversial since its inception. Cuvier was the first scientist to deeply analyze their possible taxonomic placement through Mosasaurus. While his original 1808 hypothesis that the genus was a lizard with affinities to monitor lizards remained the most popular, Cuvier was uncertain, even at the time, about the accuracy of this placement. He simultaneously proposed a number of alternative hypotheses, with one such alternative suggesting that Mosasaurus instead had closer affinities with iguanas due to their shared presence of pterygoid teeth. With the absence of sufficient fossil evidence, researchers during the early and mid-19th century had little to work with. Instead, they primarily relied on stratigraphic associations and Cuvier's 1808 research on the holotype skull. Thus, in-depth research on the placement of Mosasaurus was not undertaken until the discovery of more complete mosasaur fossils during the late 19th century, which reignited research on the placement of mosasaurs among squamates.[43]

In a span of about 30 to 40 years during the late 19th to early 20th centuries, paleontologists fiercely debated the issue, which created two major schools of thought: one that supported a monitor lizard relationship and one that supported a closer relationship with snakes.[43] The proposition of a snake relationship was spearheaded by Cope, who first published such a hypothesis in 1869 by proposing that mosasaurs, which he classified under a group called the Pythonomorpha, was the sister group of snakes. Some scientists went as far as to interpret mosasaurs as direct ancestors of snakes.[44] Opponents supporting a relationship with monitor lizards argued that mosasaurs should be placed within the infraorder Anguimorpha, with lower classifications varying from placing mosasaurs within Varanoidea or its sister taxa, or as true monitor lizards within Varanidae.[43] These debates spawned higher taxonomic groups that were erected in attempts to classify the placement of mosasaurs (although not all are compatible). One of these was the Mosasauria, a loosely-defined group erected by Marsh in 1880 but is still used by some researchers today that includes all descendants of the last common ancestor of Mosasaurus and some of its ancestral relatives, including the marine squamates dolichosaurs Dolichosaurus and Coniasaurus and Adriosaurus.[45]

In 1923, Charles Lewis Camp published Classification of the Lizards, in which he proposed through the review and rebuttal of previous arguments using his own anatomical observations that all taxa more closely related to Mosasaurus than Dolichosaurus should be classified into a superfamily called the Mosasauroidea, which would be a sister superfamily to the Varanoidea.[43][46] Camp's take on the subject virtually ended the snake-monitor lizard debate for approximately 70 years, with nearly all subsequent studies supporting a relationship with monitor lizards. However, many studies continued to support going further than Camp in the monitor lizard relationship, placing mosasaurs within the Varanoidea.[43]

The debate was reignited with the publication of a 1997 cladistical study by Michael S. Y. Lee, which recovered the Mosasauroidea as a sister taxon to the snake suborder Serpentes and resurrected the argument for a snake relationship.[43][47] Prior, there had never been a modern phylogenetic study specifically testing the relationships between mosasaurs or snakes. Lee also resurrected the defunct Pythonomorpha and redefined it to unify the Mosasauroidea and Serpentes under one clade.[47] Multiple subsequent studies conducted by scientists such as Lee, Caldwell, and Alessandro Palci refined this hypothesis. In some studies, the Mosasauria clade was used to represent mosasaurs.[43][44] However, there still remained little consensus. For example, a large-scale 2008 phylogenetic study by paleontologist Jack Conrad of the American Museum of Natural History recovered the Mosasauria clade in a polytomy, or a clade with unresolved relationships, with monitor lizards and beaded lizards;[45] and a 2012 study by Gauthier et al. recovered Mosasauria as a clade basal to both monitor lizards and snakes.[48]

With the advent of molecular genetics during the 2010s, some scientists argued for the combining of molecular and morphological data to examine relationships between mosasaurs and living squamates.[49][50] An early study was Lee (2009), which based on nuclear and mitochondrial DNA in living squamates and morphological data recovered mosasaurs as a stem group to snakes,[51] which some later authors interpreted as placement of snakes within the Mosasauria clade itself.[49] However, a 2010 study by Weins et al. attempted to replicate Lee (2009) using a larger dataset but instead yielded results that recovered the Mosasauria as a sister clade to the monitor lizards.[49] The frequent discrepancies are due to the high prevalence of convergent evolution in squamates, which creates much room for interpreting molecular and morphological data; many of these studies had results that often contradicted each other, for example in completely different phylogenetic results by simply adding more datasets or varying which taxa are represented, which resulted in various conflicts creating even more uncertainty. Due to this, some scientists argued that a molecular perspective should be abandoned entirely.[50][52] Nevertheless, other scientists have attempted to resolve these problems. One approach was utilized by a 2015 study by Reeder et al.: it closely integrated morphological, molecular, and paleontological data in a large dataset to overcome previous conflicts, which revealed new morphological support for molecular results that recovered Mosasauria as a sister clade to Serpentes.[50] Another approach was developed by R. Alexander Pyron in a 2016 study, which also recovered Mosasauria as a sister clade to the Serpentes.[53]

Restoration of Opetiosaurus bucchichi, a basal mosasauroid

Life restoration of a mosasaurine, Globidens alabamaensis

Life restoration of a mosasaurine, Plotosaurus bennisoni

Restoration of a tylosaurine, Tylosaurus pembinensis

The traditional view of mosasaur evolution held that all paddle-limbed (hydropedal) mosasaurs originated from a single common ancestor with functional legs (plesiopedal). However, this was shaken with the discovery of Dallasaurus, a plesiopedal mosasauroid that was found by multiple phylogenetic analyses to be an ancestor of the Mosasaurinae but unrelated to other mosasaurs. Bell and Polycn (2005) grouped these outside mosasaurs into two clades: the Russellosaurina, whose basal members include plesiopedal genera (Tethysaurinae) of their own and derived members consisting of the Plioplatecarpinae and Tylosaurinae; and the Halisauromorpha, containing the Halisaurinae. The placement of Dallasaurus suggested that the Russellosaurina and Halisauromorpha may have evolved a hydropedal form independently, the former through the tethysaurines, meaning that their placement within the Mosasauridae creates an unnatural polyphyly and thus potentially invalid.[54][43] Caldwell informally proposed in a 2012 publication that the definition of a mosasaur must thus be redefined into one that does not consider russellosaurines and halisauromorphs as true mosasaurs, but as an independent group of marine lizards.[43]

However, phylogenetic studies of mosasaurs can be fickle, especially when wild card taxa like Dallasaurus remain poorly understood. For example, some studies such as a 2009 analysis by Dutchak and Caldwell instead found that Dallasaurus was ancestral to both russellosaurines and mosasaurines,[55] although results were inconsistent in later studies.[56] A 2017 study by Simoes et al. noted that utilization of different methods of phylogenetic analyses can yield different findings and ultimately found a consistent indication that tethysaurines were a case of hydropedal mosasaurs reversing back to a plesiopedal condition rather than an independent ancestral feature.[56]

Cladogram of the Mosasauroidea from Madzia and Cau (2017):[57]