Sarmientosaurus
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''Sarmientosaurus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
titanosaurian Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...
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 bo ...
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 ...
belonging to the
Titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...
. It lived in what is now
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
, specifically
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, during the
Upper Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
about 95 million years ago. 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 specimen ...
is ''Sarmientosaurus musacchioi''.


Discovery

In 1997,
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Rubén D.F. Martínez, at the ''Estancia Laguna Palacios'' of the Goicoechea family in
Chubut province Chubut ( es, Provincia del Chubut, ; cy, Talaith Chubut) is a province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa Cruz Province), the Andes ra ...
, discovered a sauropod skull. This proved to be connected to the first few cervical vertebrae. In 2016, the type species ''Sarmientosaurus musacchioi'' was named and described by Rubén Darío Francisco Martínez, Matthew Carl Lamanna,
Fernando Emilio Novas Fernando Emilio Novas (born 1960) is an Argentine paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural Sciences Museum in Buenos Aires, Argentina.
, Ryan C. Ridgely, Gabriel Andrés Casal, Javier E. Martínez, Javier R. Vita and Lawrence M. Witmer. The generic name refers to the town of
Sarmiento Sarmiento may refer to: Places Argentina *Sarmiento Department, San Juan, a subdivision of the San Juan Province *Sarmiento Department, Santiago del Estero, a subdivision of the Santiago del Estero Province *Sarmiento Department, Chubut, a subdivi ...
. The specific name honours the late Eduardo Musacchio, an educator at the ''
Universidad Nacional de la Patagonia San Juan Bosco The National University of the Patagonia San Juan Bosco (Spanish: ''Universidad Nacional de la Patagonia San Juan Bosco'') is a higher education establishment in Patagonia, southern Argentina. It was created on February 25, 1980, by law 22.713, as ...
''. The
Life Science Identifiers Life Science Identifiers are a way to name and locate pieces of information on the web. Essentially, an LSID is a unique identifier for some data, and the LSID protocol specifies a standard way to locate the data (as well as a standard way of descr ...
are 537DFE26-54EC-4978-AC86-E83A04FA74DE for the genus and C1090B8D-D051-44F3-B869-8B4A0C802176 for the species. The
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 several ...
, MDT-PV 2, was found in the upper Lower Member of the
Bajo Barreal Formation The Bajo Barreal Formation is a geological formation in the Golfo San Jorge Basin of Chubut and Santa Cruz, Argentina whose strata date back to the Middle Cenomanian to Late Turonian. The formation was first described by Teruggi & Rossetto in ...
, dating from the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
to
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
ages. It consists of an almost complete skull with lower jaws, articulated with the first seven vertebrae of the front neck. Several neck parts, among them the entire atlas and fourth neck vertebra, were too eroded to be salvaged. The specimen represents an elderly individual. It is one of the few titanosaurs for which skull material has been found. Uniquely, at the side of the neck an elongated structure was discovered that was identified as an ossified tendon. From the Bajo Barreal Formation another titanosaur sauropod is known, ''
Epachthosaurus ''Epachthosaurus'' (meaning "heavy lizard") was a genus of titanosaurian sauropod dinosaur from the Late Cretaceous. It was a basal lithostrotian titanosaur. Its fossils have been found in Central and Northern Patagonia in South America. Discov ...
''. It cannot be determined whether both taxa are identical because the material of their holotypes is not overlapping. However, the authors considered an identity as improbable because in their
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 both genera occupied different positions in the evolutionary tree. Also, fragmentary fossils, of postcranial bones that differ from those of ''Epachthosaurus'' and skull bones that are dissimilar to the ''Sarmientosaurus'' cranium, show that in any case several titanosaur species were present in the habitat.


Description


Size and distinguishing traits

''Sarmientosaurus'' has an estimated length of twelve metres and a weight of ten tonnes. The describing authors indicated nine unique distinguishing traits,
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
. The eye socket is large, equalling 40% of the length of the skull. The ascending branch of the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
has a complex connection with a top process of the
lacrimal bone The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
, being wedged between its outer side and inner side. The inner edge of the rear part of the ascending branch of the maxilla touches the rim of the bony nostril with a low but distinct ridge. The ascending branch of the quadratojugal has at its lower rear a tongue-shaped process overlapping the rear of the quadrate. In the braincase there are three separate exits for the '' nervus trigeminus''. An inner vein channel connecting the
infundibulum An infundibulum (Latin for ''funnel''; plural, ''infundibula'') is a funnel-shaped cavity or organ. Anatomy * Brain: the pituitary stalk, also known as the ''infundibulum'' and ''infundibular stalk'', is the connection between the hypothalamus an ...
with the brains stem, is lacking. The premaxillary teeth are positioned vertically, the maxillary teeth are inclining to the front and the dentary teeth are inclining to the rear. The middle neck vertebrae have strut-like, instead of plate-shaped, ridges between the front joint processes and the vertebral centrum. A long and thin ossified tendon is running along the low side of the series of neck vertebrae and neck ribs.


Skeleton

The skull has a length of forty-three centimetres. In top view the skull is more or less tongue-shaped. The
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
is small but the eye socket is exceptionally large. In side view the snout is flat with a concave upper profile and surface. The
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
touches the prefrontal. The
jugal bone The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anato ...
has an unusual L-shape with a very long front branch and an almost absent rear branch. The fifth
cranial nerve Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and ...
, the ''nervus trigeminus'', has extra exits for the branches towards the maxilla and the lower jaw, whereas other sauropods possess but single exit. The front of the lower jaw has an almost constant height. The praemaxilla bears four teeth, the maxilla eleven (right side) or twelve (left), and the dentary thirteen. The premaxillary teeth are positioned vertically, the maxillary teeth incline to the front while the teeth of the lower jaw incline to behind, a unique configuration. The build of the teeth is in-between the more spatulate form of basal sauropods and the pencil shape of derived species. The teeth are moderately elongated. They each have sharply-angled wear facets in a high and a low position which, together with their strange orientation, indicates some special, as yet not fully understood, way of cropping vegetation. The neck vertebrae are long and elongated. Their internal structure is camellated, i.e. with many small air spaces inside. The middle neck vertebrae have oval, narrow and deep pleurocoels in their sides, pneumatic excavations that nearly touch each other on the midline, separated by a narrow bone plate. The rear joint processes are uncommonly long, reaching beyond the edge of the vertebral body. The front joint processes are supported from below by struts with an oval cross-section, apparently formed by a perforation of the normally plate-shaped ridges in this position. The neck ribs are delicate, thin and rod-shaped.


Ossified tendon

Parallel to the ribs, on the outer side of the neck a cable-shaped structure was discovered with a constant diameter of three millimetres. It had an oval cross-section and a rough and striated surface. The structure originated directly behind the skull and continued over a length of several vertebrae, thus of some metres. It was interpreted by the describing authors as an ossified
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
. The alternative hypothesis that it might be a neck rib was rejected because the ribs are thicker and should have a different position. Such tendons might have been a continuation of the neck ribs, but again, its position did not confirm this. Instead, it was assumed to have been internal to some neck muscle. Such ossified tendons have never before been found in any fossil dinosaur but some extant bird groups such as the cranes show them, though they are relatively shorter, at most two vertebrae long. Possible muscles, where it could have been located, are the '' Musculus rectus capitis anterior ventralis'', the '' Musculus longus colli ventralis'' or the ''Musculi intertransversarii''. The internal structure of the tendon, with much reworked bone tissue, indicated a swift ossification at a young age.


Classification

Martínez and colleagues placed ''Sarmientosaurus'' in a basal position within the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Lithostrotia Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of ''Malawisaurus'' and ''Saltasaurus'' an ...
, above ''
Malawisaurus ''Malawisaurus'' (meaning "Malawi lizard") is an extinct genus of titanosaurian sauropod dinosaur. It is known from the Dinosaur Beds of northern Malawi, which probably date to the Aptian stage of the Early Cretaceous. The type species is ''M. di ...
'' in the evolutionary tree. However, in 2021, Stephen Poropat and colleagues instead identified it as part of the new clade Diamantinasauria, along with ''
Savannasaurus ''Savannasaurus'' is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Winton Formation of Queensland, Australia. It contains one species, ''Savannasaurus elliottorum'', named in 2016 by Stephen Poropat and colleagues. The holot ...
'' and ''
Diamantinasaurus ''Diamantinasaurus'' is a genus of titanosaurian sauropod from Australia that lived during the early Late Cretaceous, about 94 million years ago. The type species of the genus is ''D. matildae'', first described and named in 2009 in paleontology, ...
''.


Palaeobiology

It had very large eye sockets, meaning that it may have had better vision than other titanosaurs. Based on the ear and neck tendon, ''Sarmientosaurus'' most likely hung its head and neck down "like an enormous
Eeyore Eeyore ( ) is a fictional character in the ''Winnie-the-Pooh'' books by A. A. Milne. He is generally characterized as a pessimistic, gloomy, depressed, anhedonic, old grey stuffed donkey who is a friend of the title character, Winnie-th ...
". This posture implies that ''Sarmientosaurus'' may have eaten much lower-lying plants than other sauropods. The correlation between inner ear structure and head posture has been questioned in previous studies.


See also

*
2016 in paleontology Flora Plants Fungi Cnidarians Research * '' Yunnanoascus haikouensis'', previously thought to be a member of Ctenophora, is reinterpreted as a crown-group medusozoan by Han ''et al.'' (2016). * A study on the fossil corals from the Late Tri ...


References


External links

Videos accompanying research article:
Sarmientosaurus musacchioi, new titanosaurian sauropod dinosaur Sarmientosaurus skull & brain - titanosaurian sauropod from Argentina Sarmientosaurus skull - titanosaurian sauropod from Argentina
{{Taxonbar, from=Q23927079 Lithostrotians Late Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Fossil taxa described in 2016 Taxa named by Matt Lamanna