Tylosaurinae
Encyclopedia
Tylosaurinae is a subfamily of mosasaur
Mosasaur
Mosasaurs are large extinct marine lizards. The first fossil remains were discovered in a limestone quarry at Maastricht on the Meuse in 1764...

s, a diverse group of Late Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 marine lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s.

Russell (1967, pp. 170 ) defined the Tylosaurinae as follows: "Large rostrum
Rostrum (anatomy)
The term rostrum is used for a number of unrelated structures in different groups of animals:*In crustaceans, the rostrum is the forward extension of the carapace in front of the eyes....

 present anterior to premaxilla
Premaxilla
The incisive bone is the portion of the maxilla adjacent to the incisors. It is a pair of small cranial bones at the very tip of the jaws of many animals, usually bearing teeth, but not always. They are connected to the maxilla and the nasals....

ry teeth. Twelve or more teeth in dentary and maxilla
Maxilla
The maxilla is a fusion of two bones along the palatal fissure that form the upper jaw. This is similar to the mandible , which is also a fusion of two halves at the mental symphysis. Sometimes The maxilla (plural: maxillae) is a fusion of two bones along the palatal fissure that form the upper...

. Cranial nerves X, XI, and XII leave lateral wall of opisthotic through a single foramen. No canal in basioccipital or basispehnoid for basilar artery. Suprastapedial process of quadrate
Quadrate bone
The quadrate bone is part of a skull in most tetrapods, including amphibians, sauropsids , and early synapsids. In these animals it connects to the quadratojugal and squamosal in the skull, and forms part of the jaw joint .- Evolutionary variation :In snakes, the quadrate bone has become elongated...

 moderately large, distally pointed. Dorsal
Dorsum (biology)
In anatomy, the dorsum is the upper side of animals that typically run, fly, or swim in a horizontal position, and the back side of animals that walk upright. In vertebrates the dorsum contains the backbone. The term dorsal refers to anatomical structures that are either situated toward or grow...

 edge of surangular rounded and longitudinally horizontal...Twenty nine presacral vertebrae present. Length of presacral series less than that of postsacral series in Tylosaurus
Tylosaurus
Tylosaurus was a mosasaur, a large, predatory marine lizard closely related to modern monitor lizards and to snakes.-Paleobiology:...

, neural spines of posterior caudal vertebrae at most only slightly elongated, do not form an appreciable fin. Haemal arch
Haemal arch
A haemal arch is a bony arch on the underside of a tail vertebra of a vertebrate.The hole so formed is the haemal canal.It sometimes has a haemal spine on.The blood vessels to and from the tail run through the arch....

es unfused to caudal centra. Appendicular elements lack smoothly finished articular surfaces."
Genera referrable to the Tylosaurinae (informally and collectively known as "tylosaurines" or "tylosaurs") have been found on all continents except Australia and South America. The etymology
Etymology
Etymology is the study of the history of words, their origins, and how their form and meaning have changed over time.For languages with a long written history, etymologists make use of texts in these languages and texts about the languages to gather knowledge about how words were used during...

 for the subfamily is derived from the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

, Tylosaurus. In general, tylosaurs were large-bodied marine lizards armed with sturdy, conical teeth and an elongated premaxilla and extensions of the dentaries that do not bear teeth to the very end such as is found in other genera of mosasaurs. Cope's original concept of a "battering ram" snout is not supported by fossil evidence. Stomach contents from a tylosaur recovered in South Dakota
South Dakota
South Dakota is a state located in the Midwestern region of the United States. It is named after the Lakota and Dakota Sioux American Indian tribes. Once a part of Dakota Territory, South Dakota became a state on November 2, 1889. The state has an area of and an estimated population of just over...

 included remains of another mosasaur, a bony fish, the large, flightless seabird
Seabird
Seabirds are birds that have adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same environmental problems and feeding niches have resulted in similar adaptations...

 Hesperornis
Hesperornis
Hesperornis is a genus of flightless aquatic birds that spanned the first half of the Campanian age of the Late Cretaceous period . One of the lesser-known discoveries of the paleontologist O. C. Marsh in the late 19th century Bone Wars, it was an important early find in the history of avian...

, and possibly a shark
Shark
Sharks are a type of fish with a full cartilaginous skeleton and a highly streamlined body. The earliest known sharks date from more than 420 million years ago....

, indicating that tylosaurs were generalists. Another specimen collected by Charles Sternberg included the bones of a small plesiosaur (see also ).

Lingham-Soliar suggested that tylosaurines were not among the fastest swimming nor the strongest mosasaurids. However, they are lightly built, having greatly reduced the weight of their bodies and possessing relatively small pectoral and pelvic girdles, fore- and hindlimbs. Their bone is highly cancellous and may have been impregated with fat
Fat
Fats consist of a wide group of compounds that are generally soluble in organic solvents and generally insoluble in water. Chemically, fats are triglycerides, triesters of glycerol and any of several fatty acids. Fats may be either solid or liquid at room temperature, depending on their structure...

 cells during life, adding buoyancy. These traits suggest that tylosaurs may have been ambush predators. Tylosaurs were among the largest mosasaurs, with some species of Tylosaurus and Hainosaurus reaching lengths of 9-15+ meters, making them among the largest of all marine reptiles. A small species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 of Tylosaurus reported by Russell , T. "zangerli" has since proven to be a juvenile individual of T. proriger . Everhart erected a third species (T. kansasensis) from the Smoky Hill Chalk
Smoky Hill Chalk
The Smoky Hill Chalk Member of the Niobrara Chalk formation is a Cretaceous conservation Lagerstätte, or fossil rich geological formation, known primarily for its exceptionally well-preserved marine reptiles. The Smoky Hill Chalk Member is the uppermost of the two structural units of the Niobrara...

.

Polcyn and Bell (2005, p. 322 ) have erected a more inclusive clade, the parafamily Russellosaurina, which includes the "subfamilies Tylosaurinae and Plioplatecarpinae
Plioplatecarpinae
Plioplatecarpinae is a subfamily of mosasaurs, a diverse group of late Cretaceous marine squamates.Russell defined the Plioplatecarpinae as follows: Small rostrum present or absent anterior to premaxillary teeth. Cranial nerves X, XI, XII leave lateral wall of opisthotic through single foramen...

 and their sister-clade containing the genera Tethysaurus, Russellosaurus
Russellosaurus
Russellosaurus is a basal mosasauroid from the Upper Cretaceous of North America. The genus was described from a skull discovered in an exposure of the Arcadia Park Shale at Cedar Hill, Dallas County in south-central Texas...

, and Yaguarasaurus."

Tylosaurs first appear in the fossil record in the early Coniacian  and persist well into the Maastrichtian
Maastrichtian
The Maastrichtian is, in the ICS' geologic timescale, the latest age or upper stage of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned from 70.6 ± 0.6 Ma to 65.5 ± 0.3 Ma...

, a period of approximately twenty million years.

Species and Taxonomy

  • Tylosaurinae
    • Tylosaurus
      Tylosaurus
      Tylosaurus was a mosasaur, a large, predatory marine lizard closely related to modern monitor lizards and to snakes.-Paleobiology:...

      (?paraphyletic)
      • T. proriger
      • T. nepaeolicus
      • T. kansasensis
      • T. pembinensis
      • T. saskatchewanensis
      • T. ivoensis (Possibly a species of Taniwhasaurus )
    • Hainosaurus
      Hainosaurus
      Hainosaurus is an extinct genus of marine lizard belonging to the mosasaur family. It was one of the largest mosasaurs, though its size has been revised more than once. At first it was estimated to be , and the largest mosasaurid. During the 1990s, its size was revised to long; more recently,...

      • H. bernardi
      • H. gaudryi
    • Taniwhasaurus
      Taniwhasaurus
      Taniwhasaurus is an extinct genus of mosasaur, a carnivorous, marine reptile which inhabited New Zealand, Japan and Antarctica. The genus was a close relative of the genera Tylosaurus and Hainosaurus.-Species:-T...

      • T. oweni (=Tylosaurus haumuriensis)
      • T. mikasaensis
      • T. antarcticus

Further reading

  • Bell, G. L. Jr., 1997. A phylogenetic revision of North American and Adriatic Mosasauroidea. pp. 293–332 In Callaway J. M. and E. L Nicholls, (eds.), Ancient Marine Reptiles, Academic Press, 501 pp.
  • Lindgren, J. et. Siverson, M. 2002.Tylosaurus ivoensis: a giant mosasaur from the early Campanian of Sweden. Royal Society of Edinburgh Transactions: Earth Sciences Vol. 93(1):73-93.
  • Russell, D. A. 1970. The vertebrate fauna of the Selma Formation of Alabama, Part VII, The mosasaurs, Fieldiana, Geology Memoirs 3(7):369-380.
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