Bunostegos

Bunostegos ("knobbly [skull] roof"[1]) is an extinct genus of pareiasaur parareptile from the Late Permian of the Agadez Region in Niger. The type species, Bunostegos akokanensis, was named from the Moradi Formation in 2003.[2] It was a cow-sized animal with a distinctive skull that had large bony knobs, similar in form to those of other pareiasaurs but far larger. The species appears to have lived in a desert in the centre of the supercontinent of Pangaea.

Analysis of the limb bones (including the scapulocoracoid, humerus, radius, ulna, pelvis, and femur) was published in 2015, and revealed that Bunostegos walked upright on four limbs, with the body held above ground.[3] This new information directly suggests that it could be the first tetrapod with a fully erect gait.

Description
The animal has been described as about the size of a modern cow with a knobbly skull and bony plate armor on its back."[1] Its teeth show it to have been a plant eater. It lived in an isolated desert region of the supercontinent of Pangaea some 260 million years ago. Its home region appears to have supported a distinctive fauna, in contrast to the rest of the supercontinent, where species were broadly distributed. It is particularly notable for the large bony knobs on its head, bigger than any seen in other species of pareiasaur. In life they were probably skin-covered horns or ossicones similar to those of modern giraffes. They are thought not to have served a protective function but were probably purely ornamental, perhaps aiding recognition between or within particular species.[5]

Bunostegos may have been part of a relict population that clung on in central Pangaea, isolated from other more advanced species by the hyperarid conditions in which it lived. It is more closely related to older and more primitive pareiasaurs. The centre of the supercontinent appears to have been a very dry desert, which prevented population exchanges between the interior and exterior and kept Bunostegos in reproductive isolation.[5] Only a few million years later, however, Bunostegos and most of the other pareiasaurs died out in the Permian–Triassic extinction event of 252 million years ago.[1]

The evidence Bunostegos walked upright:
 * the shoulder part (scapulocoracoid) where the upper arm bone moved is oriented more downward and to the back than off to side as in other pareiasaurs, allowing a greater vertical movement.
 * the humerus (upper arm bone) does not fit into a spread out posture
 * the elbow joint allows a front to back movement as in an animal standing upright.

Near vertical hind limbs were usual for pareiasaurs, but Bunostegos was more advanced in having all four limbs vertical.

Discovery
Bunostegos akokanensis was named by paleontologists Christian A. Sidor, David C. Blackburn and Boubé Gado in 2003. Remains of Bunostegos were uncovered from the Moradi Formation near the town of Akokan in 2003 and 2006. The genus name means "knobby roof" in Greek as a reference to the bony knobs on its skull and the species name akokanensis references Akokan. Bunostegos is currently known from several skulls and postcranial remains. The holotype specimen MNN-MOR72, which served as the basis for the initial description of Bunostegos, is a weathered skull lacking the lower jaw. MNN-MOR86, a better-preserved skull also lacking the lower jaw, MNN-MOR28, a less deformed but heavily weathered skull, and MNN-MOR47, a partial skull preserving the palate and braincase, served as the basis for a 2013 description of the skull anatomy of Bunostegos.