Massospondylus

Massospondylus (/ˌmæsoʊˈspɒndɪləs/ mas-oh-SPON-di-ləs; from Greek, μάσσων (massōn, 'longer') and σπόνδυλος (spondylos, 'vertebra')) is a genus of sauropodomorph dinosaur from the Early Jurassic Period (Hettangian to Pliensbachian ages, ca. 200–183 million years ago). It was described by Sir Richard Owen in 1854 from remains discovered in South Africa, and is thus one of the first dinosaurs to have been named. Fossils have since been found at other locations in South Africa, Lesotho, and Zimbabwe. Material from Arizona's Kayenta Formation, India, and Argentina has been assigned to this genus at various times, but the Arizonan and Argentinian material are now assigned to other genera. The type species is M. carinatus; seven other species have been named during the past 150 years, but only M. kaalae among these is still considered valid. Early sauropodomorphs systematics have undergone numerous revisions during the last several years, and many scientists disagree where exactly Massospondylus lies on the dinosaur evolutionary tree. The family name Massospondylidae was once coined for the genus, but because knowledge of early sauropodomorph relationships is in a state of flux, it is unclear which other dinosaurs—if any—belong in a natural grouping of massospondylids; several 2007 papers support the family's validity. Although Massospondylus was long depicted as quadrupedal, a 2007 study found it to be bipedal. It was probably a plant eater (herbivore), although it is speculated that the early sauropodomorphs may have been omnivorous. This animal, which was 4–6 metres (13–20 ft) long, had a long neck and tail, with a small head and slender body. On each of its forefeet, it bore a sharp thumb claw that was used in defense or feeding. Recent studies indicate that Massospondylus grew steadily throughout its lifespan, possessed air sacs similar to those of birds, and may have cared for its young. The first fossils of Massospondylus were described by paleontologist Sir Richard Owen in 1854. Originally, Owen did not recognize these finds as those of a dinosaur; instead he attributed them to 'large, extinct, carnivorous reptiles' that were related to today's lizards, chameleons and iguanas. This material, a collection of 56 bones, was found in 1853 by the government surveyor Joseph Millard Orpen in the Upper Elliot Formation at Harrismith, South Africa and was donated to the Hunterian Museum at the Royal College of Surgeons in London. Among the remains were vertebrae from the neck, back, and tail; a shoulder blade; a humerus; a partial pelvis; a femur; a tibia; and bones of the hands and feet. All these bones were found disarticulated, making it difficult to determine if all material belongs to a single species or not. However, Owen was able to distinguish three different types of caudal vertebrae, which he attributed to three different genera: Pachyspondylus, Leptospondylus and Massospondylus. Massospondylus was separated from the other two genera on the basis of its much longer caudal vertebrae, which also led to the scientific name that has been derived from the Greek terms masson/μάσσων 'longer' and spondylos/σπόνδυλος 'vertebra', explained by Owen as 'because the vertebrae are proportionally longer than those of the extinct Crocodile called Macrospondylus'. However, later it was shown that the putative caudal vertebrae of Massospondylus were actually cervical vertebrae and that all the material probably belongs only to a single species. On May 10, 1941, the Hunterian Museum was demolished by a German bomb, destroying all the fossils; only casts remain. Because the plaster casts of the lost type specimen fossils were not adequate to accurately diagnose a genus and species under modern taxonomic practices and for research purposes, Yates and Barrett (2010) designated BP/1/4934, a skull and a largely complete postcranial skeleton in the Bernard Price Institute for Palaeontological Research, as the neotype specimen. Massospondylus remains have been found in the Upper Elliot Formation, the Clarens Formation, and the Bushveld Sandstone of South Africa and Lesotho, as well as the Forest Sandstone of Zimbabwe. These remains consist of at least 80 partial skeletons and four skulls, representing both juveniles and adults. The report of Massospondylus from Arizona's Kayenta Formation is based on a skull described in 1985. The skull of the Kayenta specimen from Arizona is 25% larger than the largest skull from any African specimen. The Kayenta specimen possesses four teeth in the premaxilla and sixteen in the maxilla. Uniquely among dinosaurs, it also had tiny, one-millimetre-(0.04 in-) long palatal teeth. A 2004 restudy of African Massospondylus skulls, however, indicated that the Kayenta specimen did not pertain to Massospondylus. This Kayenta skull and associated postcranial elements, identified collectively as MCZ 8893, was referred to the new genus Sarahsaurus in 2010. Massospondylus had also been reported from Argentina, but this has been reassessed as a closely related but distinct genus. The fossils included several partial skeletons and at least one skull, found in the Lower Jurassic Canon del Colorado Formation of San Juan, Argentina. This material was named Adeopapposaurus in 2009. A specimen from South Africa previously assigned to Massospondylus, BP/1/4779, became the holotype of the new genus and species Ngwevu intloko in 2019. Many species have been named, although most are no longer considered valid. M. carinatus, named by Richard Owen, is the type species. Other named species include: M. browni Seeley, 1895, M. harriesi Broom 1911, M. hislopi Lydekker, 1890, M. huenei Cooper, 1981, M. kaalae Barrett 2009, M. rawesi Lydekker, 1890, and M. schwarzi Haughton, 1924. M. browni, M. harriesi, and M. schwarzi were all found in the Upper Elliot Formation of Cape Province, South Africa. All three are based on fragmentary material, and were regarded as indeterminate in the most recent review. M. browni is based on two cervical, two back, and three caudal vertebrae and miscellaneous hind limb elements. M. harriesi is known from a well-preserved forelimb and parts of a hindlimb. M. schwarzi is known from an incomplete hind limb and sacrum. M. hislopi and M. rawesi were named from fossils found in India. M. hislopi is based on vertebrae from the Upper Triassic Maleri Formation of Andhra Pradesh, whereas M. rawesi is based on a tooth from the Upper Cretaceous Takli Formation of Maharashtra. M. hislopi was tentatively retained as an indeterminate sauropodomorph in the latest review, but M. rawesi may be a theropod or nondinosaur. M. huenei is a combination derived by Cooper for Lufengosaurus huenei, as he considered Lufengosaurus and Massospondylus to be synonyms. This synonymy is no longer accepted.