Pentaceratops

Pentaceratops Temporal range: Late Cretaceous, 75.79–73 Ma PreЄ Є O S D C P T J K Pg N ↓
Type skull of Pentaceratops sternbergii, AMNH 6325
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Ornithischia
Family:	†Ceratopsidae
Subfamily:	†Chasmosaurinae
Genus:	†Pentaceratops Osborn, 1923
Type species
Pentaceratops sternbergii Osborn, 1923
Species
P. sternbergii Osborn, 1923 P. aquilonius Longrich, 2014[1]

Pentaceratops (five-horned face) is a genus of herbivorous ceratopsid dinosaur from the late Cretaceous Period of what is now North America.

Pentaceratops fossils were first discovered in 1921. The genus was named in 1923 when its type species Pentaceratops sternbergii was described. Pentaceratops lived around 76-73 million years ago, its remains having been mostly found in the Kirtland Formation^[2] in the San Juan Basin in New Mexico. Other dinosaurs which shared its habitat include Parasaurolophus cyrtocristatus, the pachycephalosaur Sphaerotholus, the armored dinosaur Nodocephalosaurus and the tyrannosauroid Bistahieversor. About a dozen skulls and skeletons have been uncovered, so that most bones are known. One exceptionally large specimen later became its own genus, Titanoceratops, due to its more derived morphology closer to Triceratops and lack of unique characters shared with Pentaceratops proper,^[3] although the author that originally assigned it to Pentaceratops has decided to ignore this in subsequent publications.^[4]

Pentaceratops was about six meters (twenty feet) long, and has been estimated to have weighed around five tonnes. It had a short nose horn, two long brow horns, and long horns on the jugal bones. Its skull had a very long frill with triangular hornlets on the edge.

Discoveries and species

Restored holotype, AMNH

The first specimens were collected by Charles Hazelius Sternberg in the San Juan Basin in New Mexico. Sternberg worked in commission for the Swedish Uppsala University . In 1921 he recovered a skull and a rump, specimens PMU R.200 and PMU R.286, at the Meyers Creek near the Kimbetoh Wash, in a layer of the Kirtland Formation, he sent the fossils to paleontologist Carl Wiman. In 1922 Sternberg decided to work on his own account and north of Tsaya Trading Post, in the Fossil Forest of San Juan County, he discovered a complete skeleton , which he sold to the American Museum of Natural History. The museum then sent out a team headed by Charles Mook and Peter Kaisen to assist Sternberg in securing this specimen; subsequent digging by Sternberg in 1923 brought the total of AMNH specimens to four. The rump of the main specimen was discarded by the museum because it had insufficient value as a display.

The species was named and described by Henry Fairfield Osborn in 1923, as Pentaceratops sternbergii. The generic name means "five-horned face", derived from the Greek penta (πέντα, meaning five), keras (κέρας, horn) and -ops (ὤψ, face),^[5] in reference to its two long epijugal bones, spikes which protrude out sidewards from under its eyes, in addition to the three more obvious horns as with Triceratops. Osborn obligingly gave it the specific name sternbergii honoring its discoverer as a veteran fossil hunter.^[6] The name had been suggested to Osborn by William Diller Matthew; the specific epithet served as a consolation to the almost bankrupt Sternberg whose 1923 fossils were initially not acquired by the museum that had to use its 1923/1924 budget to process the finds of the great Asian expeditions by Roy Chapman Andrews.^[7]

The holotype was the skull discovered by Sternberg in 1922, specimen AMNH 6325. It was found in a layer of the Fruitland Formation, dating from the Campanian, about seventy-five million years old. The other three AMNH specimens were AMNH 1624, a smaller skull; AMNH 1622, a pair of brow horns; and AMNH 1625, a piece of skull frill.^[6]

Skull in the New Mexico Museum of Natural History and Science

In 1930, Wiman named a second species of Pentaceratops: Pentaceratops fenestratus. It was based on Sternberg's 1921 specimens and the specific name referred to a hole in the left squamosal.^[8] This was later considered to be the same species as Pentaceratops sternbergii and thus a junior synonym, the hole being the likely effect of an injury.

In 1929 Steinberg's son, George Fryer Sternberg, discovered specimen USNM V12002, a right squamosal. Pentaceratops proved to be a quite common fossil in the Fruitland and Kirtland formations. It has even been used as a guide fossil: the appearance of Pentaceratops sternbergii in the fossil record marks the end of the Judithian land vertebrate age and the start of the Kirtlandian.^[2] Subsequent finds include specimens MNA Pl. 1668, MNA Pl. 1747, NMMNH P-27468 and USNM 2416, partial skeletons with skull; YPM 1229, a skeleton lacking the skull; UALP 13342 and UKVP 16100, skulls; UNM B-1701, USNM 12741, USNM 12743, USNM 8604, SMP VP-1596, SMP VP-1488, SMP VP-1500 and SMP VP-1712, fragmentary skulls. Apart from the San Juan Basin finds, a juvenile specimen of Pentaceratops, SDMNH 43470, has been reported from the Williams Fork Formation of Colorado in 2006.^[9]

Specimen being transported with help from the New Mexico National Guard, 2015

Sometimes the identification of a specimen as Pentaceratops has proven to be highly contentious. In 1998 Thomas Lehman described OMNH 10165, a very large skull and its associated skeleton found in New Mexico in 1941, and presently on display at the Sam Noble Oklahoma Museum of Natural History, as being the largest Pentaceratops exemplar known, with the distinction of having produced the largest known skull of any land vertebrate.^[10][11] However, in 2011, the skeleton was renamed as a separate genus, Titanoceratops, due to its more derived morphology closer to Triceratops and lack of unique characters shared with Pentaceratops proper.^[3]

In 2014 Nicholas Longrich named a new species Pentaceratops aquilonius, "the northern one", based on fragmentary fossils discovered during the 1930s near Manyberries in Canada. The species has a first epiparietal pointing upwards instead of forwards.^[12]

Description

Restoration

Pentaceratops is a large ceratopsid, Dodson estimated the body length at six meters, the skull length of AMNH 1624 at 2.3 meters while PMU R.200 has a length of 216 centimeters.^[8] The nose horn of Pentaceratops is small and pointing upwards and backwards. The brow horns are very long and curving strongly forwards. The somewhat upward tilted frill of Pentaceratops is considerably longer than that of Triceratops, with two large holes (parietal fenestrae) in it. It is rectangular, adorned by large triangular osteoderms: up to twelve episquamosals at the squamosal and three epiparietals at the parietal bone. These are largest at the rear corners of the frill, that are separated by a large U-shaped notch at the midline, a feature not recognized until 1981 when specimen UKVP 16100 was described.^[13] Within the notch the first epiparietals point forwards. The very thick jugal and the squamosal do not touch each other, a possible autapomorphy.^[3]

The torso of Pentaceratops is tall and wide. The rear dorsal vertebrae bear long spines from which perhaps ligaments ran to the front, to balance the high frill. The prepubis is long. The ischium is long and strongly curves forward. With the smaller specimens the thigh bone bows outwards.^[14]

Classification

Osborn originally assigned Pentaceratops to the Ceratopsia. Within this group Pentaceratops belonged to the Ceratopsinae or Chasmosaurinae. It appears to be most closely related to Utahceratops. Their clade was perhaps more derived than the earlier genus Chasmosaurus but more basal than Anchiceratops, the latter representing a line of which Triceratops was a member, which lived a few million years later, right at the end of the Cretaceous period, when all ceratopsians died out.

The cladogram of the phylogeny of Pentaceratops according to a study by Scott Sampson et al in 2010 found that the genus was most closely related to Utahceratops, from a similar age and region.^[15] The below cladogram follows Longrich (2014), who named a new species of Pentaceratops, and included nearly all species of chasmosaurine.^[1]

Longrich stated that the holotype and referred specimen of P. aquilonius fall within the diagnosis of Pentaceratops, and are recovered very close to the type species in the phylogeny. He noted that the placement of Utahceratops does not make the genus paraphyletic, as there is no requirement that genera are monophyletic. The Williams Fork chasmosaur differs from the Pentaceratops and Utahceratops species, and might require a new specific or generic name.^[1]

Diet

Pentaceratops, like all ceratopsians, was an herbivore. During the Cretaceous, flowering plants were "geographically limited on the landscape", and so it is likely that this dinosaur fed on the predominant plants of the era: ferns, cycads and conifers. It would have used its sharp ceratopsian beak to bite off the branches which then were shredded, leaves, needles and all, by the tooth batteries, providing a self-sharpening continuous cutting edge in both upper an lower jaws. Ultimately the plant material was digested by the large gut.

See also

• Timeline of ceratopsian research

• Ah-Shi-Sle-Pah Wilderness Study Area, the type locality for Pentaceratops fenestratus

References

<div class="reflist columns references-column-width" style="-moz-column-width: 30em ^{[2] [5] [9] [3] [11]}; -webkit-column-width: 30em ^{[2] [5] [9] [3] [11]}; column-width: 30em ^{[2] [5] [9] [3] [11]}; list-style-type: decimal;">

1. 1 2 3 Longrich, N. R. (2014). "The horned dinosaurs Pentaceratops and Kosmoceratops from the upper Campanian of Alberta and implications for dinosaur biogeography". Cretaceous Research 51: 292. doi:10.1016/j.cretres.2014.06.011. 
2. 1 2 3 4 5 Sullivan and Lucas (2006).
3. 1 2 3 4 5 6 Longrich (2011).
4. ↑ Wick, S. L.; Lehman, T. M. (2013). "A new ceratopsian dinosaur from the Javelina Formation (Maastrichtian) of West Texas and implications for chasmosaurine phylogeny". Naturwissenschaften 100 (7): 667. Bibcode:2013NW....100..667W. doi:10.1007/s00114-013-1063-0. PMID 23728202. 
5. 1 2 3 4 Liddell, et al. (1980). pp. 373, 542-43, 804.
6. 1 2 H.F. Osborn, 1923, "A new genus and species of Ceratopsia from New Mexico, Pentaceratops sternbergii, American Museum Novitates 93: 1-3
7. ↑ Sullivan, R.M. and S.G. Lucas, 2011, "Charles Hazelius Sternberg and his San Juan Basin Cretaceous dinosaur collections: Correspondence and photographs (1920-1925)", New Mexico Museum of Natural History and Science Bulletin 53: 429-471
8. 1 2 C. Wiman, 1930, "Über Ceratopsia aus der Oberen Kreide in New Mexico", Nova Acta Regiae Societatis Scientiarum Upsaliensis, Series 4 7(2): 1-19
9. 1 2 3 4 Lucas, et al. (2006).
10. ↑ Lehman, T.M., 1998, "A gigantic skull and skeleton of the horned dinosaur Pentaceratops sternbergi from New Mexico: Journal of Paleontology, 72(5): 894-906
11. 1 2 3 4 Delayed Debut for Jumbo Dino Skull
12. ↑ Longrich, N., 2014, "The horned dinosaurs Pentaceratops and Kosmoceratops from the upper Campanian of Alberta and implications for dinosaur biogeography", Cretaceous Research, 51: 292-308
13. ↑ Rowe, T., Colbert, E.H. and Nations, J.D., 1981, "The occurrence of Pentaceratops with a description of its frill", In: Lucas, S.G., Rigby, J.K. and Kues, B.S. (eds.) Advances in San Juan Basin Paleontology, University of New Mexico Press, Alburquerque p. 29-48
14. ↑ Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 272
15. ↑ Scott D. Sampson, Mark A. Loewen, Andrew A. Farke, Eric M. Roberts, Catherine A. Forster, Joshua A. Smith, and Alan L. Titus (2010). Stepanova, Anna, ed. "New Horned Dinosaurs from Utah Provide Evidence for Intracontinental Dinosaur Endemism". PLoS ONE 5 (9): e12292. doi:10.1371/journal.pone.0012292. PMC 2929175. PMID 20877459. 

Sources

• "Delayed Debut for Jumbo Dino Skull". Science 282 (5390): 871. 1998. doi:10.1126/science.282.5390.871b. 
• Dodson, P. (1996). The Horned Dinosaurs. Princeton University Press, Princeton, New Jersey. ISBN 0-691-05900-4. 
• Lehman, T. M., 2001, Late Cretaceous dinosaur provinciality: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, pp. 310–328.
• Liddell, Henry George and Robert Scott (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. ISBN 0-19-910207-4. 
• Sullivan, R.M., and Lucas, S.G. 2006. "The Kirtlandian land-vertebrate "age" – faunal composition, temporal position and biostratigraphic correlation in the nonmarine Upper Cretaceous of western North America." New Mexico Museum of Natural History and Science, Bulletin 35:7-29.
• Nicholas R. Longrich (2011). "Titanoceratops ouranous, a giant horned dinosaur from the Late Campanian of New Mexico". Cretaceous Research 32. doi:10.1016/j.cretres.2010.12.007. 
• Lucas, S.G., Sullivan, R.M., Hunt, A.P., 2006, Re-evaluation of Pentaceratops and Chasmosaurus (Ornithischia, Ceratopsidae) in the Upper Cretaceous of the Western Interior: New Mexico Museum of Natural History and Science, Bulletin 35

Wikimedia Commons has media related to Pentaceratops.

External links

• Pentaceratops sternbergii, DinoData, archived from the original on 30 September 2011, retrieved 6 January 2014 
• Pentaceratops, The Dinosaur Encyclopaedia, archived from the original on 27 May 2008, retrieved 6 January 2014 
• Kids Zone: Groups of Dinosaurs, Town of Drumheller, archived from the original on 31 May 2009, retrieved 6 January 2014 

Marginocephalia Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Order: Ornithischia   Pachycephalosauria Basal Pachycephalosaurs Wannanosaurus Pachycephalosauridae Stegoceras-line Stegoceras Hanssuesia Gravitholus Colepiocephale Texacephale Pachycephalosaurus-line Homalocephale Prenocephale Goyocephale Sphaerotholus Tylocephale Acrotholus Amtocephale Pachycephalosaurini Pachycephalosaurus Dracorex Stygimoloch Alaskacephale   Ceratopsia Basal Ceratopsians Yinlong Stenopelix Micropachycephalosaurus Albalophosaurus Chaoyangsauridae Chaoyangsaurus Xuanhuaceratops Psittacosauridae Psittacosaurus Neoceratopsia Mosaiceratops Microceratus Aquilops Liaoceratops Koreaceratops Auroraceratops Archaeoceratopsidae Archaeoceratops Yamaceratops Helioceratops Coronosauria Graciliceratops Leptoceratopsidae Leptoceratops Montanoceratops Prenoceratops Udanoceratops Zhuchengceratops Gryphoceratops Unescoceratops Cerasinops Asiaceratops Bainoceratops Bagaceratopsidae Ajkaceratops Gobiceratops Bagaceratops Platyceratops Protoceratopsidae Protoceratops Breviceratops Lamaceratops Magnirostris Ceratopsoidea Zuniceratops Turanoceratops Ceratopsidae   Ceratopsidae Ceratopsinae Ceratops Centrosaurinae Monoclonius Avaceratops Albertaceratops Diabloceratops Nasutoceratops Wendiceratops Centrosaurini Centrosaurus Coronosaurus Rubeosaurus Spinops Styracosaurus Pachyrhinosaurini Achelousaurus Einiosaurus Pachyrhinosaurus Sinoceratops Xenoceratops Chasmosaurinae Bravoceratops Agujaceratops Anchiceratops Arrhinoceratops Chasmosaurus Coahuilaceratops Pentaceratops Kosmoceratops Judiceratops Medusaceratops Mojoceratops Utahceratops Mercuriceratops Vagaceratops Triceratopsini Eotriceratops Triceratops Titanoceratops Regaliceratops Torosaurus Tatankaceratops Ojoceratops Polyonax   Related Categories Ornithischians Ceratopsians Pachycephalosaurs