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''Sarahsaurus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of basal
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
which lived during the Early
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period in what is now northeastern
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
, United States.


Discovery and naming

All specimens of ''Sarahsaurus'' were collected from the Lower Jurassic
Kayenta Formation The Kayenta Formation is a geological formation in the Glen Canyon Group that is spread across the Colorado Plateau area of the United States, including northern Arizona, northwest Colorado, Nevada, and Utah. Originally suggested as being Sinemur ...
near Gold Spring, Arizona. The genus is based on a nearly complete and articulated (with bones still connected to each other) skeleton with a fragmentary and disarticulated skull (
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, specimen number TMM 43646-2). In addition, a partial skeleton (specimen number TMM 43646-3) as well as a nearly complete
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
(specimen number MCZ 8893) was assigned to the genus. The latter specimen was originally referred to as ''Massospondylus'' sp. The complete skull is crushed and split horizontally, separating the skull roof from the palate; this split was caused by periodic swelling and shrinkage of the surrounding clay after
burial Burial, also known as interment or inhumation, is a method of final disposition whereby a dead body is placed into the ground, sometimes with objects. This is usually accomplished by excavating a pit or trench, placing the deceased and objec ...
. While the holotype individual was mature, the second skull indicates a less mature individual as indicated by open suture lines that separate skull bones of the braincase (the bones enclosing the brain). ''Sarahsaurus'' was first described by Timothy B. Rowe,
Hans-Dieter Sues Hans-Dieter Sues (born 1956) is a German-born American palaeontologist who is a Senior Research Geologist and Curator of Vertebrate Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. Career ...
and Robert R. Reisz in
2011 The year marked the start of a Arab Spring, series of protests and revolutions throughout the Arab world advocating for democracy, reform, and economic recovery, later leading to the depositions of world leaders in Tunisia, Egypt, and Yemen ...
; the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
is ''Sarahsaurus aurifontanalis''. The generic name honours Sarah (Mrs. Ernest) Butler, an Austin philanthropist. She helped fund the "Dino Pit" exhibit at the Austin Nature and Science Center, which Rowe helped create; he was quoted as saying "I told arahif she really raised a million dollars to build the Dino Pit, I'd name a dinosaur after her.". The specific name is derived from ''aurum'' (Latin), "gold", and ''fontanalis'' (Latin), "of the spring" in reference to Gold Spring, Arizona, where the holotype was found. ''Sarahsaurus'' is the fourth basal sauropodomorph dinosaur to have been officially identified in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
; the other three are '' Anchisaurus'' and '' Ammosaurus'' from the Early Jurassic of the
Connecticut River Valley The Connecticut River is the longest river in the New England region of the United States, flowing roughly southward for through four states. It rises 300 yards (270 m) south of the U.S. border with Quebec, Canada, and discharges into Long Isl ...
, and '' Seitaad'' of the later Navajo Sandstone of Early Jurassic Utah; a genus of basal sauropodmorph from Early Jurassic deposits around
Nova Scotia Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
called '' Fendusaurus'' remains undescribed.Fedak TJ. Description and evolutionary significance of the sauropodomorph dinosaurs from the Early Jurassic (Hettangian) McCoy Brook Formation. Ph.D. Dissertation, Dalhousie University. 2006. It is thought to have appeared through a dispersal event that originated in South America and was separate from those of the other two sauropodomorphs. The animal is notable for possessing very large, powerful hands, possibly suggesting that it was an omnivore.


Description

''Sarahsaurus'' was a basal (or "primitive") member of the sauropodomorph lineage, which also included the more derived (or "advanced")
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s – giant four-legged
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s (plant-eaters) with long necks and tails. Basal sauropodomorphs such as ''Sarahsaurus'' already resemble sauropods in a number of aspects, including the elongated neck and robust build, but are reminiscent of the ancestral dinosaurs in others: ''Sarahsaurus'' was
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
(two-legged), with forelimbs that were much shorter than the hind limbs and equipped with powerful grasping hands. ''Sarahsaurus'' was mid-sized for a basal sauropodomorph, with its vertebral column measuring more than in length. Paleontologist Gregory Paul, in a 2016 popular book, estimated its length at and its body weight at .


Skull

The (skull opening for the nostril) was less than half the size of the orbit (eye socket). It was bordered by the nasal, premaxilla, and, to a small extent, the maxilla, the latter two forming the upper jaw. Between the premaxilla and maxilla there was a large opening, the subnarial foramen. The antorbital fossa, an additional skull opening seen in most dinosaurs that was situated between the external naris and orbit, was less than half the length of the orbit. Much of the upper margin of the orbit was formed by the of the ; this bone, however, did not contribute to the front margin of the at the rear end of the skull roof. Instead, this front margin was formed by the and , with the front part of the parietal being forked to connect with the latter. Between the parietal and the supraoccipital, which follows behind the parietal at the back of the skull, was another opening, the postparietal fenestra, which was large in ''Sarahsaurus''. A unique feature of the genus can be found on the underside of the braincase, where a shallow ridge spanned between the , bony downwards projections of the braincase that brace the latter against the . The lower jaw curved downwards towards its tip, and the jaw joint was located well below the level of the tooth row. There is no evidence for a beak. Each side of the upper jaw was lined with 20 teeth – four in the premaxilla, the front bone of the upper jaw, and 16 in the maxilla which followed behind. The dentary bone of the lower jaw likewise had 20 teeth. The teeth were moderately
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals wher ...
, i.e., tooth shape varied within the jaw, and sat slightly obliquely in the jaw, slightly overlapping each other. Their cutting edges were with up to 20 per tooth.


Postcranium

The spine comprised 10 cervical (neck) vertebrae; 14 dorsal (trunk) vertebrae; three sacral (hip) vertebrae; and ca. 50 caudal (tail) vertebrae. In contrast to the later
sauropods Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
, the sides of the dorsal vertebrae did not show deep cavities caused by
air sacs Air sacs are spaces within an organism where there is the constant presence of air. Among modern animals, birds possess the most air sacs (9–11), with their extinct dinosaurian relatives showing a great increase in the pneumatization (presence ...
invading the bone (). The scapula (shoulder blade) was hourglass-shaped. In the holotype specimen, the scapula and
coracoid A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
were unfused, probably connected only via cartilage; this might indicate that the individual was not yet fully mature. The holotype specimen includes the
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
(breastbone) and the (collarbone), which are rarely found with dinosaur skeletons. The clavicula is a rod-like bone that articulates with the lower end of the scapula in an immobile joint. It has been controversial if the left and right clavicles of sauropodomorphs were attached to each other at their ends, thus connecting the left and right shoulder girdles, or if they were pressed against their scapulae along their lengths. The orientation of the clavicle in ''Sarahsaurus'' confirms that the former was the case; the ends of both clavicles would probably have been connected to each other by a
ligament A ligament is a type of fibrous connective tissue in the body that connects bones to other bones. It also connects flight feathers to bones, in dinosaurs and birds. All 30,000 species of amniotes (land animals with internal bones) have liga ...
, though they were not fused. In side view, the scapula was tilted between 45° and 65° relative to the horizontal. This would have allowed the left and right coracoids to almost touch each other at the body midline, while the left and right sterna, which were attached to each other at the midline, would have been located directly behind the coracoids. The
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
(upper arm bone) was more than half the length of the
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
(upper thigh bone), and had a large crest on its front side for muscle attachment, the , which extended for half the length of the bone and had an S-shaped edge. In the hand, the (claw bone) of the first finger was the largest. In the hip, the ilium had two downwards-facing bony projections, the ischial and pubic peduncle, which connected to the two lower hip bones, the ischium and the pubis, respectively, as in other dinosaurs. In ''Sarahsaurus'', the ischial peduncle was only half the length of the pubic peduncle. The (hip joint) was formed by all three hip bones, as in other dinosaurs, though in ''Sarahsaurus'' the ischium contributed less than half as much as the pubis. The pubis was unique in having two separate openings, the and the pubic foramen; the latter was lacking in most other
archosaurs Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
. The femur was straight and elliptical in cross-section. The
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
(shin bone) reached 84% of the length of the femur.


Classification

In a
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis, presented by Apaldetti and colleagues in November 2011, ''Sarahsaurus'' was found to be most closely related to '' Ignavusaurus'' within
Massopoda Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the Sauropoda, sauropods and their ancestral relatives. Sauropods generally grew to very large siz ...
. Their group was found to be intermediate between plateosaurids and massospondylids, being more derived than the former and more primitive than the latter. A 2018 cladistic analysis by Chapelle and Choiniere recovered ''Sarahsaurus'' and ''Ignavusaurus'' as massospondylids, mirroring the original assignment of ''Sarahsaurus'' to ''Massospondylus''.


Paleoecology


Age and habitat

A definitive radiometric dating of the Kayenta Formation has not yet been made, and the available stratigraphic correlation has been based on a combination of radiometric dates from vertebrate fossils, magnetostratigraphy and pollen evidence.J. M. Clark and D. E. Fastovsky. 1986. Vertebrate biostratigraphy of the Glen Canyon Group in northern Arizona. The Beginning of the Age of the Dinosaurs: Faunal change across the Triassic-Jurassic boundary, N. C. Fraser and H.-D. Sues (eds.), Cambridge University Press 285–301 It has been surmised that the Kayenta Formation was deposited during the
Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...
and Pliensbachian stages of the Early Jurassic period, approximately 199 to 182 million years ago.Padian, K (1997) Glen Canyon Group In: Encyclopedia of Dinosaurs, edited by Currie, P. J., and Padian, K., Academic Press. The Kayenta Formation is part of the Glen Canyon Group that includes formations not only in northern Arizona but also parts of southeastern Utah, western Colorado, and northwestern New Mexico. The formation was primarily deposited by rivers. During the Early Jurassic, the land that is now the Kayenta Formation experienced rainy summers and dry winters. By the Middle Jurassic period it was being encroached upon from the north by a sandy dune field that would become the Navajo Sandstone. The animals here were adapted to a seasonal climate and abundant water could be found in streams, ponds and lakes.


Paleofauna

''Sarahsaurus'' shared its
paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
with other dinosaurs, such as several theropods including ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 186 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserv ...
'', ''
Kayentavenator ''Kayentavenator'' (meaning " Kayenta hunter") is a genus of small carnivorous tetanuran dinosaur that lived during the Early Jurassic Period; fossils were recovered from the Kayenta Formation of northeastern Arizona and were described in 2010. ...
'', ''Coelophyisis'' ''kayentakatae'', and the "Shake N Bake" theropod, and the armored dinosaurs '' Scelidosaurus'' and '' Scutellosaurus''. The Kayenta Formation has yielded a small but growing assemblage of organisms. Vertebrates present in the Kayenta Formation at the time of ''Sarahsaurus'' included hybodont sharks, indeterminate
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
,
lungfish Lungfish are freshwater vertebrates belonging to the class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, inc ...
, salamanders, the frog '' Prosalirus'', the caecilian ''
Eocaecilia ''Eocaecilia'' is an extinct genus of gymnophionan amphibian from the early Jurassic Kayenta Formation of Arizona, United States. One species is described, ''Eocaecilia micropodia''. ''Eocaecilia'' shared some characteristics with salamanders a ...
'', the turtle '' Kayentachelys'', a sphenodontian reptile, various lizards, and the pterosaur '' Rhamphinion''. Also present were the
synapsids Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
'' Dinnebitodon'', ''
Kayentatherium ''Kayentatherium'' is an extinct genus of tritylodontid cynodonts that lived during the Early Jurassic. It is one of two tritylodonts from the Kayenta Formation of northern Arizona, United States. ''Kayentatherium'' means "Kayenta Beast", and ...
'', '' Oligokyphus'', morganucodontids, the possible early true mammal ''Dinnetherium'', and a haramiyid mammal. Several early crocodylomorphs were present including '' Calsoyasuchus'', '' Eopneumatosuchus'', '' Kayentasuchus'', and '' Protosuchus''.Jenkins, F. A., Jr., Crompton, A. W., and Downs, W. R. 1983. Mesozoic mammals from Arizona: new evidence in mammalian evolution. Science 222(4629):1233–1235.Jenkins, F. A., Jr. and Shubin, N. H. 1998. ''Prosalirus bitis'' and the anuran caudopelvic mechanism. Journal of Vertebrate Paleontology 18(3):495–510. Vertebrate trace fossils from this area included
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name ...
s and the tracks of therapsids, lizard-like animals, and dinosaurs, which provided evidence that these animals were also present.Hamblin, A. H., and Foster, J. R. 2000. Ancient animal footprints and traces in the Grand Staircase-Escalante National Monument, south-central Utah. Pages 557–568 ''in'' Sprinkel, D. A., Chidsey, T. C., Jr., and Anderson, P. B. editors. Geology of Utah's parks and monuments. Utah Geological Association, Salt Lake City, UT. Publication 28. Non-vertebrates in this ecosystem included microbial or "algal" limestone,Luttrell, P. R., and Morales, M. 1993. Bridging the gap across Moenkopi Wash: a lithostratigraphic correlation. Aspects of Mesozoic geology and paleontology of the Colorado Plateau. Pages 111–127 ''in'' Morales, M., editor. Museum of Northern Arizona, Flagstaff, AZ. Bulletin 59. freshwater bivalves, freshwater mussels and snails, and
ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...
s.Lucas, S. G., and Tanner L. H. 2007. Tetrapod biostratigraphy and biochronology of the Triassic-Jurassic transition on the southern Colorado Plateau, USA. Palaeogeography, Palaeoclimatology, Palaeoecology 244(1–4):242–256. The plant life known from this area included trees that became preserved as petrified wood.


References

{{Taxonbar, from=Q135107 Massopoda Fossil taxa described in 2011 Taxa named by Hans-Dieter Sues Kayenta Formation Dinosaur genera Sinemurian dinosaurs Pliensbachian dinosaurs Dinosaurs of the United States