Sarahsaurus
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''Sarahsaurus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of basal sauropodomorph
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
which lived during the Early
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
period in what is now northeastern
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
, United States.


Discovery and naming

All specimens of ''Sarahsaurus'' were collected from the
Lower Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma&nb ...
Kayenta Formation 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 is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
, specimen number TMM 43646-2). In addition, a partial skeleton (specimen number TMM 43646-3) as well as a nearly complete
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
(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 January 13, 1956) is a German-born American paleontologist who is Senior Scientist and Curator of Vertebrate Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. He receiv ...
and Robert R. Reisz in
2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrates ...
; the
type species In 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 species that contains the biological type specim ...
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 and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
; the other three are '' Anchisaurus'' and ''
Ammosaurus ''Anchisaurus'' is a genus of basal sauropodomorph dinosaur. It lived during the Early Jurassic Period, and its fossils have been found in the red sandstone of the Portland Formation, Northeastern United States, which was deposited from the ...
'' 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 at Long Island ...
, and ''
Seitaad ''Seitaad'' is a genus of sauropodomorph dinosaur which lived during the Early Jurassic period in what is now southern Utah, United States. ''Seitaad'' is known from an articulated partial postcranial holotype skeleton referred to as UMNH ...
'' of the later Navajo Sandstone of Early Jurassic Utah; a genus of basal sauropodmorph from Early Jurasssic deposits around
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
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 ...
s – giant four-legged
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
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 organism moves by means of its two rear 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'' 'double' ...
(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. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For exampl ...
, 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 o ...
invading the bone (). The scapula (shoulder blade) was hourglass-shaped. In the holotype specimen, the scapula and coracoid 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 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 blood vessels from injury. Sha ...
(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 the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal l ...
, 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 (; ) 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 and ulna, and consists of three sections. The humeral upper extremity consists of a r ...
(upper arm bone) was more than half the length of the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
(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. The femur was straight and elliptical in cross-section. The
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
(shin bone) reached 84% of the length of the femur.


Classification

In a
cladistic Cladistics (; ) 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 typically shared derived char ...
analysis, presented by Apaldetti and colleagues in November 2011, ''Sarahsaurus'' was found to be most closely related to ''
Ignavusaurus ''Ignavusaurus'' is a genus of basal sauropodomorph dinosaur that lived during the Early Jurassic in what is now Lesotho. Its fossils were found in the Upper Elliot Formation which is probably Hettangian in age (around 200 million years ago). ...
'' within
Massopoda Massopoda is a clade of sauropodomorph dinosaurs which lived during the Late Triassic to Late Cretaceous epochs. It was named by paleontologist Adam M. Yates of the University of the Witwatersrand in 2007. Massopoda is a stem-based taxon, define ...
. Their group was found to be intermediate between plateosaurids and massospondylids, being more
derived Derive may refer to: *Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguation ...
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 and
Pliensbachian The Pliensbachian is an age (geology), age of the geologic timescale and stage (stratigraphy), stage in the stratigraphic column. It is part of the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series an ...
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 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
'', ''
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.G ...
'', ''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 ''Saharasaurus'' included hybodont sharks, indeterminate
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
,
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
, salamanders, the frog '' Prosalirus'', the caecilian '' Eocaecilia'', the turtle ''
Kayentachelys ''Kayentachelys'' ("Kayenta turtle") is an extinct genus of turtle known only from the "silty facies" of the Lower Jurassic Kayenta Formation in northeastern Arizona on the lands of the Navajo Nation. History of discovery and significance Th ...
'', a sphenodontian reptile, various lizards, and the pterosaur '' Rhamphinion''. Also present were the
synapsids Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes rept ...
'' Dinnebitodon'', '' Kayentatherium'', '' Oligokyphus'', morganucodontids, the possible early true mammal ''Dinnetherium'', and a haramiyid mammal. Several early
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
were present including '' Calsoyasuchus'', ''
Eopneumatosuchus ''Eopneumatosuchus'' is an extinct genus of basal crocodyliform. Fossils have been found from two localities within the Kayenta Formation of Arizona. Both localities are around 20 miles southeast of the Grand Canyon and in close proximity to o ...
'', '' Kayentasuchus'', and ''
Protosuchus ''Protosuchus'' is an extinct genus of carnivorous crocodylomorph from the Early Jurassic. The name ''Protosuchus'' means "first crocodile", and is among the earliest animals that resemble crocodilians. ''Protosuchus'' was about in length an ...
''.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 is ...
s and the tracks of
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s, 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 of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...
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 Early Jurassic dinosaurs of North America Massopoda Fossil taxa described in 2011 Taxa named by Hans-Dieter Sues Paleontology in Arizona Kayenta Formation