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''Notocolossus'' 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
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 th ...
ian
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 ...
from 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 th ...
strata of
Mendoza Province Mendoza, officially Province of Mendoza, is a province of Argentina, in the western central part of the country in the Cuyo region. It borders San Juan to the north, La Pampa and Neuquén to the south, San Luis to the east, and the republic o ...
,
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 ...
.


Discovery and naming

A fossil of a large
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 ...
was discovered by the Argentine paleontologist Dr.
Bernardo Javier González Riga Bernardo Javier González Riga is an Argentinean palaeontologist; he is internationally recognised for his research on sauropod dinosaur evolution, and was awarded in 2019. He discovered in the Late Cretaceous strata of the Mendoza Province (Argen ...
in
Mendoza province Mendoza, officially Province of Mendoza, is a province of Argentina, in the western central part of the country in the Cuyo region. It borders San Juan to the north, La Pampa and Neuquén to the south, San Luis to the east, and the republic o ...
In 2016, 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 ...
''Notocolossus gonzalezparejasi'' was named and described by
Bernardo Javier González Riga Bernardo Javier González Riga is an Argentinean palaeontologist; he is internationally recognised for his research on sauropod dinosaur evolution, and was awarded in 2019. He discovered in the Late Cretaceous strata of the Mendoza Province (Argen ...
, Matthew Carl Lamanna, Leonardo Daniel Ortiz David,
Jorge Orlando Calvo Jorge Orlando Calvo (27 April 1961 – 10 January 2023) was an Argentine geologist and paleontologist working for "Centro de Investigaciones Paleontológicas Lago Barreales" (National University of Comahue). Dr. Jorge Orlando Calvo was born in C ...
and Juan P. Coria. The generic name combines the Greek words νότος, ''notos'', "south wind", and κολοσσός, ''kolossos'', "giant statue", in reference to the provenance from the Southern Hemisphere and the gigantic size of the animal. The specific name honours Jorge González Parejas, for having studied the dinosaur fossils of Mendoza province for two decades. 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 ...
, UNCUYO-LD 301, was found in a layer of the
Plottier Formation The Plottier Formation is a geologic formation that outcrops in the Argentine Patagonian provinces of Río Negro and Neuquén. It is the younger of two formations belonging to the Río Neuquén Subgroup within the Neuquén Group of the Neuquén ...
dating from the
Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by t ...
-
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
, about eighty-six million years old. It consists of a partial skeleton lacking the skull. It contains an anterior back vertebra, an anterior tail vertebra, a right humerus and the upper part of a left pubic bone. Although the bones have not been discovered in articulation, they were considered to represent a single individual because they were found in close association on a surface of eight by eight metres. A second partial skeleton lacking the skull was referred to ''Notocolossus'': specimen UNCUYO-LD 302 representing a smaller individual. It contains a series of five anterior tail vertebrae and a complete right foot connected to an astragalus. It was recovered on a distance of 403 metres to the holotype from a surface of five by five metres.


Description

The evidence suggests that ''Notocolossus'' was among the largest
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 th ...
s, and therefore one of the heaviest land animals, yet discovered. Although the incompleteness of the skeleton of the new sauropod has prevented scientists from making precise estimates of its size, its humerus, or upper arm bone, is in length, which is longer than that of any other titanosaur for which this bone is known, including other giants such as ''
Dreadnoughtus ''Dreadnoughtus'' is a genus of titanosaurian sauropod dinosaur containing a single species, ''Dreadnoughtus schrani''. ''D. schrani'' is known from two partial skeletons discovered in Upper Cretaceous (Campanian to Maastrichtian; approximately ...
'', ''
Futalognkosaurus ''Futalognkosaurus'' ( ; meaning "giant chief lizard") is a genus of titanosaurian dinosaur. The herbivorous ''Futalognkosaurus'' lived approximately 87 million years ago in the Portezuelo Formation, in what is now Argentina, of the Coniacian st ...
'', and ''
Paralititan ''Paralititan'' (meaning "tidal giant") was a giant titanosaurian sauropod dinosaur genus discovered in coastal deposits in the Upper Cretaceous Bahariya Formation of Egypt. It lived between 99.6 and 93.5 million years ago.Holtz, Thomas R. Jr. (2 ...
''.They also estimated the total femur length at ,by using an allometric equation. If, as is likely, the body proportions of ''Notocolossus'' were comparable to those of better preserved titanosaurs, the new dinosaur may have weighed in the range of , most likely . In 2019
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 mass of ''Notocolossus'' in the 45-55 tonne (49.6-60.6 short tons) range with the possibility that it might have been larger than that or approaching the mass of ''
Argentinosaurus ''Argentinosaurus'' is a genus of giant sauropod dinosaur that lived during the Late Cretaceous period in what is now Argentina. Although it is only known from fragmentary remains, ''Argentinosaurus'' is one of the largest known land animals o ...
,'' which was estimated at 65-75 tonnes (71.6-82.7 short tons). This claim though, is considered unlikely . In 2020 Molina-Perez and Larramendi estimated its length at and its weight at 40 tonnes (44 short tons). In 2016, several distinguishing traits of ''Notocolossus'' were determined. Nine of these were
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 ...
, unique derived qualities. At the anterior back vertebra, the depression between the front edge of the side process and the lower edge of the side process is divided by two accessory ridges, one of which runs vertically while the other crosses the first horizontally. On the front side of the neural spines of the front tail vertebrae, run vertical ridges fusing at their lower ends, above the level of the front joint processes, creating a V-shaped structure. The upper inner corner of the humerus is strongly expanded, extending far beyond the inner side of the shaft. The humerus has in general a strongly expanded upper edge, 2.9 wider than the shaft diameter. On the humerus shaft, below the scar for the '' musculus coracobrachialis'', runs a diagonal ridge, from the upper and outer side to the inner and lower side. The first metatarsal has an upper width exceeding its length. The third metatarsal is relatively short with 1.2 times the length of the first metatarsal. The upper toe phalanges are 50% wider at their tops than their respective metatarsals are long. The toe claws are reduced, rough and truncated. The foot or pes of ''Notocolossus'', a skeleton part that is not often preserved in sauropods, shows a unique build. It contains a compact, homogeneous
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
, thought to be adapted for bearing extraordinary weight. It also presents truncated
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...
s, characteristics otherwise unknown in the Sauropoda. This parallels the vertical position of the metacarpals and the finger reduction in the hand of other titanosaurs.


Phylogeny

''Notocolossus'' belongs to 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, ...
Titanosauria, a very diverse group of sauropods. In 2016, 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 recovered ''Notocolossus'' as a basal member of
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 ...
and a
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of ''
Dreadnoughtus ''Dreadnoughtus'' is a genus of titanosaurian sauropod dinosaur containing a single species, ''Dreadnoughtus schrani''. ''D. schrani'' is known from two partial skeletons discovered in Upper Cretaceous (Campanian to Maastrichtian; approximately ...
''. A 2017 study by Carballido and colleagues recovered it as the sister taxon of
Lognkosauria Lognkosauria is a clade of giant long-necked sauropod dinosaurs within the clade Titanosauria. It includes some of the largest and heaviest dinosaurs known. Description Lognkosaurians can be distinguished from other titanosaurs by the wide and u ...
and found it to be outside of Lithostrotia. In 2018, González Riga and colleagues found it to belong to
Lognkosauria Lognkosauria is a clade of giant long-necked sauropod dinosaurs within the clade Titanosauria. It includes some of the largest and heaviest dinosaurs known. Description Lognkosaurians can be distinguished from other titanosaurs by the wide and u ...
, in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with ''
Patagotitan ''Patagotitan'' is a genus of titanosaurian sauropod dinosaur from the Cerro Barcino Formation in Chubut Province, Patagonia, Argentina. The genus contains a single species known from multiple individuals: ''Patagotitan mayorum'', first announce ...
'' and ''
Puertasaurus ''Puertasaurus'' is a genus of sauropod dinosaur that lived in South America during the Late Cretaceous Period. It is known from a single specimen recovered from sedimentary rocks of the Cerro Fortaleza Formation in southwestern Patagonia, Ar ...
''. This analysis also found Longkosauria to belong to
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 ...
. The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
shows the position of ''Notocolossus'' in
Lognkosauria Lognkosauria is a clade of giant long-necked sauropod dinosaurs within the clade Titanosauria. It includes some of the largest and heaviest dinosaurs known. Description Lognkosaurians can be distinguished from other titanosaurs by the wide and u ...
according to Gonzalez Riga and colleagues, 2018.


Paleoecology

''Notocolossus'' was discovered in the
Plottier Formation The Plottier Formation is a geologic formation that outcrops in the Argentine Patagonian provinces of Río Negro and Neuquén. It is the younger of two formations belonging to the Río Neuquén Subgroup within the Neuquén Group of the Neuquén ...
of the
Neuquén Group The Neuquén Group is a group of geologic formations found in Argentina. Rocks in the Neuquén Group fall within the Cenomanian to early Campanian stages of the Late Cretaceous Period. It overlies the older Lohan Cura Formation and is itself over ...
. This formation is about thick and composed of light red
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too sm ...
s and thin bands of pink
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
. The Plottier Formation can be differentiated from the overlying
Portezuelo Formation The Portezuelo Formation is a geologic formation of Late Cretaceous ( Late Turonian to Early Coniacian) age, outcropping in the Mendoza, Río Negro and Neuquén provinces of Argentina.argillite :''"Argillite" may also refer to Argillite, Kentucky.'' Argillite () is a fine-grained sedimentary rock composed predominantly of indurated clay particles. Argillaceous rocks are basically lithified muds and oozes. They contain variable amounts ...
.
Tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s of the Plottier formation are poorly known. They are represented by
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
, a large
coelurosaur Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrann ...
, and several titanosaurs. These titanosaurs, in addition to ''Notocolossus'', include '' Petrobrasaurus'', ''"
Antarctosaurus ''Antarctosaurus'' (; meaning "southern lizard") is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now South America. The type species, ''Antarctosaurus wichmannianus'', and a second species, ''Antarctosau ...
"'' and two additional specimens, one of which appears to be an aeolosaurine. Additionally ''
Muyelensaurus ''Muyelensaurus'' (meaning "Muyelen lizard", after an indigenous name for the Colorado River in Argentina) is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous of Argentina. It was more slender than other titanosaurs. Fossils h ...
'' is sometimes stated to come from this formation, although it may actually be from the aforementioned
Portezuelo Formation The Portezuelo Formation is a geologic formation of Late Cretaceous ( Late Turonian to Early Coniacian) age, outcropping in the Mendoza, Río Negro and Neuquén provinces of Argentina.


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

{{Taxonbar, from1=Q23768183, from2=Q22226944 Lithostrotians Coniacian life Santonian life Late Cretaceous dinosaurs of South America Fossils of Argentina Cretaceous Argentina Neuquén Basin Fossil taxa described in 2016 Taxa named by Matt Lamanna Taxa named by Rodolfo Coria Taxa named by Jorge O. Calvo