Saltasaurus loricatus
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''Saltasaurus'' (which means "lizard from
Salta Salta () is the capital and largest city in the Argentine province of the same name. With a population of 618,375 according to the 2010 census, it is also the 7th most-populous city in Argentina. The city serves as the cultural and economic ce ...
") 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 saltasaurid
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 ...
of the Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
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 ...
of
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 ...
. Small among sauropods, though still heavy by the standards of modern creatures, ''Saltasaurus'' was characterized by a short neck and stubby limbs. It was the first genus of sauropod known to possess armour of bony plates embedded in its skin. Such small bony plates, called
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amphi ...
, have since been found on other
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 ...
ns.


Discovery

The fossils of ''Saltasaurus'' were excavated by
José Bonaparte José Fernando Bonaparte (14 June 1928 – 18 February 2020) was an Argentine paleontologist who discovered a plethora of South American dinosaurs and mentored a new generation of Argentine paleontologists . One of the best-known Argentine paleo ...
, Martín Vince and Juan C. Leal between 1975 and 1977 at the Estancia "El Brete". The find was in 1977 reported in the scientific literature. ''Saltasaurus'' was named and described by Bonaparte and Jaime E. Powell in 1980. 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 specime ...
is ''Saltasaurus loricatus''. Its generic name is derived from Salta Province, the region of north-west Argentina where the first fossils were recovered. The specific name means "protected by small armoured plates" in
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
. 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 sever ...
, PVL 4017-92, was found in a layer of the Lecho Formation dating from the early
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...
stage of 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'', th ...
period, about seventy million years old. It consists of a sacrum connected to two ilia. Under the inventory number PVL 4017 over two hundred additional fossils have been catalogued. These include rear skull elements, teeth,
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
e of the neck, back, hip and tail, parts of the shoulder girdle and the pelvis, and limb bones — plus various pieces of armour. These bones represent a minimum of five individuals, two adults and three juveniles or subadults.Powell, J.E., 1992, "Osteología de ''Saltasaurus loricatus'' (Sauropoda Titanosauridae) del Cretácico Superior del noroeste Argentino" In: Sanz, J., Buscalioni, A. (Eds.), ''Los dinosaurios y su entorno biótico: Actas del Segundo Curso de Paleontología in Cuenca'', pp. 165-230 Currently the only recognised
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of ''Saltasaurus'' is ''S. loricatus''. A ''S. robustus'' and a ''S. australis'' have been suggested but these are now considered to belong to a separate
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 ...
, '' Neuquensaurus''. Earlier, armour plates from the area had been named as '' Loricosaurus'' by Friedrich von Huene who assumed them to be from an armoured ankylosaurian. It has been suggested these plates were in fact from ''Saltasaurus''.


Description

''Saltasaurus'' is very small compared to most other members of the Sauropoda. Powell estimated the adult length at six metres. In 2010,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
estimated the maximum length at 8.5 metres (28 ft), the weight at 2.5 tonnes (2.8 short tons). The teeth of ''Saltasaurus'' were cylindrical, with spatulate points. ''Saltasaurus'' had a relatively short neck with shortened neck vertebrae. The vertebrae from the middle part of its tail had elongated
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different ...
. ''Saltasaurus'' had vertebral lateral fossae, pleurocoels, that resembled shallow depressions. Fossae that similarly resemble shallow depressions are known from ''
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 ...
'', '' Alamosaurus'', ''
Aeolosaurus ''Aeolosaurus'' (; "Aeolus' lizard") is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now South America. Like most sauropods, it would have been a quadrupedal herbivore with a long neck and tail. ''Aeolos ...
'', and '' Gondwanatitan''. ''Venenosaurus'' also had depression-like fossae, but its pleurocoels penetrated deeper into the vertebrae, were divided into two chambers, and extend farther into the vertebral columns. In ''Saltasaurus'', the vertebral bone was generally
cancellous A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, an ...
and there were larger air chambers present as well. The limbs were short and stubby with especially short hands and feet. ''Saltasaurus'' had more robust radii than ''
Venenosaurus ''Venenosaurus'' ( ) was a sauropod dinosaur. The name literally means "poison lizard", and it was named so after the Poison Strip Member of the Cedar Mountain Formation in Utah, United States, where the fossils were discovered by a Denver Museum ...
''. The belly was extremely wide. The osteoderms came in two types. There were larger oval plates with a length of up to twelve centimetres. These were keeled or spiked and perhaps were ordered in longitudinal rows along the back. The second type consists of small ossicles, rounded or pentagonal, about seven millimetres in diameter, that formed a continuous armour between the plates. A study in 2010 concluded that the larger plates had cancellous bone but that the ossicles had a denser bone tissue.Ignacio A. Cerda and Jaime E. Powell, 2010, "Dermal Armor Histology of ''Saltasaurus loricatus'', an Upper Cretaceous Sauropod Dinosaur from Northwest Argentina", ''Acta Palaeontologica Polonica'', 55(3): 389-398


Palaeobiology

Like all sauropods, ''Saltasaurus'' was
herbivorous 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 ...
. Because of its barrel-like rump, shaped like a
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
, Powell suggested that ''Saltasaurus'' was aquatic. Despite its small stature, ''Saltasaurus'' was still graviportal like other sauropods, meaning it could not run because its hindlimbs had to be held straight at the load-bearing phase of their walking cycle. Powell assumed adult individuals were protected against predators by their body armour, while juveniles were protected by the herd as a whole. In the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
Period, sauropods in North America were no longer the dominant group of herbivorous dinosaurs, with the ornithopod and ceratopsian dinosaurs, such as '' Edmontosaurus'' and ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivorous chasmosaurine ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous period, about 68 million years ago in what is now North America. It is one ...
'', becoming the most abundant (this being most evident by the Late Cretaceous epoch). However, on other landmasses such as
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 sout ...
and
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
(which were island
continents A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions are commonly regarded as continents. Ordered from largest in area to smallest, these seven ...
much like modern Australia) sauropods, in particular the
titanosaur 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 ...
s, continued to be the dominant herbivores. ''Saltasaurus'' was one such titanosaur sauropod, and lived around 70 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago) ...
. When it was first discovered, in 1975, it forced
palaeontologist 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 ...
s to reconsider some assumptions about sauropods as ''Saltasaurus'' possessed crocodile-like armour (osteoderms) 10 to 12 centimetres (4 to 5 in) in diameter. Previously, it had been assumed that size alone was sufficient defence for the massive sauropods. Since then, palaeontologists have investigated the possibility that other sauropods may also have had armour; for example, '' Laplatasaurus''. A new discovery, from another formation, may shed light on the nesting habits of ''Saltasaurus''. A large titanosaurid
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materi ...
ing ground was discovered in Auca Mahuevo, in
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
,
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 ...
(another titanosaur nesting site has reportedly been discovered in
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
). Several hundred female saltasaurines dug holes with their back feet, laid eggs in clutches averaging around 25 eggs each, and buried the nests under dirt and vegetation. The small
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s, about 11–12 cm (4–5 in) in
diameter In geometry, a diameter of a circle is any straight line segment that passes through the center of the circle and whose endpoints lie on the circle. It can also be defined as the longest chord of the circle. Both definitions are also valid fo ...
, contained
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
ised embryos, complete with
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different de ...
impressions showing a mosaic armour of small bead-like scales. The armour pattern resembled that of ''Saltasaurus''.


Footnotes


References

* Coria, R.A. and Chiappe, L.M. 200
Embryonic Skin From Late Cretaceous Sauropods (Dinosauria) of Auca Mahuevo, Patagonia, Argentina.
Journal of Paleontology v81(6):1528-1532 * Tidwell, V., Carpenter, K. & Meyer, S. 2001. New Titanosauriform (Sauropoda) from the Poison Strip Member of the Cedar Mountain Formation (Lower Cretaceous), Utah. In: Mesozoic Vertebrate Life. D. H. Tanke & K. Carpenter (eds.). Indiana University Press, Eds. D.H. Tanke & K. Carpenter. Indiana University Press. 139–165.


Further reading

* ''Walking on Eggs: The Astonishing Discovery of Thousands of Dinosaur Eggs in the Badlands of Patagonia'', by Luis Chiappe and Lowell Dingus. June 19, 2001, Scribner. .


External links



by M. Alan Kazlev, from Palæos.
The late Cretaceous nesting grounds of Patagonia
by Luis V. Rey, from his art gallery. {{Taxonbar, from=Q131111 Saltasaurids Late Cretaceous dinosaurs of South America Fossil taxa described in 1980 Taxa named by José Bonaparte Taxa named by Jaime Powell Lecho Formation Allen Formation Cretaceous Argentina