Meraxes Gigas Ilustración Científica Realizada Por Carlos Papolio
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''Meraxes'' is a genus of large
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur from the Late Cretaceous
Huincul Formation The Huincul Formation is a geologic formation of Late Cretaceous (Cenomanian to Turonian, Early Turonian) age of the Neuquén Basin that outcrops in the Mendoza Province, Mendoza, Rio Negro Province, Río Negro and Neuquén Provinces of northern ...
of
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
,
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 ...
. The genus contains a single species, ''Meraxes gigas''.


Discovery and naming

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 ...
of ''Meraxes'', MMCh-PV 65, was discovered in 2012. Known from a nearly complete
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
,
pectoral Pectoral may refer to: * The chest region and anything relating to it. * Pectoral cross, a cross worn on the chest * a decorative, usually jeweled version of a gorget * Pectoral (Ancient Egypt), a type of jewelry worn in ancient Egypt * Pectora ...
and
pelvic The pelvis (: pelves or pelvises) is the lower part of an anatomical trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis or pelvic skeleton). ...
elements, partial
forelimb A forelimb or front limb is one of the paired articulated appendages ( limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used inst ...
s, complete
hindlimb A hindlimb or back limb is one of the paired articulated appendages ( limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-H ...
s, fragmentary
ribs The rib cage or thoracic cage is an endoskeletal enclosure in the thorax of most vertebrates that comprises the ribs, vertebral column and sternum, which protect the vital organs of the thoracic cavity, such as the heart, lungs and great vessels ...
and cervical and
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
vertebrae, a
sacrum The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
, and several complete
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into th ...
vertebrae, it represents the most complete carcharodontosaurid skeleton known from the Southern Hemisphere. The specimen was referred to as the "Campanas carcharodontosaurid" before its description as a new species in 2022. ''Meraxes gigas'' was described in 2022 by Canale et al. based on these remains. The generic name, "''Meraxes''", honors a female dragon ridden by Queen Rhaenys Targaryen in the
George R. R. Martin George Raymond Richard Martin (born George Raymond Martin; September 20, 1948) also known by the initials G.R.R.M. is an American author, television writer, and television producer. He is best known as the author of the unfinished series of Hi ...
novel series, ''
A Song of Ice and Fire ''A Song of Ice and Fire'' is a series of high fantasy novels by the American author George R. R. Martin. Martin began writing the first volume, ''A Game of Thrones'', in 1991, and published it in 1996. Martin, who originally envisioned the ser ...
''. The specific name, "''gigas''", means "giant" in
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
, in reference to its large size.


Description

''Meraxes'' is one of the largest theropods, weighing approximately . Henderson (2023) listed a body length estimate of , referencing Canale et al. (2022), but also estimated a body length of using the pelvic area. Its skull is long, similar to that of ''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaurs that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous, from 113 to 110 million years ago. Like most dinosaur genera, ...
'', which has a skull length of . The shapes and proportions of various bones, including the skull, scapula, metacarpals, ischial shaft, and foot, indicate that ''Meraxes'' and ''Acrocanthosaurus'' had similar proportions and body size. ''Meraxes'' possessed reduced forelimbs, an instance of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
that occurred independently in four different lineages:
Carcharodontosauridae Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosaurida ...
,
Abelisauridae Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are fou ...
,
Tyrannosauridae Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
, and
Alvarezsauridae Alvarezsauridae is a family of small, long-legged dinosaurs. Although originally thought to represent the earliest known flightless birds, they are now thought to be an early diverging branch of maniraptoran theropods. Alvarezsaurids were highly ...
. Additionally, the second toes possess an enlarged claw, almost twice as long as the claw on the fourth toe. Osteohistological analysis of the holotype suggests the individual could have been between 39 and 53 years old when it died, having reached skeletal maturity approximately 4 years prior to its death (between 35 and 49 years old), making it the longest-lived non-avian theropod currently known. ''Meraxes'' was determined to have grown to large size by extending its growth period ( hypermorphosis), rather than increasing its relative growth rate (
acceleration In mechanics, acceleration is the Rate (mathematics), rate of change of the velocity of an object with respect to time. Acceleration is one of several components of kinematics, the study of motion. Accelerations are Euclidean vector, vector ...
) through development as in ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
'', to which it was compared.


Classification

Canale et al. (2022) recovered ''Meraxes'' as the earliest diverging member of the tribe
Giganotosaurini Carcharodontosauridae (carcharodontosaurids; from the Greek language, Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carch ...
within the Carcharodontosauridae. The results of their
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
are displayed in the
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 ...
below:


Paleoenvironment

The fossil remains of ''Meraxes'' were recovered from the
Huincul Formation The Huincul Formation is a geologic formation of Late Cretaceous (Cenomanian to Turonian, Early Turonian) age of the Neuquén Basin that outcrops in the Mendoza Province, Mendoza, Rio Negro Province, Río Negro and Neuquén Provinces of northern ...
. A substantial number of taxa are known to have inhabited this paleoenvironment. Theropods from the formation include the paravian ''
Overoraptor ''Overoraptor'' (, meaning "piebald thief") is an extinct genus of paravian theropod of uncertain affinities from the Late Cretaceous Huincul Formation of Argentinian Patagonia. The genus contains a single species, ''O. chimentoi'', known fro ...
'', the elaphrosaurine ''
Huinculsaurus ''Huinculsaurus'' (meaning "Huincul Formation lizard") is a genus of Noasauridae, noasaurid dinosaur from the Huincul Formation in Neuquén Province, Argentina. The type species, type and only species is ''Huinculsaurus montesi''. The immature ho ...
'', the abelisaurs ''
Skorpiovenator ''Skorpiovenator'' ("scorpion hunter") is a genus of abelisaurid theropod dinosaur from the Late Cretaceous (Cenomanian to Turonian) Huincul Formation of Argentina. The Monotypic taxon, sole species of ''Skorpiovenator'', ''S. bustingorryi'', was ...
'', ''
Tralkasaurus ''Tralkasaurus'' (meaning "thunder lizard") is a genus of abelisaurid dinosaur from the Huincul Formation from Río Negro Province in Argentina. The type and only species is ''Tralkasaurus cuyi'', named in 2020 by Mauricio Cerroni and colleagues ...
'', and '' Ilokelesia'', the megaraptoran ''
Aoniraptor ''Aoniraptor'' is an extinct genus of megaraptoran theropod dinosaurs from the Late Cretaceous Huincul Formation of Argentina. The genus contains a single species, ''Aoniraptor libertatem'', known from a partial skeleton. Discovery and naming ...
'', and the fellow giant carcharodontosaurids ''Mapusaurus'' and ''
Taurovenator ''Taurovenator'', from Latin ''taurus'', meaning "bull", and ''venator'', meaning "hunter", is an extinct genus of large carcharodontosaurid theropod dinosaurs that lived in what is now Argentina during the Cenomanian age of the Late Cretaceous, ...
''. ''Meraxes'' was found in an older rock layer than either ''Mapusaurus'' and ''Taurovenator'', so it is unlikely they coexisted. The herbivores of the area are represented by the
rebbachisaurid Rebbachisauridae is a Family (biology), family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, North America, Europe and possibly Central Asia. Taxonomy In 1990 sauropod specialist Jack M ...
sauropods ''
Cathartesaura ''Cathartesaura'' is a genus of rebbachisaurid sauropod dinosaur hailing from the Late Cretaceous strata of the Huincul Formation, at the "La Buitrera" locality, in the Neuquén Basin of Río Negro Province, Argentina. The fossil remains, describ ...
'' and ''
Limaysaurus ''Limaysaurus'' ("Limay lizard") is a genus represented by a single species of rebbachisaurid sauropod dinosaurs, which lived during the mid-Cretaceous period, about 99.6 to 94 million years ago, in the Cenomanian, in what is now South America (n ...
'', the titanosaurs ''
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 ...
'', ''
Choconsaurus ''Choconsaurus'' (" El Chocón lizard") is an extinct genus of herbivorous sauropod dinosaur belonging to the group Titanosauriformes, which lived in the area of present-day Argentina at the end of the Cretaceous. Discovery and naming The holo ...
'', and '' Chucarosaurus'', and indeterminate
iguanodont Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivore, herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and rel ...
s.


References

{{Taxonbar, from= Q112966897, from2= Q112969374 Carcharodontosauridae Dinosaur genera Cenomanian dinosaurs Huincul Formation Fossil taxa described in 2022 Dinosaurs of Argentina