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''Mapusaurus'' () was a giant
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
carnosauria Carnosauria is an extinct large group of predatory dinosaurs that lived during the Jurassic and Cretaceous periods. Starting from the 1990s, scientists have discovered some very large carnosaurs in the carcharodontosaurid family, such as '' G ...
n dinosaur from the early
Late 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'', ...
(early
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 b ...
stage), approximately 93.9 to 89.6 million years ago, of what is now
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

''Mapusaurus'' was excavated between 1997 and 2001, by the Argentinian-Canadian Dinosaur Project, from an exposure of the
Huincul Formation The Huincul Formation is a geologic formation of Late Cretaceous ( Early Cenomanian to Late Turonian) age of the Neuquén Basin that outcrops in the Mendoza, Río Negro and Neuquén Provinces of northern Patagonia, Argentina.
( Rio Limay Subgroup, Cenomanian) at Cañadón del Gato. It was described and named by
paleontologists 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 foss ...
Rodolfo Coria Rodolfo Aníbal Coria (born in Neuquén June 1, 1959), is an Argentine paleontologist. He is best known for having directed the field study and co-naming of '' Argentinosaurus'' (possibly the world's largest land animal ever) in 1993, and ''Giga ...
and
Phil Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
in 2006. The name ''Mapusaurus'' is derived from the
Mapuche The Mapuche ( (Mapuche & Spanish: )) are a group of indigenous inhabitants of south-central Chile and southwestern Argentina, including parts of Patagonia. The collective term refers to a wide-ranging ethnicity composed of various groups who s ...
word ''Mapu'', meaning 'of the Land' or 'of the Earth' and the
Greek Greek may refer to: Greece 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 ...
''sauros'', meaning ' lizard'. 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 ...
, ''Mapusaurus roseae'', is named for both the
rose A rose is either a woody perennial flowering plant of the genus ''Rosa'' (), in the family Rosaceae (), or the flower it bears. There are over three hundred species and tens of thousands of cultivars. They form a group of plants that can be ...
-colored rocks, in which the fossils were found and for Rose Letwin, who sponsored the expeditions which recovered these
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 ...
s. The designated
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 ...
for the
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 ...
and
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 ...
, ''Mapusaurus roseae'', is an isolated right nasal (MCF-PVPH-108.1, Museo Carmen Funes, Paleontología de Vertebrados, Plaza Huincul, Neuquén). Twelve paratypes have been designated, based on additional isolated skeletal elements. Taken together, the many individual elements recovered from the ''Mapusaurus'' bone bed represent most of the skeleton.


Description

''Mapusaurus'' was a large theropod, but smaller in size than its close relative ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period, approximately 99.6 to 95 million years ago. The holotype specimen was discovered in th ...
'', reaching in length and in body mass. Some specimens noted by Coria and Currie may have belonged to an individual comparable in size to the ''Giganotosaurus'' holotype, reaching in length, but these dinosaurs may not have the exact same proportions, so this should be regarded as a "very rough" estimate. It has been determined that ''Mapusaurus'' was diagnosed on autapomorphies, or unique traits, in regions of the skeleton that ''Giganotosaurus'' does not preserve. ''Mapusaurus'' only differs from ''Giganotosaurus'' in lacking a second opening on the middle quadrate, and in some details of the topology of the nasal rugosities.


Paleobiology

The
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 ...
remains of ''Mapusaurus'' were discovered in a
bone bed A bone bed is any geological stratum or deposit that contains bones of whatever kind. Inevitably, such deposits are sedimentary in nature. Not a formal term, it tends to be used more to describe especially dense collections such as Lagerstätte. ...
containing at least seven individuals of various growth stages. Coria and Currie speculated that this may represent a long term, possibly coincidental accumulation of carcasses (some sort of
predator trap A predator trap is a natural hazard where prey animals become trapped or incapacitated, and the attracted predators suffer the same fate. More predators, scavengers, insects and birds become attracted to this mounting accumulation of carrion, until ...
) and may provide clues about ''Mapusaurus'' behavior. Other known
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
bone beds and fossil graveyards include those of ''
Velociraptor ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in the p ...
'' and other
dromaeosaurids Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
around the planet, the ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
''-dominated
Cleveland-Lloyd Dinosaur Quarry Jurassic National Monument, at the site of the Cleveland-Lloyd Dinosaur Quarry, well known for containing the densest concentration of Jurassic dinosaur fossils ever found, is a paleontological site located near Cleveland, Utah, in the San Raf ...
of
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
, an ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of tyrannosaurid theropod dinosaurs that lived in western North America during the Late Cretaceous Period, about 71 million years ago. The type species, ''A. sarcophagus'', wa ...
'' bone bed from
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
and a ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 79.5 and 74 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three species ...
'' bone bed from
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columb ...
, and even a ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
'' bone bed from ''
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columb ...
'', as well. Paleontologist Rodolfo Coria, of the
Museo Carmen Funes Museo Municipal Carmen Funes, or, the Carmen Funes Municipal Museum, is a museum of paleontology in Plaza Huincul, Neuquén Province, Argentina. It is best known for its collection of dinosaur fossils, including the only specimen of the largest re ...
, contrary to his published article, repeated in a press-conference earlier suggestions that this congregation of fossil bones may indicate that ''Mapusaurus'' like ''Giganotosaurus'' also hunted in groups and worked together to take down large prey, such as the immense sauropod ''
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 animal ...
''. If so, this would be the first substantive evidence of gregarious behavior by large theropods other than ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
'', although whether they might have hunted in organized packs (as wolves and lions do) or simply attacked in a mob, is unknown. The authors interpreted the
depositional environment In geology, depositional environment or sedimentary environment describes the combination of physical, chemical, and biological processes associated with the deposition of a particular type of sediment and, therefore, the rock types that will be ...
of the Huincul Formation at the Cañadón del Gato locality as a freshwater paleochannel deposit, "laid down by an ephemeral or seasonal stream in a region with arid or semi-arid climate". This bone bed is especially interesting, in light of the overall scarcity of fossilized bone within the Huincul Formation. An ontogenetic study by Canale ''et al.'' (2014) found that ''Mapusaurus'' displayed heterochrony, an evolutionary condition in which the animals may retain an ancestral characteristic during one stage of their life, but lose it as they develop. In ''Mapusaurus'', the maxillary fenestrae are present in younger individuals, but gradually disappear as they mature.


Classification

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 carried out by Coria and Currie definitively showed that ''Mapusaurus'' is nested within the clade
Carcharodontosauridae Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
. The authors noted that the structure of the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
suggests a closer relationship with ''Giganotosaurus'' than either
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
shares with ''
Carcharodontosaurus ''Carcharodontosaurus'' (; ) is a genus of large carcharodontosaurid theropod dinosaur that existed during the Cenomanian age of the Late Cretaceous in Northern Africa. The genus ''Carcharodontosaurus'' is named after the shark genus '' Carc ...
''. They created a new monophyletic taxon based on this relationship, the subfamily
Giganotosaurinae Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
, defined as all carcharodontosaurids closer to ''Giganotosaurus'' and ''Mapusaurus'' than to ''Carcharodontosaurus''. They tentatively included the genus ''
Tyrannotitan ''Tyrannotitan'' (; ) is a genus of huge bipedal carnivorous dinosaur of the carcharodontosaurid family from the Aptian stage of the early Cretaceous period, discovered in Argentina. It is closely related to other giant predators like ''Carchar ...
'' in this new subfamily, pending publication of more detailed descriptions of the known specimens of that form. The following cladogram after Novas ''et al.'', 2013, shows the placement of ''Mapusaurus'' within Carcharodontosauridae.


Paleoecology

As previously mentioned, the Huincul Formation is thought to represent an arid environment with ephemeral or seasonal streams. The age of this formation is estimated at 97 to 93.5 Mya.Huincul Formation
at Fossilworks.org
The dinosaur record is considered sparse here. ''Mapusaurus'' shared its environment with the
sauropods 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 ...
''
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 animal ...
'' (one of the largest sauropods, if not the largest), ''
Choconsaurus ''Choconsaurus'' 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 holotype (specimen MMCh-P ...
'', and '' Cathartesaura''. Another carcharodontosaurid known as '' Meraxes'' was found in the same formation, but on older rocks than ''Mapusaurus'', so they probably didn't coexist.
Abelisaurid 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 foun ...
theropods ''
Skorpiovenator ''Skorpiovenator'' ("scorpion hunter") is a genus of abelisaurid theropod dinosaur from the Late Cretaceous (Cenomanian to Turonian) Huincul Formation of Argentina. It is one of the most complete and informative abelisaurids yet known, describe ...
'' and '' Ilokelesia'' also lived in the region. Fossilized pollen indicates a wide variety of plants was present in the Huincul Formation. A study of the El Zampal section of the formation found hornworts,
liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ...
s,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s,
Selaginellales ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
, possible
Noeggerathiales Noeggerathiales is a now- extinct order of vascular plants. The fossil range of the order extends from the Upper Carboniferous to the upper Permian (Lopingian). Due to gaps in the fossil record, the group is incompletely known and poorly defined, ...
,
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
s (including
gnetophyte Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three re ...
s and conifers), and angiosperms (flowering plants), in addition to several pollen grains of unknown affinities. The Huincul Formation is among the richest Patagonian vertebrate associations, preserving fish including dipnoans and
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
, chelid turtles,
squamate Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest order of reptiles, comprising lizards, snakes, and amphisbaenians (worm lizards), which are collectively known as squamates or scaled reptiles. With over 10,900 species, it ...
s,
sphenodont Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a diverse gro ...
s,
neosuchian Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to ''Crocodylus nilotic ...
crocodilians, and a wide variety of dinosaurs. Vertebrates are most commonly found in the lower, and therefore older, part of the formation.


References


External links


Meat-Eating Dinosaur Was Bigger Than T. Rex.
National Geographic News *

. Mike Taylor. The Dinosaur FAQ. August 27, 2002. (Named as Unnamed Argentinian Carcharodontosaurine) * " nd the Largest Theropod is... http://dml.cmnh.org/2003Jul/msg00355.html. The Dinosaur Mailing List Archives. Retrieved March 21, 2010 (Named as Undescribed Carcharodontosaurine) {{Taxonbar, from=Q131176 Apex predators Carcharodontosaurids Cenomanian life Turonian life Late Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Huincul Formation Fossil taxa described in 2006 Taxa named by Rodolfo Coria Taxa named by Philip J. Currie