Nyasasaurus

Nyasasaurus (meaning "Lake Nyasa lizard") is an extinct genus of dinosauriform reptile from the Middle Triassic Manda Beds of Tanzania that may be the earliest known dinosaur. The type species N. parringtoni was first described in 1956 in the doctoral dissertation of English paleontologist Alan J. Charig, but it was not formally described until 2013. Previously, the oldest record of dinosaurs was from Argentina and dated back to the late Carnian stage, about 231.4 million years ago (Ma). Nyasasaurus comes from a deposit that dates back to the Anisian Stage, meaning that it predates other early dinosaurs by about 16 million years.

Description
The type specimen, NHMUK R6856, is a partial skeleton belonging to an individual estimated to have been two to three metres in length. It consists of a right humerus, three partial presacral vertebrae and three sacral vertebrae (of the back, above the location of the hip bones). A second specimen, SAM-PK-K10654 consisting of three cervical vertebrae (neck vertebrae) and two posterior presacral vertebrae, is also known. It was attributed to the same species as NHMUK R6856 because the dorsal or back vertebrae of the two specimens are nearly identical. However, the vertebral features that link NHMUK R6856 and SAM-PK-K10654, including a connection between two bony projections called the hyposphene and hypantrum, are also found in other Triassic archosaurs. Because these characteristics are not unique to the two specimens (in other words, they are not autapomorphies), they do not by themselves provide sufficient evidence for grouping NHMUK R6856 and SAM-PK-K10654 under the same species. The authors of the 2013 description of Nysasaurus used a second line of evidence, the similar positions of the two specimens on evolutionary trees, to justify their placement in the same species.[1]

Nesbitt et al. (2013) also mentioned the similarity between the presacral vertebrae of both specimens of N. parringtoni and those of "Teleocrater rhadinus" (another nomen nudum from Charigs' dissertation, based on NHMUK R6795 from Lifua Member). Additionally, the anterior cervical vertebra attributed to NHMUK R6795 is extremely elongated relative to that of the middle dorsal vertebrae with a low centrum to neural arch ratio and a significant displacement between the two sides of the articular facet of the centrum. However, it is probable that the limb bones and other elements included in NHMUK R6795 do not belong to the same individual. Therefore, it is possible that the vertebrae of "T. rhadinus" are also referable to N. parringtoni.[1]

An analysis of the interior structure of the humerus indicates that bone growth was rapid, with interwoven bone fibers, many channels for blood vessels that radiate in all directions, and few lines of arrested growth. This structure more closely matches that of the early dinosaur Megapnosaurus than it does of dinosaur ancestors, suggesting that Nyasasaurus was closer to the ancestry of dinosaurs than were other archosaurs at the time.

History of study
In the 1930s, the holotype of Nyasasaurus was collected in Parrington’s locality B36 from the Lifua Member of the Manda beds, Ruhuhu Basin near Lake Nyasa in southern Tanzania by Francis Rex Parrington. Other fossils from the same locality included those of cynodonts, dicynodonts, and rhynchosaurs. Most, including those of Nyasasaurus, consist only of fragments of bone. The remains were first described in English paleontologist Alan J. Charig's 1956 doctoral dissertation and referred to as "Specimen 50b".[2] In 1967 Charig published the name Nyasasaurus parringtoni, in a review of the Archosauria,[3] but without description so that it was commonly considered a nomen nudum; the dissertation also was never published. The generic name referred to Lake Nyasa and the specific name honouring Parrington. In 2013 a new description was published by Sterling Nesbitt, Paul Barrett, Sarah Werning and Christian Sidor, including the late Charig as posthumous co-author, ensuring the validity of the name Nyasasaurus parringtoni.[1] The generic name is occasionally misspelled as "Nyasaurus",[1] as by Theodore Elmer White in 1973.[4]

The referred specimen of Nyasasaurus, SAM-PK-K10654, was collected by G. M. Stockley in the early 1930s in the western portion of the Manda beds at Stockley’s locality B27.[5] This locality is listed as a locality from the "Upper Bone Bed" of the Manda beds (currently understood to be from the Lifua Member) by Haughton (1932). The specimen was collected under a single field number, S507, presumably from a small area. The specimen was probably associated as evidenced by the bone quality, color and surrounding matrix (dark gray to black carbonate). The consistent sizes of the remains indicate that they represent a single individual. Stockley’s locality B27 is located near the village of Gingama and it was probably the only specimen found at this locality, although a nearby locality B26, also listed as Gingama, produced cynodonts, lungfishes, amphibians, and a shark. Dicynodonts, cynodonts and archosaurs such as Asilisaurus were also found nearby in the Lifua Member.[1] The name ?Thecodontosaurus alophos was coined for this specimen by Haughton (1932).[6] Its holotype consists of three cervical vertebrae and two middle to posterior dorsal vertebrae that are poorly preserved as they are highly fractured and parts of the bone and bone surfaces are eroded. Originally, a comparison between ?T. alophos was made only with Coelophysis longicollis. Since then, the species has been largely ignored by all subsequent vertebrate workers and no formal diagnosis of the specimen was ever provided. Nesbitt et al. (2013) found the specimen to be not diagnostic because it does not have any autapomorphic features or a unique combination of character states. Therefore, they suggested to abandon the name ?T. alophos and to refer its specimen to N. parringtoni.