Research history of Tylosaurus

987:. Based on information about the circumstances and locality of the discovery given the museum, it was recognized that the rest of the skeleton may have remained intact. In February, excavations were made under the authorization of an industrialist named Leopold Bernard, who managed the quarry the fossil resided in. The rest of the skeleton was recovered after a month of excavating between 500–600 cubic meters (650–780 cu yd) of phosphate, although a section of the tail was found to have been destroyed by erosion from an overlying deposit. The skeleton went to the museum, which was subsequently studied by Dollo, who recognized that it belonged to a new type of mosasaur. By instruction of the museum, he named it 415: 668: 1560: 774: 1081: 762:. The restoration carried a number of erroneous features, such as a baggy throat, bloated belly, and inaccurate paddles and dorsal fin. But a seminal feature was the addition of a dorsal crest (known as a fringe) lining the mosasaur's back. This was based on another 1898 description of a mosasaur fossil by Williston that erroneously identified tracheal rings as remains of a dorsal fringe, which he corrected in 1902. Knight updated his 652: 887: 556: 457:. He depicted the head as akin to a giant flat cone supporting eyes directed partially upwards that is capable of swallowing entire prey whole like a snake, with a loose and baggy pelican-like throat to allow entry of such prey. Like a whale, one pair of paddle-like flippers was described to be present in the animal and functional hind limbs were thought to be absent. The tail was imagined as long and flat, propelling 704: 201: 1289: 1173: 587:, but they nevertheless provided enough material to represent nearly an entire skeleton. These three specimens include one consisting of a posterior fragment of the skull and an "absolute complete series of vertebrae, connected from head to tip of tail" collected in an unspecified date by a fossil collector named Elias Putnam West (KUVP 1048); one containing a complete skull and 20: 69: 684: 443:

of the tail was minimal between vertebrae; thus, in order for the vertebrae in the column to approach the size of the tail vertebrae based on the observed rate of size decrease per vertebrae, plentiful amounts of vertebrae would have been needed. By measuring the proportions of the change in dimensions between the vertebrae, Cope estimated that

754:, a condition in a calcium-rich environment like the Kansas chalk that causes soft tissue to harden, allowing it to better fossilize. The completeness of AMNH FR 221 allowed Osborn to accurately measure the proportions of each body part; these ratios remain in use by modern paleontologists when extrapolating 1499:. Despite such recognition, however, no paleontologist investigated this further and the fossil remained undescribed during the remainder of the 20th century. In a 2004 meeting, Everhart suggested that the new species should finally be given a scientific name and formal description, to which paleontologist 1100:

but stated that a better description of MT 2 was needed to be certain. In 2010, paleontologists Timon Bullard and Michael Caldwell redescribed such specimen. They argued that the pygal vertebrae reported by Nicholls (1988) actually belonged to another individual mosasaur, reducing the vertebral count

817:

in 1917, which contained digested remains of a plesiosaur within its ribcage. The specimen was sold the following year to the Smithsonian, and the mosasaur has been permanently displayed as USNM 8898. The plesiosaur remains, despite their significance, were stashed as a separate specimen and promptly

442:

of the specimen were less than a fifth in diameter than one more proximal, or closer to, the base of the tail, and there was little dimensional variation between the four back tail vertebrae. To Cope, this indicated that the enormous decrease in the size of the tail vertebrae from the base to the tip

452:

it possessed, which would increase the vertebral count in the tail to an unknown but monstrous amount. The resulting animal became that akin to an enormous sea serpent-like reptile reaching lengths rivaling that of the largest whales. In an 1872 publication, Cope described in detail his imagination

192:

fossils were actually recovered from somewhere in Western Kansas and were mislabeled as being collected by Lyon in New Mexico (who was shipping a different fossil from Fort McRae around the same time) during transit. The type specimen of the taxon is split between two museums; some of the vertebrae

1237:

supporting the suture between the premaxilla and maxilla in MNHN 1896–15, a trait he claimed that was unique to the latter species, and estimations that the rostrum may have been more elongated than preserved. This was elaborated on a publication of the same year by Nathalie Bardet of the Natural

603:

specimens informed minor missing body parts such as the hind limbs. The resulting animal was correctly depicted as a four-flipper reptile with a short neck and a tail making up around half the body length. Rather unrealistically, the vertebral column was drawn straight and lacking in structural

608:

would have possessed in life. Commentating on this issue, Everhart noted that Williston was already aware of the curvature of a mosasaur's spinal column and suggested that he had a straight spine drawn to save space on the plate the reconstruction was published on. Nevertheless, apart from the

447:

had at minimum over sixty vertebrae between the described proximal tail vertebra and a hypothetical vertebra measuring half the diameter of the four back tail vertebrae. Cope speculated that there were even more tail vertebrae ahead of the proximal vertebra, justified by the small

1578:. The skeleton was found articulated and consists of a "moderately complete" skull, a largely complete vertebral column, limb bones, and bones of a smaller mosasaur as stomach content; it was curated into the museum as RSM P2588.1 and nicknamed "Omācīw" (meaning "hunter" in 1598:

as a valid species despite technically never having been formally described. 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 a distinct species.

1335:- in the species name referring to Ivö Klack. Russell (1967) amended this identification by elevating the taxon to a distinct species. He also identified a mosasaur fossil from the Niobrara Formation in Kansas consisting of a partial skull, some vertebrae, and a 372:

was not preoccupied, arguing that while it was indeed used as a genus of lizard, it was subsequently synonymized with other genera before 1872 and was thus a vacant taxon. Cope's persistence can be seen in his 1874 description of the fourth described species of

618:

grew no larger than 35 feet (11 m) in length and lived as a highly mobile predator "at the expense of strength" that was best adapted for preying on small fish and occasionally an "animal of their own kind". In a review of the taxonomy of the

1109:

and additionally argued that the femur being longer than the humerus was an insufficient characteristic to diagnose the species at the generic level. Based on these and other characteristics, the study found that the species cannot be referred to

481:

specimen with fossils relatively smaller than that of the taxon's type specimen had a skull measuring about 5 feet (1.5 m) in length; the whole animal was estimated to measure 75 feet (23 m) in total length. In 1874, Cope estimated that

183:

found that the chemical composition of the fossil did not match any known geological deposit in New Mexico and is instead identical to that of the Niobrara Formation; the study speculated from examinations of multiple letters between Lyon and the

1515:

was not adequate enough and should be readdressed in a followup paper. In 2013, Caldwell and Jiménez-Huidobro presented an abstract at the annual Society of Vertebrate Paleontology meeting casting doubt on the distinction of the species from

613:

is still considered accurate by modern standards, and adaptations of the skeleton remain in use by paleontologists and museum workers. In his own interpretation of the available fossil evidence, Williston proposed that

1532:

claimed that the study was "poorly researched and written". However, he did not provide any comments on the topic, claiming that such would merely be considered "sour grapes". Nevertheless, Everhart maintained that

1184:

In 1896, paleontologist Armand Thevenin published a formal description of a poorly-preserved skull consisting of the majority of the maxilla, premaxilla, and dentary bones that was found in a phosphate deposit in

832:

skeleton he found to the Smithsonian 1926, but the museum was satisfied with his father's earlier specimen and declined the offer. This skeleton was ultimately transferred to the Sternberg Museum as FHSM VP-3.

734:, a specimen measuring 8.83 meters (29.0 ft) long that resides in the American Museum of Natural History as AMNH FR 221. Discovered in Kansas by a local named W. O. Bourne and initially identified as 1315:. Such teeth and various other teeth pertaining to the same taxon were recovered and described from this area numerous times in the past since 1836, but they have always been misidentified as either an 738:, the fossil has been considered remarkable in that it contained not only every bone apart from a few in the tail's tip, but also had exceptional preservation of the animal's soft tissue, including the 1130:

to be a distinct genus but acknowledged that further research may provide additional characteristics. This further research was done in 2016 by Paulina Jimenez-Huidobro and Caldwell, which reexamined

1586:

thesis. At the time, the Omācīw fossil was still in preparation and Bullard was only able to examine the right side of the skull. However, he identified that the fossil represented a new species of

1537:

remains a distinct species "until proven otherwise". A 2018 study by Robert F. Stewart and Jordan Mallon also rejected the synonymy based on inconsistencies when comparing the ontogenetic trends of

1363:

which examined the fossils referred to the species as well as new material consisting of pterygoid teeth, partial jawbones, and some vertebrae. It was found that the Niobrara fossil identified as

1154:

should remain a distinct genus, justified by differences in tooth morphology unexplored by Jimenez-Huidobro & Caldwell (2016) such as the presence of an advanced cutting-based dentition in

1624:

fossil in the collections of the Royal Tyrrell Museum consisting of a partial snout and a fragment of the right mandible. The specimen was found around 55 kilometers (34 mi) northeast of

575:'s 1898 publication of a comprehensive study of the skeletal anatomy of Kansas mosasaurs, which among many things corrected many of the erroneous beliefs held by earlier paleontologists about 521:). The appearance of this mosasaur was influenced by Cope, who was an acquaintance of Webb, as evidenced by its excessively elongated tail and serpentine appearance. Additionally, the 3564:

Jiménez-Huidobro, P.; Caldwell, M.W.; Paparella, I.; Bullard, T.S. (2018). "A new species of tylosaurine mosasaur from the upper Campanian Bearpaw Formation of Saskatchewan, Canada".

1652:

species), a trait likely retained from juveniles. The uniqueness of the specimen led to the conclusion that it was a new species, which Garvey subsequently proposed that it be named

1023:

was in recognition of Bernard, who made the excavation of the skeleton possible. The skeleton is now on display at the Royal Belgian Institute under the catalog number IRSNB R23.

1469:

fossil consisting of fragmentary skull elements was recovered during an expedition to the lost holotype's locality, although the specimen was not figured or formally described.

583:, which was drawn by paleoartist Sydney Prentice under Williston's instructions. This reconstruction was almost entirely informed by three specimens in the collections of the 874:

that are still used by scientists today, and Everhart remarked that the work "remains the single best reference regarding the skeletons of the various mosasaur genera" like

1549:. Another ontogenetic study published in 2020 by Amelia R. Zietlow came in support of the synonymy based on results from ontograms constructed from a cladistical analysis. 2517: 949:. These additional species were identified as different genera during their times of discovery and it was not until the early 21st century when they were reclassified as 3643: 1041:

based on a partial skeleton consisting of a partial skull, lower jaw, a near-complete vertebral column, and some limb bones recovered from the Pembina Member of the

107:. The type specimen was described by Cope in 1869 based on a fragmentary skull measuring nearly 5 feet (1.5 m) in length and thirteen vertebrae lent to him by 3297: 2554: 3017:

Hornung, J.J.; Reich, M.; Frerichs, U. (2018). "A mosasaur fauna (Squamata: Mosasauridae) from the Campanian (Upper Cretaceous) of Hannover, northern Germany".

381:. This type specimen of this species consists of a right lower jaw, a partial upper jaw, a quadrate bone, and a dorsal vertebra and was discovered by geologist 945:. However, throughout the 19th and 20th centuries, numerous species of mosasaurs were described from around the world that are now recognized within the genus 1491:

In the 1990s, a well-preserved mosasaur skull of uncertain identity, which was discovered in an outcrop of the Niobrara Formation in the northwestern part of

1507:

specimens also from Kansas were identified as conspecific with FHSM VP-2295, and in reference to their shared localities the new species was given the name

1299:

In 1963, paleontologist Per Ove Persson identified a new taxon of mosasaur based on isolated teeth from a deposit located in an area called Ivö Klack near

584: 3754: 3713: 3665: 1222: 1503:

responded with "Well, get it done." In 2005, Everhart published a formal description of the new species represented by FHSM VP-2295. Thirteen different

1246:. This was agreed on by Lingham-Soliar in a 1992 publication. In 2005, Lindgren pointed out that the characteristics used to justify the placement in 3062:"Mosasaurid and plesiosaurian remains from marginal facies of the lower Campanian (Upper Cretaceous) Bottrop and Vaals formations of western Germany" 1205:. Through correspondence with Dollo and a reexamination of the skull, Thevenin noted that the rostrum was more elongated than those found in typical 3515: 1461:

without a catalog number, and it was later reported in 2006 that the specimen may have been lost during a fire. In a 2012 multi-author study led by

2102:

Contributions to the extinct vertebrate fauna of the western interior territories: Report of the United States Geological Survey of the Territories

150:(I bear). In 1870, Cope published a more thorough description of MCZ 4374. Without explanation, he moved the species into another European genus 3222:

Louis L. Jacobs; Octávio Mateus; Michael J. Polcyn; Anne S. Schulp; Miguel Telles Antunes; Maria Luísa Morais; Tatiana da Silva Tavares (2006).

473:

was the largest species of the genus was also described as the longest of all reptiles. Its maximum length was estimated to equal that of the

3738: 3697: 3269: 2323: 824: 95:. The early history of the genus as a taxon was subject to complications spurred by the infamous rivalry between American paleontologists 766:

restorations with the dorsal fringes removed to reflect this correction. However, his original depiction of a fringe was picked up as a

296:. This name means "knob lizard" in another reference to the elongated rostrum characteristic of the genus. It is derived from the Latin 194: 185: 2186:

Davidson, J.P. (2015). "Misunderstood Marine Reptiles: Late Nineteenth-Century Artistic Reconstructions of Prehistoric Marine Life".

1495:, Kansas in 1968 and cataloged in the Sternberg Museum as FHSM VP-2295, was recognized by multiple authors as being a new species of 3223: 547:

skeleton measuring 80 feet (24 m) in total length with a long and slender neck that is 20 feet (6.1 m) long is uncovered.

264:, meaning "lizard"). Marsh also described a third species based on a partial skeleton he collected near the southern portion of the 142:

means "prow-bearing", which is in reference to the specimen's unique prow-like elongated rostrum and is derived from the Latin word

2421: 1648:

in its teeth, which were slender and designed for piercing prey (contrary to the more robust cutting-specialized teeth of typical

3777: 1367:

by Russell (1967) is actually of a different species. Examinations of the Swedish teeth found possible similarities with that of

2527: 1574:

excavated a partial skeleton first discovered a year prior by a local farmer from a hill located northwest of Herbert Ferry,

1355:. The next year in 1999, Lindgren amended this in another abstract and instead suggested that the species should be moved to 1065:

based on a higher count of pygal vertebrae, longer external nares, and femur longer than the humerus. The species was named

389:. The fossils are stored in the American Museum of Natural History under the catalog number AMNH 1565. The specific epithet 983:. The fossil was initially discovered as eight vertebrae by a worker in the January of the same year, who sold them to the 364:. An exception to this adoption was Cope, who refused to accept Marsh's new genus and continued to refer to its species as 1632:; during the Cretaceous this locality was located at around 62° N, making the fossil the northernmost known occurrence of 1134:. They found that the unique characteristics of the type species are indeed insufficient to warrant a distinction between 854: 112: 2646: 1783: 1594:

in reference to its type locality. Bullard's thesis was never published but subsequently published studies recognized

1061:

was previously only known in Europe, Nicholls (1988) argued that the new species differed from the more locally known

115:. The fossil, which remains in the same museum under the catalog number MCZ 4374, was recovered from a deposit of the 2577: 3296:

Mateus, O.; Polcyn, M.J.; Jacobs, L.L.; Araújo, R.; Schulp, A.S.; Marinheiro, J.; Pereira, B.; Vineyard, D. (2012).

818:

forgotten until their redescription by Everhart in 2004. This rediscovery formed the basis for the plot of the 2007

292:

was preoccupied as a genus of lizard named in 1843. As a result, he suggested a move to a newly erected genus named

3689: 1989: 1457:. However, Lingham-Soliar did not figure the holotype skull, which he identified as residing in the collections of 984: 536: 319: 1221:; the specific epithet recognizing Gaudry's previous work on the fossil skull. The skull is now on display at the 228:

are taxonomically distinct from the European genus and must be assigned a new one. For this, he erected the genus

3673: 3537: 1571: 1403:, which was of similar latitude with Ivö Klack during the Cretaceous, and reasserted dental similarities between 591:

collected in 1898 by colleague Handel T. Martin (KUVP 28705); and one collected in 1877 by university chancellor

3224:"The occurrence and geological setting of Cretaceous dinosaurs, mosasaurs, plesiosaurs, and turtles from Angola" 2966:

Jiménez-Huidobro, P.; Caldwell, M.W. (2016). "Reassessment and reassignment of the early Maastrichtian mosasaur

2898: 2805: 2340: 2277:"XI.—An Account of the Great Finner Whale (Balænoptera Sibbaldii) stranded at Longniddry Part I. The Soft Parts" 2085: 3730: 1617: 1528:, thus making the former a junior synonym of the latter. This was criticized by Everhart, who in his 2017 book 572: 513:

of marine reptiles and pterosaurs from the Niobrara Formation. At the center of the illustration rises a giant

171:. The fossils came to Cope under the label as having been collected by an army doctor named William B. Lyon at 3197: 3184:(Mosasauridae, Tylosaurinae) from the upper Santonian-lower Campanian (Upper Cretaceous) of Hokkaido, Japan". 1201:. However, Dollo thought that the characteristics of skull supported an alternative placement as a species of 3600: 1949:"On the structure of the skull and limbs in mosasaurid reptiles, with descriptions of new genera and species" 1873: 1005:

means "lizard from the Haine"; Dollo wrote that this was erected specifically to complement the etymology of

1016: 866:. Many of the complete skeletons formerly assigned to the former taxon were reassigned to the type species. 382: 180: 136: 120: 414: 1458: 1217:. He suggested that this represented an intermediate trait for a new species, which he subsequently named 727: 498: 423: 124: 3523: 284:. He disagreed with Marsh's arguments but proposed that in case Marsh was indeed correct, the genus name 3772: 1422: 1126:

at the time was a higher count of pygal vertebrae. It was doubted as to if this would be sufficient for

1118:. With this rediagnosis, the study also noted that the only confirmed characteristic that distinguished 339: 100: 3340:, a new species of tylosaurine (Squamata, Mosasauridae) from the Niobrara Chalk of western Kansas, USA" 1360: 797:

were uncovered from the Niobrara Formation between the 1900s to 1920s by Charles Sternberg and his son

3726: 3396: 3351: 3073: 3026: 2983: 2929: 2852: 2725: 2616: 2469: 1511:. However, during a 2007 meeting, Caldwell approached Everhart and suggested that his description of 1492: 814: 343: 2100: 2806:"The Tylosaurine Mosasaurs (Reptilia, Mosasauridae) from the Upper Cretaceous of Europe and Africa" 819: 798: 667: 269: 96: 3723:

Oceans of Kansas, Second Edition: A Natural History of the Western Interior Sea (Life of the Past)

2714:"A New Hypothesis of the Phylogenetic Relationships of the Tylosaurinae (Squamata: Mosasauroidea)" 3748: 3707: 3659: 3633: 3581: 3275: 3201: 3159: 3089: 3042: 2999: 2945: 2912:

Bullard, T.S.; Caldwell, M.W. (2010). "Redescription and rediagnosis of the tylosaurine mosasaur

2743: 2296: 2245: 2203: 1926: 1304: 1194: 1038: 972: 848:. In a taxonomic examination, Russell found little justification for recognizing the validity of 588: 209: 116: 3678: 3249: 2399: 1373:, but due to the poor representation of the genus and the sheer distance between the two taxa ( 3734: 3693: 3685: 3647: 3492: 3265: 3248:

Mateus, O.; Callapez, P.M.; Polcyn, M.J.; Schulp, A.S.; Gonçalves, A.O.; Jacobs, L.L. (2019).

3120: 2777: 2495: 2319: 2029: 1764: 1637: 1583: 1462: 759: 691: 3298:"Cretaceous amniotes from Angola: dinosaurs, pterosaurs, mosasaurs, plesiosaurs, and turtles" 3606: 3573: 3482: 3472: 3439: 3404: 3359: 3257: 3193: 3151: 3081: 3034: 2991: 2937: 2894: 2860: 2733: 2628: 2624: 2485: 2477: 2288: 2237: 2195: 2168: 2081: 1823: 1430: 1225:

as MNHN 1896–15. In 1990, paleontologist Theagarten Lingham-Soliar proposed reassignment of

971:

from a near-complete but poorly preserved skeleton excavated from a phosphate quarry in the

810: 490:, which would translate to around 5.08 meters (16.7 ft) when using his 1872 estimates. 265: 3108: 2765: 2156: 1948: 1640:. Moreover, the morphology of TMP 2014.011.0001 was recognized as very distinct from other 1288: 1080: 773: 2354:(Report). Ottawa: Committee on the Status of Endangered Wildlife in Canada. pp. 1–32. 2120: 1740: 1500: 1054: 770:, and many subsequent illustrations of the mosasaur continued to portray this inaccuracy. 596: 592: 462: 57: 3250:"The Fossil Record of Biodiversity in Angola Through Time: A Paleontological Perspective" 1193:

during an earlier unspecified date. Thevenin wrote that the skull was first described by

631:

and went as far as to suggest the latter species was merely a juvenile representative of

3400: 3355: 3077: 3030: 2987: 2933: 2856: 2729: 2620: 2473: 2161:

Annual Report of the United States Geological and Geographical Survey of the Territories

1913:

Parris, D.C.; Grandstaff, B.S.; Denton, R.K. Jr.; Lucas, S.G. (1997). "Type Locality of

1559: 1445:. In 1992, Lingham-Soliar argued that the features of the skull disagreed with those of 651: 595:

that contains a near-complete set of front paddles along with some ribs, vertebrae, and

559:

Williston (1898) presented the earliest anatomically correct skeletal reconstruction of

3487: 3460: 1625: 886: 555: 2810:

Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre

1254:. He also noted that the dentition of MNHN 1896-15 morphologically overlaps more with 1105:. They also noted that the length of the external nares was within the known range of 3766: 3205: 3163: 3093: 3061: 3046: 2225: 2065: 1579: 1375: 1369: 1186: 751: 386: 108: 3585: 3335: 3279: 3003: 2949: 2747: 2300: 2249: 2207: 3629: 3379:

Jiménez-Huidobro, P.; Simões, T.R.; Caldwell, M.W. (2016). "Re-characterization of

1582:). In 2006, Timon Bullard of the University of Alberta studied the specimen in his 1575: 1520:

and in 2016 the two and paleontologist Tiago Simões published a study arguing that

1407:

and the genus. This opened the possibility that the species could be reassigned to

1042: 870:

was affirmed as a distinct species. Russell (1967) provided extensive diagnoses of

841: 703: 579:. A central component of the publication was a rigorous skeletal reconstruction of 527: 315: 200: 3577: 3408: 3085: 3038: 2995: 2457: 1811: 1069:, the specific epithet referring to the Pembina Member the skeleton was found in. 393:

is in reference to the Nepaholla, the Native American name for the Solomon River.

385:

in an unspecified deposit of Cretaceous grey shale about a half-mile south of the

1380: 1316: 1300: 964: 758:

fossils. Osborn (1899) included a life impression of AMNH FR 221 by paleoartist

525:

was given a rather long neck. Although not as long as those of plesiosaurs like

233: 3252:. In Huntley, Brian J; Russo, Vladimir; Lages, Fernanda; Ferrand, Nuno (eds.). 3180:

Caldwell, M.W.; Konishi, T.; Obata, I.; Muramoto, K. (2008). "A new species of

2481: 1172: 3364: 3261: 3155: 3144:

Earth and Environmental Science Transactions of the Royal Society of Edinburgh

2941: 2292: 2281:

Earth and Environmental Science Transactions of the Royal Society of Edinburgh

1387:

belongs to it. Instead, the study concluded that the species should belong to

1007: 782: 474: 176: 172: 131:, Kansas. Cope's first publication of the fossil was very brief and was named 83:

was the third new genus of mosasaur to be described from North America behind

41: 25: 19: 3124: 2781: 2738: 2713: 2604: 1768: 434:

can be traced back to Cope's 1870 description of MCZ 4374. He suggested that

3639: 2670:

The mosasaurs of George F. Sternberg, paleontologist and fossil photographer

2345:

in Canada, in COSEWIC assessment and update status report on the Blue Whale

2172: 1828: 1442: 477:, which reaches lengths of up to nearly 30 meters (98 ft) or more. One 128: 104: 68: 3496: 2654: 2499: 1791: 541:

The Young Fossil Hunters: A True Story of Western Exploration and Adventure

3651: 3302:

V Jornadas Internacionales sobre Paleontología de Dinosaurios y su Entorno

2589: 1676:

to that of a "great finner whale", which was the common name of the taxon

635:. He also remarked that there exist little anatomical differences between 3610: 2241: 2226:"Fictionalized facts; "The Young Fossil Hunters" by Charles H. Sternberg" 2199: 1438: 1336: 1234: 1142:

and declared the former as a junior synonym of the latter, thus renaming

1046: 747: 510: 449: 439: 50: 3477: 2070:(Squamata; Mosasauridae), from the Niobrara Formation of Western Kansas" 1997: 1242:

but as a separate species, thus changing the skull's scientific name to

3444: 3423: 2678:, Fort Hays State University Special Issue Number 3 (M.J. Everhart, ed) 1930: 1881: 1812:"Synopsis of the extinct Batrachia, Reptilia and Aves of North America" 1645: 1629: 1411:, but the authors considered this to be a topic for a different study. 980: 802: 743: 683: 53: 3109:"Mosasauriens de la Craie Grise de Vaux Eclusier prés Peronne (Somme)" 535:

in art. An example of this can be seen in the famous fossil collector

306:. Despite coining the new genus, Marsh never formally transferred the 2817: 2490: 1434: 1312: 1308: 1238:

History Museum, who concluded that MNHN 1896-15 belongs to the genus

1190: 1050: 976: 750:, and cartilage in the throat, chest, and shoulders. This was due to 739: 213: 197:

as AMNH 1580 while the rest were sent to the Smithsonian as USNM 41.

152: 92: 2864: 2276: 2157:"On the geology and paleontology of the Cretaceous strata of Kansas" 1158:

as opposed to a more general smashing-based dentition seen in other

2066:"New Data on Cranial Measurements and Body Length of the Mosasaur, 643:; the only real distinction between the two species being in size. 609:

straight spine, Williston's century-old skeletal reconstruction of

1558: 1400: 1323:. Persson proposed that the teeth belonged to a new subspecies of 1287: 1171: 1079: 1012: 996: 885: 772: 623:

genus, Williston (1898) expressed doubt regarding the validity of

554: 413: 199: 67: 46: 18: 999:, a river located nearby the Ciply Basin, and thus combined with 1441:

age but this was later revised to about 88 mya during the later

767: 3602:

Anatomy and systematics of North American tylosaurine mosasaurs

438:

was an "excessively elongate reptile" because four of the back

1919:

Proceedings of the Academy of Natural Sciences of Philadelphia

1757:

Proceedings of the Academy of Natural Sciences of Philadelphia

1680:

at the time. This taxon is a synonym of the modern blue whale

167:

based on several vertebrae and limb fragments, which he named

1636:

and one of the few known occasions of a mosasaur existing in

1453:. He subsequently moved the species into the genus, becoming 1000: 301: 258: 251: 244: 237: 135:, the genus being a preexisting European mosasaur taxon. The 422:(center) with an unrealistically elongated neck and tail by 352:

subsequently became the near-universally accepted genus for

163:

In 1871, Cope identified a second species of North American

2899:

10.1666/0022-3360(2005)079[1157:tfrohr]2.0.co;2

2314:

Mann, J.; Conner, R.C.; Tyack, P.L.; Whitehead, H. (2000).

2086:

10.1660/0022-8443(2002)105[0033:NDOCMA]2.0.CO;2

3317: 3315: 3313: 3311: 2523:(Squamata, Mosasauridae) from Canada with unique dentition 1620:

was published. The thesis focused on TMP 2014.011.0001, a

3198:

10.1671/0272-4634(2008)28[339:ansotm]2.0.co;2

2839:

Nicholls, E.L. (1988). "The first record of the mosasaur

2458:"A complete mosasaur skeleton, osseous and cartilaginous" 2011: 2009: 2007: 1968: 1966: 1841: 1839: 1707: 1705: 1703: 1701: 1197:, who initially thought that it belonged to a species of 501:

for paleontologist William E. Webb's 1872 fictional book

3428:(Squamata: Mosasauridae) and its taxonomic implications" 3175: 3173: 3142:: a giant mosasaur from the early Campanian of Sweden". 2316:

Cetacean Societies: Field Studies of Dolphins and Whales

1893: 1891: 1383:

at the time), it was expressed that it is unlikely that

1343:. In an abstract presented in 1998, Lindgren doubted if 933:

By Russell (1967), the only recognized valid species of

720:

specimen AMNH FR 221 and developments in its restoration

2676:

Proceeding of the Second Mosasaur Meeting, Hays, Kansas

2422:"Kansas University Vertebrate Paleontology Collections" 1942: 1940: 1351:. He instead proposed that the species was actually of 531:, this detail would influence subsequent depictions of 2961: 2959: 2712:

Paulina Jiménez-Huidobro; Michael W. Caldwell (2019).

2426:

KU Biodiversity Institute & Natural History Museum

2219: 2217: 1856: 1854: 1359:. In 2002, he co-authored a study with paleontologist 1266:. With these characteristics, Lindgren concluded that 1092:

In 2005, paleontologist Johan Lindgren suspected that

828:. George Sternberg attempted to sell another complete 801:, which were sold to museums in the United States and 338:

was formally transferred to the same genus in 1894 by

288:

should be used. Marsh later discovered that the taxon

179:. However, a 1997 study led by David C. Parris of the 39:

documents the historical developments in the study of

1734: 1732: 497:

comes from an illustration by a popular artist named

3019:

Alcheringa: An Australasian Journal of Palaeontology

2799: 2797: 2795: 2793: 2791: 1250:, such as the double buttress, can also be found in 1395:. However, a 2008 study led by Caldwell discovered 469:measured at minimum 50 feet (15 m) in length. 3677: 3559: 3557: 3555: 1816:Transactions of the American Philosophical Society 3644:Bulletin of the Peabody Museum of Natural History 2462:Memoirs of the American Museum of Natural History 1209:skulls but not as long as those in the skulls of 844:performed a complete review and redescription of 3680:Sea Dragons: Predators of the Prehistoric Oceans 3635:Systematics and morphology of American mosasaurs 2393: 2391: 2389: 2387: 2385: 1805: 1803: 1801: 838:Systematics and Morphology of American Mosasaurs 2876: 2874: 2640: 2638: 2511: 2509: 2341:Update COSEWIC status report on the Blue Whale 2150: 2148: 2146: 2144: 2142: 2140: 2138: 2059: 2057: 2055: 2053: 2051: 1983: 1981: 1616:In 2020, the MS thesis of Samuel Garvey of the 1101:in MT 2 to one that was indistinguishable with 1053:. This skeleton resides in the Miami Museum in 2114: 2112: 551:Increased understanding and complete skeletons 3291: 3289: 3217: 3215: 3060:Sachs, S.; Hornung, J.J.; Scheer, U. (2018). 2451: 2449: 2447: 2230:Transactions of the Kansas Academy of Science 2188:Transactions of the Kansas Academy of Science 2074:Transactions of the Kansas Academy of Science 1908: 1906: 8: 2843:(Reptilia: Lacertilia) from North America". 2759: 2757: 2559:Lost its Fringe, and Other Squamate Stories" 2318:. University of Chicago Press. p. 343. 1011:, which was similarly named in reference to 3432:Vertebrate Anatomy Morphology Palaeontology 3113:Bulletin de la Société Géologique de France 2605:"An extinct sea-lizard from western Kansas" 2526:(MS). University of Alberta. Archived from 2121:"Über die Pythonomorphen der Kansas-Kreide" 1656:in reference to its northernly occurrence. 1033:A second species historically pertained to 985:Royal Belgian Institute of Natural Sciences 585:University of Kansas Natural History Museum 2881:Lindgren, J. (2005). "The first record of 694:, with dorsal fringe and single-lobed tail 3510: 3508: 3506: 3486: 3476: 3443: 3363: 2737: 2489: 1827: 1437:. The deposit was initially dated to the 1223:French National Museum of Natural History 789:Additional largely complete skeletons of 730:described the first complete skeleton of 3387:Everhart, 2005: Ontogeny or sympatry?". 3321: 2699: 2540: 2438: 2376: 2364: 2262: 1845: 1711: 1429:from a partial skull excavated from the 809:skeleton collected by C. Sternberg from 674:Skeletal reconstruction by Osborn (1899) 3461:"Craniofacial ontogeny in Tylosaurinae" 2885:(Reptilia: Mosasauridae) from Sweden". 2687: 2629:10.1038/scientificamerican04021921-273a 2224:Davidson, J.P.; Everhart, M.J. (2014). 2015: 1972: 1897: 1874:"Sebastes proriger, Redstripe rockfish" 1697: 1665: 543:, which contains a scene where a giant 72:The skull of MCZ 4374, the holotype of 3753:: CS1 maint: ref duplicates default ( 3746: 3712:: CS1 maint: ref duplicates default ( 3705: 3664:: CS1 maint: ref duplicates default ( 3657: 3540:. Regina Leader-Post. October 14, 2010 2404:University Geological Survey of Kansas 1150:. However, some scientists argue that 571:as scientists do today beginning with 397:was formally transferred to the genus 1860: 1723: 1465:, it was reported that an additional 825:Sea Monsters: A Prehistoric Adventure 507:The sea which once covered the plains 505:. This illustration, which is titled 280:was already a preoccupied synonym of 236:derived from the Ancient Greek words 232:, which means "nose lizard" and is a 113:Harvard Museum of Comparative Zoology 16:Studies of an extinct aquatic reptile 7: 3138:Lindgren, J.; Siverson, M. (2002). " 995:- in the generic name refers to the 567:Paleontologists began to understand 493:One of the earliest restorations of 3566:Journal of Systematic Palaeontology 3424:"Allometric growth in the skull of 2339:Sears, R.; Calambokidis, J (2002). 1644:species. For example, it exhibited 1449:and were instead characteristic of 1233:based on the discovery of a double 486:was around one-third the length of 3422:Stewart, R.F.; Mallon, J. (2018). 3344:Netherlands Journal of Geosciences 3186:Journal of Vertebrate Paleontology 2976:Journal of Vertebrate Paleontology 2922:Journal of Vertebrate Paleontology 2845:Canadian Journal of Earth Sciences 2105:. U.S. Government Printing Office. 710:2015 restoration with bilobed tail 405:19th and 20th century developments 195:American Museum of Natural History 14: 2770:Revue des Questions Scientifiques 314:; this was first done in 1873 by 276:. Cope responded by arguing that 3231:Paleontological Society of Korea 1917:Cope (Reptilia: Mosasauridae)". 702: 682: 666: 650: 599:(KUVP 1075). Various additional 465:locomotion. Cope estimated that 1590:and suggested that it be named 1270:is most certainly a species of 658:Photograph of complete skeleton 401:in Merriam's 1894 publication. 272:as YPM 1268, which Marsh named 1545:between the better-understood 1524:was merely a juvenile form of 1329:Mosasaurus hoffmannii ivoensis 781:skeleton which was found with 509:, represents one of the first 1: 3721:Everhart, Michael J. (2017). 3605:(MS). University of Alberta. 3578:10.1080/14772019.2018.1471744 3409:10.1016/j.cretres.2016.04.008 3086:10.1016/j.cretres.2017.05.026 3039:10.1080/03115518.2018.1434899 2996:10.1080/02724634.2016.1096275 1672:Cope compared the length of 1592:Tylosaurus saskatchewanensis 1057:cataloged as MT 2. Although 1037:was described by in 1988 by 539:'s 1889 fictional narrative 259: 245: 3538:"Royal Saskatchewan Museum" 2804:Lingham-Soliar, T. (1992). 2038:American Journal of Science 1953:American Journal of Science 430:The earliest depictions of 220:In 1872, Marsh argued that 119:located in the vicinity of 3794: 3690:University Press of Kansas 2718:Frontiers in Earth Science 2482:10.1126/science.10.260.919 1563:Reconstructed skeleton of 1477: 1229:to a possible specimen of 1096:was actually a species of 1001: 836:In his landmark 1967 work 805:. A particular find was a 537:Charles Hazelius Sternberg 368:. Cope also insisted that 342:, a paleontologist of the 320:University of Pennsylvania 302: 252: 238: 156:and declared his original 3520:Royal Saskatchewan Museum 3365:10.1017/S0016774600021016 3262:10.1007/978-3-030-03083-4 3156:10.1017/s026359330000033x 2942:10.1080/02724631003621870 2816:: 171–194. Archived from 2293:10.1017/S0080456800026454 1572:Royal Saskatchewan Museum 1473:21st century developments 1379:was primarily known from 858:, along with recognizing 300:(knob) and Ancient Greek 3731:Indiana University Press 2739:10.3389/feart.2019.00047 2398:Williston, S.W. (1898). 1618:University of Cincinnati 1433:near the town of Iembe, 573:Samuel Wendell Williston 208:were recovered from the 204:The earliest fossils of 3778:History of paleontology 3334:Everhart, M.J. (2005). 2914:Hainosaurus pembinensis 2887:Journal of Paleontology 2668:Everhart, M.J. (2008), 2645:Everhart, M.J. (2014). 2576:Everhart, M.J. (2008). 2173:10.5962/bhl.title.15810 2064:Everhart, M.J. (2002). 1988:Everhart, M.J. (2000). 1829:10.5962/bhl.title.60499 1782:Everhart, M.J. (2005). 1602: 1094:Hainosaurus pembinensis 1067:Hainosaurus pembinensis 1019:. The specific epithet 862:as a junior synonym of 383:Benjamin Franklin Mudge 186:Smithsonian Institution 181:New Jersey State Museum 3599:Bullard, T.S. (2006). 3459:Zietlow, A.R. (2020). 3385:Tylosaurus kansasensis 3381:Tylosaurus nepaeolicus 3338:Tylosaurus kansasensis 3254:Biodiversity of Angola 2918:Tylosaurus pembinensis 2603:Gilmore, C.W. (1921). 2119:Merriam, J.C. (1894). 2068:Tylosaurus nepaeolicus 1784:"Macrosaurus proriger" 1678:Balaenoptera sibbaldii 1567: 1509:Tylosaurus kansasensis 1459:NOVA University Lisbon 1296: 1181: 1116:Tylosaurus pembinensis 1089: 930: 786: 785:remains in its stomach 728:Henry Fairfield Osborn 564: 427: 250:, meaning "nose") and 217: 125:Union Pacific Railroad 77: 56:that lived during the 29: 23:Mounted skeleton of a 3107:Thevenin, A. (1896). 2516:Garvey, S.T. (2020). 2456:Osborn, H.F. (1899). 2347:Balaenoptera musculus 2343:Balaenoptera musculus 2099:Joseph Leidy (1873). 1990:"Tylosaurus proriger" 1747:Ornithotarsus immanis 1682:Balaenoptera musculus 1562: 1455:Tylosaurus iembeensis 1427:Mosasaurus iembeensis 1423:Miguel Telles Antunes 1327:and gave it the name 1321:Mosasaurus hoffmannii 1291: 1175: 1083: 889: 776: 558: 453:of the appearance of 418:Early restoration of 417: 340:John Campbell Merriam 203: 101:Othniel Charles Marsh 71: 22: 3611:10.7939/r3-c3qm-y670 2968:Hainosaurus bernardi 2820:on November 12, 2021 2592:on October 20, 2020. 2519:A new high-latitude 2242:10.1660/062.117.0106 2200:10.1660/062.118.0107 2194:(1 & 2): 53–67. 2028:Marsh, O.C. (1872). 2000:on October 16, 2012. 1947:Marsh, O.C. (1872). 1884:on October 19, 2015. 1810:Cope, E. D. (1870). 1751:Macrosaurus proriger 1739:Cope, E. D. (1869). 1596:T. saskatchewanensis 1565:T. saskatchewanensis 1554:T. saskatchewanensis 1391:and thus renamed it 1084:Mounted skeleton of 989:Hainosaurus bernardi 967:described the genus 921:T. saskatchewanensis 815:Logan County, Kansas 690:1899 restoration by 344:University of Munich 336:Rhinosaurus micromus 328:Rhinosaurus dyspelor 324:Rhinosaurus proriger 274:Rhinosaurus micromus 158:Macrosaurus proriger 133:Macrosaurus proriger 64:Discovery and naming 34:research history of 3478:10.7717/peerj.10145 3426:Tylosaurus proriger 3401:2016CrRes..65...68J 3389:Cretaceous Research 3356:2005NJGeo..84..231E 3140:Tylosaurus ivoensis 3078:2018CrRes..87..358S 3066:Cretaceous Research 3031:2018Alch...42..543H 2988:2016JVPal..36E6275J 2934:2010JVPal..30..416B 2920:(Nicholls, 1988)". 2916:Nicholls, 1988, as 2857:1988CaJES..25.1564N 2730:2019FrEaS...7...47J 2657:on August 20, 2020. 2621:1921SciAm.124..273G 2615:(14): 273&280. 2609:Scientific American 2563:National Geographic 2553:Switek, B. (2012). 2474:1899Sci....10..919O 2275:Turner, W. (1870). 2155:Cope, E.D. (1872). 1654:Tylosaurus borealis 1393:Tylosaurus ivoensis 1341:Mosasaurus ivoensis 1244:Hainosaurus gaudryi 1148:Tylosaurus bernardi 820:National Geographic 270:Yale Peabody Museum 268:that is now in the 97:Edward Drinker Cope 49:of extinct aquatic 3445:10.18435/vamp29339 3256:. pp. 53–76. 2764:Dollo, L. (1885). 2367:, p. 316-317. 1794:on March 14, 2016. 1743:Holops brevispinus 1568: 1305:Kristianstad Basin 1297: 1292:Isolated tooth of 1276:Tylosaurus gaudryi 1219:Mosasaurus gaudryi 1195:Jean Albert Gaudry 1182: 1114:and renamed it to 1090: 1039:Elizabeth Nicholls 931: 882:Additional species 852:and declared it a 787: 589:cervical vertebrae 565: 450:transverse process 428: 395:Liodon nepaeolicus 379:Liodon nepaeolicus 218: 210:Niobrara Formation 146:(prow) and suffix 117:Niobrara Formation 78: 30: 3740:978-0-253-02632-3 3699:978-0-7006-1394-6 3383:(Cope, 1874) and 3271:978-3-030-03082-7 3072:(2018): 358–367. 2851:(10): 1564–1570. 2325:978-0-226-50341-7 2125:Palaeontographica 1925:(1997): 193–203. 1741:"[Remarks on 1584:Master of Science 1347:was a species of 1339:bone as being of 1274:, renaming it to 975:near the town of 890:Fossil skulls of 760:Charles R. Knight 711: 695: 692:Charles R. Knight 675: 659: 377:, which he named 91:and the first in 3785: 3758: 3752: 3744: 3717: 3711: 3703: 3683: 3669: 3663: 3655: 3638:. Vol. 23. 3630:Russell, Dale A. 3615: 3614: 3596: 3590: 3589: 3561: 3550: 3549: 3547: 3545: 3534: 3528: 3527: 3522:. Archived from 3512: 3501: 3500: 3490: 3480: 3456: 3450: 3449: 3447: 3419: 3413: 3412: 3376: 3370: 3369: 3367: 3331: 3325: 3319: 3306: 3305: 3293: 3284: 3283: 3245: 3239: 3238: 3228: 3219: 3210: 3209: 3177: 3168: 3167: 3135: 3129: 3128: 3104: 3098: 3097: 3057: 3051: 3050: 3014: 3008: 3007: 2970:Dollo, 1885, to 2963: 2954: 2953: 2909: 2903: 2902: 2893:(6): 1157–1165. 2878: 2869: 2868: 2836: 2830: 2829: 2827: 2825: 2801: 2786: 2785: 2761: 2752: 2751: 2741: 2709: 2703: 2697: 2691: 2685: 2679: 2673: 2672:, pp. 37–46 2665: 2659: 2658: 2653:. Archived from 2651:Oceans of Kansas 2642: 2633: 2632: 2600: 2594: 2593: 2588:. Archived from 2586:Oceans of Kansas 2573: 2567: 2566: 2550: 2544: 2538: 2532: 2531: 2513: 2504: 2503: 2493: 2453: 2442: 2436: 2430: 2429: 2418: 2412: 2411: 2395: 2380: 2374: 2368: 2362: 2356: 2355: 2353: 2336: 2330: 2329: 2311: 2305: 2304: 2272: 2266: 2260: 2254: 2253: 2221: 2212: 2211: 2183: 2177: 2176: 2152: 2133: 2132: 2116: 2107: 2106: 2096: 2090: 2089: 2061: 2046: 2045: 2025: 2019: 2013: 2002: 2001: 1996:. Archived from 1994:Oceans of Kansas 1985: 1976: 1970: 1961: 1960: 1944: 1935: 1934: 1910: 1901: 1895: 1886: 1885: 1880:. Archived from 1870: 1864: 1858: 1849: 1843: 1834: 1833: 1831: 1807: 1796: 1795: 1790:. Archived from 1788:Oceans of Kansas 1779: 1773: 1772: 1736: 1727: 1721: 1715: 1709: 1685: 1670: 1611: 1607: 1530:Oceans of Kansas 1486: 1482: 1431:Itombe Formation 1361:Mikael Siversson 1004: 1003: 811:Smoky Hill Chalk 709: 706: 689: 686: 673: 670: 657: 654: 597:shoulder girdles 410:Early depictions 322:by transferring 305: 304: 266:Smoky Hill River 262: 255: 254: 248: 241: 240: 137:specific epithet 76:, in Cope (1870) 3793: 3792: 3788: 3787: 3786: 3784: 3783: 3782: 3763: 3762: 3761: 3745: 3741: 3725:. Vol. 2. 3720: 3704: 3700: 3672: 3656: 3628: 3624: 3619: 3618: 3598: 3597: 3593: 3563: 3562: 3553: 3543: 3541: 3536: 3535: 3531: 3514: 3513: 3504: 3458: 3457: 3453: 3421: 3420: 3416: 3378: 3377: 3373: 3333: 3332: 3328: 3320: 3309: 3295: 3294: 3287: 3272: 3247: 3246: 3242: 3226: 3221: 3220: 3213: 3179: 3178: 3171: 3137: 3136: 3132: 3119:(24): 900–916. 3106: 3105: 3101: 3059: 3058: 3054: 3016: 3015: 3011: 2982:(3): e1096275. 2965: 2964: 2957: 2911: 2910: 2906: 2880: 2879: 2872: 2865:10.1139/e88-149 2838: 2837: 2833: 2823: 2821: 2803: 2802: 2789: 2766:"Le Hainosaure" 2763: 2762: 2755: 2711: 2710: 2706: 2698: 2694: 2686: 2682: 2667: 2666: 2662: 2647:"Tylosaur food" 2644: 2643: 2636: 2602: 2601: 2597: 2575: 2574: 2570: 2552: 2551: 2547: 2539: 2535: 2515: 2514: 2507: 2455: 2454: 2445: 2437: 2433: 2420: 2419: 2415: 2397: 2396: 2383: 2375: 2371: 2363: 2359: 2351: 2338: 2337: 2333: 2326: 2313: 2312: 2308: 2274: 2273: 2269: 2261: 2257: 2223: 2222: 2215: 2185: 2184: 2180: 2154: 2153: 2136: 2118: 2117: 2110: 2098: 2097: 2093: 2063: 2062: 2049: 2027: 2026: 2022: 2014: 2005: 1987: 1986: 1979: 1971: 1964: 1946: 1945: 1938: 1915:Liodon dyspelor 1912: 1911: 1904: 1896: 1889: 1872: 1871: 1867: 1859: 1852: 1844: 1837: 1809: 1808: 1799: 1781: 1780: 1776: 1738: 1737: 1730: 1722: 1718: 1710: 1699: 1694: 1689: 1688: 1671: 1667: 1662: 1638:boreal climates 1614: 1609: 1605: 1557: 1501:Louis L. Jacobs 1489: 1484: 1480: 1475: 1419: 1286: 1170: 1078: 1055:Miami, Manitoba 1031: 961: 884: 724: 723: 722: 721: 714: 713: 712: 707: 698: 697: 696: 687: 678: 677: 676: 671: 662: 661: 660: 655: 593:Francis H. Snow 553: 519:Liodon proriger 517:(identified as 461:forward via an 412: 407: 226:Liodon dyspelor 222:Liodon proriger 193:went to Cope's 190:Liodon dyspelor 169:Liodon dyspelor 66: 58:Late Cretaceous 17: 12: 11: 5: 3791: 3789: 3781: 3780: 3775: 3765: 3764: 3760: 3759: 3739: 3718: 3698: 3674:Ellis, Richard 3670: 3625: 3623: 3620: 3617: 3616: 3591: 3551: 3529: 3526:on 2020-10-01. 3502: 3451: 3414: 3371: 3350:(3): 231–240. 3326: 3324:, p. 208. 3307: 3285: 3270: 3240: 3211: 3192:(2): 339–348. 3169: 3130: 3099: 3052: 3025:(4): 543–559. 3009: 2974:Marsh, 1872". 2955: 2928:(2): 416–426. 2904: 2870: 2831: 2787: 2753: 2704: 2702:, p. 232. 2692: 2690:, p. 175. 2680: 2660: 2634: 2595: 2568: 2545: 2543:, p. 229. 2533: 2530:on 2020-07-23. 2505: 2468:(4): 167–188. 2443: 2441:, p. 228. 2431: 2413: 2381: 2379:, p. 216. 2369: 2357: 2331: 2324: 2306: 2287:(1): 197–251. 2267: 2265:, p. 226. 2255: 2236:(1–2): 41–54. 2213: 2178: 2134: 2108: 2091: 2080:(1–2): 33–43. 2047: 2020: 2018:, p. 173. 2003: 1977: 1975:, p. 174. 1962: 1959:(18): 448–464. 1936: 1902: 1900:, p. 184. 1887: 1865: 1863:, p. 208. 1850: 1848:, p. 207. 1835: 1797: 1774: 1728: 1726:, p. 207. 1716: 1714:, p. 206. 1696: 1695: 1693: 1690: 1687: 1686: 1664: 1663: 1661: 1658: 1646:paedomorphosis 1626:Grande Prairie 1613: 1601: 1556: 1551: 1543:T. nepaeolicus 1539:T. kansasensis 1535:T. kansasensis 1526:T. nepaeolicus 1522:T. kansasensis 1518:T. nepaeolicus 1513:T. kansasensis 1488: 1476: 1474: 1471: 1463:Octávio Mateus 1418: 1413: 1285: 1280: 1260:T. pembinensis 1180:(MNHN 1896–15) 1169: 1164: 1077: 1071: 1030: 1028:T. pembinensis 1025: 960: 955: 943:T. nepaeolicus 928:T. pembinensis 906:T. nepaeolicus 883: 880: 868:T. nepaeolicus 716: 715: 708: 701: 700: 699: 688: 681: 680: 679: 672: 665: 664: 663: 656: 649: 648: 647: 646: 645: 629:T. nepaeolicus 552: 549: 484:T. nepaeolicus 440:tail vertebrae 411: 408: 406: 403: 121:Monument Rocks 65: 62: 15: 13: 10: 9: 6: 4: 3: 2: 3790: 3779: 3776: 3774: 3771: 3770: 3768: 3756: 3750: 3742: 3736: 3732: 3728: 3724: 3719: 3715: 3709: 3701: 3695: 3691: 3687: 3682: 3681: 3675: 3671: 3667: 3661: 3653: 3649: 3645: 3641: 3637: 3636: 3631: 3627: 3626: 3621: 3612: 3608: 3604: 3603: 3595: 3592: 3587: 3583: 3579: 3575: 3571: 3567: 3560: 3558: 3556: 3552: 3539: 3533: 3530: 3525: 3521: 3517: 3511: 3509: 3507: 3503: 3498: 3494: 3489: 3484: 3479: 3474: 3470: 3466: 3462: 3455: 3452: 3446: 3441: 3437: 3433: 3429: 3427: 3418: 3415: 3410: 3406: 3402: 3398: 3394: 3390: 3386: 3382: 3375: 3372: 3366: 3361: 3357: 3353: 3349: 3345: 3341: 3339: 3330: 3327: 3323: 3322:Everhart 2017 3318: 3316: 3314: 3312: 3308: 3303: 3299: 3292: 3290: 3286: 3281: 3277: 3273: 3267: 3263: 3259: 3255: 3251: 3244: 3241: 3236: 3232: 3225: 3218: 3216: 3212: 3207: 3203: 3199: 3195: 3191: 3187: 3183: 3182:Taniwhasaurus 3176: 3174: 3170: 3165: 3161: 3157: 3153: 3149: 3145: 3141: 3134: 3131: 3126: 3122: 3118: 3114: 3110: 3103: 3100: 3095: 3091: 3087: 3083: 3079: 3075: 3071: 3067: 3063: 3056: 3053: 3048: 3044: 3040: 3036: 3032: 3028: 3024: 3020: 3013: 3010: 3005: 3001: 2997: 2993: 2989: 2985: 2981: 2977: 2973: 2969: 2962: 2960: 2956: 2951: 2947: 2943: 2939: 2935: 2931: 2927: 2923: 2919: 2915: 2908: 2905: 2900: 2896: 2892: 2888: 2884: 2877: 2875: 2871: 2866: 2862: 2858: 2854: 2850: 2846: 2842: 2835: 2832: 2819: 2815: 2811: 2807: 2800: 2798: 2796: 2794: 2792: 2788: 2783: 2779: 2775: 2772:(in French). 2771: 2767: 2760: 2758: 2754: 2749: 2745: 2740: 2735: 2731: 2727: 2723: 2719: 2715: 2708: 2705: 2701: 2700:Everhart 2017 2696: 2693: 2689: 2684: 2681: 2677: 2671: 2664: 2661: 2656: 2652: 2648: 2641: 2639: 2635: 2630: 2626: 2622: 2618: 2614: 2610: 2606: 2599: 2596: 2591: 2587: 2583: 2581: 2572: 2569: 2564: 2560: 2558: 2555:"Repost: How 2549: 2546: 2542: 2541:Everhart 2017 2537: 2534: 2529: 2525: 2524: 2520: 2512: 2510: 2506: 2501: 2497: 2492: 2487: 2483: 2479: 2475: 2471: 2467: 2463: 2459: 2452: 2450: 2448: 2444: 2440: 2439:Everhart 2017 2435: 2432: 2427: 2423: 2417: 2414: 2409: 2405: 2401: 2394: 2392: 2390: 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1766: 1762: 1758: 1754: 1752: 1748: 1744: 1735: 1733: 1729: 1725: 1720: 1717: 1713: 1712:Everhart 2017 1708: 1706: 1704: 1702: 1698: 1691: 1683: 1679: 1675: 1669: 1666: 1659: 1657: 1655: 1651: 1647: 1643: 1639: 1635: 1631: 1627: 1623: 1619: 1612: 1600: 1597: 1593: 1589: 1585: 1581: 1577: 1573: 1570:In 1995, the 1566: 1561: 1555: 1552: 1550: 1548: 1544: 1540: 1536: 1531: 1527: 1523: 1519: 1514: 1510: 1506: 1502: 1498: 1494: 1487: 1472: 1470: 1468: 1467:T. iembeensis 1464: 1460: 1456: 1452: 1448: 1444: 1440: 1436: 1432: 1428: 1424: 1417: 1416:T. iembeensis 1414: 1412: 1410: 1409:Taniwhasaurus 1406: 1402: 1398: 1397:Taniwhasaurus 1394: 1390: 1386: 1382: 1378: 1377: 1376:Taniwhasaurus 1372: 1371: 1370:Taniwhasaurus 1366: 1362: 1358: 1354: 1350: 1346: 1342: 1338: 1334: 1331:, the prefix 1330: 1326: 1325:M. hoffmannii 1322: 1318: 1314: 1310: 1306: 1302: 1295: 1290: 1284: 1281: 1279: 1277: 1273: 1269: 1265: 1261: 1257: 1253: 1249: 1245: 1241: 1236: 1232: 1228: 1224: 1220: 1216: 1212: 1208: 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The prefix 990: 986: 982: 978: 974: 970: 966: 959: 956: 954: 952: 948: 944: 940: 936: 929: 926: 922: 919: 915: 911: 907: 904: 900: 897: 893: 888: 881: 879: 877: 873: 869: 865: 861: 857: 856: 851: 847: 843: 839: 834: 831: 827: 826: 821: 816: 812: 808: 804: 800: 796: 792: 784: 780: 777:USNM 8898, a 775: 771: 769: 765: 761: 757: 753: 752:calcification 749: 745: 741: 737: 733: 729: 719: 705: 693: 685: 669: 653: 644: 642: 638: 634: 630: 626: 622: 617: 612: 607: 602: 598: 594: 590: 586: 582: 578: 574: 570: 562: 557: 550: 548: 546: 542: 538: 534: 530: 529: 524: 520: 516: 512: 508: 504: 500: 499:Henry Worrall 496: 491: 489: 485: 480: 476: 472: 468: 464: 460: 456: 451: 446: 441: 437: 433: 425: 424:Henry Worrall 421: 416: 409: 404: 402: 400: 396: 392: 388: 387:Solomon River 384: 380: 376: 371: 370:Rhamphosaurus 367: 363: 359: 355: 351: 347: 346:at the time. 345: 341: 337: 333: 329: 325: 321: 317: 313: 309: 299: 295: 291: 290:Rhamphosaurus 287: 286:Rhamphosaurus 283: 279: 275: 271: 267: 263: 261: 249: 247: 235: 231: 227: 223: 215: 211: 207: 202: 198: 196: 191: 187: 182: 178: 174: 170: 166: 161: 159: 155: 154: 149: 145: 141: 138: 134: 130: 126: 122: 118: 114: 110: 109:Louis Agassiz 106: 102: 98: 94: 90: 86: 82: 75: 70: 63: 61: 59: 55: 52: 48: 44: 43: 38: 37: 28: 27: 21: 3773:Tylosaurinae 3722: 3679: 3634: 3622:Bibliography 3601: 3594: 3572:(10): 1–16. 3569: 3565: 3542:. Retrieved 3532: 3524:the original 3519: 3468: 3464: 3454: 3435: 3431: 3425: 3417: 3392: 3388: 3384: 3380: 3374: 3347: 3343: 3337: 3329: 3301: 3253: 3243: 3237:(1): 91–110. 3234: 3230: 3189: 3185: 3181: 3150:(1): 73–93. 3147: 3143: 3139: 3133: 3116: 3112: 3102: 3069: 3065: 3055: 3022: 3018: 3012: 2979: 2975: 2971: 2967: 2925: 2921: 2917: 2913: 2907: 2890: 2886: 2882: 2848: 2844: 2840: 2834: 2824:December 24, 2822:. Retrieved 2818:the original 2813: 2809: 2773: 2769: 2721: 2717: 2707: 2695: 2688:Russell 1967 2683: 2675: 2669: 2663: 2655:the original 2650: 2612: 2608: 2598: 2590:the original 2585: 2579: 2571: 2562: 2556: 2548: 2536: 2528:the original 2522: 2518: 2465: 2461: 2434: 2425: 2416: 2410:(5): 81–221. 2407: 2403: 2372: 2360: 2346: 2342: 2334: 2315: 2309: 2284: 2280: 2270: 2258: 2233: 2229: 2191: 2187: 2181: 2164: 2160: 2128: 2124: 2101: 2094: 2077: 2073: 2067: 2041: 2037: 2031: 2023: 2016:Russell 1967 1998:the original 1993: 1973:Russell 1967 1956: 1955:. Series 3. 1952: 1922: 1918: 1914: 1898:Russell 1967 1882:the original 1877: 1868: 1822:(1): 1–252. 1819: 1815: 1792:the original 1787: 1777: 1760: 1756: 1750: 1746: 1742: 1719: 1681: 1677: 1673: 1668: 1653: 1649: 1641: 1633: 1621: 1615: 1603: 1595: 1591: 1587: 1576:Saskatchewan 1569: 1564: 1553: 1546: 1542: 1538: 1534: 1529: 1525: 1521: 1517: 1512: 1508: 1504: 1496: 1493:Ellis County 1490: 1478: 1466: 1454: 1450: 1446: 1426: 1420: 1415: 1408: 1404: 1399:in northern 1396: 1392: 1388: 1384: 1374: 1368: 1364: 1356: 1352: 1348: 1344: 1340: 1332: 1328: 1324: 1320: 1298: 1293: 1282: 1275: 1271: 1267: 1263: 1259: 1255: 1251: 1247: 1243: 1239: 1230: 1226: 1218: 1214: 1210: 1206: 1202: 1198: 1183: 1177: 1166: 1159: 1155: 1151: 1147: 1143: 1139: 1135: 1131: 1127: 1123: 1119: 1115: 1111: 1106: 1102: 1097: 1093: 1091: 1085: 1074: 1073:Synonymy of 1066: 1062: 1058: 1043:Pierre Shale 1034: 1032: 1027: 1020: 1006: 992: 988: 968: 962: 957: 950: 946: 942: 938: 934: 932: 927: 924: 920: 917: 913: 909: 905: 902: 898: 895: 891: 875: 871: 867: 863: 859: 853: 849: 845: 842:Dale Russell 837: 835: 829: 823: 813:deposits in 806: 794: 790: 788: 778: 763: 755: 735: 731: 725: 717: 640: 636: 632: 628: 624: 620: 615: 610: 605: 604:curves that 600: 580: 576: 568: 566: 560: 544: 540: 532: 528:Elasmosaurus 526: 522: 518: 514: 506: 503:Buffalo Land 502: 494: 492: 487: 483: 478: 470: 466: 463:anguilliform 458: 454: 444: 435: 431: 429: 419: 398: 394: 390: 378: 374: 369: 365: 361: 357: 353: 349: 348: 335: 331: 327: 323: 316:Joseph Leidy 311: 307: 297: 293: 289: 285: 281: 277: 273: 257: 243: 229: 225: 221: 219: 205: 189: 168: 164: 162: 157: 151: 147: 143: 139: 132: 88: 84: 80: 79: 73: 40: 35: 33: 31: 24: 3727:Bloomington 2883:Hainosaurus 2841:Hainosaurus 2776:: 285–289. 2582:Collection" 2400:"Mosasaurs" 2167:: 318–349. 2032:Rhinosaurus 1674:T. dyspelor 1547:T. proriger 1483:kansasensis 1405:T. ivoensis 1385:M. ivoensis 1381:New Zealand 1365:M. ivoensis 1353:Hainosaurus 1345:M. ivoensis 1317:ichthyosaur 1294:T. ivoensis 1283:T. ivoensis 1264:H. bernardi 1256:T. ivoensis 1248:Hainosaurus 1240:Hainosaurus 1231:H. bernardi 1215:Hainosaurus 1156:T. bernardi 1152:Hainosaurus 1144:H. bernardi 1136:Hainosaurus 1132:H. bernardi 1128:Hainosaurus 1120:Hainosaurus 1112:Hainosaurus 1088:(IRSNB R23) 1086:T. bernardi 1075:Hainosaurus 1059:Hainosaurus 1035:Hainosaurus 973:Ciply Basin 969:Hainosaurus 965:Louis Dollo 958:T. bernardi 939:T. proriger 899:T. proriger 864:T. proriger 860:T. micromus 855:nomem vanum 850:T. dyspelor 830:T. proriger 807:T. proriger 795:T. dyspelor 791:T. proriger 779:T. proriger 736:T. dyspelor 718:T. proriger 641:T. dyspelor 637:T. proriger 633:T. proriger 625:T. micromus 611:T. proriger 581:T. proriger 488:T. proriger 479:T. dyspelor 471:T. dyspelor 467:T. proriger 391:nepaeolicus 362:T. micromus 358:T. dyspelor 354:T. proriger 310:species to 308:Rhinosaurus 278:Rhinosaurus 234:portmanteau 230:Rhinosaurus 160:a synonym. 103:during the 89:Platecarpus 74:T. proriger 3767:Categories 3471:: e10145. 2972:Tylosaurus 2724:(47): 47. 2580:Tylosaurus 2557:Tylosaurus 2521:Tylosaurus 2044:(20): 147. 1861:Ellis 2003 1724:Ellis 2003 1692:References 1650:Tylosaurus 1642:Tylosaurus 1634:Tylosaurus 1622:Tylosaurus 1588:Tylosaurus 1505:Tylosaurus 1497:Tylosaurus 1451:Tylosaurus 1447:Mosasaurus 1425:described 1389:Tylosaurus 1357:Tylosaurus 1349:Mosasaurus 1272:Tylosaurus 1268:H. gaudryi 1252:Tylosaurus 1227:M. gaudryi 1211:Tylosaurus 1207:Mosasaurus 1203:Mosasaurus 1178:T. gaudryi 1167:T. gaudryi 1160:Tylosaurus 1140:Tylosaurus 1124:Tylosaurus 1107:Tylosaurus 1103:Tylosaurus 1098:Tylosaurus 1063:Tylosaurus 1008:Mosasaurus 951:Tylosaurus 947:Tylosaurus 935:Tylosaurus 914:. bernardi 892:Tylosaurus 876:Tylosaurus 872:Tylosaurus 846:Tylosaurus 783:plesiosaur 764:Tylosaurus 756:Tylosaurus 732:Tylosaurus 621:Tylosaurus 616:Tylosaurus 606:Tylosaurus 601:Tylosaurus 577:Tylosaurus 569:Tylosaurus 561:Tylosaurus 545:Tylosaurus 533:Tylosaurus 523:Tylosaurus 515:Tylosaurus 495:Tylosaurus 475:blue whale 459:Tylosaurus 455:Tylosaurus 445:Tylosaurus 436:Tylosaurus 432:Tylosaurus 420:Tylosaurus 399:Tylosaurus 375:Tylosaurus 350:Tylosaurus 332:Tylosaurus 312:Tylosaurus 294:Tylosaurus 206:Tylosaurus 177:New Mexico 173:Fort McRae 81:Tylosaurus 42:Tylosaurus 36:Tylosaurus 26:Tylosaurus 3749:cite book 3708:cite book 3660:cite book 3640:New Haven 3438:: 75–90. 3395:: 68–81. 3304:: 71–105. 3206:129446036 3164:131368986 3125:0037-9409 3094:133843005 3047:134724144 2782:0035-2160 2491:2246/5737 2349:in Canada 2030:"Note on 1769:0097-3157 1443:Coniacian 1421:In 1964, 1176:Skull of 1162:species. 1015:near its 963:In 1885, 726:In 1899, 188:that the 129:Fort Hays 123:near the 105:Bone Wars 85:Clidastes 3686:Lawrence 3676:(2003). 3632:(1967). 3586:90533033 3544:July 23, 3516:"Omācīw" 3497:33150074 3280:67769971 3004:87315531 2950:86297189 2748:85513442 2500:17837338 2301:87196687 2250:86117382 2208:83904449 1878:FishBase 1608:borealis 1439:Turonian 1337:coracoid 1301:Ivö Lake 1235:buttress 1047:Manitoba 1021:bernardi 894:species 511:paleoart 140:proriger 51:squamate 3488:7583613 3397:Bibcode 3352:Bibcode 3074:Bibcode 3027:Bibcode 2984:Bibcode 2930:Bibcode 2853:Bibcode 2726:Bibcode 2617:Bibcode 2470:Bibcode 2131:: 1–39. 1931:4065032 1763:: 123. 1630:Alberta 1303:in the 1013:a river 981:Belgium 803:Germany 748:bronchi 744:trachea 318:of the 111:of the 54:reptile 3737: 3696: 3652:205385 3650: 3584: 3495: 3485: 3278: 3268: 3204: 3162: 3123: 3092: 3045: 3002: 2948: 2780: 2746: 2498: 2322: 2299: 2248: 2206: 1929: 1767: 1753:]" 1749:, and 1435:Angola 1313:Sweden 1309:Scania 1199:Liodon 1191:France 1051:Canada 1002:σαῦρος 977:Mesvin 923:, and 799:George 740:larynx 426:, 1872 366:Liodon 360:, and 303:σαῦρος 282:Liodon 260:saûros 253:σαῦρος 214:Kansas 165:Liodon 153:Liodon 93:Kansas 3582:S2CID 3465:PeerJ 3276:S2CID 3227:(PDF) 3202:S2CID 3160:S2CID 3090:S2CID 3043:S2CID 3000:S2CID 2946:S2CID 2744:S2CID 2674:, in 2578:"The 2352:(PDF) 2297:S2CID 2246:S2CID 2204:S2CID 1927:JSTOR 1660:Notes 1610:' 1606:' 1485:' 1481:' 1401:Japan 1262:than 1122:from 997:Haine 993:Haino 937:were 822:film 298:tylos 246:rhī́s 148:-gero 144:prōra 47:genus 32:This 3755:link 3735:ISBN 3714:link 3694:ISBN 3666:link 3648:OCLC 3546:2020 3493:PMID 3266:ISBN 3121:ISSN 2826:2020 2778:ISSN 2496:PMID 2320:ISBN 1765:ISSN 1761:1869 1580:Cree 1541:and 1258:and 1138:and 941:and 793:and 768:meme 639:and 627:and 326:and 224:and 99:and 87:and 45:, a 3607:doi 3574:doi 3483:PMC 3473:doi 3440:doi 3405:doi 3360:doi 3258:doi 3194:doi 3152:doi 3082:doi 3035:doi 2992:doi 2938:doi 2895:doi 2861:doi 2734:doi 2625:doi 2613:124 2486:hdl 2478:doi 2289:doi 2238:doi 2234:117 2196:doi 2192:118 2169:doi 2082:doi 2078:105 1923:147 1824:doi 1604:T. 1479:T. 1333:ivo 1319:or 1307:in 1213:or 1146:to 1045:in 925:(E) 918:(D) 910:(C) 903:(B) 896:(A) 330:to 239:ῥίς 212:in 127:at 3769:: 3751:}} 3747:{{ 3733:. 3729:: 3710:}} 3706:{{ 3692:. 3688:: 3684:. 3662:}} 3658:{{ 3646:. 3642:: 3580:. 3570:17 3568:. 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Knowledge

Research history of Tylosaurus

Index