Protoceratops

Protoceratops (/ˌproʊtoʊˈsɛrətɒps/; from Greek proto-/πρωτο- "first", cerat-/κερατ- "horn" and -ops/-ωψ "face", meaning "first horned face")[1] is a genus of sheep-sized (1.8 m long) herbivorous ceratopsian dinosaur, from the Upper Cretaceous Period (Campanian stage) of what is now Mongolia. It was a member of the Protoceratopsidae, a group of early horned dinosaurs. Unlike later ceratopsians, however, it was a much smaller creature that lacked well-developed horns and retained some basal traits not seen in later genera.

Protoceratops Temporal range: Late Cretaceous, 75–72 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Mounted P. andrewsi skeleton, Carnegie Museum of Natural History
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Order:	†Ornithischia
Infraorder:	†Neoceratopsia
Parvorder:	†Coronosauria
Family:	†Protoceratopsidae
Genus:	†Protoceratops Granger & Gregory, 1923
Type species
†Protoceratops andrewsi Granger & Gregory, 1923
Species
†P. andrewsi Granger & Gregory, 1923 †P. hellenikorhinus Lambert et al., 2001

Protoceratops had a large neck frill which was likely used as a display site to impress other members of the species. Other hypotheses about its function include protection of the neck and anchoring of jaw muscles, but the fragility of the frill and the poor leverage offered by possible attachment sites here makes these ideas implausible. Described by Walter W. Granger and W.K. Gregory in 1923, Protoceratops was initially believed to be an ancestor of the North American ceratopsians. Researchers currently distinguish two species of Protoceratops (P. andrewsi and P. hellenikorhinus), based in part by their respective sizes.

In the 1920s, Roy Chapman Andrews discovered fossilized eggs in Mongolia that were interpreted as belonging to this dinosaur, but which turned out to be those of Oviraptor.

Description

Size of P. andrewsi compared to a human

Protoceratops was a quadrupedal dinosaur that was partially characterized by its distinctive neck frill at the back of its skull. The frill itself contained two large parietal fenestrae (holes in the frill), while its cheeks had large jugal bones.[2] The exact size and shape of the neck frill varied by individual; some specimens had short, compact frills, while others had frills nearly half the length of the skull. The frill consists mostly of the parietal bone and partially of the squamosal. Some researchers, including Peter Dodson[3] attribute the different sizes and shapes of these bones to sexual dimorphism, as well as the age of the specimen, at the time of death.

Protoceratops was approximately 1.8 meters (5.9 ft) in length and 0.6 meters (2 ft) high at the shoulder.[4] A fully grown adult would have weighed about 82.7 kilograms (182 lb).[5] Smaller specimens are estimated at 23.7 kilograms (52 lb).[6] The large numbers of specimens found in high concentration suggest that Protoceratops lived in herds.</ref>[4]

Restoration of P. andrewsi

Protoceratops was a relatively small dinosaur with a proportionately large skull. Protoceratops appears to have had muscular jaws capable of a powerful bite. These jaws were packed with dozens of teeth, well suited for chewing tough vegetation.[4] The skull consisted of a massive frontal beak, and four pairs of fenestrae (skull openings). The foremost hole, the naris, was considerably smaller than the nostrils seen in later genera. Protoceratops had large orbits (the holes for its eyes), which measured around 50 millimeters in diameter.[3] Behind the eye was a slightly smaller fenestra, known as the "infratemporal fenestra."

Discovery and species

Skeletons of P. andrewsi and Velociraptor mongoliensis in combat

Photographer James Blaine Shackelford discovered the first specimen of Protoceratops in the Gobi desert, (Gansu, Inner Mongolia), as part of a 1922 American expedition looking for human ancestors. No early human fossils were found, but the expedition, led by Roy Chapman Andrews, collected many specimens of the genus Protoceratops, along with fossil skeletons of theropods Velociraptor, Oviraptor, and ceratopsian Psittacosaurus.[3]

Walter Granger and W. K. Gregory formally described the type species, P. andrewsi in 1923, the specific name in honor of Andrews. The fossils hail from the Djadochta Formation and date from the Campanian stage of the Upper Cretaceous (dating to between 75 and 71 million years ago).[7] Researchers immediately noted the importance of the Protoceratops finds, and the genus was hailed as the "long-sought ancestor of Triceratops".[3] The fossils were in an excellent state of preservation, with even the sclerotic rings (delicate ocular bones) preserved in some specimens.[3]

In 1971, a fossil was found that captured a Velociraptor mongoliensis clutched around a Protoceratops andrewsi in Mongolia. It is believed that they died simultaneously, while fighting, when they were either surprised by a sand storm or buried when a sand dune collapsed on top of them.[8]

P. hellenikorhinus skull

In 1975, Polish paleontologists Teresa Maryanska and Halszka Osmólska described a second species of Protoceratops, also from the Campanian stage of Mongolia, which they named P. kozlowskii.[9] However, the fossils consisted of incomplete juvenile remains, and are now considered synonymous with Bagaceratops rozhdestvenskyi.[10]

In 2001, a second valid species, P. hellenikorhinus, was named from the Bayan Mandahu Formation in Inner Mongolia, China and also dates from the Campanian stage of the Upper Cretaceous. It was notably larger than P. andrewsi, had a slightly different frill, and had more robust jugal horns. The arch of bone over its nostrils had two small nasal horns, and there were no teeth at the front of the snout.[11]

In 2011, a specimen of Protoceratops first uncovered in 1965 was found to be preserved with its own footprint. It is the first example of a dinosaur to be preserved with footprints.[12]

Paleobiology

Reproduction

The cast of the Protoceratops sp. egg at the Wrexham Museum, Wales

P. andrewsi hatchling at the American Museum of Natural History in New York City

In the 1920s, Roy Chapman Andrews discovered the first known fossilized dinosaur eggs, in the Gobi Desert of Mongolia. Each egg was about 8 inches long, and newborn hatchlings estimated to have been about a foot in length. Due to the proximity and abundance of Protoceratops, these eggs were believed at the time to belong to this genus.

The contemporary theropod Oviraptor was thought to consume Protoceratops eggs due to the discovery of an Oviraptor skeleton present at a nest. The skull was crushed, and it was speculated that the injury was received by a Protoceratops mother defending her nest from the predator. However, in 1993 Norrell et al. discovered an embryo inside a supposed Protoceratops egg. This embryo, upon close examination, turned out be an Oviraptor;[13] the original find represented Oviraptor brooding behavior rather than a failed attempt at oophagy.

The oogenus Protoceratopsidovum (meaning "Protoceratops egg") was also originally misidentified as belonging to Protoceratops; it is now known to be the eggs of a derived theropod.[14]

P. andrewsi growth series

In 2011, a nest of 15 young Protoceratops andrewsi was discovered in Mongolia.[15] Stated as being the first Protoceratops nest discovered to date, the discovery suggests that Protoceratops parents might have cared for their young at nests during at least the early stages of childhood. As Protoceratops is a relatively basal member of the ceratopsians, the finding also suggests that other ceratopsians provided care for their young as well.[15]

Studies of lines of growth (i.e. lines of von Ebner) of the teeth in embryonic P. andrewsi suggests plesiomorphically long incubation times, with a minimum incubation time of 83.16 days for P. andrewsi.[16]

Daily activity patterns

The large eyes of Protoceratops has been suggested as evidence for a nocturnal lifestyle.[17] However, subsequent comparisons between the scleral rings of Protoceratops and modern birds and reptiles have indicated a more cathemeral lifestyle, being active throughout the day during short intervals. This suggests that the fight between Protoceratops and the primarily nocturnal Velociraptor indicated by the fighting specimens may have occurred at twilight or under low-light conditions.[18]

Classification

Nest with eggs, Museum of Evolution of Polish Academy of Sciences, Warsaw

Skull of an adult P. andrewsi

Protoceratops was the first named protoceratopsian and hence gives its name to the family Protoceratopsidae, a group of herbivorous dinosaurs more derived than psittacosaurids, but less derived than ceratopsids. The group is characterized by their similarities to the Ceratopsidae but with more cursorial limb proportions, generally smaller frills, and lack of large horns.

In 1998, Paul Sereno defined Protoceratopsidae as the branch-based clade including "all coronosaurs closer to Protoceratops than to Triceratops." Some studies placed Bagaceratops, Breviceratops, Graciliceratops, Lamaceratops, Magnirostris, Platyceratops, and Serendipaceratops within Protoceratopsidae, but in 2006, Makovicky and Norell published a new phylogeny which removed several genera from Protoceratopsidae; several other phylogenies also exist. Bainoceratops may be synonymous with Protoceratops.[19]

The cladogram below follows the topology from a 2015 analysis by Yiming He, Peter J. Makovicky, Kebai Wang, Shuqing Chen, Corwin Sullivan, Fenglu Han, Xing XuMichael J. Ryan, David C. Evans, Philip J. Currie, Caleb M. Brown and Don Brinkman.[20]

Leptoceratopsidae	Asiaceratops       Cerasinops         Montanoceratops     Ischioceratops         Prenoceratops       Leptoceratops       Udanoceratops       Zhuchengceratops       Gryphoceratops     Unescoceratops
	Graciliceratops     Protoceratopsidae   Bagaceratops       Lamaceratops       Breviceratops       Magnirostris       Protoceratops hellenikorhinus     Protoceratops andrewsi                 Ajkaceratops       Zuniceratops     Ceratopsidae

Origin of griffin myths

Adrienne Mayor has speculated that the discovery of Protoceratops fossils may have inspired or influenced stories of griffins.

Restoration of P. andrewsi

Folklorist and historian of science Adrienne Mayor of Stanford University has suggested that the exquisitely preserved fossil skeletons of Protoceratops and other beaked dinosaurs, found by ancient Scythian nomads who mined gold in the Tian Shan and Altai Mountains of Central Asia, may have been at the root of the image of the mythical creature known as the griffin. Griffins were described as lion-sized quadrupeds with large claws and a raptor-bird-like beak; they laid their eggs in nests on the ground.[21]

Greek writers began describing the griffin around 675 B.C., at the same time the Greeks first made contact with Scythian nomads. Griffins were described as guarding the gold deposits in the arid hills and red sandstone formations of the wilderness. The region of Mongolia and China where many Protoceratops fossils are found is rich in gold runoff from the neighboring mountains, lending some credence to the theory that these fossils were the basis of griffin myths.[3]

In 2016 this hypothesis was contested by Mark Witton, as it ignores pre-Greek griffin art and accounts.[22]

See also

• Timeline of ceratopsian research

References

1. Liddell & Scott (1980). Greek-English Lexicon, Abridged Edition. Oxford University Press, Oxford, UK. ISBN 978-0-19-910207-5.
2. Lambert, D. (1993). The Ultimate Dinosaur Book. Dorling Kindersley, New York. pp. 152–167. ISBN 978-1-56458-304-8.
3. Dodson, P. (1996). The Horned Dinosaurs. Princeton University Press, Princeton, New Jersey. pp. 200–234. ISBN 978-0-691-05900-6.
4. "Protoceratops." In: Dodson, Peter & Britt, Brooks & Carpenter, Kenneth & Forster, Catherine A. & Gillette, David D. & Norell, Mark A. & Olshevsky, George & Parrish, J. Michael & Weishampel, David B. The Age of Dinosaurs. Publications International, LTD. p. 118-119. ISBN 0-7853-0443-6.
5. Benson, R. B. J.; Campione, N. S. E.; Carrano, M. T.; Mannion, P. D.; Sullivan, C.; Upchurch, P.; Evans, D. C. (2014). "Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage". PLOS Biology. 12 (5): e1001853. doi:10.1371/journal.pbio.1001853. PMC 4011683. PMID 24802911.
6. Seebacher, F. (2001). "A new method to calculate allometric length-mass relationships of dinosaurs". Journal of Vertebrate Paleontology. 21 (1): 53.
7. Godefroit, Pascal; Currie, Philip J.; Li, Hong; Shang, Chang Yong; Dong, Zhi-ming (2008). "A new species of Velociraptor (Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China". Journal of Vertebrate Paleontology. 28 (2): 432–438. doi:10.1671/0272-4634(2008)28[432:ANSOVD]2.0.CO;2.
8. Carpenter, Ken. (1998). "Evidence of predatory behavior by theropod dinosaurs". Gaia. 15: 135–144. Archived from the original on 17 November 2007. [not printed until 2000]
9. Maryanska, T.; Osmólska, H. (1975). "Protoceratopsidae (Dinosauria) of Asia". Palaeontologica Polonica. 33: 133–181.
10. You H. & Dodson, P. 2004. Basal Ceratopsia. In: Weishampel, D.B., Dodson, P., & Osmolska, H. (Eds.). The Dinosauria (2nd Edition). Berkeley: University of California Press. Pp. 478–493.
11. Lambert, O, Godefroit, P, Li, H, Shang, C.-Y. & Dong, Z.-M. (2001). "A new species of Protoceratops (Dinosauria, Neoceratopsia) from the Late Cretaceous of Inner Mongolia (P. R. China)". Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Science de la Terre: 5–28.CS1 maint: multiple names: authors list (link)
12. Gill, Victoria. "Protoceratops Dinosaur found with its own tracks". BBC Nature. Archived from the original on 14 September 2018.
13. "Protoceratops". Dallas Museum of Natural History. Archived from the original on 19 February 2006.
14. Zelenitsky, Darla K.; Therrien, François (July 2008). "PHYLOGENETIC ANALYSIS OF REPRODUCTIVE TRAITS OF MANIRAPTORAN THEROPODS AND ITS IMPLICATIONS FOR EGG PARATAXONOMY". Palaeontology. pp. 807–816. doi:10.1111/J.1475-4983.2008.00770.x. Archived from the original on 17 March 2020.
15. Choi, Charles (17 November 2011). "15 Infant Dinosaurs Discovered Crowded in Nest". Yahoo News. Archived from the original on 17 March 2020.
16. Erickson, G.M.; Zelenitsky, D.K.; Kay, D.I.; Norrell, M.A. (2017). "Dinosaur incubation periods directly determined from growth-line counts in embryonic teeth show reptilian-grade development" (PDF). Proceedings of the National Academy of Sciences. 114 (3): 540–545. doi:10.1073/pnas.1613716114. PMC 5255600. PMID 28049837.
17. Longrich, Nick (2010). "The Function of Large Eyes in Protoceratops: A Nocturnal Ceratopsian?". In Ryan, Michael J.; Chinnery-Allgeier, Brenda J.; Eberth, David A. (eds.). New perspectives on horned dinosaurs : the Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press. pp. 308–327. ISBN 0-253-35358-0.
18. Schmitz, L.; Motani, R. (2011). "Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology". Science. 332 (6030): 705–8. doi:10.1126/science.1200043. PMID 21493820.
19. Makovicky, P.J.; Norell, M.A. (2006). "Yamaceratops dorngobiensis, a new primitive ceratopsian (Dinosauria: Ornithischia) from the Cretaceous of Mongolia. 3530:1–42" (PDF). American Museum Novitates. 3530: 1–42. doi:10.1206/0003-0082(2006)3530[1:YDANPC]2.0.CO;2. hdl:2246/5808. Retrieved 10 August 2008.
20. Yiming He; Peter J. Makovicky; Kebai Wang; Shuqing Chen; Corwin Sullivan; Fenglu Han; Xing Xu; Michael J. Ryan; David C. Evans; Philip J. Currie; Caleb M. Brown; Don Brinkman (2015). "A New Leptoceratopsid (Ornithischia, Ceratopsia) with a Unique Ischium from the Upper Cretaceous of Shandong Province, China". PLOS One. 10 (12): e0144148. doi:10.1371/journal.pone.0144148. PMC 4689537. PMID 26701114.
21. Mayor, A. (2000). The First Fossil Hunters: Paleontology in Greek and Roman Times. Princeton University Press, Princeton, New Jersey. ISBN 978-0-691-05863-4.
22. Witton, Mark (April 2016). "Why Protoceratops almost certainly wasn't the inspiration for the griffin legend". Archived from the original on 17 March 2020.

External links

• A video of Protoceratops vs Velociraptor