Massospondylus

Massospondylus (/ˌmæsɵˈspɒndɨləs/ mas-oh-SPON-di-ləs; from Greek, μάσσων (massōn, "longer") and σπόνδυλος (spondylos, "vertebra")) is a genus of prosauropod 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 found 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. Prosauropod 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 prosauropod 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 prosauropods 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.

Description
Massospondylus was a mid-sized prosauropod that was around 4 metres (13 ft) in length[1][2][3] and weighed approximately 135 kilograms (300 lb),[3] although a few sources have estimated its length at up to 6 metres (20 ft).[4][5] It was a typical prosauropod, with a slender body, a long neck and a proportionally very small head. The vertebral column was composed of nine cervical (neck) vertebrae, 13 dorsal (back) vertebrae, three sacral (hip) vertebrae, and at least 40 caudal (tail) vertebrae. The pubis faced forward, as with most saurischians. It had a slighter build than that of Plateosaurus, an otherwise similar prosauropod dinosaur.[6] The neck was proportionally longer than in most other prosauropods, with the foremost cervicals being four times the length of their width.[7] The forelimbs were only half the length of the hindlimbs[8] but quite powerful, as indicated by the broad upper end of the humerus that provided attachment areas for a large arm musculature.[7] Like Plateosaurus, it had five digits on each hand and foot. The hand was short and wide, with a large sickle shaped thumb claw used for feeding or defense against predators. The thumb was the longest finger in the hand, while the fourth and fifth digits were tiny, giving the forepaws a lopsided look.

Cranial anatomy
The small head of Massospondylus was approximately half the length of the femur. Numerous openings, or fenestrae, in the skull reduced its weight and provided space for muscle attachment and sensory organs. These fenestrae were present in pairs, one on each side of the skull. At the front of the skull were two large, elliptical nares, which were roughly half the size of the orbits. The orbits were proportionally larger in Massospondylus than in related genera, such as Plateosaurus. The antorbital fenestrae, smaller than those seen in Plateosaurus, were situated between the eyes and the nose. At the rear of the skull were two more pairs of temporal fenestrae: the lateral temporal fenestrae immediately behind the eye sockets, which were shaped like an inverted "T" in Massospondylus,[10] and the supratemporal fenestrae on top of the skull. Small fenestrae also penetrated each mandible.[6] The shape of the skull is traditionally restored as wider and shorter than that of Plateosaurus, but this appearance may be due just to differential crushing experienced by the various specimens.[6] Some features of the skull are variable between individuals; for example, the thickness of the upper border of the orbit and the height of the posterior maxilla. These differences may be due to sexual dimorphism[11] or individual variation.

Tooth count is variable between individuals and increases with skull size.[11] The premaxilla shows the constant number of four teeth per side in all known skulls; however, in the maxilla, the tooth count ranges from 14 to 22. There are 26 teeth in each side of the lower jaw in the largest known skull.[10][11] The height of the teeth crowns decreases from front to back in the upper jaw, but was more or less constant in the lower jaw.[10][11] The lack of pronounced tooth wear and the variable height of the crowns suggests that the teeth were replaced by succeeding new ones in relatively short time intervals.[11] Notably, there was variation in the tooth morphology based on the position of the teeth in the jaw.[11] The heterodonty present in Massospondylus is greater than that present in Plateosaurus, although unsurprisingly not as pronounced as the specialization of teeth in Heterodontosaurus.[13] Teeth closer to the front of the snout had round cross-sections and tapered to points, unlike the back teeth, which were spatulate and had oval cross-sections.[6][13]

As with other prosauropods, it has been proposed that Massospondylus had cheeks. This theory was proposed because there are a few large holes for blood vessels on the surfaces of the jaw bones, unlike the numerous small holes present on the jaws of cheekless reptiles. The cheeks would have prevented food from spilling out when Massospondylus ate.[6] Crompton and Attridge (1986) described skulls of Massospondylus as possessing pronounced overbites and suggested the presence of a horny beak on the tip of the lower jaw to make up the difference in length and account for tooth wear on the teeth at the tip of the snout.[14] However, the difference in length may be a misinterpretation based on crushing in a top–bottom plane, and the possession of a beak is considered unlikely in recent studies.

Discovery
The first fossils of Massospondylus were described by paleontologist Sir Richard Owen in 1854.[16] Originally, Owen did not recognize these finds as those of a dinosaur; instead he attributed them to "large, extinct, carnivorous reptiles"[9] that were related to today's lizards, chameleons and iguanas.[16] 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.[16][17][18] 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.[19] 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 lead to the scientific name that has been derived from the Greek terms masson/μάσσων 'longer' and spondylos/σπόνδυλος 'vertebra'.[16][17] 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.[17] On May 10, 1941, the Hunterian Museum was demolished by a German bomb, destroying all the fossils; only casts remain.[17]

Possible Massospondylus remains have been found in the Upper Elliot Formation, the Clarens Formation, and the Bushveld Sandstone of South Africa and Lesotho; the Forest Sandstone and the Upper Karroo Sandstone of Zimbabwe; and the Kayenta Formation of Arizona. These remains consist of at least 80 partial skeletons and four skulls, representing both juveniles and adults.[6] 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.[13] 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.[20] Recent restudy of African Massospondylus skulls, however, indicates that the Kayenta specimen does not pertain to Massospondylus.[10] This Kayenta skull and associated postcranial elements, identified collectively as MCZ 8893, has been recently referred to the newly described genus Sarahsaurus.[21]

Massospondylus had also been reported from Argentina,[6] 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.

Species
Many species have been named, although most are no longer considered valid. M. carinatus, named by Richard Owen, is the type species.[23] Other named species include: M. browni (Seeley, 1895),[24] M. harriesi (Broom 1911),[18] M. hislopi (Lydekker, 1890),[25] M. huenei (Cooper, 1981),[9] M. kaalae (Barrett 2009),[26] M. rawesi (Lydekker, 1890),[25] and M. schwarzi (Haughton, 1924).[27]

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.[6] M. browni is based on two cervical, two back, and three caudal vertebrae and miscellaneous hind limb elements.[24] M. harriesi is known from a well preserved forelimb and parts of a hindlimb.[18] M. schwarzi is known from an incomplete hind limb and sacrum.[27] M. hislopi and M. rawesi were named from fossils found in India.[25] 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.[28] M. hislopi was tentatively retained as an indeterminate sauropodomorph in the latest review,[6] but M. rawesi may be a theropod[29] or nondinosaur.[19] M. huenei is a combination derived by Cooper for Lufengosaurus huenei, as he considered Lufengosaurus and Massospondylus to be synonyms.[9] This synonymy is no longer accepted.[6]

M. kaalae was described in 2009 on the basis of a partial skull from the Upper Elliot Formation in Eastern Cape of South Africa. This species is known from the same time and region as some specimens of M. carinatus. It differs from the type species in the morphology of the braincase, as well as in several other characters of the skull such as the proportions of the premaxilla.

Dubios names
Several dinosaurs are often considered synonymous with Massospondylus. These include the above mentioned Leptospondylus and Pachyspondylus, as well as Aristosaurus, Dromicosaurus, Gryponyx taylori and Hortalotarsus, which are dubious names of little scientific value.[9] Hortalotarsus skirtopodus was named by Harry Seeley in 1894. According to Broom (1911), "Originally most of the skeleton was in the rock, and it was regarded by the farmers as the skeleton of a Bushman, but it is said to have been destroyed through fear that a Bushman skeleton in the rock might tend to weaken the religious belief of the rising generation." Some partial leg bones were salvaged. Together with Massospondylus carinatus, Owen named Leptospondylus capensis and Pachyspondylus orpenii. Aristosaurus erectus was named by E.C.N. van Hoepen in 1920 based on a nearly complete skeleton. Hoepen also named Dromicosaurus gracilis, which consisted of a partial skeleton. Gryponyx taylori was named by Sidney H. Haughton in 1924. It consists of hip bones. All of the above fossils come from the Hettangian or Sinemurian faunal stages of South Africa, where Massospondylus has been found.[30][31] Under the rules of zoological nomenclature, these names are junior synonyms. They were named after Massospondylus was described in a scientific paper; the name Massospondylus thus takes priority.

Ignavusaurus, known from a young specimen, may also be synonymous with Massospondylus.