Nedoceratops


Nedoceratops is a controversial genus of ceratopsid herbivorous dinosaur from the Late Cretaceous period Lance Formation of North America. It is known only from a single skull discovered in Wyoming. Its status is the subject of ongoing debate among paleontologists: some authors consider Nedoceratops a valid, distinct taxon, while others consider it to represent an ontogenetic stage of Triceratops and thus a synonym.

History of discovery

The paper that described Nedoceratops was originally part of O. C. Marsh's magnum opus, his Ceratopsidae monograph. Unfortunately, Marsh died before the work was completed, and John Bell Hatcher endeavored to complete the Triceratops section. However, he died of typhus in 1904 at the age of 42, leaving the paper still incomplete. It fell to Richard Swann Lull to complete the monograph in 1905, publishing Hatcher's description of a skull separately and giving it the name Diceratops hatcheri; Diceratops means "two horned face."
Since the Diceratops paper had been written by Hatcher, and Lull had only contributed the name and published the paper after Hatcher's death, Lull was not quite as convinced of the distinctiveness of Diceratops, thinking it primarily pathological. By 1933, Lull had had second thoughts about Diceratops being a distinct genus and he put it in a subgenus of Triceratops: Triceratops hatcheri, including T. obtusus; largely attributing its differences to being that of an aged individual.
Because the Diceratops name was already in use for a hymenopteran, Andrey Sergeevich Ukrainsky gave the animal its current name Nedoceratops in 2007. Unaware that Ukrainsky had already renamed the animal, Octávio Mateus coined another new name for it in 2008, Diceratus. Diceratus is thus considered a junior synonym of Nedoceratops.
Nedoceratops means "insufficient horned face". The "nedo" is the Russian prefix meaning "insufficient". The suffix common among ceratopsians, "ceratops", means "horned face". It was named in reference to its lack of a nasal horn.

Description

The nearly complete skull known as "USNM 2412" is the only fossil of Nedoceratops. Like Hatcher's other Triceratops skulls, it was found in eastern Wyoming, i.c. in 1891 in Niobrara County near Lightning Creek. Superficially, it resembles that of Triceratops, but on closer examination, it differs: there is just a rounded stump where the nasal horn should be and the occipital horns stand almost vertically. Compared to other Triceratops skulls, it is slightly larger than average, but its face is rather short. There also are large holes in the frill, unlike other Triceratops skulls known. Some of these may be pathological, others seem to be genetic. Lull suggested that the holes in the frill were the result of "accidental gorings" from another dinosaur. However, this issue is still up for debate.
The following traits of USNM 2412 have been tentatively proposed by Andrew Farke as distinguishing Nedoceratops from Triceratops: It lacks a recognizable nasal horn.
It has almost vertical postorbital horncores. It has small parietal fenestrae.

Classification

The type species is Nedoceratops hatcheri. Nedoceratops belonged to the Ceratopsia, a group of herbivorous dinosaurs with parrot-like beaks which thrived in North America and Asia during the Cretaceous Period, which ended roughly 66 million years ago. All ceratopsians became extinct at the end of this era.
Several authors have suggested that Nedoceratops may be directly ancestral to Triceratops, or perhaps its nearest relative. An ongoing debate concerns the status of Triceratops, Torosaurus, and Nedoceratops. In a series of publications, John B. Scannella and John R. Horner and claimed that the USNM 2412 skull belonged to a "young adult" Triceratops. Evidence for this hypothesis included the shapes of the epoccipital and squamosal bones, and a neck frill that had "incipient" openings. The authors were of the opinion that the nasal horn of the USNM 2412 skull could have been lost when the animal was alive or when it became fossilized., and that all three "genera" actually represent different aged individuals of Triceratops. In 2012, Farke proposed a counter argument, and suggested that the bone surface texture and shape of the horns of Nedoceratops indicate an "old adult". A follow-up study by Leonardo Maiorino and colleagues in 2013 using morphometrics found support for Triceratops and Torosaurus being distinct, valid taxa, with Nedoceratops occupying variable positions with respect to the other two but generally outside the range of variation, concluding that "the size of USNM 2412 is a plausible intermediate, but the shape is not."
However, it has also been noted that many of the features that separate Nedoceratops from Triceratops, and specifically Triceratops horridus, may be the result of pathology and/or deformation of the skull after burial. Two of the features that have been used to diagnose Nedoceratops- the position of the squamosal, and the upright brow horns- are seen on one side of the skull, but not the other. This- along with the fact that the entire skull can be seen to be twisted when seen head-on- had been used to argue that these features result from post-mortem distortion of the fossil, rather than reflecting the anatomy of the animal during life. Furthermore, the presence of numerous holes in the frill suggests pathology due to injury or illness, the 'parietal fenestrae' may therefore be an artifact. Unfortunately, the frill is only preserved on one side, which makes it difficult to test this hypothesis. The one unquestionable difference between Nedoceratops and fossils referred to Triceratops horridus is the remarkably short nasal horn, although the horns of ceratopsids show marked intraspecific and ontogenetic variation. In most features- the short, saddle-shape frill, the s-shaped snout- the animal closely resembles Triceratops horridus.