Thalassotitan

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Thalassotitan
Temporal range: Late Maastrichtian,
~67–66  Ma
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Thalassotitan atrox skull.jpg
Syntype skull and jaws (MNHM.KH.231) of T. atrox from Ouled Abdoun Basin, Morocco
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Clade:† Mosasauria
Family:† Mosasauridae
Tribe:† Prognathodontini
Genus:† Thalassotitan
Longrich et al., 2022
Type species
†Thalassotitan atrox
Longrich et al., 2022

Thalassotitan ("titan of the seas") is an extinct genus of large mosasaurs (a group of extinct marine lizards) that lived during the late Maastrichtian of the Cretaceous period in what is now Morocco, around 67 to 66 million years ago. The only known species is T. atrox, described in 2022 from fossils discovered in the Ouled Abdoun Basin, initially identified as coming from other genera such as Mosasaurus or Prognathodon . Hypothetical Thalassotitan specimens may have been found in other corners of the world, although researchers also note the possibility that they come from distinct, related taxa. It is considered to be close to the genera Prognathodon and Gnathomortis , together forming the tribe Prognathodontini. The prognathodontines are separated from other mosasaurs based on their massive jaws and robust teeth.

Contents

Thalassotitan is one of the largest known mosasaurs, having an estimated size of around 9 to 10 m (30 to 33 ft) long. This genus shows definitely that mosasaurs evolved to take over the apex predator niche in the oceans of the Late Cretaceous which is now filled by sharks and orcas. Heavy wear on its teeth and fossils found in the vicinity of the holotype etched by acid wear from partial digestion suggest that this mosasaur had a diet consisting of smaller mosasaur species, plesiosaurs, large predatory fish, and sea turtles.

Discovery and naming

Map of the Ouled Abdoun basin and other major phosphate basins of Morocco shown in yellow. The Upper Couche III, the area from which all recognized specimens of Thalassotitan were discovered, is illustrated by the black star at the top right of the table opposite Ocepechelon bouyai geography stratigraphy.png
Map of the Ouled Abdoun basin and other major phosphate basins of Morocco shown in yellow. The Upper Couche III, the area from which all recognized specimens of Thalassotitan were discovered, is illustrated by the black star at the top right of the table opposite

The phosphate deposits of the Ouled Abdoun Basin in Morocco have been known since the beginning of the 20th century to yield various fossils of numerous aquatic vertebrates dating from the Maastrichtian stage of the Upper Cretaceous. [2] : 1  [3] In a work published in 1952, Camille Arambourg carried out a broad revision of this area, where he described a certain number of these same fossil vertebrates. Among them, he describes a mosasaur taxon under the name Mosasaurus (Leiodon) cf. anceps, based on numerous fossil teeth of different morphologies. [2] : 279–282  It is now accepted that the smaller, slender teeth formerly referred to this taxon actually belong to Eremiasaurus , while the larger ones would likely have belonged to Thalassotitan. Many other fossils now attributed to Thalassotitan were then assigned to the related genus Prognathodon in later works. [1]

It is on the basis of these numerous anatomical differences that Nicholas R. Longrich and his colleagues erected in 2022 a new genus and a new species of mosasaurids under the name of Thalassotitan atrox. Although many specimens are referred to this taxon, the authors designate two partial skeletons as syntypes, which are cataloged MNHM.KH.231 and OCP DEK-GE 417. All formally known specimens of this taxon have been more precisely discovered at the level of Upper Couche III, dating from approximately 67 to 66 million years ago. The genus name Thalassotitan is a portmanteau of the Ancient Greek θάλασσα (thálassa, "sea") and τιτάν (tītā́n, "giant"), referring to the mosasaur's large size. The specific epithet atrox is a Latin word translating to "cruel" or "merciless", which references the species' trophic position as an apex predator and frequency of intraspecific bite marks on fossils. [1]

Although all fossil remains formally attributed to Thalassotitan come from the Ouled Abdoun Basin, the authors note that teeth discovered elsewhere in the world could belong to this genus, or at least to a related taxon like Prognathodon saturator. These locations include the Ganntour Basin in Morocco, Jordan, Egypt, Israel, Poland, Angola and Brazil. [1] In 2024, Trevor H. Rempert and his colleagues note the fairly abundant presence of fossil teeth comparable to those of Thalassotitan having been discovered in the Pee Dee Formation in North Carolina, USA. [4]

Description

Restoration of T. atrox with human scale Thalassotitan atrox MP.png
Restoration of T. atrox with human scale

Thalassotitan was one of the largest mosasaurs. Its skull measured up to 1.5 meters (4.9 ft) in length, corresponding to a total length of 9–10 meters (30–33 ft). [1]

Skull

Annotated schematic of a mosasaur skull Mosasaurus hoffmannii skull schematic.png
Annotated schematic of a mosasaur skull

Like all prognathodontines, the skull of Thalassotitan is blunt and robust. The premaxilla, the bone bearing the tip of the skull, is very short in a lateral view but broad and convex when viewed dorsally. The body of the premaxilla contains numerous pits called neurovascular foramina, which are believed to house tactile nerves that are very sensitive to touch. The internarial bar, a long extension of the premaxilla reaching up to the frontal bone, is broad as it passes between the maxilla (the main tooth-bearing upper jaw bone) and external nares (the openings that house the nostrils) but narrows into a slender rod as it contacts with the frontal. Between the maxilla, the internarial bar forms a distinct low and short keel. The maxilla is short, robust, and deep. Its surface is flat except for a low and broad ridge lining just above the teeth. [1]

Neurovascular foramina line this margin, increasing in size as they progress towards the back of the skull. The texture of the maxilla's surface is rough, which is especially apparent in larger individual, caused by a network of veined grooves to house blood vessels. The external nares extend from and to above the fourth and twelfth maxillary teeth. The jugal bone, which is located just below the eye, is broad and robust. The frontal bone is short and broad, shaped almost like an isosceles triangle, with large neurovascular foramina at the center. The pineal foramen, which contains the parietal eye, is small and long. The supratemporal fenestrae, large openings between the eyes and the back end of the skull, take up nearly a quarter of the entire skull length and are somewhat triangular. The dentary, the tooth-bearing bone of the lower jaw, is short, wide, robust, and curved concave towards the upper jaws. Many bones of the upper jaw are tightly sutured together; its two tooth-bearing bones the premaxilla and maxilla were connected through interlocking joints containing an unusual series of flanges and grooves while an interdigitating set of tongue-and-groove joints secure the maxilla and prefrontal bone. [1]

Teeth

Juvenile skull with teeth shown Thalassotitan referred by Nick Longrich.jpg
Juvenile skull with teeth shown

Thalassotitan teeth are roughly conical in shape, lightly curved, large in size, and robust in build. They are most similar to the teeth of P. saturator except in being slightly shorter and stockier. Tooth crowns are slightly swollen around its base next to the root, but they do not form a round circumference. The surfaces of the crown are generally smooth but may sometimes have faint ridges depending on individual or ontogenetic variation. The enamel at the tip contain veinous ridges and coarse bumps. Cutting edges are well-developed and finely serrated. Each tooth has two cutting edges, but their positions differ depending on the tooth's position in the jaw. Towards the front of the jaw, the front-facing cutting edges are more pronounced than the diminished back-facing edges. In the middle and near the end of the jaw, both edges are of equal development and located diametrically opposite to each other. At the end of the jaw, the back-facing edges become more pronounced. The tooth roots are massive and barrel-shaped. Deep pits occur within the roots, from which new replacement teeth are formed. [1]

Like all mosasaurs, Thalassotitan had four types of teeth, corresponding to the jaw bones they are located on. On the upper jaw were the premaxillary teeth, maxillary teeth, and pterygoid teeth (located separate from the main jawline near the rear of the skull); while on the lower jaw only the dentary teeth were present. Thalassotitan had in each jaw row from front to back: two premaxillary teeth, twelve maxillary teeth, at least six pterygoid teeth (the pterygoids were not fully preserved), and fourteen dentary teeth. The dentary teeth are generally flatter by the side than the maxillary teeth. Heterodonty is present, meaning that tooth shape changes down the jawline. The first four or five teeth are tall, narrow, and slightly curved, which become stockier, erect, and more robust around the middle of the jawline, then become shorter (as broad as they are tall), hooked, and flatter by the side. The pterygoid teeth are strongly hooked but are also large and robust, nearly approaching the size of the teeth on the main jawlines. [1]

Postcranial skeleton

The postcranial skeleton is not fully known, only fossils representing a little more than the front half of the body have been found. The general shape of the vertebrae are typical for mosasaurines. They are procoelous, meaning that the front side is deeply cupped concavely and the back side is bulged convexly. The cervical (neck) vertebrae are slightly wider than long. Its atlas holds rectangular or triangular neural arches; another single tall neural arch is also present at the top of the vertebra. The articular surfaces, which atach to the cartilage that connect vertebrae together, is initially heart-shaped but becomes rounded at the rearmost cervicals. The dorsal (back) vertebrae are slightly longer than wide with tall neural arches, rounded articular surfaces, and large rectangular transverse processes. The ribs are short and robust. [1]

The pectoral girdle is robust and most similar with that in P. overtoni and Mosasaurus conodon , albeit more square-shaped than the latter. The two bones making up the girdle, the scapula and coracoid, are similar in sizes. They loosely contact with each other, but their contact point is nevertheless wider than the glenoid fossa. The scapula is shaped like a square, being as long as it is wide. It lacks a defined scapular neck but expands from front to back, forming a fan-like convex blade. The coracoid is also somewhat squarish and lacks a well-defined neck. Its margins are weakly concave in the front and back but very convex at the bottom. [1]

The forelimbs formed long paddles that resembled mosasaurin mosasaurs like Mosasaurus and Plotosaurus but more primitive in possessing longer but fewer phalanges. The humerus is very stocky and resemble that in P. overtoni except in the expansion of the glenoid condyle beyond the postglenoid process. The radius is unusually shaped for a mosasaur. It is as large as the humerus and much larger than the ulna and takes on a crescent-like or subrectangular form, unlike smaller hourglass-shaped radii in typical mosasaurs. [1]

Classification

Prognathodon saturator.jpg
Israeli Prognathodon.jpg
T. atrox is most closely related to Prognathodon saturator (left) and P. currii (right), and the three species may be congeneric.

Thalassotitan is a member of the Prognathodontini tribe of the Mosasaurinae subfamily, with other members including Prognathodon and Gnathomortis . Morphologically, it is most similar to the giant mosasaurs P. currii and P. saturator, and a phylogenetic analysis by Longrich et al. (2022) recovered Thalassotitan in a clade between the two. This creates an unnatural paraphyletic relationship that reflects a wider issue with the genus Prognathodon as a whole. Several studies over the past decade found that Prognathodon is in general not monophyletic and in need of revision. Longrich et al. (2022) suggested such a revision may include an expansion of the Thalassotitan genus to include P. currii and P. saturator. [1] However, due to a high degree of convergent evolution in the relationship-determining traits among many mosasaurs (especially among prognathodontines), phylogenetic results between each study are seldom consistent, mystifying exactly which species must be revised to stabilize the Prognathodontini. For example, some studies recovered P. currii and P. saturator as phylogenetically unrelated species with either falling outside a monophyletic Prognathodon, while other studies yield variable placements for the type species P. solvayi as either outside a monophyletic P. currii-P. saturator clade in support of Longrich et al. (2022) or within it, which would in theory invalidate Thalassotitan as a junior synonym under the principle of priority. [1]

The following cladogram is modified from Longrich et al. (2022). [1]

Mosasaurinae

Kourisodon

Clidastes

Globidensini

Globidens simplex

Globidens schumani

Globidens phosphaticus

Prognathodon rapax (=Ancylocentrum hungerfordi)

Globidens alambamensis

Globidens dakotensis

Prognathodontini

Gnathomortis

Prognathodon overtoni

Prognathodon saturator

Thalassotitan atrox

Prognathodon currii

Prognathodon giganteus

Prognathodon lutugini

Prognathodon solvayi

Mosasaurini

Mosasaurus missouriensis

Mosasaurus lemonnieri

Mosasaurus hoffmannii

Mosasaurus beaugei

Mosasaurus maximus

Liodon

Mosasaurus sp. (MGGC 21876)

"Magahouanga mosasaurine"

Paleoecology

Life restoration of Thalassotitan (left) and other animals of the Ouled Abdoun Basin Ouled Abdoun Basin.png
Life restoration of Thalassotitan (left) and other animals of the Ouled Abdoun Basin

The phosphate deposits of Morocco have revealed an extremely diverse environment of late maastrichtian age. [1] [5] The oceans of the area were full of an abundance of fish, from bony fish like Enchodus and Stratodus to cartilaginous fish like Cretalamna, Squalicorax and Rhombodus. [1] There was also an abundance of marine reptiles, most notably the mosasaurs, with more than 10 genera alone known from this single site. [6] This possibly suggests that niche partitioning took place here, in which predators take on different niches to avoid competition with one another (for example, mosasaurs like Carniodens and Globidens had blunt teeth for crushing shellfish while Thalassotitan and Mosasaurus hunted much larger food). [1] [7] Other marine reptiles include the elasmosaurid plesiosaur Zarafasaura , the sea turtle Alienochelys and the gavialoid crocodilian Ocepesuchus . [8] [9]

It seems that Thalassotitan was an apex predator in its ecosystem, with evidence being digestive damage found on some of the fossils in the nearby vicinity including those of plesiosaurs, turtles, and large fish. [1] In the skies flew multiple species of pterosaurs including the azhdarchid Phosphatodraco , the nyctosauromorph Alcione and Simurghia , the nyctosaurid Barbaridactylus , and the possible pteranodontid Tethydraco . [10] [11] On land several species of dinosaurs are known, these being the abelisaur Chenanisaurus, the small lambeosaurine hadrosaurs Ajnabia and Minqaria and a so far unnamed titanosaur. [12] [13] [14] There are also several unnamed lambeosaurines and abelisaurs currently awaiting formal description. [14] Thalassotitan lived alongside other giant mosasaurs like Prognathodon, Mosasaurus, Khinjaria and Gavialimimus , as well as smaller mosasaurs like Xenodens , Halisaurus and Pluridens . [15] [1] In 2024, Rempert and colleagues note that the possible presence of teeth of Thalassotitan and other Moroccan mosasaurs in the southeastern United States suggests that the Atlantic Ocean and the Tethys Ocean shared similar fauna during the late Maastrichtian. [4]

See also

Related Research Articles

<span class="mw-page-title-main">Mosasaur</span> Extinct marine lizards of the Late Cretaceous

Mosasaurs 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 a limestone quarry at Maastricht on the Meuse in 1764. They belong to the order Squamata, which includes lizards and snakes.

<i>Mosasaurus</i> Extinct genus of marine squamate reptile from the Late Cretaceous

Mosasaurus is the type genus of the mosasaurs, an extinct group of aquatic squamate reptiles. It lived from about 82 to 66 million years ago during the Campanian and Maastrichtian stages of the Late Cretaceous. The genus was one of the first Mesozoic marine reptiles known to science—the first fossils of Mosasaurus were found as skulls in a chalk quarry near the Dutch city of Maastricht in the late 18th century, and were initially thought to be crocodiles or whales. One skull discovered around 1780 was famously nicknamed the "great animal of Maastricht". In 1808, naturalist Georges Cuvier concluded that it belonged to a giant marine lizard with similarities to monitor lizards but otherwise unlike any known living animal. This concept was revolutionary at the time and helped support the then-developing ideas of extinction. Cuvier did not designate a scientific name for the animal; this was done by William Daniel Conybeare in 1822 when he named it Mosasaurus in reference to its origin in fossil deposits near the Meuse River. The exact affinities of Mosasaurus as a squamate remain controversial, and scientists continue to debate whether its closest living relatives are monitor lizards or snakes.

<span class="mw-page-title-main">Mosasaurinae</span> Subfamily of reptiles

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 for South America.

<i>Globidens</i> Extinct genus of lizards

Globidens is an extinct genus of mosasaurid oceanic lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily. Globidens belongs to the family Mosasauridae, which consists of several genera of predatory marine lizards of various sizes that were prevalent during the Late Cretaceous. Specimens of Globidens have been discovered in Angola, Brazil, Morocco, Syria and the United States. Among mosasaurs, Globidens is probably most well known for the highly rounded, globe-like teeth that give it its name.

<i>Prognathodon</i> Extinct genus of lizards

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, New Zealand, and North America.

<i>Phosphatodraco</i> Late Cretaceous genus of pterosaur

Phosphatodraco is a genus of azhdarchid pterosaur that lived during the Late Cretaceous of what is now Morocco. In 2000, a pterosaur specimen consisting of five cervical (neck) vertebrae was discovered in the Ouled Abdoun Phosphatic Basin. The specimen was made the holotype of the new genus and species Phosphatodraco mauritanicus in 2003; the genus name means "dragon from the phosphates", and the specific name refers to the region of Mauretania. Phosphatodraco was the first Late Cretaceous pterosaur known from North Africa, and the second pterosaur genus described from Morocco. It is one of the only known azhdarchids preserving a relatively complete neck, and was one of the last known pterosaurs. Additional cervical vertebrae have since been assigned to the genus, and it has been suggested that fossils of the pterosaur Tethydraco represent wing elements of Phosphatodraco.

<i>Pluridens</i> Extinct genus of lizards

Pluridens is an extinct genus of marine lizard belonging to the Mosasauridae. Pluridens is placed in the subfamily Halisaurinae with the genera Phosphorosaurus, Eonatator and Halisaurus. Compared to related halisaurines, Pluridens had longer jaws with more teeth, and smaller eyes. It also grew large size, measuring 5–6 m (16–20 ft) long and perhaps over 9 m (30 ft) 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.

Goronyosaurus is an extinct genus of marine lizard belonging to the mosasaur family. Fossils of Goronyosaurus are exclusively known from the Late Maastrichtian of the Iullemmeden Basin in West Africa, specifically the Dukamaje Formation of Niger and Nigeria and Farin Doutchi Formation of Niger. The type specimen was first described in 1930 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, although most authors agree that Goronyosaurus belonged to Mosasauridae.

<i>Liodon</i> Extinct genus of lizards

Liodon is a dubious genus of mosasaur from the Late Cretaceous, known from fragmentary fossils discovered in St James' Pit, England and possibly also the Ouled Abdoun Basin of Morocco. 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.

<i>Plesiotylosaurus</i> Extinct genus of lizards

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.

Eremiasaurus is a genus of mosasaurs, an extinct group of marine reptiles. It lived during the Maastrichtian stage of the Late Cretaceous in what is now North Africa. Only one species is known, E. heterodontus, described in 2012 from two remarkably complete fossil specimens discovered in the Ouled Abdoun Basin, Morocco. This site is known to have delivered a significant number of other related mosasaurs.

<span class="mw-page-title-main">Ouled Abdoun Basin</span> Phosphate basin in Morocco

The Oulad Abdoun Basin is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and at least 26.8 billion tons of phosphate. It is also known as an important site for vertebrate fossils, with deposits ranging from the Late Cretaceous (Cenomanian-Turonian) to the Eocene epoch (Ypresian), a period of about 25 million years.

<span class="mw-page-title-main">Timeline of mosasaur research</span>

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.

<span class="mw-page-title-main">Globidensini</span> Tribe of lizards

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, Europe, Africa and Asia. The tribe contains the genera Globidens, Carinodens, Igdamanosaurus, Harranasaurus and Xenodens. Features of the maxilla and digits make the placement of Carinodens and Xenodens in the tribe uncertain; some researchers have suggested that they may be more appropriately placed in the Mosasaurini.

<i>Chenanisaurus</i> Extinct genus of dinosaurs

Chenanisaurus is a genus of predatory abelisaurid dinosaur, with a single known species C. barbaricus. It comes from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa. The animal is known from a holotype, consisting of a partial jaw bone, and several isolated teeth found in the same beds. Chenanisaurus is one of the largest members of the Abelisauridae, and one of the last, being a contemporary of the North American Tyrannosaurus. It would have been among the dinosaur species wiped out by the Chicxulub asteroid impact and the Cretaceous-Paleogene mass extinction that followed.

<i>Ajnabia</i> Extinct genus of reptiles

Ajnabia is a genus of lambeosaurine hadrosaur from the Late Cretaceous (Maastrichtian) of Morocco. It is the first definitive hadrosaur from Africa, and is thought to be related to European dinosaurs like Arenysaurus. The discovery of Ajnabia came as a surprise to the paleontologists who found it, because Africa was isolated by water from the rest of the world during the Cretaceous, such that hadrosaurs were assumed to have been unable to reach the continent. Ajnabia is relatively small and similar in size to its contemporary relative Minqaria, which is estimated to have reached 3.5 metres (11 ft) in total body length. Assuming that the holotype represents an adult, Ajnabia would be one of the smallest if not the smallest known hadrosaurids.

<i>Gavialimimus</i> Extinct genus of lizards

Gavialimimus is an extinct genus of plioplatecarpine mosasaur from the Maastrichtian of Morocco. The holotype MHNM.KHG.1231, an articulated skull and associated fragmentary postcrania, was found in the Ouled Abdoun Basin.

<i>Stelladens</i> Extinct genus of mosasaurine

Stelladens is an extinct genus of mosasaurine. Fossils of Stelladens have been recovered from the Late Cretaceous Ouled Abdoun Basin of Morocco. The genus contains a single species, S. mysteriosus. It was a medium-sized mosasaur, measuring about 5 metres (16 ft) long.

<i>Minqaria</i> Extinct genus of hadrosaurid dinosaurs

Minqaria is a genus of arenysaurinin lambeosaurine hadrosaur from the Late Cretaceous (Maastrichtian) Ouled Abdoun Basin of Morocco. The genus contains a single species, M. bata, known from a partial skull.

Khinjaria is an extinct genus of plioplatecarpine mosasaurid from the Late Cretaceous Ouled Abdoun Basin of Morocco. The genus contains a single species, K. acuta, known from a partial skull and vertebra. Khinjaria was likely an apex predator in its environment, as its large body size, blade-like teeth, and unusual skull morphology would have allowed it to attack large prey animals.

References

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  2. 1 2 Camille Arambourg (1952). Les vertébrés fossiles des gisements de phosphates (Maroc–Algérie–Tunisie) [Fossil vertebrates from phosphate deposits (Morocco–Algeria–Tunisia)](PDF). Notes et Mémoires du Service Géologique (in French). Vol. 92. Paris: Typographie Firmin-Didot. Archived from the original (PDF) on 2022-11-27.
  3. ↑ Aaron R. H. Leblanc; Michael W. Caldwell; Nathalie Bardet (2012). "A new mosasaurine from the Maastrichtian (Upper Cretaceous) phosphates of Morocco and its implications for mosasaurine systematics". Journal of Vertebrate Paleontology . 32 (1): 82–104. Bibcode:2012JVPal..32...82L. doi:10.1080/02724634.2012.624145. JSTOR   41407709. S2CID   130559113.
  4. 1 2 Trevor H. Rempert; Brennan P. Martens; Alexander P. M. Vinkeles Melchers (2024). "Mosasaurs (Squamata: Mosasauridae) from the Late Cretaceous (Late Maastrichtian) of North Carolina, USA". Proceedings of the Zoological Institute RAS. 328 (3): 384–391. doi: 10.31610/trudyzin/2024.328.3.384 . S2CID   272074521.
  5. ↑ Johan Yans; M'Barek Amaghzaz; Baadi Bouya; Henri Cappetta; Paola Iacumin; László Kocsis; Mustapha Mouflih; Omar Selloum; Sevket Sen; Jean-Yves Storme; Emmanuel Gheerbrant (2014). "First carbon isotope chemostratigraphy of the Ouled Abdoun phosphate Basin, Morocco; implications for dating and evolution of earliest African placental mammals". Gondwana Research . 25 (1): 257–269. Bibcode:2014GondR..25..257Y. doi:10.1016/j.gr.2013.04.004. S2CID   129475565.
  6. ↑ Catherine R. C. Strong; Michael W. Caldwell; Takuya Konishi; Alessandro Palci (2020). "A new species of longirostrine plioplatecarpine mosasaur (Squamata: Mosasauridae) from the Late Cretaceous of Morocco, with a re-evaluation of the problematic taxon 'Platecarpus' ptychodon". Journal of Systematic Palaeontology . 18 (21): 1769–1804. doi:10.1080/14772019.2020.1818322. ISSN   1477-2019. S2CID   224978215.
  7. ↑ Dale A. Russell (1967). Systematics and morphology of American mosasaurs. Vol. 23. New Haven: Bulletin of the Peabody Museum of Natural History. p. 240. OCLC   205385.
  8. ↑ Nathalie Bardet; Nour-Eddine Jalil; France de Lapparent de Broin; Damien Germain; Olivier Lambert; Mbarek Amaghzaz (2013). "A giant chelonioid turtle from the Late Cretaceous of Morocco with a suction feeding apparatus unique among tetrapods". PLOS ONE . 8 (7): e63586. Bibcode:2013PLoSO...863586B. doi: 10.1371/journal.pone.0063586 . PMC   3708935 . PMID   23874378.
  9. ↑ Peggy Vincent; Nathalie Bardet; Xabier Pereda Suberbiola; Baâdi Bouya; Mbarek Amaghzaz; Saïd Meslouh (2011). "Zarafasaura oceanis, a new elasmosaurid (Reptilia: Sauropterygia) from the Maastrichtian Phosphates of Morocco and the palaeobiogeography of latest Cretaceous plesiosaurs". Gondwana Research. 19 (4): 1062–1073. Bibcode:2011GondR..19.1062V. doi:10.1016/j.gr.2010.10.005. S2CID   129404886.
  10. ↑ Nicholas R. Longrich; David M. Martill; Brian Andres (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary". PLOS Biology . 16 (3): e2001663. doi: 10.1371/journal.pbio.2001663 . PMC   5849296 . PMID   29534059.
  11. ↑ Fernandes, Alexandra E.; Mateus, Octávio; Andres, Brian; Polcyn, Michael J.; Schulp, Anne S.; Gonçalves, António Olímpio; Jacobs, Louis L. (2022). "Pterosaurs from the Late Cretaceous of Angola". Diversity. 14 (9). 741. doi: 10.3390/d14090741 . hdl: 10362/145845 .
  12. ↑ Nicholas R. Longrich; Xabier Pereda Suberbiola; R. Alexander Pyron; Nour-Eddine Jalil (2020). "The first duckbill dinosaur (Hadrosauridae: Lambeosaurinae) from Africa and the role of oceanic dispersal in dinosaur biogeography". Cretaceous Research . 120: 104678. doi: 10.1016/j.cretres.2020.104678 . S2CID   228807024.
  13. ↑ Nicholas R. Longrich; Xabier Pereda-Suberbiola; Nour-Eddine Jalil; Fatima Khaldoune; Essaid Jourani (2017). "An abelisaurid from the latest Cretaceous (late Maastrichtian) of Morocco, North Africa". Cretaceous Research . 76: 40–52. doi: 10.1016/j.cretres.2017.03.021 . S2CID   133063691.
  14. 1 2 Nicholas R. Longrich; Xabier Pereda-Suberbiola; Nathalie Bardet; Nour-Eddine Jalil (2024). "A new small duckbilled dinosaur (Hadrosauridae: Lambeosaurinae) from Morocco and dinosaur diversity in the late Maastrichtian of North Africa". Scientific Reports. 14 (1). 3665. doi: 10.1038/s41598-024-53447-9 . PMC   10864364 . PMID   38351204.
  15. ↑ Trevor Rempert; Alexander Vinkeles Melchers; Ashley Rempert; Muhammad Haque; Andrew Armstrong (2022). "Occurrence of Mosasaurus hoffmannii Mantell, 1829 (Squamata, Mosasauridae) in the Maastrichtian Phosphates of Morocco" (PDF). The Journal of Paleontological Sciences. 22: 1–22.
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