Tylosaurus

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Extinct genus of marine squamate reptiles

title: "Tylosaurus" type: doc version: 1 created: 2026-02-28 author: "Wikipedia contributors" status: active scope: public tags: ["apex-predators", "tylosaurinae", "mosasaurs-of-north-america", "fossil-taxa-described-in-1872", "turonian-genus-first-appearances", "maastrichtian-genus-extinctions", "taxa-named-by-othniel-charles-marsh", "mooreville-chalk", "symbols-of-kansas", "coniacian-genera", "santonian-genera", "campanian-genera", "paleontology-in-alberta"] description: "Extinct genus of marine squamate reptiles" topic_path: "science/earth-science" source: "https://en.wikipedia.org/wiki/Tylosaurus" license: "CC BY-SA 4.0" wikipedia_page_id: 0 wikipedia_revision_id: 0

::summary Extinct genus of marine squamate reptiles ::

T. bernardi (Dollo, 1885)
T. gaudryi (Thevenin, 1896)
T. ivoensis (Persson, 1963)
T. iembeensis (Antunes, 1964)
T. pembinensis (Nicholls, 1988)
T. saskatchewanensis Jiménez-Huidobro et al., 2018
T. kansasensis Everhart, 2005
T. "borealis" Garvey, 2020 | synonyms = {{collapsible list|title=List of synonyms|
Elliptonodon Emmons, 1858
Rhamphosaurus Cope, 1872
Rhinosaurus Marsh, 1872
Hainosaurus(?) Dollo, 1885
Macrosaurus proriger Cope, 1869
Macrosaurus pririger Cope, 1869
Liodon proriger Cope, 1869
Rhinosaurus proriger Marsh, 1872
Rhinosaurus micromus Cope, 1872
Tylosaurus dyspelor Leidy, 1873
Tylosaurus micromus Merriam, 1894
Liodon nepaeolicus Cope, 1874
Tylosaurus kansasensis? Everhart, 2005
Hainosaurus bernardi Dollo, 1885
Mosasaurus gaudryi Thevenin, 1896
Hainosaurus gaudryi Lingham-Soliar, 1992
Leiodon lundgreni Schröder, 1885
Mosasaurus hoffmanni ivoensis Persson, 1963
Mosasaurus ivoensis Russell, 1967
Hainosaurus ivoensis Lindgren, 1998
Mosasaurus iembeensis Antunes, 1964
Hainosaurus pembinensis Nicholls, 1988

Tylosaurus (; "knob lizard") is a genus of russellosaurine mosasaur (an extinct group of predatory marine lizards) that lived about 92 to 66 million years ago during the Turonian to Maastrichtian stages of the Late Cretaceous. Its fossils have been found primarily around North Atlantic Ocean including in North America, Europe, and Africa.

Research history

Main article: Research history of Tylosaurus

Possible first finds

The earliest Tylosaurus fossils were likely discovered by various Native American peoples and may have been the source of much of their folklore, with the earliest known ones dating back to well before the arrival of European settlers, around the 1500s. More recent accounts from peoples living in the Great Plains even speak of an ancient era ruled by massive aquatic creatures that were in constant combat with thunderbirds and were petrified by them. The considerable presence of fossils of large mosasaurs such as Tylosaurus and pterosaurs such as Pteranodon in this region may have been the origins of these myths.

In 1804, the Lewis and Clark Expedition discovered a now-lost fossil skeleton alongside the Missouri River, which was identified as a 45 ft long fish. In 2003, Richard Ellis speculated that the remains may have belonged to Mosasaurus missouriensis. Alternatively, a 2007 study led by Robert W. Meredith and colleagues suggested that the fossils would possibly come from a tylosaurine mosasaur based on the measurements cited by Clark and Gass and the evidence of Tylosaurus fossils that have been found in the Missouri River. However, the authors also mentioned the possibility that the remains would also come from an elasmosaurid plesiosaur, which are also known from the river, although being rarer.

First formal discoveries

::figure[src="https://upload.wikimedia.org/wikipedia/commons/b/bb/MCZ_4374_skull.png" caption="The skull of MCZ 4374, the holotype of ''Tylosaurus proriger'' and generic type of ''Tylosaurus'', in Cope (1870)"] ::

Tylosaurus was the third new genus of mosasaur to be described from North America behind Clidastes and Platecarpus and the first in Kansas. The early history of the genus as a taxon was subject to complications spurred by the infamous rivalry between American paleontologists Edward Drinker Cope and Othniel Charles Marsh during the Bone Wars. The type specimen was described by Cope in 1869 based on a fragmentary skull measuring nearly 5 ft in length and thirteen vertebrae lent to him by Louis Agassiz of the Harvard Museum of Comparative Zoology. The fossil, which remains in the same museum under the catalog number MCZ 4374, was recovered from a deposit of the Niobrara Formation located in the vicinity of Monument Rocks near the Union Pacific Railroad at Fort Hays, Kansas. Cope's first publication of the fossil was very brief and was named Macrosaurus proriger, the genus being a preexisting European mosasaur taxon. In 1870, Cope published a more thorough description of MCZ 4374. Without explanation, he moved the species into another European genus Liodon and declared his original Macrosaurus proriger a synonym.

In 1872, Marsh argued that Liodon proriger is taxonomically distinct from the European genus and must be assigned a new one. For this, he erected the genus Rhinosaurus, which means "nose lizard" and is a portmanteau derived from the Ancient Greek words ῥίς (rhī́s, meaning "nose") and σαῦρος (saûros, meaning "lizard"). Cope responded by arguing that Rhinosaurus was already a preoccupied synonym of Liodon. He disagreed with Marsh's arguments but proposed that in case Marsh was indeed correct, the genus name Rhamphosaurus should be used. Marsh later discovered that the taxon Rhamphosaurus was preoccupied as a genus of lizard named in 1843. As a result, he suggested a move to a newly erected genus named Tylosaurus. This name means "knob lizard" in another reference to the elongated rostrum characteristic of the genus. It is derived from the Latin tylos (knob) and Ancient Greek σαῦρος. Despite coining the new genus, Marsh never formally transferred this Rhinosaurus species to Tylosaurus; this was first done in 1873 by Joseph Leidy. Tylosaurus subsequently became the almost universally accepted genus to include this species, the exception to this adoption being Cope, who refused to accept Marsh's new genus and continued to refer to its species as Liodon. Cope's persistence can be seen in his 1874 description of another species of Tylosaurus, which he named Liodon nepaeolicus. The type specimen of this species was discovered by geologist Benjamin Franklin Mudge near the Solomon River, and consists of several cranial fragments and a dorsal vertebra now catalogued as AMNH 1565. This species, whose specific epithet refers to Nepaholla, the Native American name for the Salomon River

Later discoveries and other species

::figure[src="https://upload.wikimedia.org/wikipedia/commons/5/53/Tylosaurus_spp._skulls.jpg" caption="Fossil skulls of ''Tylosaurus'' species '''(A)''' ''T. proriger'', '''(B)''' ''T. nepaeolicus'', '''(C)''' ''T. bernardi'', '''(D)''' ''T. saskatchewanensis'', and '''(E)''' ''T. pembinensis''"] ::

In his major work published in 1967, Dale A. Russell recognized only two valid species in Tylosaurus, namely T. proriger and T. nepaeolicus. However, throughout the 19th and 20th centuries, many species of mosasaurs coming from around the world, originally described as being from separate genera, were now recognized as belonging to Tylosaurus.

In 1885, Louis Dollo described the genus and species Hainosaurus bernardi from an almost complete but poorly preserved skeleton discovered in a phosphate quarry in the Ciply Basin in Belgium, the specimen having since been catalogued as IRSNB R23.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/d/d7/Tylosaurus_gaudryi_MNHN_1896–15_(2).png" caption="Holotype skull of ''T. gaudryi'' (MNHN 1896–15)"] ::

The fourth recognized species of the genus was described in 1896 by Armand Thevenin on the basis of a partial skull discovered at Éclusier-Vaux, in Somme, France. In his description, Thevenin thinks that this specimen, since catalogued as MNHN 1896–15, would represent a species of Mosasaurus, naming it Mosasaurus gaudryi. The specific epithet is named in honor of his mentor Jean Albert Gaudry, the latter having previously studied the skull and thinking that it would come from a species of Liodon. In 1992, Theagarten Lingham-Soliar reassigned the species to Hainosaurus, the latter having previously interpreted the holotype of this taxon as an additional specimen of H. bernardi. In 2005, Lindgren moved this species to Tylosaurus, notably due to its dental characteristics being closer to other lineages of the genus.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/1/17/Tylosaurus_ivoensis_tooth.png" caption="Isolated tooth of ''T. ivoensis''"] ::

In 1963, Per Ove Persson identified a new mosasaurid on the basis of isolated teeth discovered in a deposit located in an area called Ivö Klack, near Ivö Lake in the Kristianstad Basin in Scania, Sweden. Fossils from this same mosasaurid have been documented in this area since 1836, but it is from that year onwards that they are described as coming from a subspecies of Mosasaurus hoffmannii, being then named M. hoffmannii ivoensis, the second specific epithet referring to the type locality. In 1967, Russell elevated the taxon to a separate species within the genus, and assigned to it fossils from the Niobrara Formation of Kansas, including a partial skull. When the taxon was significantly revised in a in 2002 study, being reassigned to Tylosaurus, Lindgren and Mikael Siverson referred additional fossils to this latter that had been discovered at Ivö Klack, including cranial and vertebral remains. In their study, the authors also found that Russell's attributions of the Kansas fossils to this species were erroneous, the remains coming from a distinct taxon. In a 2008 paper, Caldwell and colleagues suggested that T. ivoensis might belong in the related genus Taniwhasaurus based on its dental features and its fossils distribution, but the authors see this as a subject for another study.

In 1964, Miguel Telles Antunes described the species Mosasaurus iembeensis from a partial skull excavated from the Itombe Formation near the town of Iembe (hence the name), Angola. In 1992, Lingham-Soliar argued that the cranial features were not consistent with those of Mosasaurus and were more characteristic of Tylosaurus, the species being renamed as T. iembeensis. However, the author did not identify the holotype skull, which he considered to reside in the collections of the NOVA University Lisbon without a catalogue number,

::figure[src="https://upload.wikimedia.org/wikipedia/commons/4/4c/Tylosaurus_(side_angle).jpg" caption="Reconstructed skeleton of ''T. saskatchewanensis''"] ::

In 2005, Michael J. Everhart described the species T. kansasensis based on several specimens that had been discovered in Kansas, again in the fossil record of the Niobrara Formation. The holotype specimen consists of a well-preserved skull and six cervical vertebrae cataloged as FHSM VP-2295, which was discovered in 1968 in Ellis County. The validity of this species was questioned as early as 2007 by Caldwell, to the point that in a 2016 study led with Jiménez-Huidobro and other authors, the latter considers it a juvenile form of T. nepaeolicus, thus making the first name a junior synonym of the second. This is disputed by Everhart himself in a 2017 book, but he only comments on the study as "poorly researched and written" without detailing how. An ontogenetic review of Tylosaurus conducted by Robert F. Stewart and Jordan Mallon in 2018 favors maintaining T. kansasensis as valid, while another conducted in 2020 by Amelia R. Zietlow prefers to follow the advice set out in the 2016 revision.

In 2006, Bullard wrote a Master of Science thesis describing the species T. saskatchewanensis from a partial skeleton catalogued as RSM P2588.1, known from the Bearpaw Formation. This specimen, nicknamed "Omācīw" (meaning "hunter" in Cree), was discovered in 1994 near Herbert Ferry, at the Lake Diefenbaker, Saskatchewan. Although originally described informally and via incompletely prepared fossils, the proposed taxon was nevertheless recognized as valid in some subsequent studies. In 2018, Bullard co-authored a multi-author study led by Jiménez-Huidobro which formally described Omācīw, which by then was more fully prepared, and confirmed its identity as belonging to a distinct species.

In 2020, Samuel Garvey wrote a thesis on a partial skull of Tylosaurus catalogued as TMP 2014.011.0001, known from the Puskwaskau Formation. With visibly distinct features from other species and having been discovered approximately 55 km northeast of Grande Prairie, Alberta, this makes the specimen the northernmost known occurrence of the genus, being then named T. borealis, in reference to its northernly presence.

Depiction history

::figure[src="https://upload.wikimedia.org/wikipedia/commons/f/f2/The_sea_which_once_covered_the_plains.jpg" caption="Henry Worrall]], 1872"] ::

When Cope described the holotype specimen of T. proriger in 1870, he visualized it as an "excessively elongated reptile", due to the morphology of the caudal vertebrae which suggested this. The following year, Cope added more details to his visualization of the animal. For him, the head of Tylosaurus would be conical in shape, with eyes on top, and having a jaw connected to a throat similar to that of a pelican, thus facilitating the entry of its prey. Still according to Cope, the animal would have had only the flippers located at the front of the body, those at the back being absent. The tail is seen as long and flat, used in eel-like locomotion. This depiction was followed in various works published during the late 19th century, although some depictions also depict the animal with a long neck.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/d/d4/Williston1898Mosasaurs.jpg" caption="Williston]]'s 1898 paper showing the first correct skeletal reconstruction of ''Tylosaurus'' (bottom)"] ::

In a major revision of mosasaurs published in 1898, Samuel Wendell Williston provided a new anatomical description of Tylosaurus that corrected many of the misconceptions of earlier paleontologists about the genus. Specifically, his paper included a rigorous skeletal reconstruction of T. proriger based on three partial specimens from the collections of the University of Kansas Natural History Museum. Thus, this reconstruction depicts the animal as very mobile marine predator with four flippers, a short neck and a much shorter tail than previous depictions, Williston also fixing a maximum body measurement close to those still cited today, i. e. 11 m long.

Description

Tylosaurus was a type of derived mosasaur, or a latecoming member with advanced evolutionary traits such as a fully aquatic lifestyle. As such, it had a streamlined body, an elongated tail ending with a downturn supporting a two-lobed fin, and two pairs of flippers. While in the past derived mosasaurs were depicted as akin to giant flippered sea snakes, it is now understood that they were more similar in build to other large marine vertebrates such as ichthyosaurs, marine crocodylomorphs, and archaeocete whales through convergent evolution.

Size

::figure[src="https://upload.wikimedia.org/wikipedia/commons/d/d7/Tylosaur_Size.svg" caption="Estimated size range of ''Tylosaurus'' compared with a human"] ::

Some species of Tylosaurus are among the largest known mosasaurs. The largest well-known specimen, a skeleton of T. proriger from the University of Kansas Natural History Museum nicknamed "Bunker" (KUVP 5033), has been estimated to measure between 12-15.8 m long. A fragmentary skeleton of another T. proriger from the Sternberg Museum of Natural History (FHSM VP-2496) may be from an even larger individual; Everhart estimated the specimen to come from a 14 m individual compared to his 12 meter estimate for Bunker. The genus exhibits Cope's rule, in which its body size has been observed to generally increase over geologic time. and Coniacian (90-86 mya), which included early T. nepaeolicus and its precursors, typically measured 5-7 m long During the Santonian (86-83 mya), T. nepaeolicus and newly-appearing T. proriger were 8-9 m long By the Early Campanian, T. proriger attained lengths of 13-14 m. Everhart speculated that because mosasaurs continuously grew throughout their lifetime, it would have been possible for some extremely old Tylosaurus individuals to reach 20 m in absolute maximum length. However, he stressed the lack of fossil evidence suggesting such sizes and the odds against any being preserved.

Other Campanian-Maastrichtian species were similarly large. The most recent maximum estimate for T. bernardi is 12.2 m by Lindgren (2005); historically the species was erroneously estimated at even larger sizes of 15-17 m. A reconstruction of T. saskatchewanensis by the Royal Saskatchewan Museum estimated a total length of over 9.75 m. A mounted skeleton of T. pembinensis, nicknamed "Bruce," at the Canadian Fossil Discovery Centre measures at 13.05 m long and was awarded a Guinness World Records for "Largest mosasaur on display" in 2014. However, the skeleton was assembled for display prior to Bullard and Caldwell (2010)'s reassessment that found the species' number of vertebrae to be exaggerated. T. "borealis" is estimated at 6.5-8 m in total length.

Skull

::figure[src="https://upload.wikimedia.org/wikipedia/commons/6/6c/FHSM-VP-3.jpg" caption="Profile view of a ''Tylosaurus proriger'' skull (FHSM VP-3)"] ::

The largest known skull of Tylosaurus is T. proriger KUVP 5033 (the "Bunker" specimen), estimated at 1.7 m long. Depending on age and individual variation, The head was strongly conical and the snout proportionally longer than most mosasaurs, with the exception of Ectenosaurus.

Cranium

::figure[src="https://upload.wikimedia.org/wikipedia/commons/c/c8/Tylosaurus_saskatchewanensis_holotype_RSM_P2588.1.png" caption="''T. saskatchewanensis'' holotype skull showing the elongated rostrum"] ::

The most recognizable characteristic of Tylosaurus is the elongated edentulous rostrum that protrudes from its snout, for which the genus is named. This is formed by the elongation of the front end of the premaxilla and dentary. The rostrum was small and acutely angled at birth, but rapidly developed into a blunt, elongated "knob." The snout is heavily built, supported by a broad and robust internarial bar (comprising the posterodorsal process of the premaxilla, nasals, and anterior process of the frontal). The snout holds the terminal branches for the trigeminal nerves through randomly scattered foramina on the rostrum and along the ventral margin of the maxilla, above the gum line.

| align = right | total_width = 440 | image1 = Tylosaurus proriger skull (CMN 8162).png | alt1 = A photograph of a mosasaur skull | image2 = Tylosaurus proriger skull ventral labeled.svg | alt2 = Drawing of a mosasaur skull's underside | footer = T. proriger skull (left) in dorsal view with mandibles; (right) in ventral view of cranium. Individual bones are labeled in the descriptions. The premaxilla, maxilla, and frontal bones border the external nares, or body nostril openings; unlike other mosasaurs, the prefrontal bones are excluded from the border of the nares by a long posterodorsal process of the maxilla.}} between the third and fourth tooth in T. nepaeolicus, and posterior to the fourth tooth in T. bernardi. Nare length relative to skull length varied between species: it is proportionally short in T. proriger (20-27% skull length

The external nares lead to the choanae (internal nares) in the palate, which provide passage from the nostrils to the throat. In Tylosaurus, they are shaped like a compressed teardrop and bordered by the vomers, palatines, and the maxilla. Anterior to the choanae, each vomer borders the fenestra for the Jacobson's organ, which is involved in the tongue-based sense of smell. It begins opposite of the fourth maxillary tooth in Tylosaurus, and also ends immediately past the fifth maxillary tooth in T. bernardi. The exit point for the veins leading to sinuses inside the palatine occur right in front of the Jacobson's organ between the vomers and maxilla. This differs from living varanids, where the exit occurs behind the organ.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/2/2c/Hainosaurus_bernardi_skull_4.JPG" caption="''T. bernardi'' holotype skull with its [[jugal bone"] ::

The frontal bone in Tylosaurus usually, but not always, possesses a low midline crest. It is most prominent in T. proriger, and is moderately developed in T. saskatchewanensis and T. bernardi, extending onto the premaxilla in the latter. The frontal crest is present but poorly developed in most T. nepaeolicus skulls, and occasionally lost in some mature individuals. The frontal overlaps the prefrontals and postorbitofrontals above the orbits (eye sockets), and the parietal posteriorly. The position of the pineal eye on the parietal is variable, either appearing close to the frontoparietal suture or contacting it. The orbits are bordered by the prefrontal, lacrimal, postorbitofrontal, and jugal bones. A diagnostic feature of Tylosaurus is that the prefrontals and postorbitofrontals overlap above the orbits, preventing contribution of the frontal. The jugal forms the bottom of the orbit; in Tylosaurus, it is L-shaped and has a distinctive serif-like extension at the lower back corner of the junction between the horizontal and vertical rami (arms) called the posteroventral process. The vertical ramus is overlapped by the postorbitofrontal in most species, and the horizontal ramus overlaps the maxilla. In T. bernardi, the vertical ramus is not overlapped but joins with the postorbitofrontal by a suture, and is much thicker than the horizontal ramus.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/8/84/Tylosaurus_spp._cranial_elements.jpg" caption="'''(A)''' quadrates, '''(B)''' skull roofs, and '''(C)''' teeth of '''(i)''' ''T. nepaeolicus'', '''(ii)''' ''T. proriger'', '''(iii)''' ''T. bernardi'', '''(iv)''' ''T. pembinensis'', and '''(v)''' ''T. saskatchewanensis''"] ::

The quadrate bones (homologous to the incus in mammals) are located at the back of the skull, articulating the lower jaw to the cranium and holding the eardrums. The complex anatomy of the bone renders it extremely diagnostic, even to the species level. In lateral view, the quadrate resembles a hook in immature T. nepaeolicus and T. proriger individuals, but in adult forms for both species and in T. bernardi, T. pembinensis, and T. saskatchweanensis it takes on a robust oval-like shape. The eardrum (tympanum) attached to the lateral surface of the bone within a bowl-like depression called the alar conch. The conch is shallow in T. nepaeolicus, T. proriger, and T. bernardi, and deep in T. pembinensis and T. saskatchewanensis. The alar rim is thin in T. nepaeolicus, T. proriger, and T. bernardi, and thick in T. bernardi, T. pembinensis, and T. saskatchewanensis. The suprastapedial process is a hook-like extension of bone that curves posteroventrally from the apex of the shaft into an incomplete loop, and it likely served as the attachment point for the depressor mandibulae muscles that opened the lower jaw. The process is slender and proportionally long in immature T. nepaeolicus and T. proriger, and thickened as the animals matured. The process is of similar length to T. proriger in T. saskatchwanensis and shorter in T. bernardi. In T. pembinensis, it abruptly turns medially at a 45° downward angle. A similar deflection appears in some juvenile T. nepaeolicus quadrates. Emerging from the posteroventral margin of the alar conch is the infrastapedial process. Its shape appears to changes ontogenetically in T. nepaeolicus and T. proriger; in the former, the process is absent in juveniles but appears as a small bump in adults, while in T. proriger, it is present as a subtle point in juveniles of and becomes a distinctively broad semicircle in adults. The process is small in T. bernardi, and in T. pembinensis and T. saskatchewanensis, it is rounded. In T. saskatchewanensis, the suprastapedial process almost touches the infrastapedial process. At the bottom of the shaft is the mandibular condyle, which forms the joint between the quadrate and the lower jaw. It is rounded in shape in adults. On the medial surface of the bone, a thick, pillar-like vertical ridge often protrudes beyond the dorsal margin of the quadrate so that it is visible in lateral view.

Jaws and teeth

The upper jaws include the premaxilla and maxilla, and the lower jaws include the dentary, splenial, coronoid, angular, surangular, and prearticluar-articular (like other squamates, the prearticular is fused to the articular). The premaxilla, maxilla, and dentary house the marginal dentition, and the pterygoids house palatal dentition. On each side of the skull, Tylosaurus had 2 premaxillary teeth, 12 to 13 maxillary teeth, 13 dentary teeth, and 10 to 11 pterygoid teeth.

Tylosaurine dentaries were elongate; the dentary is between 56 and 60% of total length of the entire lower jaw in adult T. nepaeolicus and T. proriger, about 55% in T. pembinensis, and 62% in T. saskatchwanensis. The dentary is robust, though not as strongly built as it is in Mosasaurus, Prognathodon, or Plesiotylosaurus. The ventral margin of the dentary ranges from straight to slightly concave. A small dorsal ridge appears anterior to the first dentary tooth in mature individuals of T. proriger.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/b/b8/Tylosaurus_proriger_skull_brightened.jpg" caption="Mounted ''T. proriger'' skull (a cast of KUVP 5033) showing both marginal and pterygoid teeth"] ::

The marginal dentition of most species is adapted for cutting large marine vertebrates, while those in T. ivoensis and T. gaudryi appear more optimized for piercing or smashing prey, and T. "borealis" in both piercing and cutting. Marginal teeth are triangular with a slight recurve towards the back of the jaws so that the lingual (tongue-facing) side forms a U-shaped curve. From top view, they are compressed at the lingual and labial (lip-facing) sides to form an oval-like shape. Teeth of immature T. proriger are initially compressed, but become conical in adulthood. Carinae (cutting edges) are finely serrated with small denticles except in juvenile T. nepaeolicus. In T. pembinensis, they are faint. The teeth generally have both anterior and posterior carinae, but some anterior teeth may have only anterior carinae. The placement of carinae, if paired, is not always equal; in at least T. proriger, T. ivoensis, T. gaudryi, and T. pembinensis, they are positioned such that the surface area of the tooth's lingual side is greater than the labial side. Both sides are always balanced in area in T. bernardi. The enamel surface is lined with thin fine ridges called striations that run vertically from the tooth's base. The surface is also either smooth or faintly faceted, in which it is flattened into multiple sides to form a prism-like geometry.

| align = right | direction = vertical | total_width = 220 | image1 = Tylosaurus nepaeolicus AMNH 124, 134.png | alt1 = A photograph of a partial mosasaur skull | image2 = Tylosaurus gaudryi MNHN 1896–15 (1).png | alt2 =A photograph of a partial mosasaur skull | footer = Jaws showing teeth of proriger group T. nepaeolicus (top) and ivoensis group T. gaudryi (bottom) Bardet et al. (2006) classified Tylosaurus species into two morphological groups based on marginal dentition. The North American proriger group includes T. proriger and T. nepaeolicus and is characterized by teeth with smooth or faint facets, less prominent carinae, and a vein-like network of primitive striations extending to near the tip. The group was originally defined as having slender teeth, Marginal teeth in T. gaudryi are virtually indistinguishable from those in T. ivoensis. T. iembeensis was not placed within either group; no further description is known of its teeth other than having striations and no facets. The distinction of an ivoensis group is contentious. Caldwell et al. (2008) argued that T. pembinensis cannot be compared with T. ivoensis as the former's teeth are not fluted, and that T. ivoensis is more allied with the distinctively fluted teeth of Taniwhasaurus. Jiménez-Huidobro and Caldwell (2019) listed the absence of marginal fluting as a diagnostic (taxon-identifying) trait that differentiates Tylosaurus from Taniwhasaurus.

The pterygoid teeth may have enabled ratchet feeding, in which the upper teeth held prey in place as the lower jaw slides back and forth via a streoptostylic jaw joint. The bases of the pterygoid teeth are nearly circular, and each tooth is divided into front and back-facing sides of near-equal surface area via a pair of faint buccal and lingual carinae, except in T. gaudryi, in which the teeth are mediolaterally compressed. Carinae are not serrated. The anterior surface tends to be either smooth of faintly faceted, while the posterior surface is striated.

Postcranial skeleton

Both pectoral and pelvic girdles are unfused in adult Tylosaurus, in contrast to other taxa (e.g., Prognathodon overtoni). Tylosaurus is also distinguished from other mosasaurs by a scapula that is significantly smaller than the coracoid and the absence of the anterior emargination of the coracoid, as well as the absence of a well-developed pubic tubercle.

Tylosaurus limbs are primitive relative to other mosasaurs; their stylopodia (humeri and femora) lack both the complex muscle attachment sites and extreme proximodistal shortening present in other derived taxa. Both carpals and tarsals in tylosaurines are mostly unossified; while other mosasaurs typically have between three and five carpals and tarsals, adult Tylosaurus never possess more than two ossified carpal bones (usually only the ulnare, sometimes the ulnare and distal carpal four) and two ossified tarsal bones (usually only the astragalus, sometimes the astragalus and distal tarsal four). Hyperphalangy (increased number of phalanges relative to the ancestral condition) is present in both fore- and hindlimbs, and the phalanges are spindle-shaped, unlike the short, blocky hourglass-shaped phalanges possessed by mosasaurines. The pisiform appears to be either unossified or absent in tylosaurines. The functional consequences of differences in limb anatomy across different mosasaur clades is unclear.

Tylosaurus had 29 to 30 presacral vertebrae, 6 to 7 pygal vertebrae, and 89 to 112 caudal vertebrae; due to the lack of a bony articulation between the ilium and vertebral column, it is unclear whether any mosasaurs possessed true sacral vertebrae. In all tylosaurines, like in plioplatecarpines, the chevrons articulate to the caudal vertebrae, and are not fused to them, as they are in mosasaurines. The tail possesses a distinct downward curve, suggesting the presence of a tail fluke.

Soft tissue

Skin and coloration

| align = left | direction = horizontal | total_width = 400 | image_gap = 1 | image1 = Tylosaurus sp skin 1.JPG | image2 = Tylosaurus nepaeolicus NT.png | footer = Preserved skin of Tylosaurus (left); restoration of T. nepaeolicus with coloration based on melanosomes found in preserved skin (right)

Fossil evidence of the skin of Tylosaurus in the form of scales has been described since the late 1870s. These scales were small and diamond-shaped and were arranged in oblique rows, comparable to that found in modern rattlesnakes and other related reptiles. However, the scales in the mosasaur were much smaller in proportion to the whole body. An individual measuring 5 m in total body length had dermal scales measuring 3.3 x, and in each square inch (2.54 cm) of the mosasaur's underside an average of ninety scales were present.

Microscopic analysis of scales in a T. nepaeolicus specimen by Lindgren et al. (2014) detected high traces of the pigment eumelanin indicative of a dark coloration similar to the leatherback sea turtle in life. This may have been complemented with countershading, present in many aquatic animals, though the distribution of dark and light pigments in the species remains unknown. A dark-colored form would have provided several evolutionary advantages. Dark coloration increases absorption of heat, allowing the animal to maintain elevated body temperatures in colder environments. Possession of this trait during infancy would in turn facilitate fast growth rates. Unreflective dark coloring and countershading would have provided the mosasaur with increased camouflage. Additional speculative functions includes increased tolerance to solar ultraviolet radiation, strengthened integuments. The study remarked that certain melanism-coding genes are pleiotropic for increased aggression.

Respiratory system

| image1 = Tylosaurus windpipe AMNH.jpg | image2 = Tylosaurus trachea AMNH.jpg | footer = Preserved bronchi (left) and trachea (right) in AMNH FR 221 AMNH FR 221 preserves parts of the cartilaginous respiratory system. This includes parts of the larynx (voice box), trachea (windpipe), and bronchi (lung airways). They were however only briefly described in the preserved position by Osborn (1899). The larynx is poorly preserved; a piece of its cartilage first appears below just between the pterygoid and quadrate and extends to behind the latter. This connects to the trachea, which appears below the atlas vertebra but is not preserved afterwards. The respiratory tract reappears below the fifth rib as a pair of bronchi and extends to just behind the as-preserved coracoids where preservation is lost. The pairing is suggestive of two functional lungs like modern limbed lizards but unlike snakes. Similar branching is also found in Platecarpus while in Mosasaurus the organ is dislocated. A bifurcation point's position ahead of the forelimbs would be unlike terrestrial lizards, whose point is within the chest region, but similar to the short trachea and parallel bronchi of whales.

Classification

Taxonomy

::figure[src="https://upload.wikimedia.org/wikipedia/commons/e/e4/Tylosaurus_pembinensis_1DB.jpg" caption="Restoration]] of ''T. pembinensis''"] ::

Tylosaurus is classified within the family Mosasauridae in the superfamily Mosasauroidea. The genus is the type genus of its own subfamily, the Tylosaurinae. Other members of this group include Taniwhasaurus and possibly Kaikaifilu, and the subfamily is defined by a shared feature of an elongated premaxillary rostrum that does not bear teeth.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/4/40/SGM-M1.jpg" caption="The [[Turonian]]-aged skull of ''T. sp. aff. kansasensis'' (SGM-M1) is one of the oldest known fossils of ''Tylosaurus''."] ::

Tylosaurus was among the earliest derived mosasaurs. The oldest fossil attributable to the genus is a premaxilla (TMM 40092-27) recovered from Middle Turonian deposits of the Arcadia Park Shale in Texas, which is dated between 92.1 and 91.4 million years old based on correlations with index fossils. Although formally referred to as Tylosaurinae incertae sedis during its first description, it was remarked to probably belong to T. kansasensis. The specimen was later listed within said species in a 2020 reexamination. A slightly younger specimen is of a skull (SGM-M1) of an indeterminate Tylosaurus species similar to T. kansasensis from the Ojinaga Formation in Chihuahua, Mexico, dated around ~90 million years old at earliest. A tooth from a Late Maastrichtian deposit in Nasiłów, Poland dating close to the Cretaceous–Paleogene boundary has been attributed to Hainosaurus sp. With the incorporation of Hainosaurus as a synonym of Tylosaurus, this also makes the genus one of the last mosasaurs. Currently, eight species of Tylosaurus are recognized by scientists as taxonomically valid. They are as follow: T. proriger, T. nepaeolicus, T. bernardi, T. gaudryi, T. ivoensis, T. iembeensis, T. pembinensis, and T. saskatchewanensis. The validity of two additional taxa remain unsettled; there is still debate whether T. kansasensis is synonymous with T. nepaeolicus, and T. "borealis" has yet to be described in a formal publication.

Phylogeny and evolution

In 2020, Madzia and Cau performed a Bayesian analysis to better understand the evolutionary influence on early mosasaurs by contemporaneous pliosaurs and polycotylids by examining the rates of evolution in mosasauroids like Tylosaurus (specifically T. proriger, T. nepaeolicus, and T. bernardi). A Bayesian analysis in the study's implementation can approximate numerically defined rates of morphological evolution and ages of divergence of clades. The Tylosaurinae was approximated to have diverged from the Plioplatecarpinae around 93 million years ago; the divergence was characterized by the highest rate of evolution among all mosasaurid lineages. This trend of rapid evolution coincided with the extinction of the pliosaurs and a decrease in polycotylid diversity. The study noted converging traits between Tylosaurus, pliosaurs, and some polycotylids in tooth morphology and body size. However, there was no evidence to suggest that Tylosaurus or its precursors evolved as a result of out-competing and/or driving to extinction the pliosaurs and polycotylids. Instead, Madiza and Cau proposed that Tylosaurus may have taken advantage of the extinction of the pliosaurs and decline of polycotylids to quickly fill the ecological void they left behind. The Bayesian analysis also approximated a divergence of T. nepaeolicus from the rest of the genus around 86.88 million years ago and a divergence between T. proriger and T. bernardi around 83.16 million years ago. The analysis also generated a paraphyletic status of the genus, approximating Taniwhasaurus to have diverged from Tylosaurus around 84.65 million years ago, but this result is not consistent with previous phylogenetic analyses.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/1/1d/Tylosaurus_ontogram.jpg" caption="Ontogram demonstrating the evolution of ''T. nepaeolicus'' into ''T. proriger'' through [[peramorphosis"] ::

In the Western Interior Seaway, two species—T. nepaeolicus and T. proriger—may represent a chronospecies, in which they make up a single lineage that continuously evolves without branching in a process known as anagenesis. This is evident by how the two species do not stratigraphically overlap, are sister species, share minor and intermediate morphological differences such as a gradual change in the development of the quadrate bone, and lived in the same locations. The means by which this lineage evolved has been hypothesized to be through one of two evolutionary mechanisms related to changes in ontogeny. First, Jiménez-Huidobro, Simões, and Caldwell proposed in 2016 that T. proriger evolved as a paedomorph of T. nepaeolicus, in which the descendant arose as a result of morphological changes through the retention of juvenile features of the ancestor in adulthood. This was based on the presence of a frontal crest and convex borders of the parietal bone of the skull shared in both juvenile T. nepaeolicus and all T. proriger but lost in adult T. nepaeolicus. However, an ontogenetic study by Zietlow (2020) found that it was unclear whether this observation was a result of paedomorphosis, although this uncertainty may have been due that the sample size of mature T. nepaeolicus was too low to determine statistical significance. Second, the same study proposed an alternative hypothesis of peramorphosis, in which T. proriger evolved by developing traits found in mature T. nepaeolicus during immaturity. Based on results from a cladistical ontogram developed using data from 74 Tylosaurus specimens, the study identified a multitude of traits that were present in all T. proriger and mature T. nepaeolicus but absent in juvenile T. nepaeolicus: the skull size and depth are large, the length of the elongated rostrum exceeds 5% of the total skull length, the quadrate suprastapedial processes are thick, the overall quadrate shape converges, and the posteroventral process is fan-like.

The following cladogram is modified from a phylogenetic analysis by Jiménez-Huidobro & Caldwell (2019) using Tylosaurus species with sufficiently known material to model accurate relationships; T. gaudryi, T. ivoensis, and T. iembeensis were excluded from the analysis due to extensive missing data (i.e., lack of material with scoreable phylogenetic characters).

|1={{clade |1=Aigialosaurus |2={{clade |1=Komensaurus carrolli |2={{clade |label1=Russellosaurina |1={{clade |1={{clade |1={{clade |1=Tethysaurus nopcsai |2=Pannoniasaurus inexpectatus |2={{clade |1=Yaguarasaurus columbianus |2={{clade |1=Russellosaurus coheni |2=Romeosaurus fumanensis |2={{clade |2={{clade |1={{clade |1=Angolasaurus bocagei |2={{clade |1=Ectenosaurus clidastoides |2=Selmasaurus johnsoni |2={{clade |1=Plesioplatecarpus planifrons |2={{clade |1=Latoplatecarpus willistoni |2={{clade |1=Plioplatecarpus |2=Platecarpus tympaniticus |label1=Tylosaurinae |1={{clade |1={{clade |1=Taniwhasaurus oweni |2=Taniwhasaurus antarcticus |2={{clade |1=Tylosaurus nepaeolicus [[File:Tylosaurus nepaeolicus AMNH 124, 134.png|80px]] |2={{clade |1={{clade |1=Tylosaurus proriger [[File:Tylosaurus proriger (USNM V6086).png|80px]] |2=Tylosaurus bernardi [[File:Tylosaurus bernardi IRSNB R23 Skull.png|80px]] |2={{clade |1=Tylosaurus pembinensis [[File:Tylosaurus pembinensis MMV 95 2.png|80px]] |2=Tylosaurus saskatchewanensis [[File:Tylosaurus saskatchewanensis holotype flipped RSM P2588.1.png|80px]] |2={{clade |1=Halisaurinae |2=Mosasaurinae

Paleobiology

Function of edentulous rostrum

::figure[src="https://upload.wikimedia.org/wikipedia/commons/0/08/Taniwhasaurus_antarcticus_skull_diagram.png" caption="Reconstruction of [[CT scan]]s of a ''[[Taniwhasaurus]]'' skull, with the [[ophthalmic nerve]] (''ramus ophthalmicus'') in red"] ::

Russell (1967) first speculated the elongated rostrum was used to "stun" prey or defend against sharks, though provided no further explanation. This combat hypothesis was further elaborated in Lingham-Soliar's (1992) review of Tylosaurus biomechanics. He observed that the internarial bar was unusually robust for a mosasaur, and sutured to the frontal with deep interdigitations such that there is a large interfacial shear area. This probably allowed for greater resistance to bending, shearing, and breaking forces though more effective shock absorption and stress transfer from a blunt force to the rostrum. and orcas to attack prey. Lingham-Soliar (1998) would later suggest possible evidence of a ram-attack in action in the skull of a subadult Mosasaurus whose braincase was fractured in life by a concentrated force, which he argued was rammed by a T. bernardi due to lack of damage elsewhere in the skull that would indicate other causes of injury. Lingham-Soliar doubted that the elongated rostrum was involved in sexual selection, such as male-male combat, as previous surveys of Tylosaurus skulls found no evidence of sexual dimorphism in the length of the rostrum. This was supported by Konishi et al.'s (2018) finding that the edentulous rostrum began to grow exceptionally early in development, which should not occur if the trait was sexual in nature but reasonable for general-purpose use like ramming. Likewise, Zietlow (2020) found no evidence of sexual dimorphism relating to rostrum growth.

An alternate hypothesis holds that the rostrum was used as a sensory tool. It was long known since Russell (1967) that the edentulous region of the rostrum contains numerous foramen that housed exits for the ophthalmic nerve, which transmits sensory information from the skin to the brain. But that the elongated rostrum specifically functioned for sensation was first advanced by Álvarez–Herrera et al. (2020) based on CT scans of a Taniwhasaurus skull. They discovered a large network of internal canals of the ophthalmic nerve along the rostrum with up to 16 branching terminals inside the front end of the premaxilla, half of which exit into external foramina that dot the front and dorsal surfaces of the bone's edentulous region. This pattern was later found in preliminary CT scans of a T. nepaeolicus skull by Paulina-Carabajal et al. (2023), which similarly observed profuse branching of the ophthalmic nerve inside the premaxilla exiting through many foramina on the dorsal tip of the rostrum. The high concentration of sensory nerves within the edentulous portion of the rostrum suggests that it housed a mechanosensory or electrosensory organ. Similar adaptations were previously found in plesiosaurs and ichthyosaurs. It was also suggested that the heavy branching inside the rostrum is associated with pathways for blood supply, and therefore a thermoregulation mechanism. Paleontologist Gordon Bell remarked in 2002 that any injuries to the snout, such as if inflicted via ramming, would have been extremely painful due to the nerve endings.

Growth and development

The ontogenetic history of Tylosaurus remains poorly researched despite the wealth of specimens including of juveniles. Several insights have been made nonetheless. Mosasaurs like Tylosaurus likely gave birth to live young. The youngest known Tylosaurus individual (FHSM VP-14845) is a newborn (neonate) estimated to have measured 30 cm in skull length and 2.23 m in total length, about 17.2% the size of Bunker and 24.8–27.9% of the estimated maximum lengths for T. nepaeolicus. The premaxilla of this specimen lacked an edentulous elongation of the rostrum; this trait already appears in juvenile specimens only slightly larger than FHSM VP-14845, including T. nepaeolicus and T. proriger specimens with estimated skull lengths of 40 cm and 60 cm respectively.

::figure[src="https://upload.wikimedia.org/wikipedia/commons/7/7c/Tylosaurus_growth_curve_from_LAGs.svg" caption="Growth curve and sexual maturity based on the histology of LAGs"] ::

A 2007 study by Pellegrini applied skeletochronology on the humeri of two juvenile Tylosaurus sp. specimens to measure the growth rates based on lines of arrested growth (LAGs). In poikilothermic sauropsids, bone growth tends to slow or freeze seasonally, either due to physiological cycles or in response to seasonal change in their environment. This creates a thin layer of avascular tissue called an annulus that, when deposited as a single line due to near-complete cease of growth, is called an LAG. The seasonal nature of LAG deposition can therefore proxy as annual growth rings, and measuring the amount of bone deposition during growing seasons via the distance of each LAG from the center of the bone can infer the animal's growth rate each year starting from the first year of life. The older humerus in Pellegrini's sample indicated the individual was 14 years old at death while the younger was aged 7 years. Both showed similar logarithmic growth curves that grew rapidly in the first few years but reduced to a steadier rate starting year 5 in the older specimen and year 6 in the younger. This may represent the ages when the individuals reached sexual maturity. The distance of first-year LAGs to the bone center in both specimens were longer than those in Platecarpus and Clidastes humeri, which suggests that Tylosaurus was larger at birth rather than being born at a similar size and then growing more rapidly. Like the other two mosasaurs, the overall growth rate was similar to modern monitor lizards but accelerated, which Pellegrini attributed to the need for increased metabolic rates to adapt to an aquatic environment.

A 2018 study by Stewart and Mallon attempted to assess the degree of allometry in the growth of the T. proriger skull based on fifteen specimens. Their analysis of covariance test failed to reject isometry from five of the seven skull variables tested. The two definitively shown to be allometric were length of the edentulous rostrum (grew proportionally smaller with age) and height of the quadrate (grew proportionally larger with age). The isometric results were interpreted as most likely an artifact of low sample size, also known as "soft isometry", and the authors maintained that a larger sample size could reveal allometry in the five other variables. The negative allometry found for the edentulous rostrum was also unexpected, as it contradicted prior observations that juvenile Tylosaurus tended to have shorter rostrums than adults.

| align = right | total_width = 400 | image1 = Tylosaurus proriger ontogram.png | image2 = Tylosaurus nepaeolicus kansasensis ontogram.png | footer = Ontograms of T. proriger (left) and T. nepaeolicus/kansasensis (right) A 2020 study by Zietlow used cladistic analysis to reconstruct ontograms (growth series charts) of the skull in T. proriger, T. nepaeolicus, and T. kansasensis based on 79 specimens. This approach tracks morphological changes with maturity although, it does not correlate them to particular ages. The resulting ontogram for T. proriger yielded 17 growth series. The most notable changes included the appearance of bony processes on the several bones of the skull, shrinking of the premaxillary foramina, ossification of the basioccipital and quadrate, reshaping of the premaxilla-maxilla suture to an M-shape, and increase in tooth size relative to the skull. T. nepaeolicus and T. kansasensis was coalesced into a single ontogram to test whether they were a single species. The result showed no evidence of separation; instead, the T. kansasensis specimens were recovered at the earlier stages and T. nepaeolicus at the later stages over a total of 12 growth series. The most notable changes were the enlargement of bony processes in several bones, enlargement of the premaxillary foramina, change in pterygoid shape, ossification of the quadrate and change from a hook-like to elongate shape, development of the M-shape in the premaxilla-maxilla suture, development of an extra dentary tooth, and increase in tooth size relative to the skull. Skull size in both ontograms as well as skull depth in T. proriger progressively increased through all growth stages. Statistical tests showed that both total skull length and quadrate height are reliably correlated with maturity in T. proriger and T. nepaeolicus/kansasensis. Neither ontogram showed evidence of sexual dimorphism, although Zietlow held that this is either an artifact of small sample size or because sex-differentiation does not manifest in the skull.

Metabolism

| align = left | total_width = 400 | image1 = Double-crested cormorant at Sutro Baths-6460.jpg | image2 = Sea Turtle.jpg | footer = Tylosaurus was warm-blooded and maintained a body temperature closer to seabirds (left) than cold-blooded sea turtles (right). Nearly all squamates are characterized by their cold-blooded ectothermic metabolism, but mosasaurs like Tylosaurus are unique in that they were likely endothermic, or warm-blooded. The only other known lizard with such a trait is the Argentine black and white tegu, though only partially. Endothermy in Tylosaurus was demonstrated in a 2016 study by Harrell, Pérez‐Huerta, and Suarez by examining δ18O isotopes in Tylosaurus bones. δ18O levels can be used to calculate the internal body temperature of animals, and by comparing such calculated temperatures between coexisting cold-blooded and warm-blooded animals, the type of metabolism can be inferred. The study used the body temperatures of the cold-blooded fish Enchodus and sea turtle Toxochelys (correlated with ocean temperatures) and warm-blooded seabird Ichthyornis from the Mooreville Chalk as a proxy. Analyzing the isotope levels of eleven Tylosaurus specimens an average internal body temperature of 34.3 C was calculated. This was much higher than the body temperature of Enchodus and Toxochelys (28.3 C and 27.2 C respectively) and similar to that of Ichthyornis (38.6 C). Harrell, Pérez‐Huerta, and Suarez also calculated the body temperatures of Platecarpus and Clidastes with similar numbers, 36.3 C and 33.1 C respectively. The fact that the other mosasaurs were much smaller in size than Tylosaurus and yet maintained similar body temperatures made it unlikely that Tylosauruss body temperature was the result of another metabolic type like gigantothermy. Endothermy would have provided several advantages to Tylosaurus such as increased stamina for foraging larger areas and pursuing prey, the ability to access colder waters, and better adaptation to withstand the gradual cooling of global temperatures during the Late Cretaceous.

Mobility

Scientists previously interpreted Tylosaurus as an anguilliform swimmer that moved by undulating its entire body like a snake due to its close relationship with the animal. However, it is now understood that Tylosaurus actually used carangiform locomotion, meaning that the upper body was less flexible and movement was largely concentrated at the tail like in mackerels. This is corroborated by a statistical reconstruction of the tail fin by Song and Lindgren (2025), which predicted an outline resembling those in carangiform sharks. A BS thesis by Jesse Carpenter published in 2017 examined the vertebral mobility of T. proriger spinal columns and found that the dorsal vertebrae were relatively rigid but the cervical, pygal, and caudal vertebrae were more liberal in movement, indicating flexibility in the neck, hip, and tail regions. This contrasted with more derived mosasaurs like Plotosaurus, whose vertebral column was stiff up to the hip. Interestingly, an examination of a juvenile T. proriger found that its cervical and dorsal vertebrae were much stiffer than those in adult specimens. This may have been an evolutionary adaptation among young individuals; a more rigid tail-based locomotion is associated with faster speed, and this would allow vulnerable juveniles to better escape predators or catch prey. Older individuals would see their spine grow in flexibility as predator evasion becomes less important for survival.

Tylosaurus likely specialized as an ambush predator. It was lightweight for a mosasaur of its size, having a morphological build designed to vastly reduce body mass and density. Its pectoral and pelvic girdles and paddles, which are associated with weight, are proportionally small. Its bones were highly cancellous and were likely filled with fat cells in life, which also increased buoyancy. It is unlikely that the latter trait was evolved in response to increasing body size as the similarly sized Mosasaurus hoffmannii lacked highly cancellous bone. These traits allowed Tylosaurus to be more conservative in its energy requirements, which is useful when traveling between ambush sites over large distances or through stealth. In addition, a reduced body density likely helped Tylosaurus to rapidly accelerate during an attack, assisted with the long and powerful tail of the mosasaur.

A 1988 study by Judith Massare attempted to calculate the sustained swimming speed, the speed at which the animal moves without tiring, of Tylosaurus through a series of mathematical models incorporating hydrodynamic characteristics and estimations of locomotive efficiency and metabolic costs. Using two T. proriger specimens, one 6.46 m long and the other 6.32 m, she calculated a consistent average maximum sustained swimming speed of 2.32 m/s. However, when testing whether the models represented an accurate framework, they were found to exaggerated. This was primarily because the variables accounting for drag may have been underestimated; estimation of drag coefficients for an extinct species can be difficult as it requires a hypothetical reconstruction of the morphological dimensions of the animal. Massare predicted that the actual sustained swimming speed of Tylosaurus was somewhere near half the calculated speed.

Feeding

::figure[src="https://upload.wikimedia.org/wikipedia/commons/c/cc/Tylosaurus_gut_contents_(SDSM_10439).svg" caption="Schematic of the gut content in a single ''Tylosaurus'' (SDSM 10439)"] ::

One of the largest marine carnivores of its time, Tylosaurus was an apex predator that exploited the wide variety of species in the marine fauna of its ecosystem. Stomach contents are well documented in the genus, which includes other mosasaurs, plesiosaurs, turtles, birds, bony fish, and sharks. Additional evidence from bite marks suggests the animal also preyed on giant squid and ammonites.

The enormous and varied appetite of Tylosaurus can be demonstrated in a 1987 find that identified fossils of a mosasaur measuring 2 m or longer, the diving bird Hesperornis, a Bananogmius fish, and possibly a shark all within the stomach of a single T. proriger skeleton (SDSM 10439) recovered from the Pierre Shale of South Dakota.{{efn|Identification of the mosasaur and shark vary. Scientists have identified the mosasaur as either a Platecarpus, or Latoplatecarpus. or of uncertain identity. Other records of stomach contents include a sea turtle in a T. bernardi-like species,{{efn|Usually identified as Hainosaurus sp.; Puncture marks on fossils of ammonites, the carapace of a Protostega, and the gladius of an Enchoteuthis have been attributed to Tylosaurus. ::figure[src="https://upload.wikimedia.org/wikipedia/commons/b/b8/Tylosaurus_hunting_Xiphactinus.jpg" caption="Skeletal reconstruction of ''Tylosaurus'' hunting a ''Xiphactinus'' at the [[Academy of Natural Sciences of Drexel University]], Philadelphia"] ::

Pasch and May (2001) reported bite marks from a dinosaur skeleton known as the Talkeetna Mountains Hadrosaur, which was found in marine strata of the Turonian-age Matanuska Formation in Alaska. The features of these marks were found to closely match that of the teeth of T. proriger. Because the fossil's locality was of marine deposits, the study reasoned that the dinosaur must have drifted offshore as a bloat-and-float carcass that was subsequently scavenged by the mosasaur. It was unlikely that the marks were a result of predation, as that would have led to a puncture, preventing the buildup of the bloating gases that allowed the corpse to drift out to sea in the first place. Garvey (2020) criticized the lack of conclusive evidence to support this hypothesis and ruled out T. proriger as a possible culprit, given that the species did not appear until the Santonian and is exclusive to the Western Interior Seaway.

Social behavior

::figure[src="https://upload.wikimedia.org/wikipedia/commons/0/06/Tylosaurus_attack_diagram.jpg" caption="Reconstructed scenario of the attack (FHSM VP-2295); the red circles represent bite marks on the skull roof"] ::

The behavior of Tylosaurus towards each other may have been mostly aggressive, evidenced by fossils with injuries inflicted by another of their own kind. Such remains were frequently reported by fossil hunters during the late 19th and early 20th centuries, but few examples reside as specimens in scientific collections. Many of these fossils consist of healed bite marks and wounds that are concentrated around or near the head region, implying that there were the result of non-lethal interaction, but the motives of such contact remain speculative. In 1993, Bruce M. Rothschild and Larry D. Martin noted that some modern lizards affectionately bite their mate's head during courtship, which can sometimes result in injuries. Alternatively, they also observed that some males lizards also employ head-biting as territorial behavior against rivals in a show of dominance by grappling the head to turn over the other on its back. It is possible that Tylosaurus behaved in similar ways.

At least one fatal instance of intraspecific combat among Tylosaurus is documented in the holotype of the discussed species T. kansasensis (FHSM VP-2295), representing a 5 m long animal, which possesses numerous injuries that indicate it was killed by a larger Tylosaurus. The skull roof and surrounding areas exhibit signs of trauma in the form of four massive gouges, and the dentary contains at least seven puncture wounds and gouges. These pathologies are characteristic of bite marks from a larger Tylosaurus that measured around 7 m in length. The largest of the marks are about 4 cm in length, matching the size of large mosasaur teeth, and they are positioned along two lines that converge close to 30°, matching the angle that each jaw converges towards in a mosasaur skull. In addition, FHSM VP-2295 suffered damage to its neck: the cervical vertebrae were found articulated at an unnatural angle of 40° relative to the long axis of the skull. The pattern of preservation makes it unlikely that the condition of the vertebrae was a result of disturbances by scavengers and instead indicates damage caused by a violently twisted neck during life. In a reconstructed scenario, the larger Tylosaurus would have first attacked at an angle slightly below the left side of the victim's head. This impact would cause the victim's skull to roll to the right side, allowing the aggressor to sink its teeth into the skull roof and right lower jaw, crushing the jaw and causing further breaks of nearby bones, such as the pterygoid, and the twisting of the jaw outwards, which would cause the quadrate to detach from its position and for the spinal cord to twist and sever at the skull's base, leading to a swift death.

Paleopathology

Examining 12 North American Tylosaurus skeletons and one T. bernardi skeleton, Rothschild and Martin (2005) identified evidence of avascular necrosis in every individual. For aquatic animals, this condition is often a result of decompression illness, which is caused when bone-damaging nitrogen bubbles build up in inhaled air that is decompressed either by frequent deep-diving trips or by intervals of repetitive diving and short breathing. The studied mosasaurs likely gained avascular necrosis through such behaviors, and given its invariable presence in Tylosaurus it is likely that deep or repetitive diving was a general behavioral trait of the genus. The study observed that between 3-15% of vertebrae in the spinal column of North American Tylosaurus and 16% of vertebrae in T. bernardi were affected by avascular necrosis. Carlsen (2017) posited that Tylosaurus gained avascular necrosis because it lacked the necessary adaptations for deep or repetitive diving, although noted that the genus had well-developed eardrums that could protect themselves from rapid changes in pressure.

Unnatural fusion of some vertebrae in the tail has been reported in some Tylosaurus skeletons. A variation of these fusions may concentrate near the end of the tail to form a single mass of multiple fused vertebrae called a "club tail." Rothschild and Everhart (2015) surveyed 23 North American Tylosaurus skeletons and one T. bernardi skeleton and found that five of the North American skeletons exhibited fused tail vertebrae. The condition was not found in T. bernardi, but this does not rule out its presence due to the low sample size. Vertebral fusion occurs when the bones remodel themselves after damage from trauma or disease. However, the cause of such events can vary between individuals and/or remain hypothetical. One juvenile specimen with the club tail condition was found with a shark tooth embedded in the fusion, which confirms that at least some cases were caused by infections inflicted by predator attacks. The majority of vertebral fusion cases in Tylosaurus were caused by bone infections, but some cases may have alternatively been caused by any type of joint disease such as arthritis. However, evidence of joint disease was rare in Tylosaurus when compared to mosasaurs such as Plioplatecarpus and Clidastes. Similar amassing of remodeled bone is also documented in bone fractures in other body parts. One T. kansasensis specimen possesses two fractured ribs that fully healed. Another T. proriger skull shows a fractured snout, probably caused by ramming into a hard object such as a rock. Presence of some healing indicates that the individual survived for some extended time before death, albeit under extreme pain.

Paleoecology

::figure[src="https://upload.wikimedia.org/wikipedia/commons/5/59/LateCretaceousMap.jpg" caption="30°N]]."] ::

Throughout its ~20 million-year history, Tylosaurus was endemic to the North Atlantic Circle Basin, a geographic region comprising the North Atlantic and neighboring waters including the Western Interior Seaway in North America and Mediterranean Tethys in Europe above 30°N. There are a few exceptions of occurrences in warmer low-latitude zones in scientific literature. This includes T. iembeensis from the Coniacian of Angola, an isolated tooth from Maastrichtian deposits in the Democratic Republic of the Congo suggested as a possible Hainosaurus,

Habitat preference

Stratigraphic evidence in the WIS is inconsistent on whether Tylosaurus preferred a specific habitat. Russell (1967) suggested that Tylosaurus in the Niobrara Formation frequented open waters far offshore, potentially up to 400 km from the nearest coastline in the WIS. This is supported by evidence in bone structure indicative of adaptation towards deeper waters. This includes osteoporosis, which naturally occurs in modern deep-diving or fast-swimming tetrapods due to bone growth in low-gravity environments both lifestyles create, and high fat deposition within bony microstructures, which together with osteoporosis would have allowed Tylosaurus to achieve neutral buoyancy in deep water. However, later work found that Tylosaurus is found throughout both shallow and open water deposits of the Niobrara Formation. On the other hand, Kiernan (2002) found that Tylosaurus comprised 57.7% of all mosasaurs found in Santonian to early Campanian deposits in Alabama that correspond to shallow continental shelves, and only 8.2% of mosasaurs in offshore deposits of similar ages. This implied that, at least in this region, Tylosaurus preferred nearshore waters.

Biogeochemical analyses of fossils instead support Tylosaurus as a spatial generalist that inhabited a wide range of marine habitats with little specific preference. Analysis of rare earth element (REE) ratios, which preserve information about the habitat an animal was buried in, from T. proriger and T. sp. bones from Alabama and South Dakota by Harrell and Pérez-Huerta (2014) found a widely distributed range of ratios indicating habitation across waters with depths from 50 m to below 150 m. This range was wider than all other mosasaurs examined, including Clidastes, Mosasaurus, Platecarpus, and Plioplatecarpus. Carbon-13 isotope analysis (δ13C) of tooth enamel by Robbins (2010) and Polcyn et al. (2025) also support a generalized habitat distribution. Unlike REEs, δ13C is correlated with the habitat the animal fed or foraged in, with lower values associated with more offshore foraging habitats. Both studies together found a δ13C range of -8.2‰ to -13.3‰ for Campanian T. proriger-like teeth from Texas (T. sp. aff. T. proriger) and -7.6‰ to 12.6‰ (all but one within -7.6‰ to -10.6‰) for T. ivoensis from Sweden. Isotope analyses of modern marine mammal teeth previously found that δ13C ratios between roughly -8‰ to -12.5‰ correspond to nearshore marine habitats and 11.5‰ to 15‰ to offshore marine habitats.{{efn|The study found an overlap from 11.5‰ to 12.5‰. Based on this, Polcyn et al. (2025) inferred that the T. proriger affiliates foraged in both nearshore and offshore marine habitats, while T. ivoensis foraged primarily nearshore with occasional offshore excursions. Both studies also examined a yet-undescribed Turonian specimen belonging to a 3-4 m long individual and found a δ13C ratio of -7.0‰. This corresponded to shallow waters analogous to modern kelp beds. Polcyn et al. (2025) suggested that early representatives of Tylosaurus were restricted to shallower habitats but expanded to more offshore zones over time as it evolved into larger sizes.

Notes

References

Bibliography

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