''Notocolossus'' 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 ...

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 ...

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 b ...

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 ...

from late

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

strata of

Mendoza Province Mendoza (), officially the 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 r ...

Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...

Discovery and naming

A fossil of a large

was discovered by the Argentine paleontologist Dr. Bernardo Javier González Riga in

Mendoza province Mendoza (), officially the 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 r ...

. In 2016, 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 ...

''Notocolossus gonzalezparejasi'' was named and described by Bernardo Javier González Riga, Matthew Carl Lamanna, Leonardo Daniel Ortiz David,

Jorge Orlando Calvo Jorge Orlando Calvo (27 April 1961 – 10 January 2023)Southern Hemisphere and the gigantic size of the animal. The

specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...

honours Jorge González Parejas, for having studied the dinosaur fossils of Mendoza province for two decades. The

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 ...

, 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 ...

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 m ...

, 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

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. ...

''. 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. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...

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'' (meaning "lizard from Argentina") is a genus of giant sauropod dinosaur that lived during the Late Cretaceous Period (geology), period in what is now Argentina. Although it is only known from fragmentary remains, ''Argentinos ...

,'' 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). Notocolossus NT small

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 ...

, 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 (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...

, 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 sauropod Sauropoda (), whose members are known as sauropods (; ...

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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

Titanosauria, a very diverse group of sauropods. In 2016, 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 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 ...

of ''

''. A 2017 study by Carballido and colleagues recovered it as the sister taxon of Lognkosauria and found it to be outside of Lithostrotia. In 2018, González Riga and colleagues found it to belong to Lognkosauria, 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 tree ...

with ''

Patagotitan ''Patagotitan'' is a genus of titanosaurian sauropoda, sauropod dinosaur from the Cerro Barcino Formation in Chubut Province, Patagonia, Argentina. The genus contains a single species known from at least six young adult individuals, ''Patagotita ...

'' and '' Puertasaurus''. This analysis also found Longkosauria to belong to

. The following

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

shows the position of ''Notocolossus'' in Lognkosauria according to Gonzalez Riga and colleagues, 2018.

Paleoecology

''Notocolossus'' was discovered in the

of the

Neuquén Group The Neuquén Group is a geologic unit, 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 ...

. 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 and shale. Most of the particles of which the stone is composed are less than and are too small to ...

s and thin bands of pink

sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...

. The Plottier Formation can be differentiated from the overlying

Portezuelo Formation The Portezuelo Formation is a geologic formation of Late Cretaceous (Turonian, Late Turonian to Coniacian, Early Coniacian) age, outcropping in the Mendoza Province, Mendoza, Río Negro Province, Río Negro and Neuquén Province, Neuquén provinc ...

by the former's higher content of

argillite Argillite () is a fine-grained sedimentary rock composed predominantly of Friability, indurated clay particles. Argillaceous rocks are basically lithified muds and Pelagic sediment, oozes. They contain variable amounts of silt-sized particles. T ...

Tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...

s of the Plottier formation are poorly known. They are represented by

mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...

, 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, ty ...

, 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 (geology), Period of what is now South America. The type species, ''Antarctosaurus wichmannianus'', and a second speci ...

"'' and a specimen that appears to be an oldest member of the group

Aeolosaurini Aeolosaurini is an extinct clade of titanosaurian dinosaurs known from the Cretaceous period of Argentina and Brazil. Rodrigo M. Santucci and Antonio C. de Arruda-Campos (2011) in their cladistic analysis found ''Aeolosaurus'', '' Gondwanatitan'' ...

. '' Muyelensaurus'' is also discovered in this formation, although it was originally assigned to the

References

{{Taxonbar, from1=Q23768183, from2=Q22226944 Lithostrotia Dinosaur genera Santonian dinosaurs Taxa named by Matt Lamanna Taxa named by Rodolfo Coria Taxa named by Jorge O. Calvo Fossil taxa described in 2016 Dinosaurs of Argentina

Discovery and naming

Description

Phylogeny

Paleoecology

See also

References